THE FAMILIES
AND GENERA
OF VASCULAR PLANTS
Edited by K. Kubitzki
Volumes published in this series
Volume I
Pteridophytes and Gymnosperms
Edited by K.U. Kramer and P.S. Green (1990)
Date of publication: 28.9.1990
Volume II
Flowering Plants. Dicotyledons. Magnoliid, Hamamelid
and Caryophyllid Families
Edited by K. Kubitzki, J.G. Rohwer, and V. Bittrich (1993)
Date of publication: 28.7.1993
Volume III
Flowering Plants. Monocotyledons: Lilianae (except Orchidaceae)
Edited by K. Kubitzki (1998)
Date of publication: 27.8.1998
Volume IV
Flowering Plants. Monocotyledons: Alismatanae and Commelinanae
(except Gramineae)
Edited by K. Kubitzki (1998)
Date of publication: 27.8.1998
Volume V
Flowering Plants. Dicotyledons: Malvales, Capparales and Non-betalain
Caryophyllales
Edited by K. Kubitzki and C. Bayer (2003)
Date of publication: 12.9.2002
Volume VI
Flowering Plants. Dicotyledons: Celastrales, Oxalidales, Rosales, Cornales,
Ericales
Edited by K. Kubitzki (2004)
Date of publication: 21.1.2004
Volume VII
Flowering Plants. Dicotyledons: Lamiales (except Acanthaceae including
Avicenniaceae)
Edited by J.W. Kadereit (2004)
Date of publication: 13.4.2004
Volume VIII Flowering Plants. Eudicots: Asterales
Edited by J.W. Kadereit and C. Jeffrey (2007)
The Families
and Genera
of Vascular Plants
Edited by K. Kubitzki
VIII
Flowering Plants · Eudicots
Asterales
Volume Editors:
J.W. Kadereit and C. Jeffrey
With 131 Figures
123
Professor Dr. Klaus Kubitzki
Universität Hamburg
Biozentrum Klein-Flottbek und Botanischer Garten
Ohnhorststraße 18
22609 Hamburg
Germany
Professor Dr. Joachim W. Kadereit
Johannes Gutenberg-Universität Mainz
Institut für Spezielle Botanik und Botanischer Garten
55099 Mainz
Germany
Charles Jeffrey
flat 91, block 5, pr. Morisa Toreza 102
194017 St. Petersburg
Russia
Library of Congress Control Number: 2006924681
ISBN-10 3-540-31050-9 Springer Berlin Heidelberg New York
ISBN-13 978-3-540-31050-1 Springer Berlin Heidelberg New York
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Preface
It is a great pleasure to introduce this volume of the “Families and Genera of Vascular
Plants”, containing the treatments of Compositae and all other families of the Asterales.
In these treatments, the immense amount of evidence recently accrued has been taken
into account to present an up-to-date picture of the systematics of these groups. This
fully meets the aim of this series to distil and organise knowledge.
Compositae have always been in the focus of plant systematists, and here more
than elsewhere it is obvious how much we owe to our predecessors, of which Cassini
and Bentham may be singled out. Note, for instance, that as early as 1816 Calyceraceae
and Campanulaceae were suggested to be the closest relatives of Compositae, a concept
very similar to our present understanding. Although most of what is known about
interrelationships among organisms is based on comparative morphology, we have also
learned that morphology alone is unable to resolve all problems in systematics; for
example, the placement of Roussea or the recognition of the sister-group relationship
between Barnadesioideae and the other Compositae would never have been possible
without molecular data.
I am highly indebted to the editors of this volume, Joachim W. Kadereit and Charles
Jeffrey, for their Herculean effort in bringing the book to a successful end, and this
despite several obstacles. Moreover, deep appreciation is due to those who have provided
the scholarly and meticulous treatments assembled in this volume. Kåre Bremer is
acknowledged for invaluable advice on the selection of potential authors given during
early stages of this work. We are grateful to Linda Klöckner for the editing of the figures,
and to Sabine von Mering, Miriam Repplinger and Christian Uhink for their assistance
in the assembly of the final manuscript of Compositae. Our thanks also go to Monique
Delafontaine who so ably copy-edited the book. Special thanks are due to the copyright
holders of published illustrations who so generously permitted the inclusion of their
valuable material in the present volume.
Finally, it is a pleasure to acknowledge the agreeable collaboration with the staff of
Springer-Verlag who so willingly responded to all requests raised in connection with
planning and production.
Hamburg, August 2006
K. Kubitzki
Contents
Asterales: Introduction and Conspectus
J.W. Kadereit . . . . . . . . . . . . . . . . . . . . .
1
Alseuosmiaceae
J. Kårehed . . . . . . . . . . . . . . . . . . . . . . . .
7
Argophyllaceae
J. Kårehed . . . . . . . . . . . . . . . . . . . . . . . .
13
Calyceraceae
F.H. Hellwig . . . . . . . . . . . . . . . . . . . . . .
19
Campanulaceae
T.G. Lammers . . . . . . . . . . . . . . . . . . . . .
26
Carpodetaceae
M.H.G. Gustafsson . . . . . . . . . . . . . . . .
57
Compositae
A.A. Anderberg, B.G. Baldwin,
R.G. Bayer, J. Breitwieser, C. Jeffrey,
M.O. Dillon, P. Eldenäs, V. Funk,
N. Garcia-Jacas, D.J.N. Hind, P.O. Karis,
H.W. Lack, G. Nesom, B. Nordenstam,
Ch. Oberprieler, J.L. Panero,
C. Puttock, H. Robinson, T.F. Stuessy,
A. Susanna, E. Urtubey, R. Vogt,
J. Ward and L.E. Watson . . . . . . . . . . . . 61
Introduction with Key to Tribes
C. Jeffrey . . . . . . . . . . . . . . . . . . . . . . . . .
61
I. Tribe Barnadesieae
T.F. Stuessy and E. Urtubey . . . . . . . . .
87
II. Tribe Mutisieae
D.J.N. Hind . . . . . . . . . . . . . . . . . . . . . . . .
90
III. Tribe Cardueae
A. Susanna and N. Garcia-Jacas . . . . . 123
Carduoid Genera
of Uncertain Placement
C. Jeffrey . . . . . . . . . . . . . . . . . . . . . . . . . 146
IV. Tribe Gymnarrheneae
C. Jeffrey . . . . . . . . . . . . . . . . . . . . . . . . . 147
V. Tribe Moquinieae
H. Robinson . . . . . . . . . . . . . . . . . . . . . . 148
VI. Tribe Vernonieae
H. Robinson . . . . . . . . . . . . . . . . . . . . . . 149
VII. Tribe Liabeae
V.A. Funk, H. Robinson and M.O. Dillon 175
VIII. Tribe Cichorieae
H.W. Lack . . . . . . . . . . . . . . . . . . . . . . . . . 180
IX. Tribe Gundelieae
C. Jeffrey . . . . . . . . . . . . . . . . . . . . . . . . . 199
X. Tribe Arctotideae
P.O. Karis . . . . . . . . . . . . . . . . . . . . . . . . . 200
XI. Tribe Corymbieae
B. Nordenstam . . . . . . . . . . . . . . . . . . . . 207
XII. Tribe Senecioneae
B. Nordenstam . . . . . . . . . . . . . . . . . . . . 208
XIII. Tribe Calenduleae
B. Nordenstam . . . . . . . . . . . . . . . . . . . . 241
viii
Contents
XIV. Tribe Gnaphalieae
R.J. Bayer, I. Breitwieser, J. Ward
and C. Puttock . . . . . . . . . . . . . . . . . . . . 246
XV. Tribe Astereae
G. Nesom and H. Robinson . . . . . . . . . . 284
XVI. Tribe Anthemideae
Ch. Oberprieler, R. Vogt
and L.E. Watson . . . . . . . . . . . . . . . . . . . 342
XVII. Tribe Inuleae
A.A. Anderberg and P. Eldenäs . . . . . 374
Key to the Tribes
of the Heliantheae Alliance
J.L. Panero . . . . . . . . . . . . . . . . . . . . . . . . 391
XVIII. Tribe Athroismeae
J.L. Panero . . . . . . . . . . . . . . . . . . . . . . . . 395
XIX. Tribe Helenieae
J.L. Panero . . . . . . . . . . . . . . . . . . . . . . . . 400
XX. Tribe Coreopsideae
J.L. Panero . . . . . . . . . . . . . . . . . . . . . . . . 406
XXI. Tribe Neurolaeneae
J.L. Panero . . . . . . . . . . . . . . . . . . . . . . . . 417
XXII. Tribe Tageteae
J.L. Panero . . . . . . . . . . . . . . . . . . . . . . . . 420
XXIII. Tribe Chaenactideae
J.L. Panero . . . . . . . . . . . . . . . . . . . . . . . . 431
XXIV. Tribe Bahieae
J.L. Panero . . . . . . . . . . . . . . . . . . . . . . . . 433
XXV. Tribe Polymnieae
J.L. Panero . . . . . . . . . . . . . . . . . . . . . . . . 439
XXVI. Tribe Heliantheae
J.L. Panero . . . . . . . . . . . . . . . . . . . . . . . . 440
XXVII. Tribe Millerieae
J.L. Panero . . . . . . . . . . . . . . . . . . . . . . . . 477
XXVIII. Tribe Madieae
B.G. Baldwin and J.L. Panero . . . . . . . . 492
XXIX. Tribe Perityleae
J.L. Panero . . . . . . . . . . . . . . . . . . . . . . . . 507
XXX. Tribe Eupatorieae
D.J.N. Hind and H. Robinson . . . . . . . . 510
Asteroid Genus
of Uncertain Placement
C. Jeffrey . . . . . . . . . . . . . . . . . . . . . . . . . 574
Selected Bibliography
to Compositae
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 576
Goodeniaceae
R.C. Carolin . . . . . . . . . . . . . . . . . . . . . . 589
Menyanthaceae
G. Kadereit . . . . . . . . . . . . . . . . . . . . . . . 599
Pentaphragmataceae
T.G. Lammers . . . . . . . . . . . . . . . . . . . . . 605
Phellinaceae
G. Barriera, V. Savolainen
and R. Spichiger . . . . . . . . . . . . . . . . . . . 608
Rousseaceae
J.A. Koontz, J. Lundberg and D.E. Soltis 611
Stylidiaceae
R.C. Carolin . . . . . . . . . . . . . . . . . . . . . . 614
Index to Scientific Names
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 621
List of Contributors
Anderberg, A.A.
Department of Phanerogamic Botany, Swedish Museum of
Natural History, P.O. Box 50007, 10405 Stockholm, Sweden
Baldwin, B.G.
Jepson Herbarium & Dept. of Integrative Biology,
1001 Valley Life Sciences Bldg. #2465,
University of California, Berkeley, CA 94720-2465, USA
Barriera, G.
Conservatoire et Jardin botaniques de la Ville de Genève,
1 ch. de l’Impératrice, Case postale 60, 1292 Chambésy,
Switzerland
CSIRO – Plant Industry, Australian National Herbarium,
GPO Box 1600, Canberra, ACT 2601, Australia
Bayer, R.J.
Breitwieser, I.
Biosystematics of New Zealand Plants, Manaaki Whenua –
Landcare Research, P.O. Box 69, Lincoln 8152, New Zealand
Carolin, R.C.
Pulman’s Cottage, 30 Pulman Street, Berry, N.S.W. 2535,
Australia
Department of Botany, Field Museum of Natural History,
1400 South Lake Shore Drive, Chicago, IL 60605-2496, USA
Dillon, M.O.
Eldenäs, P.
Molecular Systematics Laboratory, Swedish Museum of
Natural History, P.O. Box 50007, 10405 Stockholm, Sweden
Funk, V.A.
US National Herbarium, Department of Botany,
Smithsonian Institution, MRC 166, Washington, DC 20560,
USA
Botanic Institute of Barcelona, Passeig del Migdia s.n., Parc
de Montjuic, 08038 Barcelona, Spain
Garcia-Jacas, N.
Gustafsson, M.H.G.
Institute of Biological Sciences, University of Aarhus, Ny
Munkegade, Building 540, 8000 Århus C, Denmark
Hellwig, F.H.
Institut für Spezielle Botanik mit Botanischem Garten und
Herbarium Haussknecht, Friedrich-Schiller-Universität
Jena, Philosophenweg 16, 07743 Jena, Germany
Hind, D.J.N.
The Herbarium, Royal Botanic Gardens, Kew, Richmond,
Surrey TW9 3AE, UK
Jeffrey, C.
Flat 91, Block 5, pr. Morisa Toreza 102,
194017 St. Petersburg, Russia
Kadereit, G.
Institut für Spezielle Botanik und Botanischer Garten,
Johannes Gutenberg-Universität, 55099 Mainz, Germany
Kadereit, J.W.
Institut für Spezielle Botanik und Botanischer Garten,
Johannes Gutenberg-Universität, 55099 Mainz, Germany
x
List of Contributors
Kårehed, J.
Department of Systematic Botany, Evolutionary Biology
Centre, Norbyvägen 18D, Uppsala University,
75236 Uppsala, Sweden
Karis, P.O.
Department of Botany, Stockholm University,
10691 Stockholm, Sweden
Department of Biology, Augustana College, 639 38th Street,
Rock Island, IL 61201, USA
Botanischer Garten und Botanisches Museum BerlinDahlem, Freie Universität Berlin, Königin-Luise-Str. 6–8,
14195 Berlin, Germany
Koontz, J.A.
Lack, H.W.
Lammers, T.G.
Lundberg, J.
Nesom, G.
Nordenstam, B.
Department of Biology and Microbiology, University of
Wisconsin Oshkosh, Oshkosh, WI 54901, USA
Department of Systematic Botany, Evolutionary Biology
Centre, Uppsala University, Norbyvägen 18D,
75236 Uppsala, Sweden
Botanical Research Institute of Texas, 509 Pecan Street,
Fort Worth, TX 76102-4060, USA
Department of Phanerogamic Botany, Swedish Museum of
Natural History, P.O. Box 50007, 10405 Stockholm, Sweden
Oberprieler, Ch.
Institute of Botany, University of Regensburg,
Universitätsstr. 31, 93040 Regensburg, Germany
Panero, J.L.
Section of Integrative Biology, 1 University Station C0930,
The University of Texas, Austin, TX 78712, USA
Puttock, C.
Bishop Museum, Department of Botany, 1525 Bernice
Street, Honolulu, HI 96817-2704, USA
US National Herbarium, Department of Botany,
Smithsonian Institution, MRC 166, Washington, DC 20560,
USA
Molecular Systematics Section, Royal Botanic Gardens,
Kew, Richmond, Surrey TW9 3DS, London, UK
Robinson, H.
Savolainen, V.
Soltis, D.E.
Spichiger, R.
Stuessy, T.F.
Susanna, A.
Department of Botany, University of Florida, Gainesville,
FL 32611, USA
Conservatoire et Jardin botaniques de la Ville de Genève,
1 ch. de l’Impératrice, Case postale 60, 1292 Chambésy,
Switzerland
Department of Systematic and Evolutionary Botany,
Institute of Botany, University of Vienna, Rennweg 14,
1030 Vienna, Austria
Botanic Institute of Barcelona, Passeig del Migdia s.n., Parc
de Montjuic, 08038 Barcelona, Spain
Urtubey, E.
Division Plantas Vasculares, Museo de La Plata,
Universidad Nacional de La Plata, Paseo del Bosque s.n.,
La Plata, Argentina
Vogt, R.
Botanischer Garten und Botanisches Museum BerlinDahlem, Freie Universität Berlin, Königin-Luise-Str. 6–8,
14191 Berlin, Germany
List of Contributors
Ward, J.
School of Biological Sciences, University of Canterbury,
Private Bag 4800, Christchurch, New Zealand
Watson, L.E.
Department of Botany, Miami University, Oxford, OH
45056, USA
xi
Asterales: Introduction and Conspectus
J.W. Kadereit
Asterales (incl. Campanulales of many authors),
with Alseuosmiaceae, Argophyllaceae, Compositae (= Asteraceae), Calyceraceae, Campanulaceae
(incl. Cyphiaceae, Lobeliaceae, Nemacladaceae),
Carpodetaceae (included in Rousseaceae by
APG II 2003), Goodeniaceae, Menyanthaceae,
Pentaphragmataceae, Phellinaceae, Rousseaceae
and Stylidiaceae (incl. Donatiaceae), contain about
26,300 species in c. 1,720 genera. The large majority of species and genera belong to Compositae
and Campanulaceae. The order is well supported
in all major molecular phylogenetic analyses
(APG II 2003), and is part of the Euasterids II or
Campanulids sensu Bremer et al. (2002).
Phylogenetic structure within Campanulids
(also containing Apiales, Aquifoliales, Dipsacales
and several families of uncertain ordinal placement; APG II 2003) is not sufficiently well resolved
to identify the sister group of Asterales. It appears
to be evident, however, that of all representatives
of the Campanulids, Aquifoliales are least closely
related to Asterales (Savolainen et al. 2000a, b;
Soltis et al. 2000; Albach et al. 2001; Bremer et al.
2001, 2002).
Although several of the constituent families
of the order had been recognized to be closely
related to one another long ago (for discussion, see
Lammers 1992), the recognition of the relationship
of others to Asterales (Lundberg and Bremer
2003) is the result mainly (but not only) of recent
molecular phylogenetic work. This applies particularly to Alseuosmiaceae (Backlund and Bremer
1997; Gustafsson and Bremer 1997; Kårehed
et al. 1999; Cronquist 1981: Rosales; Thorne 1992:
Saxifragales; Takhtajan 1997: Hydrangeales), Argophyllaceae (Kapil and Bhatnagar 1992; Gustafsson
et al. 1996; Kårehed et al. 1999; Olmstead et al.
2000; Cronquist 1981: Rosales; Takhtajan 1997:
Hydrangeales), Carpodetaceae (Gustafsson and
Bremer 1997; Lundberg 2001; Takhtajan 1997: Hydrangeales), Phellinaceae (Backlund and Bremer
1997; Gustafsson and Bremer 1997; Kårehed et al.
1999; Cronquist 1981: Celastrales; Thorne 1992:
Theales; Takhtajan 1997: Icacinales) and Rousseaceae (Lundberg 2001; Takhtajan 1997: Brexiales), and partly also to Menyanthaceae (Downie
and Palmer 1992; Olmstead et al. 1992; Cronquist
1981: Solanales; Thorne 1992: Campanulales;
Takhtajan 1997: Menyanthales) and Stylidiaceae
(Cronquist 1981: Campanulales; Thorne 1992: Saxifragales; Takhtajan 1997: Stylidiales). Further sampling may identify other taxa from distant corners
of the traditional angiosperm system which should
be included in the order. On the other hand, Sphenocleaceae, as a family often associated with Asterales/Campanulales (e.g. Lammers 1992), do not
belong here but rather in Solanales (APG II 2003).
Members of Asterales are mostly herbaceous
and in most cases have alternate leaves without
stipules. Flowers are very rarely solitary but mostly
aggregated in sometimes axillary but more commonly terminal inflorescences which are capitulate
and involucrate in most of the closely related Goodeniaceae, Calyceraceae and Compositae, and also
in some Campanulaceae. The mostly zoophilous
flowers typically are tetracyclic and pentamerous
but variation of organ number per whorl is
known from several families. Flower symmetry is
actinomorphic or zygomorphic with bilabiate or
unilabiate flowers – actinomorphic and zygomorphic flowers are both found in the capitula of many
Compositae – and resupination of flowers is known
from Campanulaceae-Lobelioideae and some Stylidiaceae. The sepals are commonly fused (not in
Alseuosmiaceae and some Menyanthaceae), and
in Compositae the calyx commonly is replaced by
a pappus of variable structure assisting in fruit dispersal. Petals are free only in Carpodetaceae, Phellinaceae and some Argophyllaceae, Pentaphragmataceae and Stylidiaceae (Donatia). The androecium
normally is isomerous with calyx and corolla, and
the stamens alternate with the petals. Reduction of
stamen number is largely limited to Stylidiaceae.
Stamens can be inserted on the corolla or not, and
2
J.W. Kadereit
anthers are mostly tetrasporangiate, basifixed and
commonly introrse. Pollen grains are mostly tricolporate, but both colpate or porate pollen grains
with an increased number of apertures are known.
Carpodetus (Carpodetaceae) and Lechenaultia
(Goodeniaceae) are unusual in having pollen
tetrads. The pluri- to unilocular ovary is commonly
inferior (or semi-inferior) but superior ovaries
are found in some Carpodetaceae, some Goodeniaceae, some Campanulaceae, and in Menyanthaceae, Phellinaceae and Rousseaceae. Ovules
usually are anatropous (hemi- to campylotropous
in Phellinaceae), unitegmic and tenuinucellate
and, where known, endosperm formation is mostly
cellular, but nuclear in some Compositae. Fruits
are commonly capsules or achenes (= cypselae),
rarely berries or drupes. Inulin is found in several
families (Calyceraceae, Campanulaceae, Compositae, Goodeniaceae, Menyanthaceae and Stylidiaceae), and iridoids or seco-iridoids are common,
but absent from Campanulaceae and Compositae, and apparently also from Alseuosmiaceae,
Phellinaceae and Rousseaceae.
A tight integration of stamens and style is
found in several families. In most Stylidiaceae,
the two stamens are fused with the style to form
a pressure-sensitive gynostemium. In Calyceraceae, Campanulaceae, Compositae and Goodeniaceae, the interaction of style and either fused or
free anthers results in various forms of secondary
pollen presentation (Carolin 1960; Leins and Erbar
1990, 2003; Erbar and Leins 1995). Erbar and Leins
(1995) classified these as (1) brushing or pump
mechanism in Compositae and CampanulaceaeLobelioideae (pollen is removed from an anther
tube by the elongating style which is hairy or not),
(2) deposition (or rarely brushing) mechanism
in Campanulaceae-Campanuloideae (pollen from
free anthers is deposited on hairs on the outside of
the style, these hairs can invaginate or not), (3) cup
and cup/brushing mechanism in Goodeniaceae
(pollen is deposited in a cup-like outgrowth below
the stigma, the indusium; in addition to this cup,
hairs can be present on the style) and (4) deposition
mechanism of Goodeniaceae (deposition of pollen
grains on top of the style). Detailed summaries
of character distribution in Asterales have been
provided by Lammers (1992; excl. Alseuosmiaceae,
Argophyllaceae, Carpodetaceae, Phellinaceae,
Rousseaceae) and, covering the entire order,
particularly by Lundberg and Bremer (2003).
In spite of the very high molecular support for
the order, it is difficult to identify synapomorphies.
Following Lundberg and Bremer (2003), valvate
corolla aestivation and the absence of apotracheal
wood parenchyma can be identified as synapomorphic. Both these characters, however, are not
unique for the order and are variable within it.
Previously identified synapomorphies, such as secondary pollen presentation (which is present in the
form of different mechanisms and is likely to have
arisen more than once; see above) and the presence of inulin, are characteristic only of subgroups
of Asterales.
Relationships within the order are clear and
well supported in some parts but not in others
(Lundberg and Bremer 2003). One well-supported
clade identified in several analyses (Chase et al.
1993; Morgan and Soltis 1993; Cosner et al. 1994;
Gustafsson and Bremer 1995; Olmstead et al. 2000;
Soltis et al. 2000; Bremer et al. 2001; Lundberg and
Bremer 2003) consists of Menyanthaceae, Goodeniaceae, Calyceraceae and Compositae (MGCA
clade; Fig. 1). This clade is characterized by the
presence of petal lateral veins (Gustafsson 1995),
the loss of micropylar endosperm haustoria (Cosner et al. 1994), and a thick and multilayered (> 10
cells) integument (Inoue and Tobe 1999). Within
this clade, the sister-group relationship between
Calyceraceae and Compositae is supported by several potential synapomorphies in wood anatomical
(Carlquist and De Vore 1998), inflorescence, flower
and fruit morphological and anatomical (Hansen
1992; Gustafsson 1995), and pollen (Hansen
1992) characters. Goodeniaceae are sister to
these two families, and the clade consisting of
Goodeniaceae/Calyceraceae/Compositae may be
supported by pollen grains with a prominent layer
with branched columellae and secondary pollen
presentation involving fused anthers (Lundberg
and Bremer 2003). Lundberg and Bremer (2003)
suggested that Stylidiaceae incl. Donatiaceae,
a strongly supported clade in their study, are sister
to the MGCA clade. A close relationship between
Donatiaceae and Stylidiaceae, however, was not
found in other analyses (Albach et al. 2001; Bremer
et al. 2002), and neither Donatiaceae nor Stylidiaceae were sister to the MGCA clade in these two
analyses. Instead, Stylidiaceae were sister to Campanulaceae (Albach et al. 2001; Bremer et al. 2002),
and Donatiaceae sister to Alseuosmiaceae/Argophyllaceae/Phellinaceae (Bremer et al. 2002) or
to all families except Stylidiaceae/Campanulaceae
(Albach et al. 2001). A second possible clade of
the order consists of Alseuosmiaceae, Phellinaceae
and Argophyllaceae (APA clade; Fig. 1), where
Asterales: Introduction and Conspectus
Fig. 1. A phylogenetic hypothesis for the families of Asterales. (Modified from Lundberg and Bremer 2003)
the latter two families probably are sister to each
other (Lundberg and Bremer 2003). This clade
had already been identified in earlier analyses
(Gustafsson et al. 1996; Backlund and Bremer
1997; Gustafsson and Bremer 1997; Källersjö et al.
1998; Kårehed et al. 1999; Savolainen et al. 2000b;
Lundberg 2001) and may be supported by pollen
being 3-celled at anthesis and the presence of
ellagic acid (not known in all groups; Lundberg
and Bremer 2003). Stevens (2001 onwards) further
records the presence of subepidermal cork as well
as serrate and gland-toothed leaf blades as possible
synapomorphies. In the analysis of Lundberg and
Bremer (2003), the APA clade is sister to the Sty-
3
lidiaceae/MGCA clade. All three groups together
constitute the “Core Asterales” of these authors
and are characterized by having a non-intrusive
placenta (Lundberg and Bremer 2003). Sister to
this in the analysis by Lundberg and Bremer (2003)
is a clade consisting of Rousseaceae (incl. Carpodetaceae), Pentaphragmataceae and Campanulaceae.
This clade was resolved as a basal grade (incl.
Stylidiaceae as sister to Campanulaceae) by Bremer et al. (2002). The close relationship between
Roussea and Carpodetaceae is well supported
(Savolainen et al. 2000b; Lundberg 2001; Bremer
et al. 2002). The possible sister-group relationship
between Pentaphragmataceae and Campanulaceae
found by Lundberg and Bremer (2003) but not
in several other analyses (Cosner et al. 1994;
Jansen and Kim 1996; Backlund and Bremer 1997;
Olmstead et al. 2000; Savolainen et al. 2000b) may
be supported (Lundberg and Bremer 2003) by the
presence of a free hypanthium and petal veins
which form a dense reticulum (Gustafsson 1995).
In summary, relationships within the order
should be viewed (Fig. 1), as by Stevens (2001 onwards), as a polytomy consisting of four lineages.
These are (1) Campanulaceae, (2) Pentaphragmataceae, (3) Rousseaceae/Carpodetaceae and
(4) a trichotomy of the APA clade, Stylidiaceae
(incl. Donatiaceae), and the MGCA clade.
Although the earliest fossils of the order are of
Oligocene (c. 29 Ma b.p.) age (Magallón et al. 1999),
consideration of phylogenetic relationships and
molecular evidence led to the conclusion that the
order must have originated c. 100 Ma b.p. in the Cretaceous (Bremer and Gustafsson 1997; Wikström
et al. 2001). Stem node and crown node ages of 112
and 93 Ma b.p. respectively were recently estimated
by Bremer et al. (2004). The notion of a Cretaceous
origin of Asterales certainly requires revision of
the observation by Magallón and Sanderson (2001)
that Asterales have the highest diversification rate
of all angiosperm orders. This inference was based
on the assumption of an Oligocene age of Asterales.
Apart from the cosmopolitan Campanulaceae,
Compositae and Menyanthaceae, of which Compositae have been postulated to have originated
in South America (Bremer 1994) and Campanulaceae which have centres of diversity in southern
Africa and Andean South America but also in
Eurasia between the Mediterranean region and
the Himalayas, all other families of the order have
an almost exclusively southern hemispherical
distribution, mostly in Australasia and partly in
South America. Based on an analysis of ancestral
4
J.W. Kadereit
areas, Bremer and Gustafsson (1997) concluded
that the order originated in Australasia. Although
this interpretation was based on a rather terminal
position of the cosmopolitan Campanulaceae in
the phylogeny of the order these authors used, the
placement of this family in a basal polytomy (see
above) probably will not change the outcome of an
ancestral area analysis. Many species of the small
families of the order are found in either temperate
forest or more open, often humid to wet habitats.
By far the largest amount of generic and species
diversity is found in Campanulaceae and Compositae. Interestingly, these are the two major families
of the order lacking iridoids or secoiridoids. In
Compositae, the biosynthetic pathway producing
iridoids has been blocked and diverted to the
production of sesquiterpene lactones (Zdero
and Bohlmann 1990), and the diversification of
secondary compounds in the family has been
held responsible for its great success in terms of
species diversity (Cronquist 1977; Lammers 1992).
In Campanulaceae, iridoids are replaced by polysterols (particularly Campanuloideae), acetylenes
and/or alkaloids (particularly Lobelioideae) which,
however, have a biosynthetic origin unrelated to
the iridoid pathway (Lammers 1992). It has not
been claimed that the success of Campanulaceae is
related to their biochemical diversification.
Conspectus of families as treated in this volume
1.
1.
Stamens as many as corolla lobes
2. Corolla lobes with distinct wings or appendages
3. Corolla zygomorphic; herbs, shrubs or scramblers with zygomorphic flowers, fruit a drupe, nut or capsule;
11/400, southern hemisphere, mainly Australia
Goodeniaceae
3. Corolla actinomorphic
4. Plants herbaceous, from wet habitats; flowers actinomorphic, petal lobes often fimbriate or crested;
fruit a capsule or rarely a berry; 5/c. 60, subcosmopolitan
Menyanthaceae
4. Plants woody
5. Sepals free, fruit a berry; shrubs or subshrubs, leaf axils with tufts of hairs;
flowers actinomorphic; 4/9, Australia, New Zealand, New Guinea and New Caledonia
Alseuosmiaceae
5. Sepals fused, fruit a capsule or drupe; shrubs or small trees with actinomorphic flowers; 2/c. 20,
Australia, New Zealand, Lord Howe and Rapa Islands, New Caledonia
Argophyllaceae
2. Corolla lobes without distinct wings or appendages
6. Petals free
7. Fruit a drupe; shrubs or small trees with actinomorphic flowers; 1/11,
New Caledonia
Phellinaceae
7. Fruit a berry or capsule; shrubs or trees with actinomorphic flowers; 3/5, Australia, New Zealand,
New Guinea and Solomon Islands
Carpodetaceae
6. Petals fused, sometimes corolla tube short
8. Ovary unilocular with one ovule, inflorescence capitulate
9. Calyx mostly modified, anthers connate, ovule insertion apical; 1,621/c. 23,300, cosmopolitan
Compositae
9. Calyx not modified, anthers free, ovule insertion basal; annual or perennial herbs with actinomorphic flowers in involucrate head, fruit an achene; 4/c. 60, South America and Falkland Islands
Calyceraceae
8. Ovary two- to multilocular, rarely unilocular with only one ovule, then inflorescence not capitulate
10. Climbing shrub with opposite or verticillate leaves; flowers actinomorphic, fruit a berry;
1 sp., Mauritius
Rousseaceae
10. Not as above
11. Shrub, flowers inclined, corolla tube short, stamens sessile, fruit a 2-locular capsule;
1 sp., New Caledonia
Platyspermation (Alseuosmiaceae)
11. Not as above
12. Leaf bases asymmetrical, plants without milky latex; mostly fleshy perennial herbs with
asymmetrical leaf blades and actinomorphic flowers, fruit a berry; 1/c. 30, SE Asia
Pentaphragmataceae
12. Leaf bases not asymmetrical, plants with milky latex; herbs, lianas, rosette plants,
subshrubs, shrubs, treelets or trees with actinomorphic or zygomorphic flowers,
fruit a capsule or berry; 84/c. 2,400, cosmopolitan
Campanulaceae
Stamens fewer than corolla lobes
13. Corolla lobes free, gynoecium with separate stylodia; perennial herbs with solitary, actinomorphic flowers
and capsular fruits; 1/2, South America, Tasmania and New Zealand
Donatia (Stylidiaceae)
13. Corolla lobes fused, gynoecium with one style; herbs or subshrubs with mostly zygomorphic flowers, filaments
and style fused into a column in most genera, fruits capsular; 6/c. 160, southern hemisphere, mainly Australia
Stylidiaceae
Asterales: Introduction and Conspectus
References
Albach, D.C., Soltis, P.S., Soltis, D.E., Olmstead, R.G. 2001.
Phylogenetic analysis of Asterids based on sequences
of four genes. Ann. Missouri Bot. Gard. 88: 163–212.
APG II 2003. An update of the Angiosperm Phylogeny Group
classification for the orders and families of flowering
plants: APG II. Bot. J. Linn. Soc. 141: 399–436.
Backlund, A., Bremer, B. 1997. Phylogeny of the Asteridae
s.str. based on rbcL sequences, with particular reference to the Dipsacales. Pl. Syst. Evol. 207: 225–254.
Bremer, K. 1994. Asteraceae. Cladistics and classification.
Portland, OR: Timber Press.
Bremer, K., Gustafsson, M.H.G. 1997. East Gondwanan ancestry of the sunflower alliance of families. Proc. Natl
Acad. Sci. U.S.A. 94: 9188–9190.
Bremer, K., Backlund, A., Sennblad, B., Swenson, U.,
Andreasen, K., Hjertson, M., Lundberg, J., Backlund, M., Bremer, B. 2001. A phylogenetic analysis
of 100+ genera and 50+ families of euasterids based
on morphological and molecular data with notes on
possible higher level morphological synapomorphies.
Pl. Syst. Evol. 229: 137–169.
Bremer, B., Bremer, K., Heidari, N., Olmstead, R.G., Anderberg, A.A., Källersjö, M., Barkhordarian, E. 2002.
Phylogenetics of asterids based on 3 coding and 3 noncoding chloroplast DNA markers and the utility of noncoding DNA at higher taxonomic levels. Mol. Phylog.
Evol. 24: 274–301.
Bremer, K., Friis, E.-M., Bremer, B. 2004. Molecular phylogenetic dating of Asterid flowering plants shows early
Cretaceous diversification. Syst. Biol. 53: 496–505.
Carlquist, S., De Vore, M.L. 1998. Wood anatomy of Calyceraceae with reference to ecology, habit, and systematic
relationships. Aliso 17: 63–76.
Carolin, R.C. 1960. The structures involved in the presentation of pollen to visiting insects in the order Campanulales. Proc. Linn. Soc. New South Wales 85: 197–207.
Chase, M.W. et al. 1993. Phylogenetics of seed plants: an
analysis of nucleotide sequences from the plastid gene
rcbL. Ann. Missouri Bot. Gard. 80: 528–580.
Cosner, M.E., Jansen, R.K., Lammers, T.G. 1994. Phylogenetic relationships in the Campanulales based on rbcL
sequences. Pl. Syst. Evol. 190: 79–95.
Cronquist, A. 1977. The Compositae revisited. Brittonia 29:
137–153.
Cronquist, A. 1981. An integrated system of classification
of flowering plants. New York: Columbia University
Press.
Downie, S.R., Palmer, J.D. 1992. Restriction site mapping
of the chloroplast DNA inverted repeat: a molecular
phylogeny of the Asteridae. Ann. Missouri Bot. Gard.
79: 266–283.
Erbar, C., Leins, P. 1995. Portioned pollen release and
the syndromes of secondary pollen presentation
in the Campanulales-Asterales-complex. Flora 190:
323–338.
Gustafsson, M.H.G. 1995. Petal venation in Asterales and
related orders. Bot. J. Linn. Soc. 118: 1–18.
Gustafsson, M.H.G., Bremer, K. 1995. Morphology and phylogenetic interrelationships of the Asteraceae, Calyceraceae, Campanulaceae, Goodeniaceae, and related
families (Asterales). Amer. J. Bot. 82: 250–265.
5
Gustafsson, M.H.G., Bremer, K. 1997. The circumscription
and systematic position of Carpodetaceae. Austral. J.
Bot. 10: 855–862.
Gustafsson, M.H.G., Backlund, A., Bremer, B. 1996. Phylogeny of the Asterales sensu lato based on rbcL sequences with particular reference to the Goodeniaceae.
Pl. Syst. Evol. 199: 217–242.
Hansen, H.V. 1992. Studies in the Calyceraceae with a discussion of its relationships to Compositae. Nordic J.
Bot. 12: 63–75.
Inoue, N., Tobe, H. 1999. Integumentary studies in
Menyanthaceae (Campanulales sensu lato). Acta
Phytotax. Geobot. 50: 75–79.
Jansen, R.K., Kim, K.-J. 1996. Implications of chloroplast
DNA data for the classification and phylogeny of the
Asteraceae. In: Hind, D.J.N., Beentje, H.J. (eds) Compositae: systematics. Proceedings of the International
Compositae Conference, Kew, 1994, vol. 1. Royal
Botanic Gardens, Kew, pp. 317–339.
Källersjö, M., Farris, J.S., Chase, M.W., Bremer, B., Fay, M.F.,
Humphries, C.J., Petersen, G., Seberg, O., Bremer, K.
1998. Simultaneous parsimony jackknife analysis of
2538 rbcL DNA sequences reveals support for major
clades of green plants, land plants, seed plants and
flowering plants. Pl. Syst. Evol. 213: 259–287.
Kapil, R.N., Bhatnagar, A.K. 1992. Embryology and systematic position of Corokia A. Cunn. In: Proceedings
of the 11th International Symposium on Embryology
and Seed Reproduction, Leningrad, 1990. St. Petersburg: Nauka, pp. 246–247.
Kårehed, J., Lundberg, J., Bremer, B., Bremer, K. 1999. Evolution of the Australasian families Alseuosmiaceae,
Argophyllaceae, and Phellinaceae. Syst. Bot. 24: 660–
682.
Lammers, T.G. 1992. Circumscription and phylogeny of
the Campanulales. Ann. Missouri Bot. Gard. 79: 388–
413.
Leins, P., Erbar, C. 1990. On the mechanisms of secondary
pollen presentation in the Campanulales-Asteralescomplex. Bot. Acta 103: 87–92.
Leins, P., Erbar, C. 2003. The pollen box in Cyphiaceae (Campanulales). Intl J. Pl. Sci. 164 suppl. 5: S321–S328.
Lundberg, J. 2001. The asteralean affinity of the Mauritian
Roussea (Roussaceae). Bot. J. Linn. Soc. 137: 267–276.
Lundberg, J., Bremer, K. 2003. A phylogenetic study of the
order Asterales using one morphological and three
molecular data sets. Intl J. Pl. Sci. 164: 553–578.
Magallón, S., Sanderson, M.J. 2001. Absolute diversification
rates in angiosperm clades. Evolution 55: 1762–1780.
Magallón, S., Crane, P.R., Herendeen, P.S. 1999. Phylogenetic
pattern, diversity and diversification of eudicots. Ann.
Missouri Bot. Gard. 86: 297–372.
Morgan, D.R., Soltis, D.E. 1993. Phylogenetic relationships
among members of Saxifragaceae sensu lato based on
rbcL sequence data. Ann. Missouri Bot. Gard. 80: 631–
660.
Olmstead, R.G., Michaels, H.J., Scott, K.M., Palmer, J.D.
1992. Monophyly of the Asteridae and identification
of their major lineages inferred from DNA sequences
of rbcL. Ann. Missouri Bot. Gard. 79: 249–265.
Olmstead, R.G., Kim, K.-J., Jansen, R.K., Wagstaff, S.J. 2000.
The phylogeny of the Asteridae sensu lato based on
chloroplast ndhF gene sequences. Mol. Phylog. Evol.
16: 96–112.
6
J.W. Kadereit
Savolainen, V., Chase, M.W., Hoot, S.B., Morton, C.M.,
Soltis, D.E., Bayer, C., Fay, M.F., De Bruijn, A.Y., Sullivan, S., Qiu, Y.-L. 2000a. Phylogenetics of flowering
plants based on combined analysis of plastid atpB and
rbcL gene sequences. Syst. Biol. 49: 306–362.
Savolainen, V., Fay, M.F., Albach, D.C., Backlund, A.,
van der Bank, M., Cameron, K.M., Johnson, S.A.,
Lledo, M.D., Pintaud, J.-C., Powell, M., Sheahan, M.C.,
Soltis, D.E., Soltis, P.S., Weston, P., Whitten, W.M.,
Wurdack, K.J., Chase, M.W. 2000b. Phylogeny of the
eudicots: a nearly complete familial analysis based on
rbcL gene sequences. Kew Bull. 55: 357–309.
Soltis, D.E., Soltis, P.S., Chase, M.W., Mort, M.E., Albach, D.C., Zanis, M., Savolainen, V., Hahn, W.H.,
Hoot, S.B., Fay, M.F., Axtell, M., Swensen, S.M.,
Prince, L.M., Kress, W.J., Nixon, K.C., Farris, J.S. 2000.
Angiosperm phylogeny inferred from 18S rDNA, rbcL
and atpB sequences. Bot. J. Linn. Soc. 133: 381–461.
Stevens, P.F. 2001 onwards. Angiosperm Phylogeny website,
version 5, May 2004 (and more or less continuously updated since). http://www.mobot.org/MOBOT/research
/APweb/
Takhtajan, A. 1997. Diversity and classification of flowering
plants. New York: Columbia University Press.
Thorne, R.F. 1992. An updated phylogenetic classification
of flowering plants. Aliso 13: 365–389.
Wikström, N., Savolainen, V., Chase, M.W. 2001. Evolution
of the angiosperms: calibrating the family tree. Proc.
Roy. Soc. London ser. B 268: 2211–2220.
Zdero, C., Bohlmann, F. 1990. Systematics and evolution
within the Compositae, seen with the eyes of a chemist.
Pl. Syst. Evol. 171: 1–14.
Alseuosmiaceae
Alseuosmiaceae Airy Shaw, Kew Bull. 18: 249 (1965).
Platyspermatiaceae Doweld (2001).
J. Kårehed
Shrubs, sometimes creeping or epiphytic subshrubs. Leaves alternate, sub-opposite or in
pseudo-whorls, simple, entire or serrate, estipulate. Multicellular, uniseriate hairs present in leaf
axils, rarely also on leaves and stems, and erect unicellular hairs sometimes present on both leaves and
stems. Flowers regular, hermaphroditic or functionally unisexual, tetra- or pentamerous, rarely up
to hexamerous, fascicled in the leaf axils or terminally, sometimes solitary, rarely racemose. Calyx
with free lobes. Corolla funnel-shaped or campanulate to urn-shaped, sympetalous, sometimes
only shortly tubular (Platyspermation), with more
or less lobed petal wings (not Platyspermation),
sometimes carunculate inside the lobes. Aestivation valvate. Stamens isomerous, attached to the
corolla, sometimes inserted at the very base of the
corolla tube, alternating with the corolla lobes,
sometimes sessile (Platyspermation). Anthers
introrse, longitudinally dehiscent. Disc present or
absent. Style single with capitate or discoid stigma,
often more or less bi- or trilobed. Ovary inferior or
sometimes semi-inferior, with two to three locules,
each containing two to many anatropous ovules.
Placentation axile. Fruits berries or (Platyspermation) capsules with one to several seeds with
minute embryo and copious endosperm.
Ten species classified into five genera in eastern
Australia, New Zealand, New Caledonia and New
Guinea.
Vegetative Morphology. Alseuosmiaceae are
shrubs, ranging from the creeping or epiphytic subshrubs of Wittsteinia to the sometimes 6-m-tall
Periomphale. The simple leaves are either entire
or serrate, lack stipules, and are alternate, subopposite, or in pseudo-whorls. Venation is pinnate,
very faint in Crispiloba. Leaves size varies between
3 and 20 cm.
Vegetative Anatomy (Gardner 1976; Dickison
1986, 1989; Platyspermation not investigated).
Rusty brown, multicellular uniseriate hairs are
present in the leaf axils. In Platyspermation
uniseriate hairs with persistent reddish bases are
found also on other parts of the plant (Stevens
2001). Erect unicellular hairs may be present on
both leaves and stems. The latter type is especially
abundant in Wittsteinia vacciniacea, forming
an indumentum on stems, petioles and basal
portions of the leaves. In contrast, Crispiloba and
Periomphale are (almost) completely devoid of this
hair type.
The leaf epidermis is thin and consists of one
cell layer. In transectional view, the epidermal cells
are square or rectangular. Their anticlinal walls are
usually undulate and deeply lobed in surface view.
The anomocytic stomata are level with the epidermis and have prominent outer cuticular ledges. The
one-layered palisade cells are poorly differentiated
from the lacunose spongy mesophyll. In Crispiloba, numerous long sclereids with thick lignified
walls form an “interwoven mass . . . that permeates
the mesophyll” (Dickison 1989). More or less rodshaped sclereids may be found around the midvein
in Periomphale. The latter are lobed or armed, have
thinner walls, and are not as elongate as the sclereids of Crispiloba. Sclerenchyma is present either
as bundle sheaths or as idioblastic sclereids in the
leaves of all taxa except Wittsteinia vacciniacea,
which completely lacks foliar sclerenchyma.
The nodes are trilacunar with three traces. In
Alseuosmia the traces fuse about halfway up the
petiole, in Wittsteinia papuana and Periomphale
they fuse at the base of the lamina, and in Crispiloba
and W. vacciniacea they remain separated throughout the petiole and into the lamina. Fibrous bundle
caps develop distally in the petiole, except in W.
vacciniacea.
All genera have an endodermis with prominent
Casparian banding present in young stems and surrounding the petiolar vascular bundles in all genera
but Periomphale. Calcium oxalate crystals have not
been detected.
8
J. Kårehed
The wood has very faint or no growth rings. The
narrow vessels are mostly solitary, sometimes in radial multiples, or rarely clustered. The mean number of bars of the scalariform perforation plates
ranges from 20 (Crispiloba) to 43 (Periomphale).
Intervessel pits commonly have circular borders
and are opposite and transitional to alternate, or
sometimes scalariform. In Alseuosmia and Crispiloba the vessel elements have fine helical thickenings. The imperforate elements are living, store
starch at maturity, and have indistinctly bordered
or simple pits. Both septate and non-septate fibres are present in all genera. Periomphale has tall
(> 1.5 cm) and wide, multiseriate rays, whereas the
rays of Crispiloba are heterocellular, shorter and
narrower. No rays are present in the other genera.
Axial parenchyma is very sparse or absent.
Inflorescences. Alseuosmia and Wittsteinia
have few-flowered fascicles or solitary flowers in
the leaf axils. In Periomphale the inflorescences are
predominantly terminal. Commonly, they consist
of fascicled flowers but racemes are sometimes
found. Terminal, umbel-like, mostly pedunculate
inflorescences of one to five flowers are found
in Crispiloba. Platyspermation has few-flowered
inflorescences with inclined flowers. Pedicels
have few bracts which are very early caducous in
Periomphale.
Flower Morphology. The flowers are regular
with whorls of normally four or five flower
parts; hexamerous flowers are sometimes encountered (septamerous flowers in Periomphale
were reported by Baillon 1888). According to
Tirel and Jérémie (1996), Periomphale has both
hermaphroditic (perhaps functionally male) and
functionally female flowers. In the other genera,
the flowers are hermaphroditic. The calyx lobes are
valvate, free, more or less triangular, and persistent
or circumscissile-caducous (Alseuosmia). The
corolla tube of Platyspermation is short whereas
in the other genera the clearly sympetalous corolla
is funnel-shaped or campanulate to urn-shaped.
The colour of the corolla ranges from dull red in
Alseuosmia macrophylla over various pale shades
of pink, yellow and green, with or without red
markings, to pure white in Crispiloba. The valvate
corolla lobes have appendages, so-called petal
wings, which are more or less lobed (Fig. 2A–E). In
Crispiloba they are conspicuously fringed whereas
in Platyspermation the corolla lobes lack evident
petal wings but are papillate. Petal wings remi-
niscent of those in Alseuosmiaceae are also found
in Goodeniaceae and Menyanthaceae (Gustafsson
1995). In Crispiloba, the base of the midrib on the
inside of the lobes is fringed in a similar way. Also,
there are irregular appendages at the throat of
the corolla tube. In Wittsteinia and Periomphale,
a caruncle at the base of the lobes forms a ‘corona’
which may cover the tube (Fig. 2D). The length
of the tube varies from c. 0.5 cm (Wittsteinia
papuana and small-flowered Periomphale) to
4.5 cm (Alseuosmia macrophylla and Crispiloba).
The stamens alternate with the corolla lobes, and
are inserted either in the throat of the corolla tube
(Alseuosmia and Crispiloba) or at the very base of
the tube. The introrse anthers open by longitudinal
slits. In Platyspermation the anthers are sessile
and extrorse (?), brown hairy below, and with
a large flat connective appendage at the apex (van
Steenis 1982). A disc is mostly present. It is very
reduced or missing in Wittsteinia, and inhabited
by parasitic insects in flowers of Periomphale
(see below). The single style has a capitate or
discoid, often bilobed stigma, sometimes trilobed
in Wittsteinia vacciniacea. The ovary is inferior
or sometimes initially semi-inferior in Periomphale and two-locular; Wittsteinia vacciniacea
commonly has three locules. Each locule contains two to many anatropous ovules with axile
placentation, in Platyspermation the placenta is
brown and very thick (van Steenis 1982). Besides
the normal type of flowers, Tirel (1996) and Tirel
and Jérémie (1996) described flowers inhabited
by parasitic insects in Periomphale. These are
globose or obconical, do not open, and are often
borne on elongated pedicels (up to 6 cm long).
The stamens and the style are frequently only
weakly developed, as is the corolla, which is shed
prematurely or withers. A disc is usually lacking
and the ovules, if present, do not develop into
seeds. Similar flowers are reportedly present also in
Wittsteinia and Platyspermation (van Steenis 1984;
Stevens 2001).
Floral Anatomy. The floral anatomy of Alseuosmia and Periomphale was investigated by Gardner (1976). According to him, the anatomical features are uniform within Alseuosmia, and essentially agree with those in Periomphale. Notably, at
the top of the ovary the locules interconnect above
the placental region for about 50–100 µm, due to
the septum being transversely divided (in one examined flower of Periomphale, this resulted in parietal placentation of the uppermost ovule).
Alseuosmiaceae
9
tegillate, slightly undulating, thicker than the
nexine, sometimes with small fissures; ectosexine
thicker than endosexine, the latter only faintly
baculate. According to Hufford (1992) and Kårehed
et al. (1999), Alseuosmia pollen has well-developed
columellae. The pollen of Periomphale is similar
in appearance to that of Alseuosmia (Bortenschlager et al. 1966), as is that of Platyspermation
(3-colpor(oid)ate, oblate, spheroidal, c. 31 × 36 µm
and with the sexine thicker than the nexine;
Erdtman 1952).
Fruit and Seed. The fruits of Alseuosmiaceae
are two-locular berries, two- to three-locular in
Wittsteinia, or two-locular capsules (Platyspermation). Their shape varies from globose (W. vacciniacea) to narrowly ellipsoid. Each fruit contains one
to many ellipsoid or ovoid, more or less compressed
seeds with a brown to black testa. In Platyspermation, seeds are sculptured and have fine, inflated
hairs on the margin (van Steenis 1982). The seed
coat structure and the lignified exotesta cells of
Alseuosmia have been described by NemirovichDanchenko and Lobova (1998).
Fig. 2. Alseuosmiaceae. A Alseuosmia macrophylla, habit.
B A. banksii, flower with opened corolla. C Crispiloba disperma, habit. D Periomphale balansae, female flower; longitudinal section of ovary. E Wittsteinia vacciniacea, habit.
(Redrawn after A Hooker 1887, B Hooker 1853–1855, C van
Steenis 1984, D Tirel and Jérémie 1996, E Mueller 1885)
Embryology. In Alseuosmia the anther wall
consists of an epidermis, an endothecium with
fibrous thickenings when mature, two middle layers and a one-layered tapetum. The tapetum cells
are binucleate before meiosis in the pollen mother
cells, remain in place during pollen development,
and subsequently undergo nuclear fusion. The
pollen is shed in the trinucleate condition. The
ovules of Alseuosmia are unitegmic and tenuinucellate, and the embryo sac is eight-nucleate at
maturity (Gardner 1976).
Karyology. The diploid chromosome number
of Alseuosmia is 2n = 18 (Gardner 1976).
Pollen. The 3-colporate pollen of Alseuosmia
was studied by Erdtman (1952) who described
the pollen grains as often angulaperturate, oblate
spheroidal-prolate (longest axis 45–50 µm). Sexine
Reproductive Biology. According to Gardner
(1976), Alseuosmia pusilla is self-compatible,
whereas A. macrophylla is an obligate outbreeder
with an incompatibility mechanism operating at
the stylar level.
Phytochemistry. In the leaves of Alseuosmia,
Cambie and Parnell (1969) detected lupeol, lupenyl
acetate, stigmasterol, stearic acid, and triterpene
acetates. Also from Alseuosmia, Gardner (1976) reported the presence of a condensed tannin (leucocyanidin) and ellagitannin, together with simple
phenols (quercetin, caffeic acid, kaempferol, and
p-coumaric acid) and triterpenoid saponins. He
could not detect either alkaloids or iridoids.
Affinities. Alseuosmiaceae form a monophyletic group in Asterales, together with
Argophyllaceae and Phellinaceae (Gustafsson et al.
1996; Backlund and Bremer 1997; Gustafsson and
Bremer 1997; Källersjö et al. 1998; Kårehed et al.
1999; Lundberg and Bremer 2003).
Before the recent addition of Platyspermation
(see below), several studies supported Alseuosmiaceae as a well-defined family (e.g. van Steenis
1984; Dickison 1986, 1989; Kårehed et al. 1999).
Already when Cunningham (1839) described
Alseuosmia, he suggested it to be a distinct family,
10
J. Kårehed
related to Cornaceae, Caprifoliaceae and Loranthaceae. Alseuosmia was, however, mostly treated
as a more or less aberrant member of Caprifoliaceae (e.g. Hooker 1873; Fritsch 1891), as were the
two New Caledonian genera Memecylanthus and
Pachydiscus (Schlechter 1906; Guillaumin 1948)
now included in Periomphale (van Steenis 1978).
Initially, Baillon (1888) placed Periomphale in Gesneriaceae and, due to lack of detailed knowledge,
the genus was mostly kept in that family until Airy
Shaw (1965) realized its correct affinities. Airy
Shaw (1965) also gave the first formal description
of Alseuosmiaceae, indicating affinities with
Escalloniaceae. Gardner (1976, 1978a) suggested
Alseuosmiaceae to be most closely related to
Argophyllaceae, then included in Escalloniaceae
but now considered a separate family of Asterales
(e.g. Källersjö et al. 1998; Kårehed et al. 1999).
van Steenis (1978, 1984) added to the family the
newly discovered Wittsteinia papuana and W. vacciniacea, formerly thought to belong in Ericaceae
(von Mueller 1861; Bentham 1869; Drude 1889;
Stevens 1971) or Epacridaceae (Burtt 1949), and
Crispiloba, earlier misplaced in Rubiaceae (Moore
1917). van Steenis (1984) included Periomphale
in Wittsteinia but Tirel (1996) and Tirel and
Jérémie (1996) argued that it should be regarded
as a distinct genus. Alseuosmiaceae obtained their
present circumscription when molecular studies
showed the little known genus Platyspermation
to be the sister to Alseuosmiaceae (Lundberg and
Bremer 2003). Here, Platyspermation is included
in the family, rather than recognising a monotypic
Platyspermataceae (Doweld 2001). Platyspermation was originally described as a member of
Myrtaceae (Guillaumin 1950). Airy Shaw (in Willis
1973) placed it in Rutaceae, while a relationship to
Escalloniaceae was suggested by Erdtman (1952),
Schmid (1980) and van Steenis (1982).
Distribution and Habitats. Alseuosmia is
found in lowland to montane forests in New
Zealand (Gardner 1976). Crispiloba is endemic
to rain forests of Queensland, Australia. On New
Caledonia, Periomphale inhabits humid forests
(Tirel and Jérémie 1996), as does Platyspermation
which is also found in the maquis vegetation.
Wittsteinia is a mountainous genus; the Australian
W. vacciniacea grows in crevices of rocks and on
rocky summits of Victorian mountains (Bentham
1869), and the Papua New Guinean W. papuana
has been collected in woodland at 3,000 m (van
Steenis 1978).
Key to the Genera
1. Corolla tube short; stamens sessile; fruit a capsule
5. Platyspermation
– Corolla tube well developed; stamens inserted in the
corolla tube; fruit a berry
2
2. Corolla funnel-shaped or narrowly cylindrical; stamens inserted at the apex of the corolla tube
3
– Corolla campanulate to urn-shaped or narrowly
barrel-shaped; stamens inserted at the base of the
corolla tube
4
3. Leaves alternate, (sub)serrate; flowers fascicled
or solitary in the leaf axils; calyx circumscissilecaducous; corolla funnel-shaped with ± lobed petal
wings
1. Alseuosmia
– Leaves in pseudo-whorls, entire; flowers terminal;
calyx persistent; corolla narrowly cylindrical with
strongly fringed petal wings
2. Crispiloba
4. Shrubs; leaves entire; inflorescences terminal; corolla
with entire petal wings
3. Periomphale
– Subshrubs; leaves serrate; inflorescences axile; corolla
with lobed petal wings
4. Wittsteinia
Genera of Alseuosmiaceae
1. Alseuosmia A. Cunn.
Fig. 2A, B
Alseuosmia A. Cunn., Ann. Nat. Hist. 2: 209 (1839); Merrett
& Clarkson, N. Z. J. Bot. 38: 153–164 (2000), rev.
Shrubs. Leaves alternate, simple, serrate, sometimes subentire. Flowers fascicled in the leaf axils
or solitary, regular, hermaphroditic, tetra- or pentamerous, rarely up to hexamerous. Corolla with
a funnel-shaped tube and valvate lobes with more
or less fringed petal wings. Stamens inserted at
apex of corolla tube. Disc present. Stigma capitate
or bilobed on elongate style. Ovary inferior, twolocular with two to many ovules in each locule.
Fruit a two-locular berry. n = 9. Five species in
New Zealand.
2. Crispiloba Steen.
Fig. 2C
Crispiloba Steen., Blumea 29: 391 (1984).
Shrub. Leaves in pseudo-whorls of 3–6, entire. Inflorescences terminal, predominantly pedunculate,
umbel-like with (one) up to five flowers. Flowers
(tetra-) pentamerous, with a salver-shaped corolla
and a long, narrow, cylindrical tube. Corolla
lobes fringed inside and with strongly fringed
petal wings. Stamens inserted at the throat of the
corolla tube. Ovary inferior with two locules, each
containing 2–3 ovules. Fruit a berry with black,
more or less planoconvex seeds. One species, C.
disperma (S. Moore) Steen., Queensland, Australia.
Alseuosmiaceae
3. Periomphale Baill.
Fig. 2D
Periomphale Baill., Bull. Mens. Soc. Linn. Paris 1: 731 (1888);
Tirel & Jérémie, Fl. Nouvelle-Calédonie 20: 100–106 (1996),
rev.
Memecylanthus Gilg & Schltr. (1906).
Pachydiscus Gilg & Schltr. (1906).
Shrubs with flexuose branches. Leaves alternate, often sub-opposite or in pseudo-whorls at the tip of
the branches, entire. Inflorescences predominantly
terminal, rarely racemose or more commonly of
(1)2–8(15)-fascicled flowers. Flowers tetra- or pentamerous (rarely hexamerous), hermaphroditic or
functionally unisexual. Corolla urn-shaped to campanulate; corolla lobes carunculate inside. Stamens
inserted at the very base of the corolla tube. Disc
present, rarely only weakly developed. Ovary inferior, sometimes initially semi-inferior, with two
locules, each containing 4–6 ovules. Fruit a subcylindric to ovoid-fusiform berry with more or
less compressed, ellipsoid seeds. One species, P.
balansae Baill., endemic to New Caledonia.
4. Wittsteinia F. Muell.
Fig. 2E
Wittsteinia F. Muell., Fragm. 2: 136 (1861).
Subshrubs, creeping with ascending branches, W.
papuana (Steen.) Steen. epiphytic. Leaves alternate
or in pseudo-whorls, serrate. Inflorescences of one
or two axillary flowers. Corolla more or less campanulate or narrowly barrel-shaped (W. papuana
(Steen.) Steen.); corolla lobes carunculate inside
(inconspicuously so in W. vacciniacea F. Muell.) and
with lobed petal wings. Stamens inserted on the
corolla tube at the very base. Ovary inferior, twoto three-locular, few ovules in each locule. Fruit
a globose berry with ovoid seeds. Two species; one
endemic to Victoria, Australia, and one to Papua
New Guinea.
5. Platyspermation Guillaumin
Platyspermation Guillaumin, Acta Horti Gotob. 18: 253
(1950).
Shrub up to 5 m high. Leaves alternate, pseudowhorled at the end of branches, with small
dentations on the recurved margins, sclerophyllous. Inflorescences few-flowered with inclined
flowers. Corolla with short tube. Stamens sessile.
Anthers with brown hairs below. Ovary inferior,
two-locular, with few ovules on thick, brown
placenta. Fruit a two-locular capsule with few, flat
seeds. One species, P. crassifolium Guillaumin,
endemic to New Caledonia.
11
Selected Bibliography
Airy Shaw, H.K. 1965. Diagnoses of new families, new
names, etc., for the seventh edition of Willis’ ‘Dictionary’. Kew Bull. 18: 249–273.
Backlund, A., Bremer, B. 1997. Phylogeny of the Asteridae s.
str. based on rbcL sequences, with particular reference
to the Dipsacales. Pl. Syst. Evol. 207: 225–254.
Baillon, H.E. 1888. Observations sur les Gesnériacées. Bull.
Mens. Soc. Linn. Paris 1: 731–732.
Bentham, G. 1869. Flora Australiensis, vol. IV. London:
Lovell Reeve.
Bortenschlager, S., Erdtman, G., Praglowski, J. 1966. Pollenmorphologische Notizen über einige Blütenpflanzen
incertae sedis. Bot. Notiser 119: 160–168.
Burtt, B.L. 1949. Studies in the Ericales, IX. The taxonomic
position of Wittsteinia. Kew Bull. 3: 493–495.
Cambie, R.C., Parnell, J.C. 1969. A New Zealand phytochemical survey, part 7. Constituents of some dicotyledons.
N. Z. J. Sci. 12: 453–466.
Cunningham, A. 1839. Florae Insularum Novae Zelandiae
Precursor; or a specimen of the botany of the islands
of New Zealand. Genus Corneis affine. Ann. Nat. Hist.
2: 209–210.
Dickison, W.C. 1986. Wood anatomy and affinities of the
Alseuosmiaceae. Syst. Bot. 11: 214–221.
Dickison, W.C. 1989. Stem and leaf anatomy of the Alseuosmiaceae. Aliso 12: 567–578.
Doweld, A.B. 2001. Prosyllabus Tracheophytorum. Tentamen Systematis Plantarum Vascularium (Tracheophytorum). Moscow: GEOS.
Drude, O. 1889. Ericaceae. In: Engler, A., Prantl, K., Die
natürlichen Pflanzenfamilien, IV, 1. Leipzig: W. Engelmann, pp. 15–65.
Erdtman, G. 1952. Pollen morphology and plant taxonomy.
Angiosperms. Stockholm: Almqvist & Wiksell.
Fritsch, K. 1891. Caprifoliaceae. In: Engler, A., Prantl, K.,
Die natürlichen Pflanzenfamilien, IV, 4. Leipzig: W.
Engelmann, pp. 156–169.
Gardner, R.O. 1976. Studies in the Alseuosmiaceae. Ph.D.
Thesis, University of Auckland, Auckland, New
Zealand.
Gardner, R.O. 1978a. Systematic notes on the Alseuosmiaceae. Blumea 24: 138–142.
Gardner, R.O. 1978b. The species of Alseuosmia (Alseuosmiaceae). N. Z. J. Bot. 16: 271–277.
Guillaumin, A. 1948. Flore (analytique et synoptique) de la
Nouvelle Calédonie, Phanérogams. Paris: Office de la
Recherche Scientifique Coloniale.
Guillaumin, A. 1950. Plantae Neocaledonicae a C. Skottsberg a. 1949 lectae (96ème contribution à la flore de
la Nouvelle-Calédonie). Acta Horti Gotob. 18: 247–
265
Gustafsson, M.H.G. 1995. Petal venation in Asterales and
related orders. Bot. J. Linn. Soc. 118: 1–18.
Gustafsson, M.H.G., Bremer, K. 1997. The circumscription
and systematic position of Carpodetaceae. Austral.
Syst. Bot. 10: 855–862.
Gustafsson, M.H.G., Backlund, A., Bremer, B. 1996. Phylogeny of the Asterales sensu lato based on rbcL sequences with particular reference to the Goodeniaceae.
Pl. Syst. Evol. 199: 217–242.
Hooker, J.D. 1853–1855. The botany of the Antarctic voyage of H.M. Discovery ships Erebus and Terror in
12
J. Kårehed
the years 1839–43. II. Flora Novae-Zelandiae. London:
Lovell Reeve.
Hooker, J.D. 1873. Caprifoliaceae. In: Bentham, G.,
Hooker, J.D., Genera Plantarum, vol. II, part 1.
London: Lovell Reeve, pp. 1–7.
Hooker, J.D. 1887. Curtis’s Botanical Magazine, vol. CXIII.
London: Lovell Reeve.
Hufford, L. 1992. Rosidae and their relationships to other
nonmagnoliid dicotyledons: a phylogenetic analysis
using morphological and chemical data. Ann. Missouri
Bot. Gard. 79: 218–248.
Källersjö, M., Farris, J.S., Chase, M.W., Bremer, B., Fay, M.F.,
Humphries, C.J., Petersen, G., Seberg, O., Bremer, K.
1998. Simultaneous parsimony jackknife analysis
of 2538 rbcL DNA sequences reveals support for
major clades of green plants, land plants, seed
plants and flowering plants. Pl. Syst. Evol. 213:
259–287.
Kårehed, J., Lundberg, J., Bremer, B., Bremer, K. 1999. Evolution of the Australasian families Alseuosmiaceae,
Argophyllaceae, and Phellinaceae. Syst. Bot. 24: 660–
682.
Lundberg, J., Bremer, K. 2003. A phylogenetic study of the
order Asterales using one morphological and three
molecular data sets. Intl J. Pl. Sci. 164: 553–578.
Moore, S. 1917. A contribution to the phyto-geography of
Bellenden-Ker. II. Systematic account. Phanerogams.
J. Bot. 55: 302–309.
Mueller, F. von 1861. Fragmenta Phytographiae Australiae,
vol. II. Melbourne: Auctoritate Gubern. Coloniae Victoriae.
Mueller, F. von 1885. Key to the system of Victorian plants.
Melbourne: Government Printer.
Nemirovich-Danchenko, E.N., Lobova, T.A. 1998. The seed
coat structure in some representatives of the order
Hydrangeales. Bot. Zhurn. (Moscow & Leningrad) 83:
1–9.
Schlechter, R. 1906. Beiträge zur Kenntnis der Flora von
Neu-Kaledonien. Bot. Jahrb. Syst. 39: 1–274.
Schmid, R. 1980. Comparative anatomy and morphology
of Psiloxylon and Heteropyxis, and the subfamilial and
tribal classification of Myrtaceae. Taxon 29: 559–595.
Stevens, P.F. 1971. A classification of the Ericales: subfamilies and tribes. Bot. J. Linn. Soc. 64: 1–53.
Stevens, P.F. 2001 (onwards). Angiosperm Phylogeny
website, version 4, May 2003. http://www.mobot.org/
MOBOT/research/APweb/
Tirel, C. 1996. Rétablissement de Periomphale Baill. (Alseuosmiaceae), genre endémique de Nouvelle-Calédonie.
Bull. Mus. Natl Hist. Nat. B, Adansonia 18: 155–160.
Tirel, C., Jérémie, J. 1996. Alseuosmiaceae. In: Morat, P. (ed.)
Flore de la Nouvelle-Calédonie, vol. 20. Paris: Muséum
National d’Histoire Naturelle, pp. 100–106.
van Steenis, C.G.G.J. 1978. The genus Periomphale in New
Guinea (Caprifoliaceae). Blumea 24: 480–481.
van Steenis, C.G.G.J. 1982. 157. Preliminary note on the
taxonomic disposition of Platyspermation Guillaumin
(Myrtaceae) from New Caledonia. In: van Steenis, C.G.G.J., Veldkamp, J.F., Miscellaneous botanical
notes XXVI. Reinwardtia 10: 21–26.
van Steenis, C.G.G.J. 1984. A synopsis of Alseuosmiaceae
in New Zealand, New Caledonia, Australia, and New
Guinea. Blumea 29: 387–394.
Willis, J.C. 1973. A dictionary of the flowering plants and
ferns, ed. 8, revised by H.K. Airy Shaw. Cambridge:
University Press.
Argophyllaceae
Argophyllaceae (Engl.) Takht., Systema Magnoliophytorum: 208 (1987).
Corokiaceae Kapil ex Takht. (1997).
J. Kårehed
Shrubs or small trees. T-shaped trichomes present
on stems, leaves, inflorescences, and flowers.
Leaves alternate (or in 3–4-leaved fascicles on
brachyblasts), simple, estipulate with entire or
serrate margins. Flowers borne in axile or sometimes terminal panicles or racemes, sometimes
in few-flowered fascicles or solitary, regular,
hermaphroditic, mostly pentamerous. Aestivation
valvate. Sepals 5(8), connate at base, persistent.
Petals 5(8), white or yellow, joined at their very
base or free, alternating with the sepals, with
fringed appendages (corolline ligules) on the
adaxial side. Stamens isomerous, alternipetalous,
anthers elongate-ovate or elongate. Ovary semiinferior or inferior, 1–3(6)-locular, with one to
many anatropous ovules in each locule. Placentation axile. Style with more or less capitate,
2–5-lobed stigma. Fruits loculicidal capsules
or drupes. Seeds obovate or linear-elongate
with minute or elongate embryo in fleshy endosperm.
Two genera comprising c. 20 species distributed in eastern Australia, Lord Howe Island,
New Caledonia, New Zealand, and Rapa Island.
Vegetative Morphology. Argophyllaceae are
a family of shrubs or small trees up to 7 m tall.
Argophyllum has branches with brown or blackish
(A. ellipticum) bark and yellow or reddish (A.
ellipticum) wood. In Corokia the branches are dark
brown to almost black. Zigzagging, interlacing
branches are characteristic for C. cotoneaster. The
leaves are alternate or arranged in fascicles of three
to four on brachyblasts (C. cotoneaster), simple,
entire or serrate, and exstipulate. Argophyllum has
ovate, obovate, or linear-elliptic leaves, whereas
the lamina in Corokia is elliptic or lanceolate
to obovate or oblanceolate. The leaf shape is
variable in C. cotoneaster; adult leaves are obovate
to orbicular, sometimes emarginate, and leaves
on seedlings are obovate-spathulate, often threelobed.
Vegetative Anatomy. T-shaped hairs are
present on (young) stems, leaves (depending on
species in various degrees on petioles, upper and
lower side, as well as on the margin), inflorescences,
sepals, and on the abaxial side and the margin of
the petals. Especially on the lower side of the leaves,
they may give the impression of a whitish, silvery,
or rusty indumentum. They consist of a multicellular, uniseriate stalk (in Corokia cotoneaster
the stalk sometimes consists of only one cell) and
a very elongated, T-shaped terminal cell (see, e.g.
Al-Shammary and Gornall 1994). There are small
slits at the junction of this T-cell with the stalk.
In some Argophyllum species, these are oriented
parallel to the long axis of the T-cell, whereas in
other species and in Corokia the slits cross the
long axis obliquely or at right angles. Carbonate
deposits are present on the cuticle of the T-hairs in
some Corokia species but not in Argophyllum.
The leaves have a poorly differentiated, uni- or
biseriate palisade layer and a compact or lacunose
(Corokia) spongy mesophyll. The uppermost palisade cells, as seen in cross-section, are columnarelongate or short and square (some Argophyllum
species). A one- to three-layered hypodermis is
present in Argophyllum ellipticum and A. nitidum.
The stomata are of the haplocheilic type and are
anomocytic. The nodes are trilacunar (pentalacunar in A. laxum and unilacunar in C. x virgata),
with three traces which either fuse halfway up the
petiole into a shallow arc or split into more traces
(A. ellipticum and A. laxum).
The following description of wood anatomy is
based mainly on the studies of Patel (1973a, b), Hils
(1985), and Noshiro and Baas (1998). The wood
is dense, diffuse-porous or semi-ring-porous,
whitish to yellow, pinkish to reddish, or very pale
brown. The vessels are mostly solitary, occasionally
in pairs or small groups, round-angular to angular
in outline. Perforation plates are scalariform with
rather numerous bars (6–32+). Intervessel pits and
vessel-ray pits are transitional, opposite or, more
14
J. Kårehed
commonly, alternate. Fine helical thickenings
are present in the vessel elements. The vascular
tracheids (not present in Argophyllum and Corokia
collenettei) have distinct helical thickenings. Both
genera have septate fibres with minutely, nonbordered, distinctly or indistinctly bordered pits
on the tangential and radial walls. Very fine helical
thickenings have been recorded in the fibres of
Corokia. Axial parenchyma is absent or very sparse,
scanty paratracheal and diffuse. Rays are heterocellular, one to six cells broad. Crystals are not present.
Inflorescences. The inflorescences are axile or
sometimes terminal panicles, racemes, corymbs
(in some Argophyllum), or few-flowered fascicles
(Corokia cotoneaster; along with solitary flowers).
The bracts and prophylls are mostly small and
linear.
Flower Structure. The flowers are regular,
hermaphroditic, and predominantly pentamerous.
In Corokia collenettei, hexamerous flowers are the
normal condition. Higher merism occurs occasionally; eight-merous flowers have been reported for
C. collenettei (Brown 1928). Aestivation is valvate.
The calyx is turbinate and the calyx tube is adnate
to the ovary. The sepals are 0.5–3 mm long. The
petals, which alternate with the sepals, are 2–6 mm
long, white or yellow, and have fringed appendages
adaxially. These appendages (Fig. 3C, D, F), called
corolline ligules by Eyde (1966), are only a few
cells thick and almost as long as to a little longer
than half the length of the petals. They are divided
for two-thirds of their length into about 10–20
fringes, and are found in all species except Corokia
macrocarpa. The alternipetalous, free stamens
have elongate-ovate or elongate (Corokia), longitudinally dehiscent, introrse anthers. The short style
has a capitate or inconspicuously 2–3(5)-lobed
stigma. In Argophyllum the ovary is semi-inferior
at anthesis but inferior early in development (Tobe
and Raven 1999). It has predominantly two or three
locules (occasionally four; in A. verae this number
is normal (Forster 1990); Eichler (1878) reported
of an ovary with five locules in A. nitidum). In
Corokia there is one pendulous ovule in each
locule of the inferior ovary. The number of locules
is mostly one or two, sometimes up to four in C.
buddleioides (Kapil and Bhatnagar 1992). Ovaries
of C. collenettei have regularly up to four locules,
and Eyde (1966) reported a five-locular ovary in
this species. The placentation in both genera is
axile and the ovules are anatropous, unitegmic,
Fig. 3. Argophyllaceae. Argophyllum ellipticum. A Habit. B
Bud. C Flower. D Flower opened out. E Stamens, adaxial and
abaxial view. F Corolla, laid open. G Flower with corolla
partly removed. H Transverse section of capsule. (From
Labillardière 1824)
and tenuinucellate (e.g. Mauritzon 1933; Kapil and
Bhatnagar 1992). Numerous ovules are present in
each locule in Argophyllum, whereas in Corokia
there is a single, pendulous, apically attached
ovule in each locule. Corokia has a usually bright
orange, epigynous, pulvinate nectariferous disk.
Floral Anatomy. The vascular pattern of
Corokia flowers was thoroughly described by
Eyde (1966). He noted anatomical similarities
with Argophyllum and that Corokia differs from
Cornaceae in having vascular bundles running
longitudinally through the centre of the inferior
gynoecium (Eyde 1966).
Embryology. The anthers of Corokia are
tetrasporangiate, have a four-layered wall, a secre-
Argophyllaceae
tory tapetum, and endothecial cells which develop
fibrous thickenings prior to dehiscence (Kapil and
Bhatnagar 1992).
The embryo of Argophyllum is minute and surrounded by fleshy endosperm. The following information on Corokia is extracted mainly from the
works of Mauritzon (1933), Kapil and Bhatnagar
(1992), and Lobova (1997). The layer of integument cells which border the embryo sac is uniform
and the cells are radially elongated. The embryo
sac is of the Polygonum type. The synergids are
very long and have a small vacuole below their nuclei. The egg cell is small and the polar nuclei are
positioned just below it. The antipodal cells are persistent. The pollen tube traverses through a wide
stylar canal to reach the ovule and the embryo sac.
Endosperm is cellular with aggressive micropylar
and chalazal haustoria (Kapil and Bhatnagar 1992).
Embryogeny is of the Chenopodiad type. When
mature, the cylindrical embryo has a long radiclehypocotyl axis and short cotyledons, compared to
the embryo of Argophyllum.
Karyology. The diploid chromosome number
for Corokia is 2n = 18 (Wanscher 1933; Hamel 1953;
Hair and Beuzenberg 1959).
Pollen Morphology. Hideux and Ferguson
(1976) and Ferguson and Hideux (1978) have
studied the pollen of Argophyllaceae. The pollen
is tricolporate with colpi longer than two-thirds
of the polar axis. It has a prominent, complex
H-shaped endo-aperture with thinning of the
endexine and association of the lamellation in the
apertural region (Ferguson and Hideux 1978). The
pollen grains are spheroidal; 15–20 × 15–20 µm
(polar axis × equatorial diameter) in Argophyllum,
and 25–33 × 24–30 µm in Corokia. The exine
is 1–1.5 µm thick in both genera. The tectum is
complete perforate and Corokia has supratectal
spines. The tectum:columellae ratio is less than 1,
and the solea (foot-layer):endexine ratio is equal
to or larger than 1.
Pollination. Webb (1994) studied the pollination system of Corokia cotoneaster. The flowers
seem to be functional for 5–6 days and are selfincompatible. They are not dichogamous, but are
slightly herkogamous when fully open. Twelve insect species, mostly bees and flies, have been recorded to visit the flowers, some of which, especially the bee Lasioglossum sordidum, seem to be
legitimate pollinators.
15
Fruit and Seed. The fruit of Argophyllum is a loculicidal capsule opening with as many teeth as
locules. The teeth split, however, along their midrib,
so that in the open fruit their number is twice
the number of locules. The ripe seeds are obovate,
weakly triangular, and the shiny brown surface is
large-celled. The proximal and radial walls of the
almost hexagonal exotestal cells have very strong
thickenings. Corokia has yellow, orange, bright red
to purplish red, or blackish drupes, c. 2–12 mm
long, crowned by the persistent calyx. The spherical to obovoid endocarp is thick and woody. At the
apical end of the endocarp, there are woody germination plugs formed by the lignification of cells of
the placentae. The seed surface is small-celled compared to that of Argophyllum, the exotestal layer is
thinner, and the cells are smaller and have thinner
walls. More elaborate descriptions of the fruits and
seeds of Argophyllaceae are found in, for example,
Eyde (1966) and Lobova (1997).
Phytochemistry. Two species of Argophyllum
are able to hyperaccumulate nickel (Jaffré et al.
1979). Triterpenes have been reported from both
Argophyllum and Corokia (Briggs et al. 1967;
Cambie and Parnell 1969). Flowers, fruits, and
leaves of Corokia have darkly stained cells, which
Eyde (1966) called tannin-containing cells. Similar
cells from Argophyllum have been mentioned by
Zemann (1907) and Kårehed et al. (1999), and
tannin-like substances in the testa of both genera
were noticed by Lobova (1997). Leucoanthocyanins
are present in both genera (Cambie et al. 1961;
Gibbs 1974). In addition, gallotannin (Hegnauer
1969) and a number of additional phenolic constituents have been detected in Corokia (cyanidin,
quercetin, caffeic acid, kaempferol: Gardner 1976;
gallic acid, quinic acid: Hegnauer 1969), as well
as iridoids (Wieffering 1966; Jensen et al. 1975).
Attempts to detect HCN in Corokia have given
negative results (Gibbs 1974).
Affinities. Argophyllaceae belong in Asterales
(Gustafsson et al. 1996; Backlund and Bremer 1997;
Gustafsson and Bremer 1997; Källersjö et al. 1998;
Kårehed et al. 1999). According to morphological and DNA sequence data (rbcL and ndhF), Argophyllaceae and their sister group Phellinaceae
form a monophyletic group with Alseuosmiaceae
(Kårehed et al. 1999). Hallier (1908) was the first
to suggest that the two genera of Argophyllaceae
are related to each other, emphasizing the presence of corolline ligules and T-hairs in both Ar-
16
J. Kårehed
gophyllum and Corokia. The family was erected by
Takhtajan (1987) to acknowledge the increased evidence for a close relationship between the two genera from, e.g. floral anatomy (Eyde 1966), pollen
morphology (Hideux and Ferguson 1976; Ferguson and Hideux 1978), and wood anatomy (e.g.
Hils 1985). Previously, Argophyllum had been regarded as a member of either Saxifragaceae (e.g.
Engler 1890, 1928; Schlechter 1906; Thorne 1992)
or Escalloniaceae (Willis 1966; Takhtajan 1983).
Corokia had mostly been placed either in Cornaceae (Hooker 1867; Harms 1897; Wangerin 1909;
Melchior 1964; Hutchinson 1967; Cronquist 1981)
or together with Argophyllum in the aforementioned families (e.g. Engler 1928; Takhtajan 1983;
Thorne 1983, 1992).
Takhtajan (1997) elevated Corokia to family
level, noting that it differs from Argophyllum in
the structure of fruit and seed, placentation and
the number of locules (1–2-locular ovaries). The
latter, however, is not correct (see above). The two
genera are closely related to each other and share
the presence of, for example, the easily recognized
corolline ligules and the T-shaped hairs.
Distribution and Habitats. The Australian
Argophyllum species, one in New South Wales
and about four in Queensland, inhabit more or
less mountainous rainforests, except A. verae
which grows on sandstone ledges far north on the
Cape York Peninsula (Forster 1990). The ten New
Caledonian species are more frequent in open,
sunny places (Schlechter 1906; Guillaumin and
Virot 1953).
Corokia species are found in diverse habitats,
ranging from lowland shrubland, river flats,
and rocky places (C. cotoneaster) to rainforests
(C. whiteana). The distribution of the genus is
intriguing. Corokia whiteana is known only from
one mountain range in New South Wales, on
Lord Howe Island there is another species (C. carpodetoides), three species inhabit New Zealand (C.
cotoneaster on both the North and South Island,
C. buddleioides restricted to the North Island, and
C. macrocarpa on the Chatham Islands), and yet
another species is found on the isolated, volcanic
Rapa Island (C. collenettei), more than 6,000 km
from Australia.
Economic Importance and Conservation.
The New Zealand species of Corokia and the hybrid
C. x virgata are cultivated as ornamentals. Argophyllum nullumense is cultivated in some plant
nurseries in Australia for its beautiful foliage. An
increased interest in cultivating this and the other
Australian Argophyllum species would facilitate
the conservation of these rare rainforest plants.
One Argophyllum species, A. verae, is known only
from the type locality (Forster 1990), as is the
vulnerable C. whiteana (Smith 1958).
Key to the Genera
1. Ovary semi-inferior with many ovules in each locule;
loculicidal capsules
1. Argophyllum
– Ovary inferior with one apically attached ovule in each
locule; drupes
2. Corokia
1. Argophyllum J.R. Forst. & G. Forst.
Fig. 3
Argophyllum J.R. Forst. & G. Forst., Char. Gen. 29, t. 15
(1776); Zemann, Ann. K.K. Naturhist. Hofmus. 22: 270–292
(1907), rev.
Shrubs or small trees with alternate, simple leaves,
either entire or serrate. Inflorescence a panicle
or raceme with regular pentamerous or rarely
hexamerous flowers. Stigma capitate, two- to
five-lobed, on short style. Semi-inferior ovary with
many ovules in each locule. Locules mostly two or
three, rarely four or five. Fruit a loculicidal capsule
with obovate seeds. About 15 species from eastern
Australia (five, including one undescribed) and
New Caledonia (ten).
2. Corokia A. Cunn.
Corokia A. Cunn., Ann. Nat. Hist. 3: 249 (1839).
Shrubs or small trees with alternate, simple leaves,
either entire or with a few teeth (C. carpodetoides
(F. Muell.) L.S. Sm. and C. collenettei L. Riley). Inflorescence mostly a panicle or raceme, sometimes
a few-flowered fascicle or flowers solitary. Flowers
yellow, pentamerous or rarely with higher merism.
Stigma capitate, shallowly bilobed or rarely up to
five-lobed. Inferior ovary with mostly one to three,
rarely up to five, uni-ovulate locules. Drupes containing elongated, spindle-shaped seeds. n = 9. Six
species from eastern Australia, Lord Howe Island,
New Zealand, and Rapa Island.
Selected Bibliography
Al-Shammary, K.I., Gornall, R.J. 1994. Trichome anatomy
of the Saxifragaceae s.l. from the southern hemisphere.
Bot. J. Linn. Soc. 114: 99–131.
Argophyllaceae
Backlund, A., Bremer, B. 1997. Phylogeny of the Asteridae s.
str. based on rbcL sequences, with particular reference
to the Dipsacales. Pl. Syst. Evol. 207: 225–254.
Briggs, L.H., Cambie, R.C., Couch, R.A.F. 1967. Triterpenes
from some New Zealand dicotyledons. N. Z. J. Sci. 10:
1076–1082.
Brown, F.B.H. 1928. Cornaceae and allies in the Marqesas
and neighboring islands. Bernice P. Bishop Mus. Bull.
52: 1–22.
Cambie, R.C., Parnell, J.C. 1969. A New Zealand phytochemical survey. Part 7. Constituents of some dicotyledons.
N. Z. J. Sci. 12: 453–466.
Cambie, R.C., Cain, B.F., Laroche, S. 1961. A New Zealand
phytochemical survey. Part 2. The dicotyledons. N. Z.
J. Sci. Technol. 4: 604–663.
Cronquist, A. 1981. An integrated system of classification
of flowering plants. New York: Columbia University
Press.
Eichler, A.W. 1878. Blütendiagramme, II. Leipzig: W. Engelmann.
Engler, A. 1890. Saxifragaceae. In: Engler, A., Prantl, K.,
Die natürlichen Pflanzenfamilien, III, 2a. Leipzig: W.
Engelmann, pp. 41–93.
Engler, A. 1928. Saxifragaceae. In: Engler, A., Prantl, K., Die
natürlichen Pflanzenfamilien, ed. 2, 18a. Leipzig: W.
Engelmann, pp. 74–226.
Eyde, R.H. 1966. Systematic evolution of the flower and fruit
of Corokia. Amer. J. Bot. 53: 833–847.
Ferguson, I.K., Hideux, M.J. 1978. Some aspects of the pollen
morphology and its taxonomic significance in Cornaceae sens. lat. In: Proceedings IV International Palynological Conference, Lucknow, 1976–1977, vol. 1, pp.
240–249.
Forster, P.I. 1990. Argophyllum verae (Saxifragaceae), a new
species from northern Queensland. Austrobaileya 3:
173–176.
Gardner, R.O. 1976. Studies in the Alseuosmiaceae. Ph.D.
Thesis, University of Auckland, Auckland, New
Zealand.
Gibbs, D.R. 1974. Chemotaxonomy of flowering plants, vol.
III. Montreal: McGill-Queen’s University Press.
Guillaumin, A., Virot, R. 1953. Contributions à la Flore de
la Nouvelle Calédonie, CII. Plantes récoltées par M.R.
Virot. Mém. Mus. Natl Hist. Nat. B, Bot. 4: 1–82.
Gustafsson, M.H.G., Bremer, K. 1997. The circumscription
and systematic position of Carpodetaceae. Austral.
Syst. Bot. 10: 855–862.
Gustafsson, M.H.G., Backlund, A., Bremer, B. 1996. Phylogeny of the Asterales sensu lato based on rbcL sequences with particular reference to the Goodeniaceae.
Pl. Syst. Evol. 199: 217–242.
Hair, J.B., Beuzenberg, E.J. 1959. Contributions to a chromosome atlas of the New Zealand Flora, 2. N. Z. J. Sci.
2: 148–156.
Hallier, H. 1908. Über Juliana, eine TerebinthaceenGattung mit Cupula, und die wahren Stammeltern der
Kätzchenblütler. Beih. Bot. Centralbl. 13: 81–265.
Hamel, J.L. 1953. Contribution à l’étude cyto-taxonomique
des Saxifragacées. Rev. Cytol. Biol. Vég. 14: 9–313.
Harms, H. 1897. Cornaceae. In: Engler, A., Prantl, K., Die
natürlichen Pflanzenfamilien, III, 8. Leipzig: W. Engelmann, pp. 250–270.
Hegnauer, R. 1969. Chemical evidence for the classification
of some plant taxa. In: Harborne, J.B., Swain, T. (eds.)
17
Perspectives in phytochemistry. Proceedings Phytochemical Society Symposium, Cambridge, April 1968.
London: Academic Press, pp. 121–138.
Hideux, M.J., Ferguson, I.K. 1976. The stereo-structure of
the exine and its evolutionary significance in Saxifragaceae s.l. In: Ferguson, I.K., Muller, J. (eds) The evolutionary significance of the exine. London: Academic
Press, pp. 327–377.
Hils, M.H. 1985. Comparative anatomy and systematics of
twelve woody Australian genera of the Saxifragaceae.
Ph.D. Thesis, The University of Florida.
Hooker, J.D. 1867. Cornaceae. In: Bentham, G., Hooker, J.D.,
Genera Plantarum, vol. I, part 3. London: Lovell Reeve,
pp. 947–952.
Hutchinson, J. 1967. The genera of flowering plants. Dicotyledons, vol. 2. Oxford: Clarendon Press.
Jaffré, T., Brooks, R.R., Trow, J.M. 1979. Hyperaccumulation
of nickel by Geissois species. Pl. Soil 51: 157–162.
Jensen, S.R., Nielsen, B.J., Dahlgren, R. 1975. Iridoid compounds, their occurrence and systematic importance
in the Angiosperms. Bot. Notiser 128: 148–180.
Källersjö, M., Farris, J.S., Chase, M.W., Bremer, B., Fay, M.F.,
Humphries, C.J., Petersen, G., Seberg, O., Bremer, K.
1998. Simultaneous parsimony jackknife analysis of
2538 rbcL DNA sequences reveals support for major
clades of green plants, land plants, seed plants and
flowering plants. Pl. Syst. Evol. 213: 259–287.
Kapil, R.N., Bhatnagar, A.K. 1992. Embryology and systematic position of Corokia Cunn. In: Proceedings XI International Symposium on Embryology and Seed Reproduction, Leningrad, 1990. St. Petersburg: Nauka.
Kårehed, J., Lundberg, J., Bremer, B., Bremer, K. 1999. Evolution of the Australasian families Alseuosmiaceae, Argophyllaceae, and Phellinaceae. Syst. Bot. 24: 660–682.
Labillardière, J.J.H. de 1824. Sertum Austro-Caledonicum.
Paris: Huzard.
Lobova, T.A. 1997. Seed morphology and anatomy in the
genera Argophyllum and Corokia (Argophyllaceae).
Bot. Zhurn. (Moscow & Leningrad) 82: 68–78.
Mauritzon, J. 1933. Studien über die Embryologie der Familien Crassulaceae und Saxifragaceae. Lund: Gleerupska
Univ.-Bokhandeln.
Melchior, H. 1964. A. Engler’s Syllabus der Pflanzenfamilien, Angiospermen, ed. 12, vol. 2. Berlin: Gebrüder
Bornträger.
Noshiro, S., Baas, P. 1998. Systematic wood anatomy of Cornaceae and allies. IAWA J. 19: 43–97.
Patel, R.N. 1973a. Wood anatomy of the dicotyledons indigenous to New Zealand. 1. Cornaceae. N. Z. J. Bot.
11: 3–22.
Patel, R.N. 1973b. Wood anatomy of the dicotyledons indigenous to New Zealand. 2. Escalloniaceae. N. Z. J.
Bot. 11: 421–434.
Schlechter, R. 1906. Beiträge zur Kenntnis der Flora von
Neu-Kaledonien. Bot. Jahrb. Syst. 39: 1–274.
Smith, L.S. 1958. Corokia A. Cunn. An addition to the Australian genera of Saxifragaceae. Proc. Roy. Soc. Queensland 69: 53–55.
Takhtajan, A. 1983. The systematic arrangement of dicotyledonous families. In: Metcalfe, C.R., Chalk, L. (eds.)
Anatomy of the dicotyledons, 2nd edn, vol. 2. Oxford:
Clarendon Press, pp. 180–201.
Takhtajan, A. 1987. Systema Magnoliophytorum. Leningrad: Nauka.
18
J. Kårehed
Takhtajan, A. 1997. Diversity and classification of flowering
plants. New York: Columbia University Press.
Thorne, R.T. 1983. Proposed new realignments in
angiosperms. Nordic J. Bot. 3: 85–117.
Thorne, R.T. 1992. Classification and geography of the flowering plants. Bot. Rev. 58: 225–348.
Tobe, H., Raven, P.H. 1999. Floral structures of Alseuosmiaceae, Argophyllaceae, Carpodetaceae, Phellinaceae
and Rousseaceae: additional members in Asterales.
In: Abstract Volume XVI International Botanical
Congress, St. Louis, MO, 1999.
Wangerin, W. 1909. Cornaceae. In: Engler, A., Das Pflanzenreich, IV, 229. Leipzig: W. Engelmann, pp. 1–110.
Wanscher, J.H. 1933. Studies on the chromosome numbers
of the Umbelliferae, III. Bot. Tidsskr. 42: 384–399.
Webb, C.J. 1994. Pollination, self-incompatibility, and fruit
production in Corokia cotoneaster (Escalloniaceae). N.
Z. J. Bot. 32: 385–392.
Wieffering, J.H. 1966. Aucubinartige Glucoside (Pseudoindikane) und verwandte Heteroside als systematische Merkmale. Phytochemistry 5: 1053–1064.
Willis, J.C. 1966. A dictionary of the flowering plants and
ferns, 7th edn. Revised by Airy Shaw. H.K. Cambridge:
University Press.
Zemann, M. 1907. Studien zu einer Monographie der Gattung Argophyllum Forst. Ann. K.K. Naturhist. Hofmus.
22: 270–292.
Calyceraceae
Calyceraceae R. Br. ex Rich., Mém. Mus. Hist. Nat. 6: 74 (1820), nom. cons.
F.H. Hellwig
Perennial or rarely annual herbs, occasionally
subligneous, often stemless or with scapiform
flowering shoots. Leaves alternate, often forming
a basal rosette, somewhat fleshy, without stipules, sessile or narrowed into a distinct petiole,
undivided to dissected, margin entire, sinuate,
dentate or serrate, sometimes revolute. Flowers in
capitula, surrounded by triangular to lanceolate
involucral bracts which are more or less united.
Receptacle flat to conical, often with herbaceous
to membranous, subulate to obovate or lanceolate
receptacular bracts, usually free but rarely fused.
Flowers perfect or rarely functionally male (and
then plants andromonoecious), epigynous, actinomorphic, calyx adnate to the ovary, with (4)5(6)
small, hyaline to membranous lobes or teeth,
these rounded-orbicular or flat, linear-lanceolate
and tapering. Corolla sympetalous, pentamerous
or sometimes tetramerous, only occasionally
hexamerous in individual flowers, cylindrical
to narrowly infundibuliform with (4)5 short
corolla lobes or with a slender basal tube and
a widened limb, uppermost part campanulate
with (4)5 valvate lobes, white or greenish. Stamens
enclosed in corolla or subexserted, (4)5(6), as
many as and alternating with the corolla lobes,
inserted at various levels in the corolla. Filaments
united into a tube over most of their length,
sometimes distally free; anthers connate, at least
at their bases, tetrasporangiate, dithecal, introrse,
opening by longitudinal slits, mostly without
appendages, sometimes with subsagittate bases
and inconspicuous apical appendages. Nectary
glands at base of filament tube alternating with
the vascular bundles of the filaments. Gynoecium
of two carpels, unilocular, inferior. Ovule solitary,
pendulous, anatropous, tenuinucellate. Style
strongly exserted, slender, cylindrical, stigma
capitate, dry, papillose, with two (three) vascular
bundles. Fruit an achene, often crowned by the
calyx lobes which sometimes become lignified
and spiny, often calyx also accrescent; achene
cylindrical or prismatic, more or less distinctly
ribbed. Seed with well-developed endosperm,
embryo straight.
The family comprises four genera and about 60
species and is endemic to southern South America.
One species occurs in the Falkland Islands (Islas
Malvinas).
Vegetative Morphology. Besides inconspicuous annual herbs, the family contains many species
which form vegetative rosettes until the inflorescences are formed. Growth is then continued by
stolons, rhizomes or by lateral shoots arising from
lower parts of the plant. Branching of Calyceraceae
is sympodial, and in some genera monochasia are
formed (in Acicarpha the capitula are terminal and
the shoot system is continued by secondary axes
overtopping the preceding ones). Metatopies with
re- and concaulescence can be observed in some
species (e.g. Calycera sessiliflora, C. eryngioides).
Vegetative Anatomy. Plants usually are glabrous but sometimes uniseriate filiform hairs are
present, especially on the stems (Calycera). No
external glands or secretory cavities have been observed (Hansen 1992), nor are resin-like droplets
present in the wood (Carlquist and DeVore 1998).
However, droplets of a resin-like substance seem
to be present on and in stems and involucral
bracts of Boopis gracilis (pers. obs.). Stem vascular
bundles are arranged in a circle and are separated
from each other by broad rays. The bundles are
embedded in a ring of mechanical tissue in some
species (Boopis spp.). Phloem strands of Acicarpha
are accompanied externally by sclerenchyma,
xylem includes moderately wide vessels with
simple perforation plates, with bordered pits
between vessels and ray parenchyma. Secondary
medullary rays are wanting (Cronquist 1981). Two
types of imperforate tracheary elements occur in
the family, i.e. vasicentric tracheids and libriform
fibres, the latter with small simple pits. Pits
20
F.H. Hellwig
between vessels and libriform fibres are alternate
and circular to oval (Carlquist and DeVore 1998).
Laterally elongated pits, called pseudoscalariform
pits, are present in several species of Calyceraceae
(Carlquist and DeVore 1998). The amount of wood
is always very limited, with a maximum at stem
bases and adjacent parts of the roots (Carlquist and
DeVore 1998). The wood has strong indications of
paedomorphosis with varying degrees in different
species. This is supported by the presence of
pseudoscalariform pitting and predominantly
upright ray cells (Carlquist and DeVore 1998).
Secondary xylem vessels with pseudoscalariform
pitting transitional to helical-banded patterns
support the ability of the roots to be bent or
to expand and contract in reaction to changing
water content (Boopis graminea, Nastanthus spp.).
Seasonality is mirrored in the formation of growth
rings observed in Boopis anthemoides (Carlquist
and DeVore 1998).
The family exhibits an unusually wide range
of mesomorphy values. This may be explicable
by the very different habitats the plants live in.
Wood anatomy is not yet sufficiently known to
be used systematically. It is believed to largely
reflect autapomorphic ecological adaptations
(Carlquist and DeVore 1998). The primary cortex
and the central pith are rich in clustered Ca-oxalate
crystals. There are no specialized bast strands
in the cortical parenchyma (Reiche 1901). The
fleshy stems found in various species of Boopis
result from the presence of massive medullary
and cortical parenchyma (Reiche 1901). Tuberous
roots of the fleshy species of Boopis have a strongly
transversely rugose cortex which is a feature
connected to the ability of many alpine plants to
translocate themselves into deeper strata of the
soil by root contraction (Reiche 1901).
Inflorescence and Flower Structure. The
most striking feature of many species is the condensed, often many-flowered inflorescence. This
condensation reaches several levels in the family. While Acicarpha has simple capitula with centripetal anthesis, the other genera have more complicated capitula of higher orders. It has been hypothesized that these consist of cymose units (Baillon 1880; Reiche 1901; DeVore 1994) but this has not
been confirmed by detailed analyses. Some species
of Calycera and Boopis tend to aggregate few to
many capitula.
Flowers usually are hermaphrodite but functionally male flowers occur regularly in the centre
of the capitula of Acicarpha (Miers 1870). The flowers are characterized by the transformed calyx and
a close association of corolla, androecium and the
style. The free part of the calyx is reduced to five
small lobes above the edges of the gynoecium, the
lower part being fused and closely attached to the
inferior ovary. It does not protect the flower bud
(Erbar 1993).
The filaments form a tube which consists of
their fused lower parts. In Acicarpha tribuloides,
this tube overtops the corolla tube in mature flowers. Below the separation of the filaments from the
corolla tube, there is a corolla stamen tube. Five
nectaries are present in the flower. The glandular
tissue extends from the base of the filament tube
to the top of the stamen corolla tube (Erbar 1993).
Nectar is secreted through nectar slits outside of the
filament tube into five small pockets. Form and alternate position with the filaments of the nectaries
are very distinctive for the family (Brown 1817; Reiche 1901). The style is cylindrical with a club-like
papillose head. Intercalary growth of the receptacle
results in a carpel stamen corolla tube between the
ovary and the other parts of the flower, except the
sepals. This feature seems to be unique among the
families of Asteridae (Erbar 1993). The unilocular
gynoecium usually is interpreted to consist of two
carpels (Cronquist 1981). Investigations by Erbar
(1993) and Hansen (1992), however, revealed the
existence of five ledges in the ovary which can be
interpreted as rudimentary septa. Following this
interpretation, the family has five carpels (Erbar
1993).
Floral Anatomy. The free corolla contains 10
parallel longitudinal vascular bundles. The commissural bundles are bifurcated below the corolla
lobes and anastomose with the median bundles of
the corolla lobes just below their apex (Gustafsson
1995; Gustafsson and Bremer 1995). Micromorphological characters of the corolla (petal epidermis
patterns) are rather uniform in the Calyceraceae,
the cells always being Senecioid. Cells are often long
(more than 200 µm) and confluent. A transversely
striate pattern prevails on the adaxial surface, while
the abaxial pattern is mostly longitudinally striate
although rugose, reticulate-rugose or glabrous cells
also occur. The cells are generally distinctly sculptured on both surfaces (Hansen 1992).
Embryology. An integumentary tapetum is
formed by the innermost layers of the one massive
integument (Dahlgren 1915). The mature embryo
Calyceraceae
sac has eight nuclei; the polar nuclei fuse before
fertilization. The seed, which is suspended from
the top of the locule and fixed by a short funiculus
near the upper end of the seed, contains a straight,
terete embryo and abundant endosperm (Cronquist 1981). Endosperm development is cellular,
and no haustoria are formed. The endosperm
contains protein grains, rather than starch. The
outer cells of the testa contain chloroplasts before
the seed is mature. The radicula points towards the
micropyle, i.e. the apex of the seed (Miers 1870).
The pollen grains are binucleate when released. No
periplasmodial tapetum is formed in the pollen
sacs (Dahlgren 1915).
Pollen Morphology. Pollen grains of Calyceraceae are tricolporate. The colpi are very long, rendering the grains of some species sub-syncolporate.
The colpus membrane is more or less granular. The
pores are lalongate, the ends overlapping or joined,
forming a colpus transversalis of irregular width in
some species (Heusser 1971; Markgraf and D’Antoni 1978). Minute spinules are usually present on
the pollen surface but rarely they are almost lacking. The exine has thickened ridges above the pores
and rounded depressions in the mesocolpi. A granulate colpus margin (Skvarla et al. 1977) is prominent in many species. The pollen grains rarely exceed 30 µm in length and 20 µm in width, and are
often rather irregularly shaped. The majority of
pollen grains are at first glance spheroidal but, seen
in equatorial view, they are irregularly rhomboidal
(Hansen 1992). Prolate (elliptic, ellipsoidal) grains
occur in a few species. Dimorphic grains are rare
(Boopis gracilis) but shape can vary within species.
Avetisjan (1980) distinguishes two pollen types in
the family, the Moschopsis-type and the Calyceratype. They differ by the shape of pollen grains (polar view rounded, three-lobed, elliptical in equatorial view, pores elliptical in the Moschopsis-type,
polar view irregularly hexagonal, equatorial view
transversely elliptical, pores broad-elliptical (lalongate) in the Calycera-type) and by the shape
of the colpi, especially by the ridges above the apertures, which are weakly to moderately thickened in
the Moschopsis-type and strongly developed in the
Calycera-type. The Moschopsis-type is regarded as
primitive, and the Calycera-type as derived by the
author. The pollen grains have TEM-patterns approaching the plesiomorphic Anthemoid pattern
of Compositae, i.e. with distinct infratectal bacula and an infratectum. The tectum is massive and
a cavus is lacking (Hansen 1992).
21
Karyology. Several chromosome numbers are
known in the family. Stebbins (1977) suggested
x = 7, 8 and 9 as base numbers. The only known
diploid species of the family is an Acicarpha
(DeVore 1994). All other species investigated are
interpreted as being polyploid. Accepting this,
species of Calycera and the species of Boopis based
on x = 7 are mostly hexaploid or hexaploidderived, or derived from tetraploids, and species of
Boopis with the base number x = 9 are tetraploid
or tetraploid-derived. In Calycera, n = 21 or 22,
17 and 13 are found in the Chilean species, while
Argentinean species are all polyploids with n = 21.
DeVore (1994) interprets the base number x = 8
as ancestral; x = 7 would then have originated by
descending aneuploidy, and the counts in Boopis
point to the other derived base number x = 9.
However, the interpretation of karyotype evolution
in Calyceraceae is inconclusive, due to lack of data
and a well-supported phylogenetic hypothesis.
Pollination. Calyceraceae show secondary
pollen presentation. The mechanism corresponds
to the pump mechanism (Leins and Erbar 1990)
as known from Acicarpha. This mechanism seems
to prevail in some basal lineages of Asteraceae
(Bremer 1994). Initially, the pollen grains are
released into the cavity of the anther tube which
is basally sealed by the tip of the style. The pollen
is then extruded apically by the growing style.
One major difference to the pump mechanism of
Asteraceae is that the anthers are not fused over
their entire length in many species, and are bent
outwards at least in Acicarpha tribuloides.
Fruit, Seed and Seed Dispersal. The
gynoecium consists basically of two carpels
forming a compound, inferior, unilocular, pseudomonomerous ovary (Cronquist 1981). The fruit
is an achene, crowned by the persistent lobes
of the calyx. In some species, all lobes or some
of them are enlarged and become indurated to
spine-like structures. In Calycera, capitula are
distinctly heterocarpic. In Calycera eryngioides,
the smaller achenes are dispersed individually
whereas those with elongated calyx lobes remain
attached to the receptacle and are dispersed with
the whole capitulum (DeVore 1994). Acicarpha
is characterized by partly fused achenes which
remain attached to the receptacle. The spine-like
elongated calyx lobes point to epizoochorous
dispersal. A spongy tissue in the pericarp may
facilitate hydrochory or even anemochory.
22
F.H. Hellwig
Phytochemistry. Calyceraceae commonly produce iridoid compounds, e.g. seco-loganin (Jensen
et al. 1975), but are not tanniferous. They lack proanthocyanins and presumably also ellagic acid.
No latex or secretory cavities are known (Cronquist 1981). The seeds store inulin (Cronquist
1981; Bohm et al. 1995). Acetylenes (Mabry and
Bohlmann 1977), sedoheptulose, aluminium accumulation, raphides, leucoanthocyanins, cyanogenic glycosides, saponins and l-inositol are absent
(Gibbs 1974). For flavonoids, refer to Bohm et al.
(1995).
Distribution and Habitats. The family is
endemic to southern South America. Its range
extends from the extreme south of the continent
to southern Peru and Bolivia in the west and as
far north as Bahia (Brazil) in the east. One species
occurs in the Falkland Islands (Islas Malvinas).
Most species grow in high-altitude arid habitats,
e.g. open grassland and meadows, in the Andes
of Chile and Argentina. A few species grow in
coastal sand dunes. Acicarpha tribuloides grew in
the south-western United States for some decades
in the 19th century after it was introduced by man,
but did not become naturalized there (DeVore
1991).
Subdivisions and Relationships in the Family. The generic classification of the family is unsatisfactory. Currently, six genera are widely accepted (Reiche 1901; Pontiroli 1963; Marticorena
and Quezada 1985; Chiapella 1999). Possibly apomorphic characters have been identified for Acicarpha, Calycera and Gamocarpha but not for Nastanthus, Boopis and Moschopsis (Hansen 1992), and
the diagnostic characters reported for these latter three genera proved to be unsuitable because
of erroneous observations and misinterpretations
of morphology. In the present treatment, Boopis,
Moschopsis and Nastanthus are united under the
oldest generic name, Boopis, in order to obtain
recognizable genera. The relationships among the
genera of the family are discussed in several papers
without satisfactory result. The only unanimously
accepted fact is that Acicarpha is distinct from the
other genera by centripetal anthesis within the capitula and the presence of functionally male flowers in the centre of the receptacle. It is further
believed (DeVore 1994) that Calycera and Nastanthus are closely related to each other (chromosome
numbers), while the relationships of Boopis, Gamocarpha and Moschopsis remain uncertain (Hansen
1992). No formal phylogeny has been published for
the family.
Phylogeny and Systematic Position of the
Family. Calycera herbacea, the first species of
Calyceraceae described (Cavanilles 1797), was
placed in Dipsacaceae by Ruiz and Pavón in 1798,
and the family was kept in Dipsacales by several
taxonomists throughout the last two centuries. On
the other hand, Acicarpha tribuloides, described
by Jussieu in 1803, was included, although with
doubts, in Asteraceae by the author. In 1816,
Brown and Cassini independently recognized
Calyceraceae as a separate family in oral presentations before the Linnaean Society and the French
Académie des Sciences respectively, and placed
it between Dipsacaceae and Asteraceae (see also
Cassini 1817). The nomenclaturally valid publication of Calyceraceae was provided by Richard
(1820). Miers (1870) underlined the close affinity
between Asteraceae and Calyceraceae. Cladistic
studies in Asteridae supported such relationship
(Gustafsson 1996). The family is very likely to be
the closest living relative of Compositae (Lundberg
and Bremer 2003). This position is indicated not
only by evidence from DNA sequence variation
but also by characters such as pollen morphology,
petal venation, morphology of the receptacle and
receptacular bracts, and wood anatomy (DeVore
and Stuessy 1995; Carlquist and DeVore 1998).
Similarities with Dipsacales include the pendulous
ovule, copious endosperm, morphology of capitula
in the majority of Calyceraceae, and phytochemical
features such as the absence of iridoid compounds
from Asteraceae. This may have caused Dahlgren
(1989) to place Calyceraceae in Dipsales. However,
iridoids are found also in Goodeniaceae, a family
closely related to Calyceraceae plus Asteraceae and
Menyanthaceae. Lack of iridoids in Asteraceae may
therefore be apomorphic for this family. Hansen
(1992) and Erbar (1993) provide a summary of
differences and similarities of Calyceraceae with
various possibly related groups (see also Moore
1993). The molecular phylogenetic studies of
Gustafsson and Bremer (1995) and Albach et al.
(2001) showed Calyceraceae as sister to Asteraceae,
while Calyceraceae are sister to Goodeniaceae in
the combined 18S/rbcL analysis of Bremer et al.
(2001), as they were in previous rbcL analyses
(Cosner et al. 1994; Gustafsson et al. 1996). This
latter position is supported by chemical features:
both contain bis-secoiridoids of the sylvestroside
type (Jensen et al. 1979; Capasso et al. 1996). The
Calyceraceae
combined analysis of morphology, rbcL and ndhF
sequences confirmed the sister-group relationship
between Calyceraceae and Asteraceae, while Goodeniaceae were placed as sister to these two (Bremer
et al. 2001). Takhtajan (1997) established the order
Calycerales, which is not recognized in systems
which reflect the results of cladistic studies.
Economic Importance. Some species are weeds
in South America (Boopis gracilis, Boopis anthemoides, Acicarpha tribuloides). Medicinal use of
Acicarpha tribuloides has been reported (Capasso
et al. 1996).
Key to the Genera
1. Capitula with dimorphic flowers, upper/central
flowers functionally male, lower/outer flowers
hermaphrodite; all achenes with spiny calyx lobes,
outer achenes with united bases and also fused to the
receptacle
1. Acicarpha
– Capitula with isomorphic flowers, all flowers
hermaphrodite and fertile; achenes not united
2
2. Achenes dimorphic, some with spiny calyx lobes, some
without
2. Calycera
– Achenes monomorphic, without spiny calyx lobes 3
3. Receptacular bracts fused into groups, these and
involucral bracts around groups of flowers
3. Gamocarpha
– Receptacular bracts free or absent
4. Boopis
Genera of Calyceraceae
1. Acicarpha Juss.
Acicarpha Juss., Ann. Mus. Paris 2: 347 (1803).
Annual or perennial herbs; stems more or less
branched, erect or procumbent, terminating in
an inflorescence overtopped by lateral branches.
Leaves petiolate or sessile, subamplexicaulous,
undivided to pinnatifid, margin entire to dentate,
leaf bases attenuate to auriculate. Capitula solitary,
sessile or pedunculate; involucral bracts about five
in one series, linear-oblong, unequal, similar to the
uppermost cauline leaves, fused in their lower part;
receptacle convex to conical; receptacular bracts
free, linear to lanceolate. Flowers pentamerous,
dimorphic, the outer (or lower) hermaphrodite,
the upper (or central) functionally male; corolla
with a long slender tube and an infundibuliform to
campanulate limb deeply divided into five corolla
lobes; stamens 5, enclosed, fused to the lower 2/3 of
the corolla; anthers connate from base to middle,
free in distal part and bent outwards at anthesis;
achenes cylindrical to obconical, pentasulcate
23
to prismatic, crowned by variously elongated
spiny calyx lobes. Marginal achenes fused to the
receptacle and united with each other. 2n = 16.
Three species in Brazil, Peru, Bolivia, Paraguay,
Uruguay and Argentina.
2. Calycera Cav.
Calycera Cav., Icon. Pl. 4: 14 (1797).
Annual or perennial herbs, glabrous or weekly
lanuginose; stems erect or decumbent. Leaves
alternate or in rosettes, undivided to pinnatifid, spathulate to elliptical; margin entire to
dentate-mucronate. Involucral bracts 4–7 in one
series, sometimes with additional bracts, broadly
triangular to linear-lanceolate, united to various
degrees, entire or dentate; receptacle convex or
subglobose; receptacular bracts lanceolate to linear
or absent. Flowers hermaphrodite, tetramerous or
pentamerous; corolla with a long tubular part and
an infundibuliform to campanulate limb, or limb
more or less truncate at base, abruptly narrowed
into a short slender tube, upper part of limb
divided to various degrees into four or five lobes;
stamens 4 or 5, not exserted, inserted at the base
or higher up in the corolla tube; anthers connate,
bases rounded or sagittate, connectives sometimes
with apical appendages; achenes cylindrical to
obconical or prismatic, dimorphic, some crowned
by short or long spiny calyx lobes, some with only
very short calyx lobes, not spiny. 2n = 26, 24,
42, 44. About 20 species, Andes of Bolivia, Chile
and Argentina. Two subgenera, subg. Calycera
and subg. Leucocera (Turcz.) Reiche, have been
distinguished (Miers 1870; DeVore 1994).
3. Gamocarpha DC.
Gamocarpha DC., Prodr. 5: 2 (1836).
Perennial herbs, glabrous, some species stoloniferous. Leaves generally forming a rosette, entire
or divided. Capitula surrounded by 6–12 involucral bracts, these and receptacular bracts more or
less fused, forming chambers which contain several
flowers. Bracts with triangular distal parts, fused in
the lower part, entire or weakly lobed. Flowers pentamerous (occasionally tetramerous); corolla tube
long, slender, with an infundibuliform to campanulate limb, corolla lobes long, or corolla cylindrical
to infundibuliform, base sometimes abruptly narrowed into a short filiform tube, corolla lobes short;
anthers subsagittate at base, connate in the lower
part; achenes cylindrical to prismatic, ribs weak,
24
F.H. Hellwig
crowned by ovate to triangular non-spinescent calyx lobes. Six or seven species in Chile and Argentina.
4. Boopis Juss.
Fig. 4
Boopis Juss., Ann. Mus. Natl. Hist. Nat. 2: 350 (1803).
Perennial or rarely annual herbs, glabrous; stems
erect or decumbent, scapose or ramified. Leaves
alternate or clustered in a rosette, entire to pinnatisect, linear-lanceolate, laciniate, dentate or pectinate, or spathulate with dentate or crenate lamina.
Capitula terminal, solitary or in groups of few,
then shortly pedunculate to almost sessile, sometimes densely surrounded by foliar leaves, rarely
scapose. Terminal capitula sometimes overtopped
by axillary shoots, in some species terminal and
lateral scapes forming a disk-like to hemispherical
syncephalium; involucral bracts 5–10, united from
basis to middle or beyond, exceptionally free, triangular to lanceolate, sometimes laciniate; receptacle
flat to convex or conical, inflated in some species;
receptacular bracts free, linear to lanceolate, or
absent. Flowers pentamerous or tetramerous,
hermaphrodite, uniform within species; corolla
with short or moderately long tube and cylindrical
to infundibuliform or slightly campanulate limb,
or with slender tube and limb consisting of lower
infundibuliform part and distal, more or less
campanulate part; corolla lobes mostly short, long
in flowers with long tube; stamens five or four,
anthers connate at their base, free in the upper half;
achenes prismatic, 5- or 4-ribbed, ribs crowned
by ovate or broadly lanceolate, acute, more or less
rigid but not spiny calyx lobes. 2n = 36, 40, 42.
About 30 species, some of them polymorphic, in
Chile, Argentina incl. Islas Malvinas (Falkland
Islands), very few in Brazil, Peru, Bolivia.
Selected Bibliography
Fig. 4. Calyceraceae. Boopis bupleuroides. A Habit. B Inflorescence. C Flower bud, longitudinal section. D Open flower.
E Fruit. F Fruit, longitudinal section. (Martius 1878)
Albach, D.C., Soltis, P.S., Soltis, D.E., Olmstead, R.G.
2001. Phylogenetic analysis of Asterids based on
sequences of four genes. Ann. Missouri Bot. Gard. 88:
163–212.
Avetisjan, E.M. 1980. Pollen morphology of the family Calyceraceae (in Russian). In: Sistematika i Évolyutsiya
Vysshikh Rastenii. Leningrad: Nauka, pp. 57–64.
Baillon, H. 1880. Histoire des plantes, VII. Paris: Hachette.
Bohm, B.A., Reid, A., DeVore, M., Stuessy, T.F. 1995.
Flavonoid chemistry of Calyceraceae. Canad. J. Bot.
73: 1962–1965.
Bremer, K. 1994. Asteraceae. Cladistics and classification.
Portland, OR: Timber Press.
Bremer, K., Backlund, A., Sennblad, B., Swenson, U.,
Andreasen, K., Hjertson, M., Lundberg, J., Backlund, M., Bremer, B. 2001. A phylogenetic analysis
of 100+ genera and 50+ families of euasterids based
on morphological and molecular data with notes on
possible higher level morphological synapomorphies.
Pl. Syst. Evol. 229: 137–169.
Brown, R. 1817. Observations on the natural family of plants
called Compositae. Trans. Linn. Soc. 12: 76–142.
Capasso, A., Urruñaga, R., Garofala, L. et al. 1996. Phytochemical and pharmacological studies on the medical
herb Acicarpha tribuloides. Intl J. Pharmacog. 34. 255–
261.
Carlquist, S., DeVore, M.L. 1998. Wood anatomy of Calyceraceae with reference to ecology, habit, and systematic
relationships. Aliso 17: 63–76.
Cassini, H. 1817. Boopidées (Bot.). Dict. Sci. Nat. suppl. 5:
26–28.
Calyceraceae
Cavanilles, A.J. 1797. Icones et descriptiones plantarum, IV.
Matriti: Regia Typographia.
Chiapella, J. 1999. Calyceraceae. In: Correa, M.N. (ed.) Flora
Patagonica, VI. Buenos Aires: Colección científica del
INTA, pp. 492–517.
Cosner, M.E., Jansen, R.K., Lammers, T.G. 1994. Phylogenetic relationships in the Campanulales based on rbcL
sequences. Pl. Syst. Evol. 190: 79–95.
Cronquist, A. 1981. An integrated system of classification
of flowering plants. New York: Columbia University
Press.
Dahlgren, O. 1915. Über die Embryologie von Acicarpha
tribuloides Juss. Svensk Bot. Tidskr. 9: 184–191.
Dahlgren, G. 1989. The last Dahlgrenogram. System of classification of dicotyledons. In: Tan, K. (ed.) The Davis
and Hedge Festschrift. Edinburgh: University Press,
pp. 249–260.
DeVore, M. 1991. The occurrence of Acicarpha tribuloides
(Calyceraceae) in eastern North America. Rhodora 93:
26–35.
DeVore, M. 1994. Systematic studies of Calyceraceae. Ph.D.
Thesis, Ohio State University, Columbus, OH.
DeVore, M., Stuessy, T.F. 1995. The place and time of origin of the Asteraceae, with additional comments on
the Calyceraceae and Goodeniaceae. In: Hind, D.J.N.,
Jeffrey, C., Pope, G.V. (eds) Advances in Compositae systematics. Royal Botanic Gardens, Kew, pp. 23–
40.
Erbar, C. 1993. Studies on the floral development and pollen
presentation in Acicarpha tribuloides with a discussion
of the systematic position of the family Calyceraceae.
Bot. Jahrb. Syst. 115: 325–350.
Gibbs, R.D. 1974. Chemotaxonomy of flowering plants, vol.
2. Montreal: McGill-Queen’s University Press.
Gustafsson, M.H.G. 1995. Petal venation in the Asterales
and related orders. Bot. J. Linn. Soc. 118: 1–18.
Gustafsson, M.H.G. 1996. Phylogenetic hypotheses for
Asteraceae relationships. In: Hind, D.J.N., Beentje, H.J.
(eds) Compositae: systematics. Proceedings of the
International Compositae Conference, Kew, 1994,
vol. 1. Royal Botanic Gardens, Kew, pp. 9–19.
Gustafsson, M.H.G., Bremer, K. 1995. Morphology and phylogenetic interrelationships of the Asteraceae, Calyceraceae, Campanulaceae, Goodeniaceae and related families (Asterales). Amer. J. Bot. 82: 250–265.
Gustafsson, M.H.G., Backlund, A., Bremer, B. 1996. Phylogeny of the Asterales sesu lato based on rbcL sequences with particular reference to the Goodeniaceae.
Pl. Syst. Evol. 199: 217–242.
Hansen, H.V. 1992. Studies in the Calyceraceae with a discussion of its relationship to Compositae. Nordic J. Bot.
12: 63–75.
25
Heusser, C.J. 1971. Pollen and spores of Chile. Tucson, AR:
University of Arizona Press.
Jensen, S.R., Nielsen, B.J., Dahlgren, R. 1975. Iridoid compounds, their occurrence and systematic importance
in angiosperms. Bot. Notiser 128: 148–180.
Jensen, S.R., Lyse-Pedersen, S.E., Nielsen, B.J. 1979. Novel
bis-iridoid glucosides from Dipsacus sylvestris. Phytochemistry 18: 273–277.
Jussieu, A.L. 1803. Mémoire sur l’Acicarpha et le Boopis,
deux genres nouveaux de plantes de la famille des
Cinarocéphales. Ann. Mus. Natl Hist. Nat. 2: 345–350.
Leins, P., Erbar, C. 1990. On the mechanisms of secondary
pollen presentation in the Campanulales-Asteralescomplex. Bot. Acta 103: 87–92.
Lundberg, J., Bremer, K. 2003. A phylogenetic study of the
order Asterales using one morphological and three
molecular data sets. Intl J. Pl. Sci. 164: 553–578.
Mabry, T.J., Bohlmann, F. 1977. Summary of the chemistry
of the Compositae. In: Heywood, V.H., Harborne, J.B.,
Turner, B.L. (eds) The biology and chemistry of the
Compositae, I. London: Academic Press, pp. 1097–
1104.
Markgraf, V., D’Antoni, H.L. 1978. Pollen flora of Argentina.
Tucson, AR: University of Arizona Press.
Marticorena, C., Quezada, M. 1985. Catálogo de la flora
vascular de Chile. Gayana (Bot.) 42: 5–157.
Martius, C.F.P. 1878. Flora Brasiliensis, vol. 6. London: Lovell
Reeve.
Miers, J. 1870. Contributions to Botany, II. London: Williams
and Norgate.
Moore, D.M. 1993. Calyceraceae. In: Heywood, V.H. (ed.)
Flowering plants of the world. London: B.T. Bastford.
Pontiroli, A. 1963. Flora Argentina, Calyceraceae. Revista
Mus. La Plata 9, Bot. 41: 175–214.
Reiche, K. 1901. Beiträge zur Systematik der Calyceraceen.
Bot. Jahrb. Syst. 29: 107–119.
Richard, L.C. 1820. Mémoire sur une famille de plantes,
dites les Calycérées. Mém. Hist. Nat. 6: 28–82.
Ruiz, H., Pavón, J. 1798. Systema vegetabilium florae peruvianae et chiliensis. Madrid: Gabrielis de Sancha.
Skvarla, J.J., Turner, B.L., Patel, V.C., Tomb, A.S. 1977.
Pollen morphology in the Compositae and in morphologically related families. In: Heywood, V.H.,
Harborne, J.B., Turner, B.L. (eds) The biology and
chemistry of the Compositae, I. London: Academic
Press, pp. 141–248.
Stebbins, L. 1977. Development and comparative anatomy
of the Compositae. In: Heywood, V.H., Harborne, J.B.,
Turner, B.L. (eds) The biology and chemistry of the
Compositae, I. London: Academic Press, pp. 91–109.
Takhtajan, A. 1997. Diversity and classification of flowering
plants. New York: Columbia University Press.
Campanulaceae
Campanulaceae Jussieu, Gen. Pl. 163 (1789), nom. cons.
T.G. Lammers
Herbaceous perennials, less often annuals or
biennials, herbaceous or woody lianas (sometimes
twining), pachycaul rosette plants, subshrubs,
shrubs, treelets, or trees to 15 m tall, typically
terrestrial, rarely aquatic or epiphytic, with milky
(sometimes coloured) latex; stems simple to much
branched, sometimes rhizomatous or acaulescent. Leaves estipulate, alternate, rarely opposite
or whorled, commonly simple (pinnate when
compound), entire, variously toothed, or sometimes dissected, petiolate or sessile, sometimes
sheathing. Inflorescences monotelic or polytelic,
commonly appearing racemose, paniculate, spicate, umbellate or capitate, the last sometimes
involucrate, typically terminal or sometimes
axillary, or the flowers solitary in an axillary or
rarely terminal position; bracts foliose or reduced,
rarely absent; pedicels often bracteolate. Flowers
tetracyclic, commonly perfect, rarely imperfect
and the plants dioecious or gynodioecious, with
a specialized method of proterandrous secondary
pollen presentation, rarely with extrafloral nectaries, sometimes resupinate. Calyx synsepalous,
adnate to the ovary, forming a hypanthium, rarely
free; lobes (3–)5(–10), valvate, sometimes with
a reflexed appendage in each sinus. Corolla sympetalous, radially or bilaterally symmetric, most
often some shade of blue or violet; lobes (4–)5(–10),
valvate or rarely induplicate. Stamens equalling the
number of corolla lobes, antesepalous, inserted at
base of corolla tube, on rim of hypanthium or atop
the inferior ovary, rarely epipetalous; filaments
distinct or connate; anthers tetrasporangiate,
dithecal, introrsely dehiscent by longitudinal slits,
basifixed, rarely dorsifixed, distinct, connivent, or
connate. Gynoecium syncarpous, 2–5(–10)-locular
with axile placentation, rarely 1-locular with parietal, basal, or apical placentation; ovary inferior,
rarely superior or nearly so, often crowned by an
annular nectary; style solitary, pubescent below
apex; stigma typically with as many lobes as
ovary locules. Fruit a capsule, commonly apically
loculicidal or laterally poricidal, or a berry. Seeds
usually small, numerous; embryo small, straight;
endosperm copious, cellular, oily or rarely starchy.
As circumscribed here, Campanulaceae comprise 84 genera and almost 2,400 species. The family
is cosmopolitan, with representatives on six continents and several of the world’s archipelagos.
Vegetative Morphology. Most Campanulaceae are polycarpic perennial herbs; in seasonally
cold or dry climates, they overwinter as hemicryptophytes by means of basal rosettes or basal
portions of the axis, or as geophytes by means
of the stem base, rhizomes or tuberous roots.
Among taxa which are monocarpic, an annual
habit is commonest; relatively few are biennial (e.g.
Campanula medium) or long-lived (e.g. Lobelia
wollastonii).
Though Campanulaceae are primarily an
herbaceous family, a significant number of taxa
are woody to some degree (e.g. Azorina, Centropogon, Cyanea, Heterochaenia). Woody species are
typically evergreen, branched or unbranched, and
range in stature from subshrubs a decimetre or so
tall (e.g. Merciera), to treelets and shrubs a meter
or two tall (e.g. Delissea rhytidosperma), up to
trees of 15 m (e.g. Cyanea leptostegia). Of special
interest are the so-called giant rosette plants
found in certain insular and montane habitats.
These are pachycaul plants, often long-lived and
pliestesial, of large stature and bulk, with dense
apical rosettes of typically sessile leaves, large
racemose inflorescences, and a hollow pith which
narrows basipetally (e.g. Lobelia rhynchopetalum).
A number of species are lianas, either herbaceous or woody. Many simply sprawl over surrounding vegetation. Those which climb typically
do so with twining stems (e.g. Cyphia lasiandra,
Siphocampylus convolvulaceus). In Canarina, however, support is provided by the twining petioles
and pedicels, while Cyanea copelandii climbs by
adventitious roots produced at the nodes.
Campanulaceae
Roots typically are coarse, fibrous, and of
adventitious origin, though in some genera (e.g.
Canarina, Codonopsis, Cyphia, Platycodon) the
primary root becomes fleshy, greatly enlarged and
tuberous. Stem branching may be monopodial
or sympodial. Mature herbs may have elongate
multinodal shoots, or form basal rosettes; in
seedlings of the former, the epicotyl is well
developed whereas in the latter, it is distinctly
shortened or lacking (Shulkina 1980). Many of the
woody taxa form dense apical rosettes of leaves.
Leaves are generally dorsiventral (rarely centric) and conform to a dillenid pattern (Hickey
and Wolfe 1975), with pinnate (craspedodromous
or camptodromous) venation (rarely parallellodromous, e.g. Siphocampylus smilax). They lack stipules and may be petiolate or sessile. Phyllotaxy typically is alternate, with a 2/5 sequence, though in
a few taxa the leaves are opposite (e.g. Cyclocodon)
or whorled (e.g. Ostrowskia, Siphocampylus orbignianus).
Vegetative Anatomy and Ultrastructure.
A major anatomical characteristic of the family is
the anastomosing network of articulated lactifers
associated with the phloem of leaves, stems,
and floral organs. The viscous latex produced
typically is copious and white but in some species
it is coloured, e.g. tan (Lobelia excelsa), bright
orange (L. cordifolia) or canary yellow (Cyanea
leptostegia).
In the stem, xylem and phloem typically form
a continuous cylinder, rather than discrete bundles;
medullary bundles are commonly present, cortical
bundles rarely. The xylem lacks tracheids and fibretracheids but libriform fibres are common. Vessels
typically have simple perforation plates (less often
scalariform) and simple pits; helical thickenings
are absent (Carlquist 1969, 1992). Internal phloem
is lacking, as are phloem transfer cells. Plastids
found in the sieve-elements lack proteinaceous inclusions, i.e. they are S-type. Axial parenchyma is
scanty vasicentric, and sclerenchyma is generally
lacking from the pericycle. Nodes are unilacunar.
Stomates are anomocytic and usually found on
both surfaces of the leaf, though sometimes they
are solely abaxial. Hydathodes are common, often associated with marginal teeth of leaves. Xerophytic species often have a fibrous hypoderm.
The mesophyll typically comprises one or two palisade layers. Mesophyll cells of certain Campanuloideae (Campanula, Edraianthus, Jasione, Phyteuma, Trachelium) contain protein intranuclear
27
inclusions of a unique fibrillar structure (Bigazzi
1986), though these are absent from other genera of
Campanuloideae (Asyneuma, Canarina, Legousia,
Petromarula, Platycodon, Wahlenbergia) as well as
from Lobelioideae (Downingia, Isotoma, Lobelia,
Solenopsis). Cystoliths occur occasionally.
Trichomes typically are unicellular or sometimes uniseriate. Arbusculiform (dendritic) trichomes (Batterman and Lammers 2004) characterize many species of Centropogon (e.g. sect. Siphocampyloides) and a few in other genera (e.g. Siphocampylus furax, Cyanea calycina). Some Cyanea
(e.g. C. tritomantha) have prickles on the vegetative
and floral organs which represent a unicellular
trichome with a thick secondary wall, atop an enlarged multicellular mass of epidermal cells (Carlquist 1962). Glandular trichomes are unknown.
Inflorescence Structure. Flowers of Campanulaceae may occur singly (commonly in
leaf axils, rarely at the stem apex) or may be
aggregated into diverse types of monotelic or
polytelic inflorescences, commonly in a terminal
position (Philipson 1948; Carolin 1967). Most have
a racemose, paniculate or spicate appearance but
are actually more complex and often composed of
essentially cymose subunits. Corymbose, umbellate and capitate inflorescences are not uncommon
and, in Cryptocodon, Jasione, Treichelia and some
species of Campanula (e.g. C. glomerata), the
capitate inflorescence may be subtended by an
involucre. In Ruthiella, flowers are epiphyllous.
Flowers of Lobelioideae, with few exceptions
(e.g. Downingia laeta, Monopsis stellarioides), are
resupinate through torsion of the pedicel; in genera with sessile flowers (e.g. Downingia, Grammatotheca), it is the hypanthium which undergoes torsion. As a result, the visually dorsal components of
the flower arise from the ventral portion of the
floral primordium. Throughout this treatment, the
terms “dorsal” and “ventral” refer to the apparent
orientation of fully formed structures, irrespective
of resupination.
Floral Structure and Anatomy. Flowers
are commonly complete (tetracyclic) and perfect
(bisexual), though a few species are dioecious (e.g.
Lobelia dioica) or gynodioecious (e.g. some L.
siphilitica). Proterandry characterizes the family
and is correlated with specialized mechanisms for
secondary presentation of pollen to floral visitors
(see below). Cleistogamy occurs in some genera of
Campanuloideae (Githopsis, Heterocodon, Trioda-
28
T.G. Lammers
nis, and a few Campanula, e.g. C. dimorphantha)
and Lobelioideae (Howellia, Legenere).
Typically, the calyx, corolla and androecium are isomerous and pentamerous. Certain
Campanuloideae have more than 5 units in
a whorl: 6 (Canarina, Cyclocodon lancifolius),
7 (Ostrowskia) or 8–10 (Michauxia). A few taxa are
tetramerous (e.g. Cyananthus hookeri, Cyclocodon
parviflorus, Echinocodon, Phyteuma tetramerium).
In cleistogamous flowers, sepals are often as few as
3 and the corolla is typically absent.
In Campanuloideae, the odd (unpaired) sepal
originates in a dorsal (posterior) position on the
primordium, while in the remainder of the family, it arises in a ventral (anterior) position; in
Lobelioideae, however, floral resupination brings
it into a visually dorsal position at anthesis. The
sepals are connate for a portion of their length
and (except in Cyananthus) this connate basal portion is adnate to the basal portions of the corolla
and androecium, forming a hypanthium of appendicular origin (Kaplan 1967). This hypanthium is
adnate to the ovary, typically for its entire length,
thus placing the ovary in an inferior position relative to the other whorls. In some taxa, however, the
adnate hypanthium is shorter than the ovary, i.e.
a portion of the ovary extends above the rim of the
point of insertion of the calyx lobes. In Diastatea,
Unigenes, and occasional species of Siphocampylus (e.g. S. reflexus), Lobelia (e.g. L. xalapensis) and
Wahlenbergia (e.g. W. welwitschii), the hypanthium
is reduced to such a degree that the ovary appears
nearly superior. Cyclocodon is unique in having the
calyx lobes inserted at the base of the hypanthium;
in some species of Codonopsis (e.g. C. javanica),
they are inserted at its middle. The anatomy of this
bizarre situation has not been studied in detail. In
some taxa (e.g. Campanula alliariifolia, Lobelia appendiculata, Zeugandra), the calyx bears a reflexed
appendage or auricle in each sinus.
The epigynous portions of the petals are likewise connate for some part of their length, typically
half or more. This fusion is the result of early sympetaly: the petals arise on a ring primordium or are
already connected at initiation. In some genera (e.g.
Asyneuma, Dialypetalum, Jasione, Michauxia), the
corolla tube may be so short that the flower appears choripetalous. The corollas of Physoplexis
and Phyteuma are also of this sort but the tips of
the lobes are connate or coherent apically, forming
a tube; the medial portion of the corolla is bowed
out between this apical tube and the base of the
corolla, forming five prominent fenestrations. In
some species of Lobelioideae (e.g. Lobelia fenestralis), the corolla tube bears a single lateral fenestration on each side. Venation of the petals consists of a prominent midvein (marginal veins may
also be present) which anastomoses into a dense
reticulum distally (Gustafsson 1995). Radial symmetry (actinomorphy) characterizes the corolla in
Campanuloideae, and is expressed in a broad diversity of corolla shapes, including campanulate,
infundibular, urceolate, tubular, salverform, and
rotate. Bilateral symmetry (zygomorphy) characterizes the remaining subfamilies; the degree of
zygomorphy varies from scarcely perceptible (e.g.
Dialypetalum, some Cyphioideae) to pronounced
(e.g. Cyphocarpoideae, most Lobelia). Most are bilabiate, with 3 ventral and 2 dorsal corolla lobes
(3 dorsal and 2 ventral in non-resupinate taxa);
some may form a definite palate on the ventral lip
(e.g. Lobelia coronopfolia). Unilabiate corollas also
occur, with all 5 lobes in a ventral position (e.g.
Clermontia peleana); in such flowers, the five lobes
are sometimes coherent at their apices (e.g. Lobelia tupa). Cyphocarpoideae are characterized by
a unique bilabiate pattern, with 1 cucullate dorsal
lobe and 4 ventral lobes. Nectar spurs are found in
Heterotoma and some Lobelia (e.g. L. goldmannii).
The commonest colours are various shades of blue
and violet, often with undertones of rose or mauve.
Other colours include pink (e.g. Lobelia bridgesii),
red (e.g. L. cardinalis), orange (e.g. Canarina canariensis), yellow (e.g. Musschia aurea), green (e.g.
Clermontia kakeana) and white (e.g. Hippobroma
longiflora). Bicoloured flowers are not uncommon,
particularly among Lobelioideae (e.g. Centropogon
granulosus, Lobelia laxiflora).
Stamens are antesepalous (i.e. alternating with
the corolla lobes) and inserted at the very base of
the corolla tube, on the rim of the hypanthium or
atop the inferior ovary. In a few genera (Cyphocarpus, Rhigiophyllum, Siphocodon), the filaments
are inserted higher on the corolla tube, apparently
through adnation of the basal portions of the
filaments. Among Campanuloideae, the bases of
the filaments often are expanded and elaborated
in diverse ways; though not connate, they may
cohere in such a way that a chamber is formed
over the nectary atop the ovary. In Nemacladus
and Parishella the filaments have unique multicellular appendages. Anthers are tetrasporangiate,
basifixed (dorsifixed in Berenice), and commonly
dehisce introrsely by longitudinal slits. In Lobelioideae, the anthers (and distal portions of
their filaments) are connate, forming a ventrally
Campanulaceae
oblique tube around the style into which the
pollen is shed. The two ventral (sometimes all five)
anthers typically bear a tuft of stiff trichomes at
the mouth of this tube. In Nemacladoideae and
most Cyphioideae, the filaments are connate for
part of their length but the anthers are distinct. In
many Campanuloideae, the anthers cohere around
the style but separate following pollen discharge.
In the few Campanuloideae with connate anthers
(e.g. Campanula cretica), the tube formed is
symmetrical, not oblique.
Carpels are fully connate, forming an ovary
which is characteristically inferior. Among Campanuloideae, a 3-locular ovary is most common;
some taxa have 5 locules, others have only 2 (sometimes reduced to 1); some genera (e.g. Michauxia,
Ostrowskia) appear to have more than 5 (up to
10) due to intrusive partitions from the carpellary
midribs. Among the other subfamilies, a 2-locular
ovary is the general rule, though in some taxa this
is reduced to a single locule. Each locule typically contains several to many ovules; in 2–5(–10)locular ovaries, placentation is axile whereas in 1locular ovaries (e.g. Legenere) it is parietal. In some
genera, the number of ovules per locule is reduced
to a few or even 1, in which case placentation may
be basal (e.g. Unigenes) or apical (e.g. Rhigiophyllum). In most of the subfamilies, the single style
is topped by stigmatic branches equal to the number of carpels in the gynoecium. The exception is
Cyphioideae, which feature a unique stigmatic cavity which is fluid-filled and communicates with the
outer air by means of a lateral aperture (Leins and
Erbar 2003).
Embryology. Anther wall development follows
the Dicotyledonous pattern, forming a single middle layer (Rosén 1932). Microsporogenesis is simultaneous, the tetrads tetrahedral or isobilateral.
Fibrous thickenings are found in the endothecium.
The tapetum is of the glandular (secretory) type,
and the tapetal cells are binucleate. Pollen grains
may be either binucleate or trinucleate when shed.
Ovules are anatropous, unitegmic, and tenuinucellar. Embryo sac formation is monosporic and of
the Polygonum type (Tobe and Morin 1996). Upon
coming in contact with the embryo sac, the inner
layer of the integument develops as an endothelium
(integumentary tapetum). Embryogenesis follows
the Solanad pattern. Endosperm formation is cellular ab initio (Rosén 1949); the endosperm typically is copious and oily (starchy in some Campanuloideae). Haustoria form at both the chalazal
29
and micropylar ends, and are equally aggressive.
No hypostase forms.
Pollen Morphology. Pollen grains are
spheroidal or variously compressed (oblate
or prolate). Tricolporate pollen characterizes
Cyphioideae, Cyphocarpoideae, Lobelioideae and
Nemacladoideae (6-colpate in Parishella), though
the pores may be lacking (i.e. the grain is 3-colpate)
in some Lobelioideae (Chapman 1966; Dunbar
1984). Campanuloideae show greater diversity. In
most genera, grains are 3–4(–5)-porate (6-porate in
Githopsis). Pantoporate grains with 8, 12 or 14–20
pores characterize a few North American species
of Campanula (e.g. C. americana, C. exigua). Other
genera are 3–6-colporate (Canarina, Platycodon)
or 6–10-colpate (Cyananthus, Ostrowskia). Surface
ornamentation is variable (Dunbar 1975). Spinules
are all but ubiquitous on porate grains but reduced
to verrucae in the colpate and colporate Campanuloideae. Spinules also characterize Cyphioideae
and Cyphocarpoideae but are lacking in Lobelioideae and Nemacladoideae. Pollen grains are
dispersed as monads (tetrads in Namacodon).
Karyology. Chromosome numbers have been
counted in 415 species of Campanuloideae. The
commonest by far is 2n = 2x = 34, accounting for
about 42% of the published counts; another 13%
are presumed polyploid derivatives: 2n = 4x = 68
and 2n = 6x = 102. A base number as large as
x = 17 may well have been derived from some
smaller number by aneuploidy following polyploidization, or by allopolyploidy. Other numbers
which have been reported are 2n = 12, 14, 16, 18,
20, 22, 24, 26, 28, 30, 32, 36, 38, 40, 42, 46, 48, 50,
52, 54, 56, 58, 60, 68, 70, 72, 80, 84 and 90 (Gadella
1966). Of particular significance is the occurrence
of 2n = 2x = 14 in Cyananthus, the only genus of
the subfamily characterized by a truly superior
ovary, which suggests x = 7 for the family.
Codonopsis and Echinocodon with 2n = 2x = 16,
and Platycodon with 2n = 2x = 18 may represent
lineages with x = 8 and x = 9 respectively. Ancient
hybridization of such plants could have yielded the
x = 17 found in much of the rest of the subfamily.
About 170 species of Lobelioideae have been
counted (Lammers 1993). Over three-fourths have
2n = 2x = 14 or its presumed polyploid derivatives,
2n = 4x = 28, 2n = 6x = 42, 2n = 10x = 70 and
2n = 20x = 140. The base number of the subfamily
thus appears to be x = 7. Other numbers reported
are 2n = 12, 16, 18, 20, 22, 24, 26 and 38. Diversifica-
30
T.G. Lammers
tion in some taxa has been accompanied by significant aneuploidy, e.g. Downingia with 2n = 2x = 12,
16, 18, 20, 22 and 24, and Lobelia sect. Delostemon
with 2n = 2x = 12, 14, 16, 18, 20 and 22.
Cyphioideae and Nemacladoideae have 2n =
2x = 18 in the two species of each which have been
examined, suggesting a base number of x = 9. No
chromosome numbers are known for Cyphocarpoideae.
Pollination and Reproductive Systems.
A major characteristic of the flowers of Campanulaceae is their specialized mechanism for
secondary pollen presentation, i.e. presentation of
pollen to potential pollinators on the style or other
floral structures, rather than directly from the
anther (Carolin 1960; Erbar and Leins 1989, 1996;
Leins and Erbar 1990). Key aspects of this phenomenon in Campanulaceae are the proterandry
of the flowers, the introrse discharge of pollen
from the anthers, and the gathering of the stamens
around the style by coherence or connation. The
combination of these features results in deposition
of pollen onto the style prior to its elongation and
the maturation of the stigma. Here, the pollen is
typically held by some sort of hairs. Those found
in subfamilies with merely coherent stamens are
dispersed along a significant portion of the length
of the style. In such plants, pollinators pick up
pollen from the style. The hairs of at least some
Campanuloideae are able to invaginate as a means
of dislodging adherent pollen grains (Shetler 1979).
In the syngenesious Lobelioideae, however, the stylar hairs are concentrated into a ring just below the
unreceptive stigmatic lobes. As the style lengthens
through the anther tube, these hairs sweep the
accumulated pollen load ahead of it towards the
tube’s orifice. In most lobelioid genera, the dorsal
anthers are longer than the ventral, occluding the
orifice, and the ventral anthers are tipped by tufts
of stiff trichomes. Pressure builds up on the pollen
load within the anther tube as the expanding style
pushes it forwards. When a pollinator searching for
nectar inadvertently depresses the stiff trichomes
at the orifice of the anther tube, the orifice is levered open and a load of pollen discharged onto the
foraging animal. A somewhat similar mechanism
characterizes the campanuloid genera Phyteuma
and Physoplexis, with the coherent or connate tips
of the corolla lobes replacing the anther tube.
Most species are zoophilous. Those with
actinomorphic corollas may be visited by a broad
suite of generalized insects, including bees, flies,
wasps, butterflies and settling moths. Species
with zygomorphic corollas have a more restricted
range of specialized, not infrequently vertebrate vectors. Ornithophily is well documented
among Lobelioideae, involving both nectarivorous passerines (e.g. Clermontia, Lobelia sect.
Rhynchopetalum) and hummingbirds (e.g. Centropogon, Siphocampylus). Chiropterophily has
been reported in Burmeistera and Siphocampylus,
while the long slender corollas of Brighamia,
Hippobroma and two species of Burmeistera
(Lammers 2002) appear adapted to sphingophily.
Autogamy is also an option for some taxa. Some
Campanuloideae (e.g. Githopsis, Triodanis) and Lobelioideae (e.g. Howellia, Legenere) bear reduced
cleistogamous flowers in addition to normal chasmogamous ones. Certain, otherwise zoophilous lobelioid flowers will self-pollinate if the pollen load
is not removed by the time the carpellate phase of
the proterandrous flower begins. As the stigmatic
lobes spread and recurve, they eventually come into
contact with the pollen-bearing ring of hairs on the
style just below.
Fruit and Seed. Most Campanulaceae form
capsular fruits, which typically open by valves, slits
or pores, located either at the apex of the ovary (i.e.
above the calyx lobes) or on the lateral walls of the
hypanthium (i.e. below the calyx lobes). Capsules
in a few genera (e.g. Craterocapsa, Lysipomia,
Parishella) open by an operculum.
In some genera, dehiscence is arrested. The
pericarp may become papery, forming a capsule which releases its seeds only by irregular
rupture or decomposition (e.g. Cyphocarpus,
Legenere, Peracarpa), or it may become fleshy,
forming a variously coloured berry (e.g. Canarina,
Clermontia), which sometimes may be quite
inflated (e.g. Burmeistera glabrata). The fruit is
a schizocarp in Theodoravia. Winged seeds are of
sporadic occurrence in the family (e.g. Ostrowskia,
Trematolobelia, some Cyphia). Seed germination
is phanerocotylar.
Dispersal. The genera with fleshy fruits are obviously adapted to endozoochory, but seeds from
dry fruits have no apparent adaptation for dispersal, aside from their small size. The few species
with winged seeds may be presumed to be anemochorous to some degree.
Phytochemistry. In Campanulaceae, starch is
replaced as a storage carbohydrate by inulin. Pyri-
Campanulaceae
dine alkaloids such as lobeline commonly accumulate in the latex of Lobelioideae but appear to be
all but absent from the other subfamilies. Among
Campanuloideae, polysterols apparently take their
place. Some species produce polyacetylenes. Those
isolated from Campanulaceae are aliphatic with 14
carbon atoms; the tetrahydropyranes are unique to
the family. Campanulaceae do not produce iridoids
nor sesquiterpene lactones.
Caffeic acid is a common phenolic constituent
of Campanuloideae, primarily as esters with quinic
acid, but is lacking from Lobelioideae where it is
seemingly replaced by chelidonic acid; p-coumaric
acid is likewise common in Campanuloideae
but absent from Lobelioideae. Other phenolics
reported from various Campanulaceae include
quercetin, kaempferol, ferulic acid, cyanidin,
sinapic acid and anthocyanins. Ellagic acid,
myricetin, leucodelphinidin and leucoanthocyanins are absent from the family as a whole.
A few species are cyanogenic, due to the presence
of tyrosine-derived triglochin (Tjon Sie Fat 1978).
Saponins are scarce and known primarily from
Platycodon.
Subdivision of and Relationships Within
the Family. In its broadest circumscription
(e.g. Schönland 1889; Takhtajan 1980), the family
includes not only the genera treated here but
also Pentaphragma Wallich ex G. Don (see Pentaphragmataceae) and Sphenoclea Gaertn. (see
Sphenocleaceae). In its narrowest circumscription
(e.g. Shetler and Morin 1986; Kolakovskii 1994;
Takhtajan 1997), it includes only those genera
treated here as Campanuloideae. The intermediate circumscription adopted here (following
Wagenitz 1964, and Cronquist 1981) is supported
by phylogenetic analyses of both morphological
and molecular data (Cosner et al. 1994; Gustafsson
and Bremer 1995; Gustafsson et al. 1996; Bremer and Gustafsson 1997; Kårehed et al. 1999;
Lundberg and Bremer 2003). In all these analyses,
the taxa included here as Campanulaceae form
a monophyletic group, whereas Sphenoclea and
Pentaphragma consistently fall outside this clade,
sometimes at a considerable distance.
The genera of Campanulaceae typically
have been divided among three subordinate
groups, ranked as tribes (Bentham 1876) or
more commonly subfamilies (Schönland 1889;
Wagenitz 1964; Wimmer 1968): Campanuloideae, Cyphioideae and Lobelioideae. However,
Cyphioideae as traditionally circumscribed (i.e.
31
to include Cyphia, Cyphocarpus, Nemacladus,
Parishella and Pseudonemacladus) are almost
certainly not monophyletic. Though exact relationships remain uncertain, it is clear that the traditional Cyphioideae comprise three disparate evolutionary lineages, based on morphological (Bentham 1875; Lammers 1992; Gustafsson and Bremer
1995), palynological (Dunbar 1975, 1984) and
molecular data (Cosner et al. 1994; Lundberg and
Bremer 2003). To remedy this situation, these five
genera have been divided among three subfamilies
(Lammers 1998a): Cyphioideae (Cyphia), Cyphocarpoideae (Cyphocarpus) and Nemacladoideae
(Nemacladus, Parishella and Pseudonemacladus).
Several divergent classifications have been
proposed for the genera within Campanuloideae.
Schönland (1889) divided them among three
subtribes, based on features of the ovary and
mature fruit; Yeo (1993) accepted this classification
but employed tribal rank. In Platycodoneae Yeo,
the five carpels alternate with the calyx lobes. In
the other two tribes, the carpels are fewer than
the calyx lobes or, if isomerous, are positioned
opposite the calyx lobes. In Campanuleae Dumort.,
dehiscence is lateral, i.e. the capsule opens below
the calyx lobes, whereas in Wahlenbergieae Endl.,
dehiscence is apical, i.e. the capsule opens above
the calyx lobes.
Fedorov (1957) proposed a new system which
accounted only for genera in the Soviet Union. It
was “based primarily on the different modes of dehiscence of the capsule, the shape of the corolla and
aggregate characters, determining general similarity.” In it, four tribes (Michauxieae Fed., Ostrowskieae Fed., Peracarpeae Fed., and Phyteumeae
Dumort.) were segregated from Campanuleae, and
two more (Edraiantheae Fed., Jasioneae Dumort.)
from Wahlenbergieae; in addition, Platycodoneae
were subsumed into Wahlenbergieae. This system
was applied to the species of China by Hong et al.
(1983), with the addition of one tribe, Cyanantheae
Meisn.
More recently, three additional classifications
have been proposed. Kolakovskii (1987, 1994) recognized four subfamilies within Campanulaceae
s. str.: Prismatocarpoideae Kolak., Canarinoideae
Kolak. (two tribes), Wahlenbergioideae (Endl.) Kolak. (10 tribes) and Campanuloideae (nine tribes);
only Old World genera were covered, leaving
unknown the disposition of the North American
endemics Githopsis, Heterocodon and Triodanis.
Takhtajan (1997) likewise divided Campanulaceae
s. str. into four subfamilies: “Cyananthoideae”,
32
T.G. Lammers
nom. invalid. sub Art. 36.1 (three tribes); Ostrowskioideae (Fed.) Takht.; Canarinoideae Kolak.;
and Campanuloideae (13 tribes). Hong (1995)
tentatively divided the family into six unnamed
groups: the Cyananthus group (two subgroups),
Platycodon group (two subgroups), Ostrowskia
group, Wahlenbergia group (four subgroups),
Jasione group and Campanula group (three
subgroups).
Lobelioideae have not received as much classificatory attention. Presl (1836) recognized three
tribes, based largely on fruit characteristics. Delisseeae C. Presl comprised genera with baccate fruit,
versus the dehiscent fruits of the other two. Those
of Clintonieae C. Presl (nom. illegit. sub Art. 19.5)
had a single locule, those of Lobelieae Rchb., two
locules. De Candolle (1839) modified this system
by dividing the 1-locular genera between those
with an elongate laterally dehiscent capsule (Clintonieae) and those with a more isodiametric pyxicidal capsule (Lysipomieae A. DC.). In his monograph of the subfamily, Wimmer (1943, 1953, 1968)
merged Clintonieae, Lysipomieae and Lobelieae. As
such, he recognized only two tribes: Delisseeae with
baccate, and Lobelieae with dehiscent fruits. The
former was divided into two subtribes, the latter
into nine. In as much as Lobelia and Isotoma as
here circumscribed include both baccate and capsular species, this classification is scarcely tenable.
In summary, there is a lack of consensus among
authors who have proposed classifications of Campanuloideae, compounded by a lack of attention to
this topic in Lobelioideae. None of the existing classifications has been expounded in great detail, nor
has any been tested through phylogenetic analysis
and/or the addition of molecular data. For these
reasons, no formal classification below the rank of
subfamily is employed here.
Affinities. Campanulaceae traditionally have
been assigned to Campanulales. The varying
circumscriptions and taxonomic history of this
order have been discussed in detail by Lammers
(1992). In that summary, other families assigned
to the order were Asteraceae, Brunoniaceae,
Calyceraceae, Goodeniaceae, Menyanthaceae,
Pentaphragmataceae and Sphenocleaceae. Since
that time, phylogenetic analyses of DNA sequences
(Olmstead et al. 1992, 1993; Chase et al. 1993;
Michaels et al. 1993; Cosner et al. 1994; Gustafsson
et al. 1996; Bremer and Gustafsson 1997; Kårehed
et al. 1999; Lundberg and Bremer 2003) have
consistently shown that (1) these families, with
the exception of Sphenocleaceae, do indeed
form a monophyletic group; (2) Stylidiaceae and
Donatiaceae, excluded to Ericales or Stylidiales
by Lammers (1992), belong to this lineage; and
(3) additional families traditionally assigned
to Saxifragales or Cornales also belong to this
lineage (here called Asterales, despite Rec. 16B.1).
Although a complete evaluation of this expanded
circumscription in light of non-molecular data has
not been completed, preliminary studies (Gustafsson and Bremer 1995; Lundberg and Bremer 2003)
suggest that it could be supported by morphology.
Within Asterales, relationships are not yet fully
resolved. In some studies (Cosner et al. 1994), Stylidiaceae were shown to be the sister group of Campanulaceae whereas in others (Kårehed et al. 1999;
Lundberg and Bremer 2003), Pentaphragmataceae
were sister to Campanulaceae. However, each of
these studies utilized slightly different sets of taxa.
Distribution and Habitats. Campanulaceae
taken as a whole are cosmopolitan in distribution, with representatives on all six continents and
many oceanic islands, from the tropics to the frigid
zones. However, the subfamilies are not evenly distributed. The three smaller subfamilies are endemic
to rather restricted areas: Cyphioideae to southern Africa, Cyphocarpoideae to central Chile, and
Nemacladoideae to south-western North America.
The two major subfamilies, Campanuloideae and
Lobelioideae, have much wider distributions but
are largely non-overlapping. In general, Campanuloideae are found primarily in the temperate zones
of the Old World, whereas Lobelioideae are distributed predominantly in the world’s tropical and
subtropical zones. For example, South America is
home to approximately 575 species of Lobelioideae
but only 10 Campanuloideae, while Europe harbours 275 Campanuloideae but only 8 Lobelioideae.
For the family as a whole, Africa and South
America each has about one-fourth of the species;
18% are Asian, 11% European, 11% North American, 6% Polynesian and 4% Australasian. Specifically, the greatest concentration of diversity occurs
in southern Africa, where 18 genera and nearly 400
species occur. This large total is due to the fact
that this is the only region to have significant numbers of both Campanuloideae and Lobelioideae, as
well as all Cyphioideae. Other major centres of diversity are the Andes of South America, with over
500 species of Lobelioideae, and Eurasia between
the Mediterranean and the Himalayas, where the
Campanuloideae are similarly diverse. Particularly
Campanulaceae
notable is the presence of over 130 endemic species
of Lobelioideae (almost 6% of the family) on the
small but highly isolated Hawaiian islands.
Ecologically, the family is extremely diverse.
The majority of species are mesophytes, many occurring in montane habitats. Though exceptions
are numerous, there is a discernible trend among
Campanuloideae for more open habitats, especially
meadows, while many Lobelioideae tend to be associated with forested areas, particularly rainforests
and cloud forests. Some of the latter are facultative
epiphytes (e.g. Clermontia). The species of Lobelia
which have adapted to the alpine conditions in the
mountains of East Africa (e.g. L. rhynchopetalum,
L. telekii) are particularly noteworthy.
A significant number of taxa grow under xeric
conditions, either as sclerophyllous shrubs and
subshrubs (e.g. Roella, Merciera), by developing
underground storage organs (e.g. Cyphia) or
by adopting an annual habit (e.g. Cyphocarpus,
Nemacladus). A few species are rooted freshwater
hydrophytes, growing submersed in shallow water.
Howellia is cabomboid in habit, with flimsy stems
and leaves, while Lobelia dortmanna is isoetid,
with a basal rosette of stiff linear aerenchymatous
leaves and emergent scapose inflorescences.
Downingia and its allies Legenere and Porterella
have exploited seasonally dry, vernal pool habitats. A number of species (e.g. Lobelia boykinii,
Siphocampylus verticillatus) grow in saturated soil
of marshes, bogs, and other wetland habitats.
Palaeobotany. The sole fossil remains ascribed
to Campanulaceae are seeds from the Miocene
of Poland, described by Lancucka-Srodoniowa
(1977, 1979) as Campanula palaeopyramidalis and
“Campanula sp.”.
Economic Importance. The primary economic
value of the Campanulaceae is in horticulture.
Many species are cultivated as ornamentals and
commonly available in the home gardening trade.
Those best known to the average grower are the
large perennial species of Campanula planted in
beds and borders (e.g. C. persicifolia, C. carpatica,
C. glomerata); Lobelia cardinalis, Platycodon
grandiflorus and Trachelium caeruleum are used
similarly. Lobelia erinus is a standard annual for
bedding, window boxes and hanging baskets,
while biennial Campanula medium is a staple
of old-fashioned cottage gardens. Other genera
(e.g. Adenophora, Azorina, Brighamia, Canarina,
Codonopsis, Isotoma, Jasione, Michauxia, Phy-
33
teuma, Wahlenbergia) are cultivated to a lesser
extent, primarily by those who fancy the rare
and unusual. Favratia zoysii and the many dwarf
species of Campanula (e.g. C. alpestris, C. arvatica,
C. waldsteiniana) are quite popular with rock
garden and alpine enthusiasts. Compared to many
horticultural groups (e.g. Rosa, Tulipa), Campanulaceae have been only moderately affected by
selective breeding and cultivar development, and
even less by hybridization. Most of the family’s
garden representatives are not far removed from
their wild relatives, and are readily referred to
naturally occurring species.
In addition to ornament, a few species are cultivated as comestibles. Campanula rapunculus, the
rampion or rapunzel, has been grown as a vegetable in Europe, for both its young leaves and its
roots, but the cultivars with sweet, well-developed
taproots have largely been lost. Platycodon grandiflorus has been used similarly in eastern Asia.
The family also produces several pharmaceuticals. Lobelia inflata is the commercial source of
lobeline, a pyridine alkaloid used in anti-smoking
therapy and formerly in the treatment of bronchial
asthma and as a respiratory stimulant. The roots of
Codonopsis, primarily C. pilosula, are the source
of dang-shen, a mainstay of traditional Chinese
medicine. A popular substitute for the more costly
ginseng, like that drug it is employed as a tonic for
fatigue, loss of appetite, and weakness. Research
has demonstrated that the raw extract promotes
digestion, strengthens the immune system, dilates
peripheral blood vessels and inhibits adrenal cortex activity, but the active principle has to date not
been isolated. The root of Platycodon grandiflorus
yields another important Asian drug, jie-geng or
kikyo, much used as an expectorant and antitussitive; recent studies have shown it to also have
analgesic, antipyretic, anti-inflammatory and antibacterial properties. The active principle appears
to be a saponin (platycodin or kikyosaponin). Various other species of the family have been reported
to have local usages in folk medicine, particularly
among non-industrialized populations, but have
not been researched in detail nor exploited commercially.
Classification of Campanulaceae
I. Subfamily Campanuloideae Burnett (1835).
Genera 1–50
II. Subfamily Nemacladoideae Lammers (1998).
Genera 51–53
34
T.G. Lammers
III. Subfamily Lobelioideae Burnett (1835).
Genera 54–82
IV. Subfamily Cyphocarpoideae Miers (1848).
Genus 83
V. Subfamily Cyphioideae (A. DC.) Walp. (1852).
Genus 84
Key to the Genera
1. Corolla actinomorphic; odd (unpaired) sepal in a dorsal (posterior) position; locules and stigmas (1–)3–
5(–10)
2
– Corolla zygomorphic (sometimes only slightly so);
odd (unpaired) sepal in a ventral (anterior) position
prior to any floral resupination; locules and stigmas
(1–)2
51
2. Fruit dehiscing apically (i.e. above the calyx lobes), if
indehiscent, pericarp fleshy
3
– Fruit dehiscing laterally (i.e. below the calyx lobes),
if indehiscent, then pericarp dry, membranous or papery
22
3. Ovary wholly superior, completely free from calyx
1. Cyananthus
– Ovary inferior or semi-superior, at least its base adnate
to the calyx, forming a hypanthium
4
4. Ovary 5-locular, locules alternating with the sepals 5
– Ovary 2–6-locular, if 5-locular, then locules opposite
the sepals
6
5. Perennial with tuberous roots; flowers solitary; corolla
large, bowl-shaped; pollen 5–7-colpate 2. Platycodon
– Annual; flowers in corymbs or heads; corolla small,
cylindric; pollen 3-porate
19. Microcodon
6. Calyx lobes inserted at base of hypanthium
4. Cyclocodon
– Calyx lobes inserted at summit (rarely middle) of hypanthium
7
7. Roots greatly enlarged, tuberous; stems typically scandent or climbing, rarely erect or ascending
8
– Roots fibrous; stems erect or ascending
9
8. Plant often malodorous; leaves entire or toothed; petioles and pedicels not twining; corolla lobes and stamens 5; fruit a capsule, rarely a berry partially subtended by the calyx lobes
3. Codonopsis
– Plant not malodorous; leaves hastate or sagittate; petioles and pedicels twining; corolla lobes and stamens
6; fruit a berry crowned by the calyx lobes 7. Canarina
9. Leaves pinnately lobed or parted; corolla lobes and
stamens typically 4; pollen 4–5-colpate 5. Echinocodon
– Leaves typically entire or toothed; corolla lobes and
stamens typically
5; pollen 3-porate
10
10. Capsule dehiscent by pores, an operculum, or irregularly, never by valves
11
– Capsule dehiscent by valves, rarely also by pores 16
11. Annuals; capsule dehiscent by 1 apical pore
41. Githopsis
– Perennial herbs and subshrubs; capsule dehiscent by
an operculum or irregularly (rarely by 1 apical pore)12
12. Capsule irregularly dehiscent
23. Edraianthus
– Capsule dehiscent by an operculum (rarely by 1 apical
pore)
13
13. Corolla cylindrical; filaments adnate to corolla
14
– Corolla campanulate or funnelform; filaments free
from corolla
15
14. Leaves dense; flowers in a head; ovules several to
numerous in each locule
22. Rhigiophyllum
– Leaves sparse; flowers in a raceme or panicle; ovules
2–6 per locule
21. Siphocodon
15. Ovary and capsule 2-locular
15. Roella
– Ovary (2–)3-locular, capsule 1 locular at maturity
14. Craterocapsa
16. Anthers dorsifixed
11. Berenice
– Anthers basifixed
17
17. Shrubs; flowers with conspicuous nectaries
18
– Herbs or subshrubs (rarely shrubs); flowers lacking
nectaries, or nectaries inconspicuous
19
18. Flowers in racemes or panicles; ovary topped by a
5-notched nectar disc; each valve of capsule with
a pair of pores and three more pairs of pores at base
of capsule
10. Heterochaenia
– Flowers solitary; ovary topped by 5 ellipsoid nectaries;
capsule lacking pores
9. Nesocodon
19. Inflorescence involucrate; anthers connate at base
25. Jasione
– Inflorescence not involucrate; anthers distinct or
coherent
20
20. Flowers in flat-topped corymbs; anthers apiculate,
inner pollen sacs shorter than outer
24. Feeria
– Flowers solitary or in various racemose or paniculate
inflorescences; anthers not apiculate, thecae equal 21
21. Corolla cylindric; filaments filiform; placentation
basal
12. Theilera
– Corolla campanulate, funnelform, rotate, or rarely
urceolate; filaments basally dilated; placentation axile
8. Wahlenbergia
22. Calyx lobes, corolla lobes, stamens and locules
(6–)7–10
23
– Calyx lobes, corolla lobes and stamens 3–5 (corolla
rarely lacking); locules 1–5
24
23. Biennial; leaves alternate; corolla rotate, lobes much
longer than tube; pollen 3-porate; capsule dehiscent
by 8–10 basal valves
49. Michauxia
– Perennial; leaves whorled; corolla funnelform,
lobes shorter than tube; pollen 6–7-colpate; capsule
dehiscent by (10–)14(–18) median longitudinal slits
6. Ostrowskia
24. Fruit indehiscent, or dehiscing only tardily and
irregularly
25
– Fruit dehiscent by definite valves, pores, fissures
or slits, or breaking into 3 mericarps joined by the
persistent style bases
27
25. Suffruticose perennial or subshrub; corolla tubular
or cylindric, lobes shorter than tube; ovary 1- or
imperfectly 2-locular, containing only 4 ovules on
basal placentae; mature seeds 1–2
20. Merciera
– Annual or herbaceous perennial; corolla funnelform
or campanulate, lobes equalling or longer than tube;
ovary 2–3-locular, containing 10 to many ovules on
axile placentae; mature seeds 10 to many
26
26. Annual; locules 2; ovules and seeds numerous
16. Gunillaea
– Perennial; locules (2–)3; ovules 8–10 per locule; seeds
10–16 per capsule
36. Peracarpa
27. Fruit a schizocarp, the 3 locules indehiscent but
separating from the ovary and held together by the
persistent style
33. Theodorovia
– Fruit a capsule, locules opening with valves, pores,
fissures or slits
28
Campanulaceae
28. Corolla bright yellow or reddish brown; capsule
dehiscent by numerous transverse slits 26. Musschia
– Corolla various shades of violet, lilac, blue or white,
only rarely yellowish or pinkish; capsule dehiscent by
valves, pores, or longitudinal fissures or slits
29
29. Capsule split for almost its entire length by 3–5
longitudinal fissures
30
– Capsule opening by relatively short valves, pores or
slits
33
30. Biennial; capsule splitting from base to apex; seeds
large, few
35. Sachokiella
– Suffruticose perennials and subshrubs; capsule
splitting from apex to base; seeds small, many
31
31. Capsule splitting into 3 fragments; pollen shed in
tetrads
13. Namacodon
– Capsule splitting into 5 fragments; pollen in monads
32
32. Locules 2
17. Prismatocarpus
– Locules 3
34. Muehlbergella
33. Corolla cleft nearly to base, appearing choripetalous
34
– Corolla cleft for 1/10–3/4 of its length, clearly sympetalous
40
34. Corolla lobes coherent or connate apically, creating
an apical tube separated from the basal tube by the
bowed-out lobes
35
– Corolla lobes distinct apically
36
35. Flowers distinctly pedicellate; corolla lobes connate
throughout anthesis; filaments filiform 48. Physoplexis
– Flowers sessile or nearly so; corolla lobes merely
coherent, separating after fertilization; filaments
dilated basally
47. Phyteuma
36. Capsule dehiscent by basal valves
46. Sergia
– Capsule dehiscent by subapical or medial pores or slits
37
37. Flowers solitary; hypanthium several times longer
than broad; capsule dehiscent by slits
45. Cylindrocarpa
– Flowers in racemes, panicles, spikes or heads; hypanthium slightly longer than broad; capsule dehiscent
by pores
38
38. Leaves pinnatifid; stigmatic lobes minute, stigma
capitate
44. Petromarula
– Leaves entire or toothed; stigmatic lobes filiform,
obvious
39
39. Inflorescence involucrate; sinuses of calyx appendaged; corolla tubular
43. Cryptocodon
– Inflorescence lacking an involucre; sinuses of calyx
unappendaged; corolla rotate or funnelform
42. Asyneuma
40. Rosette shrub; herbage viscid
27. Azorina
– Herbs; herbage not viscid
41
41. Flowers with a large, conspicuous, cylindrical or
tubular nectary
42
– Flowers with no obvious nectary, or nectary
inconspicuous, disciform
43
42. Anthers connate throughout anthesis
32. Hanabusaya
– Anthers distinct, or coherent in bud but distinct
in anthesis or after fertilization
31. Adenophora
43. Filaments connate; anthers apically apiculate; seeds
large, c. 15 per locule
30. Zeugandra
– Filaments distinct; anthers not apiculate apically;
seeds small, numerous
44
35
44. Corolla urceolate, each sinus with a large saccate
appendage
29. Favratia
– Corolla campanulate, funnelform, tubular, or rotate,
lacking saccate appendages
45
45. Flowers in an involucrate head; corolla narrowly cylindric above a globose base; base of style conspicuously
swollen, globose; locules 2; capsule dehiscent with
longitudinal slits, covered by the solid, conic style base
18. Treichelia
– Flowers solitary or in various inflorescences, only
rarely in an involucrate head; corolla rotate, bowlshaped, funnelform, or campanulate; base of style
filiform, confluent with ovary; locules typically 3–5,
sometimes 1 by abortion of septa, only rarely 2; capsule dehiscent with valves or pores, style deciduous 46
46. Flowers in elongate spike-like inflorescence; hypanthium elongate, 3–8 times longer than broad; corolla
rotate
47
– Flowers solitary or in various racemose, paniculate,
or corymbiform inflorescences, seldom spike-like;
hypanthium compact, twice as long as broad or less;
corolla campanulate, funnelform, or tubular, rarely
rotate
48
47. All flowers chasmogamous, with 5 calyx lobes and
well-developed corolla; filaments filiform or nearly so
38. Legousia
– Most flowers in inflorescence cleistogamous, calyx
lobes reduced to 3 and corolla rudimentary or lacking;
filaments basally dilated
39. Triodanis
48. Leaves all sessile; flowers sessile, solitary opposite
a leaf, many cleistogamous
40. Heterocodon
– Some leaves on plant petiolate; flowers sessile or
pedicellate, in various inflorescences, if solitary, then
either terminal or in leaf axil, any cleistogamous
flowers pedicellate
49
49. Stems distinctly triquetrous; flowers sessile or nearly
so, 1–3 in leaf axils
37. Homocodon
– Stems terete or shallowly winged; flowers pedicellate,
in various racemose, paniculate or corymbiform
inflorescences, if solitary, then terminal
50
50. Perennial lacking basal rosette; flowers very small,
narrowly tubular, in a pedunculate flat-topped
corymb; filaments filiform; style greatly exserted
50. Trachelium
– Annuals, biennials, or perennials with basal rosette;
flowers small to more commonly medium-sized
or large, campanulate, funnelform, bowl-shaped,
tubular, or rotate, solitary or in various racemose or
paniculate inflorescences; filaments basally dilated;
style included or only slightly exserted28. Campanula
51. Flowers oriented normally, the odd (unpaired) sepal
in a ventral (anterior) position at anthesis; anthers
distinct, all alike
52
– Flowers typically resupinate, the odd (unpaired) sepal
in a dorsal (posterior) position at anthesis; anthers
connate, the dorsal 3 longer than the ventral 2
56
52. Flowers sessile; corolla of a single cucullate dorsal
lobe bearing an apical appendage plus 4 ventral lobes;
stamens all epipetalous
83. Cyphocarpus
– Flowers pedicellate; corolla composed of 2 more or
less flat, unappendaged ventral lobes plus 3 dorsal
lobes; stamens inserted at base of corolla tube, on
the rim of the hypanthium or atop the inferior ovary,
rarely the dorsal two epipetalous
53
36
T.G. Lammers
53. Perennials with tuberous roots; style tipped by a fluidfilled stigmatic cavity with a lateral pore 84. Cyphia
– Annuals (if perennial, lacking tubers); style tipped by
a bilobed stigma
54
54. Perennials; leaves subopposite; pedicels bibracteolate
at apex
51. Pseudonemacladus
– Annuals; leaves alternate or in a basal rosette; pedicels
ebracteolate
55
55. Cauline leaves (if any) much smaller and narrower
than basal leaves; capsule apically dehiscent with 2
valves
52. Nemacladus
– Cauline leaves resembling basal leaves; capsule
circumscissile
53. Parishella
56. Corolla actinomorphic, petals free almost to base
54. Dialypetalum
– Corolla zygmorphic (sometimes only slightly so),
petals connate, forming a definite tube
57
57. Robust perennial herbs, shrubs, treelets, trees, giant
rosette plants and lianas; stems woody or suffruticose,
at least at base, (0.3–)1–8 m tall; corolla (1.4–)3–15 cm
long, lobes all more or less similar in size and shape,
or dorsal lobes larger than ventral ones
58
– Slender perennial, biennial or annual herbs; stems
herbaceous throughout, 0.03–1 m tall; corolla
0.2–2.5 cm long, dorsal lobes typically smaller than
ventral ones (rarely all more or less similar)
70
58. Inflorescence an axillary raceme (sometimes
subumbellate)
59
– Inflorescence a terminal raceme (sometimes corymbiform or capitate) or flowers solitary in leaf axils 62
59. Corolla salverform; fruit an apically dehiscent capsule
79. Brighamia
– Corolla bilabiate or unilabiate; fruit a berry
60
60. Much-branched shrubs or trees; inflorescence
typically 2-flowered; berries large
82. Clermontia
– Unbranched or sparingly branched shrubs, treelets
or trees (rarely lianas); inflorescence typically
6–40-flowered; berries small
61
61. Corolla with a knob at base of dorsal slit; seeds large,
white, transversely rugose
80. Delissea
– Corolla smooth at base of dorsal slit; seeds small,
brown or black, smooth
81. Cyanea
62. Corolla salverform
71. Hippobroma
– Corolla tubular, bilabiate or unilabiate
63
63. Hypanthium and corolla distended ventrally into
a large, broad, crescent-shaped nectar spur
72. Heterotoma
– Flower lacking a nectar spur, at most hypanthium
and corolla slightly oblique or gibbous ventrally 64
64. Ovary 1-locular, placentation parietal 77. Apetahia
– Ovary 2-locular, placentation axile
65
65. Fruit dehiscent via pores
66
– Capsule dehiscent via apical valves, or indehiscent 67
66. Pores 2, apical; flowers solitary in leaf axils
76. Sclerotheca
– Pores 3–12, lateral; flowers in a raceme, commonly
horizontally branched at base
78. Trematolobelia
67. Corolla tube cleft dorsally to base
55. Lobelia(subg.Tupa)
– Corolla tube entire
68
68. Top of ovary conical; fruit a capsule
73. Siphocampylus
– Top of ovary flat; fruit a berry
69
69. Apex of anther tube occluded; pedicels typically
bibracteolate; corolla, staminal column and style
early deciduous on the developing fruit; seeds oblong
or linear, their reticulations more or less elongate
74. Centropogon
– Apex of anther tube broadly open; pedicels typically
ebracteolate; corolla, staminal column and style
withering-persistent on the developing fruit; seeds
ellipsoid or lenticular, their reticulations more or less
isodiametric
75. Burmeistera
70. Flowers sessile, resupinate due to torsion of the
hypanthium (rarely not resupinate); hypanthium
pedicelliform, 5–10 times longer than wide
71
– Flowers pedicellate (rarely subsessile), resupinate due to torsion of the pedicel (rarely not
resupinate); hypanthium cupulate, shorter than wide
to as much as 4 times longer than wide
73
71. Corolla tube entire; ventral anthers with apical tufts
of hairs
69. Downingia
– Corolla tube dorsally cleft at least 2/3 the distance to
base; all 5 anthers with apical tufts of hairs
72
72. Leaves sessile; flowers solitary in leaf axils; stigmas
round
58. Grammatotheca
– Leaves petiolate; flowers in axillary fascicles; stigmas
filiform
59. Dielsantha
73. Ovary 1-locular, placentation parietal or basal
74
– Ovary 2-locular, placentation axile
77
74. Corolla tube entire (very rarely cleft dorsally to base);
filament tube adnate to corolla, at least at base; capsule
dehiscent by an umbonate operculum 70. Lysipomia
– Corolla tube dorsally cleft to base; filament tube free
from corolla; capsule dehiscent with apical valves
and/or irregular lateral ruptures
75
75. Plants terrestrial; leaves petiolate; hypanthium very
shallow, all but obsolete; ovary almost superior;
placentation basal; seed 1
61. Unigenes
– Plants aquatic (submersed or emergent); leaves
sessile; hypanthium 4–8 times longer than wide;
ovary inferior; placentation parietal; seeds 3–20 76
76. Plants typically emergent; seeds 18–20, 1–1.8 mm long
67. Legenere
– Plants typically submersed; seeds 3–5, 2–4 mm long
68. Howellia
77. Corolla tube cleft dorsally from apex almost to base 78
– Corolla tube entire, or cleft dorsally from base half
the distance to apex
80
78. Flowers not resupinate (rarely so); stigmas filiform
60. Monopsis
– Flowers resupinate; stigmas round
79
79. Pedicel free from subtending leaf or bract; dorsal
corolla lobes shorter than ventral or about equal
55. Lobelia (subg. Lobelia, subg. Isolobus)
– Pedicel adnate to subtending leaf, flower epiphyllous;
dorsal corolla lobes 1.5–5 times longer than ventral
63. Ruthiella
80. Filaments (at least dorsal one) adnate to corolla tube 81
– Filaments free from corolla tube
83
81. Ovary more than 4/5 superior, adnate to hypanthium
only at base; capsule dehiscent to about the middle
64. Diastatea
– Ovary less than 1/3 superior, adnate to hypanthium
for all or most of its length; capsule dehiscent only at
apex
82
Campanulaceae
82. Corolla tube entire; all filaments adnate to corolla
62. Isotoma
– Corolla tube cleft dorsally from base half the distance
to apex; only dorsal filament adnate to corolla
65. Palmerella
83. Stems fleshy; calyx lobes 3–8(–11) mm long; all 5 anthers with apical tufts of hairs; capsules 5–10(–16) mm
long; seeds c. 1 mm long
66. Porterella
– Stems herbaceous; calyx lobes 1–4 mm long; ventral
2 anthers with apical tufts of hairs; capsules 1–6 mm
long; seeds 0.3–0.5 mm long
84
84. Stems erect, flowers solitary and axillary or terminal;
pedicels with 1–3 bracteoles near the middle; filament
tube 1.5–2.5 mm long; capsules 1–3 mm long; seeds
ellipsoid, strophiolate, sulcate with keeled walls
56. Solenopsis
– Stems decumbent, flowers solitary and axillary or, if
stems erect, then flowers 2–15 in a terminal raceme;
pedicels bibracteolate at base; filaments 2–6 mm long;
capsules 2.5–6 mm long; seeds subglobose, lacking
a strophiole, sulcate with flattened walls
57. Wimmerella
Subfamilies and Genera
of Campanulaceae
I. Subfam. Campanuloideae Burnett (1835).
Herbaceous perennials, less often annuals or
biennials, herbaceous lianas, sometimes twining,
pachycaul rosette plants, subshrubs or shrubs, terrestrial, rarely epiphytic. Inflorescences monotelic
or polytelic, commonly appearing racemose, paniculate, spicate, umbellate or capitate, the latter two
sometimes involucrate, typically terminal, or the
flowers solitary in an axillary or terminal position.
Calyx with the odd (unpaired) lobe in a dorsal
(posterior) position. Corolla radially symmetric;
lobes (4–)5(–10), valvate. Anthers distinct or connivent, rarely connate, in which case the resulting
tube is symmetric. Ovary (2–)3–5(–10)-locular
with axile placentation, rarely 1-locular with parietal, basal, or apical placentation. Fruit a capsule,
commonly loculicidal or poricidal, or a berry.
Campanuloideae comprise 50 genera and 1,046
species. Though the subfamily is represented on all
six continents, 89% of the genera and 96% of the
species are endemic to the Old World. Africa and
Asia each harbour about one-third of the species,
Europe another one-quarter and Australasia about
4%; by contrast, just 3% of the species are North
American and 1% South American.
1. Cyananthus Wall. ex Benth.
Cyananthus Wall. ex Benth. in Royle, Ill. Bot. Himal. Mts.
309 (1836), nom. cons.; Hong & Ma, Acta Phytotax. Sin.
37
29: 25–51 (1991), rev.; Shrestha, Acta Phytotax. Sin. 35:
396–433 (1997), rev.
Annual or dwarf perennial herbs, often with
a large caudex. Leaves cauline, petiolate. Flowers
medium-sized, solitary and terminal (rarely in
cymes), short-pedicellate or rarely sessile. Calyx
(3–)4–5-lobed, free from ovary and stamens.
Corolla funnelform or tubular, blue (rarely yellow
or white); lobes 4–5, equalling or shorter than
the tube. Stamens 4–5; filaments slender; pollen
(6–)7–9(–12)-colpate, minutely spinulose. Ovary
superior, 3–5-locular. Fruit capsular, apically
dehiscent by 3–5 valves. 2n = 14. Twenty-three
species, eastern Asia, divided by Shrestha (1997)
into two subgenera: subg. Cyananthus (21 spp.)
with sect. Stenolobi Franch. (9 spp.), sect. Suffruticulosi K. Shrestha (2 spp.), sect. Cyananthus (7 spp.)
and sect. Annui (Y.S. Lian) D.Y. Hong & L.M. Ma
(3 spp.), and subg. Micranthus K. Shrestha (2 spp.).
2. Platycodon A. DC.
Platycodon A. DC., Monogr. Campan. 125 (1830).
Perennial herb from tuberous roots. Leaves cauline,
often subopposite or 3–4-verticillate, sessile or
short-petiolate. Flowers large, pedicellate, solitary,
terminal. Corolla bowl-shaped, blue-purple (rarely
pink or white); lobes equalling or shorter than
tube. Filaments dilated basally; anthers longer than
filaments; pollen 5–6(–7)-colpate, spinulose. Ovary
5-locular, locules antesepalous. Fruit capsular, loculicidal, apically dehiscent by 5 valves; seeds large.
2n = 18, 36. One species, P. grandiflorus (Jacq.) A.
DC., eastern Asia and common in cultivation.
3. Codonopsis Wall.
Codonopsis Wall. in Roxb., Fl. Ind. 2: 103 (1824); Shen &
Hong, Fl. Reip. Pop. Sin. 73, 2: 32–69 (1983), reg. rev.
Scandent, climbing or twining (rarely erect or
ascending) perennials, often with a skunk-like
odour; roots large, tuberous. Leaves sometimes
opposite or fasciculate, petiolate or sessile. Flowers
large, pendulous or rarely erect, pedicellate,
solitary, terminal or axillary. Calyx lobes crowning
hypanthium or inserted medially. Corolla purple,
blue, green, yellow or white, often with intricate
contrasting patterning, campanulate, infundibular, or tubular (rarely rotate); lobes shorter
than tube. Filaments dilated basally; anthers coherent; pollen (5–)7–9(–10)-colpate, spinulose
or reticulate. Ovary 3- or 5-locular, sometimes
38
T.G. Lammers
only half-inferior. Fruit capsular, loculicidal,
dehiscent by 3 or 5 apical valves, or a berry partly
subtended by the persistent calyx lobes; seeds
sometimes winged. 2n = 16. Fifty-nine species,
eastern Asia, divided by Shen and Hong (1983)
into three subgenera: subg. Codonopsis (52 spp.),
subg. Obconicapsula D.Y. Hong (1 sp.) and subg.
Pseudocodonopsis Kom. (6 spp.).
4. Cyclocodon Griff.
Cyclocodon Griff., Not. Pl. Asiat. 4: 277 (1854); Hong & Pan,
Acta Phytotax. Sin. 36: 106–110 (1998), rev.
Perennial herbs; root large, tuberous. Leaves
opposite (rarely alternate), petiolate. Flowers small
to medium-sized, erect, pedicellate, in a 3-flowered
terminal cyme and solitary in leaf axils. Calyx lobes
4 or 6, subtending hypanthium. Corolla white,
pale pink or lilac, campanulate or infundibular;
lobes 4 or 6, equalling tube. Stamens 4 or 6;
filaments distinct; anthers coherent, equalling
filaments; pollen 3-colporate, spinulose. Ovary
4- or 6-locular. Fruit a berry subtended by the
persistent calyx lobes. Two species, eastern Asia.
5. Echinocodon D.Y. Hong
Echinocodon D.Y. Hong, Acta Phytotax. Sin. 22: 183 (1984).
Small perennial herb. Leaves petiolate, pinnately
lobed or parted. Flowers very small, pedicellate,
solitary or in 2–3-flowered cymes in leaf axils.
Calyx lobes (2–)4(–5), longer than corolla, spinosemargined. Corolla purple-blue, cupular; lobes
(3–)4(–5), equalling tube. Stamens 3–5; filaments
distinct, dilated at base; anthers connivent, slightly
shorter than filaments; pollen 4–5-colpate, spinulose. Ovary 3–5-locular; stigmas filiform. Fruit
capsular, loculicidal; seeds triquetrous. 2n = 16.
One species, E. lobophyllus D.Y. Hong, eastern Asia.
6. Ostrowskia Regel
Ostrowskia Regel, Trudy Imp. S.-Peterburgsk. Bot. Sada 8:
686 (1884).
Perennial herb; roots tuberous. Leaves whorled.
Flowers large, pedicellate, in a terminal pyramidal
panicle. Calyx lobes (5–)7(–9). Corolla large, bowlshaped or funnelform, pale lilac or white; lobes
(5–)7(–9), shorter than tube. Stamens (5–)7(–9);
filaments dilated basally; anthers 3 times longer
than filaments, apically mucronate; pollen 6–7colpate, verrucose. Ovary (5–)7(–9)-locular. Fruit
capsular, laterally dehiscent by several oblong,
medial or subapical parallel slits; seeds narrowly
winged. 2n = 34. One species, O. magnifica Regel,
central Asia.
7. Canarina L.
Canarina L., Mant. Pl. 148, 588 (1771), nom. cons.; Hedberg,
Svensk Bot. Tidskr. 55: 17–62 (1961), rev.
Scandent (rarely epiphytic) sympodially branched
perennials, climbing with twining petioles and
pedicels; root tuberous. Leaves opposite or ternate,
petiolate; base cordate or hastate. Flowers large,
pendent, pedicellate, solitary, terminal in sympodial forks or on branches. Calyx 6-lobed. Corolla
orange streaked red or yellowish purple, campanulate or funnelform; lobes 6, shorter than tube.
Stamens 6; anthers distinct; pollen 3(–4)-colporate
or -colporoidate, spinulose. Fruit a yellow, pale red
or black berry, crowned by the persistent calyx
lobes. 2n = 34. Three species, one endemic to the
Canary Islands, the other two to eastern Africa.
8. Wahlenbergia Schrad. ex Roth
Fig. 5
Wahlenbergia Schrad. ex Roth, Nov. Pl. Sp. 399 (1821), nom.
cons.; Adamson, J. S. African Bot. 21: 155–218 (1955), part.
reg. rev.; Thulin, Symb. Bot. Upsal. 21: 1–223 (1975), reg.
rev.; Smith, Telopea 5: 91–175 (1992), reg. rev.; Petterson,
N. Z. J. Bot. 35: 9–54 (1997), reg. rev.
Lightfootia L’Hér. (1789), nom. illegit.
Cephalostigma A. DC. (1830).
Annual or perennial herbs, subshrubs and shrubs.
Leaves sometimes opposite, usually sessile, cauline
and often basally rosulate, rarely ericoid. Flowers
small to medium-sized, sessile or pedicellate,
terminal or axillary, solitary or in thyrses, panicles
or fascicles, very rarely involucrate. Calyx lobes
(3–)5(–6). Corolla campanulate or infundibular
(rarely tubular, urceolate, or rotate), blue or white
(rarely rose); lobes (3–)5(–7), much shorter to
much longer than tube. Stamens (3–)5; filaments
free and distinct, dilated basally; anthers coherent;
pollen (2–)3(–5)-porate, spinulose. Ovary 2–5locular, rarely superior; ovules few to many. Fruit
capsular, loculicidal, apically dehiscent by 2–5
valves. 2n = 14, 16, 18, 22, 36, 42, 54, 72, 90. Two
hundred and sixty species, circumaustral.
9. Nesocodon Thulin
Nesocodon Thulin, Kew Bull. 34: 813 (1980).
Shrub. Leaves aggregated towards stem tips.
Flowers large, pedicellate, pendent, solitary, axil-
Campanulaceae
39
a 5-notched nectar disc. Fruit capsular, loculicidal,
apically dehiscent by 3 valves, each with a pair of
pores, later forming six basal pores. Three species,
Mascarene Islands.
11. Berenice L.R. Tulasne
Berenice L.R. Tulasne, Ann. Sci. Nat., Bot. IV, 8: 156 (1857);
Badré et al., Adansonia II, 15: 139–146 (1975), rev.
Delicate shrub. Leaves cauline, petiolate. Flowers
very small, pedicillate, in terminal and axillary
open panicles. Corolla white, tipped with rose,
rotate; lobes longer than tube, recurved. Stamens
exserted; anthers free, dorsifixed, about half as
long as filament; pollen 3-porate, spinulose. Ovary
3-locular, topped by a nectar disc. Fruit capsular,
loculicidal, apically dehiscent by 3 valves. One sp.,
B. arguta L.R. Tulasne, Mascarene Islands.
12. Theilera E. Phillips
Theilera E. Phillips, Gen. S. African Fl. Pl. 606 (1926).
Fig. 5. Campanulaceae-Campanuloideae. Wahlenbergia
virgata. A Habit. B Flower (two petals and stamens
removed). C Stamen. D, E Closed and open capsule. F
Seeds. (Thulin 1976)
lary. Corolla campanulate, pale blue with darker
veins; lobes much shorter than tube. Stamens
distinct, included, appressed to corolla; anthers
a little shorter than filaments. Ovary 3-locular,
topped by 5 scarlet ellipsoid nectaries alternating
with stamens. Fruit capsular, loculicidal, apically
dehiscent by 3 valves. 2n = 34. One species, N.
mauritianus (I. Richardson) Thulin, Mauritius.
Subshrub. Leaves fascicled. Flowers solitary,
axillary, sessile. Corolla cylindric; lobes shorter
than tube. Stamens included; filaments filiform,
free from corolla; anthers shorter than filaments.
Ovary 3(–4)-locular; ovules numerous, basal; style
exserted. Fruit capsular, apically dehiscent by 3
valves. Two species, South Africa.
13. Namacodon Thulin
Namacodon Thulin, Bot. Notiser 127: 173 (1974).
Subshrub. Leaves sessile. Flowers small to mediumsized, terminal, solitary. Hypanthium elongate.
Corolla campanulate, blue; lobes as long as tube.
Stamens free; filament bases dilated; anthers
apically mucronate; pollen shed in tetrads. Ovary
3-locular; style included. Fruit capsular, septicidal,
dehiscing apically by 3 longitudinal fissures. One
sp., N. schinzianum (Markgr.) Thulin, South Africa.
14. Craterocapsa Hilliard & B.L. Burtt
10. Heterochaenia A. DC.
Heterochaenia A. DC. in Meisn., Pl. Vasc. Gen. 2: 149
(1839); Badré et al., Adansonia II, 12: 267–278 (1972), rev.
Shrubs. Leaves sessile, clustered at branch tips.
Flowers large, pedicellate, in a terminal raceme
or panicle. Corolla campanulate, violet, blue or
yellow-white; lobes equalling or shorter than tube.
Filaments distinct, the bases slender; anthers
equalling filaments. Ovary 3-locular, topped by
Craterocapsa Hilliard & B.L. Burtt, Notes Roy. Bot. Gard.
Edinburgh 32: 314 (1973).
Perennial herbs. Stems prostrate, often matted.
Leaves occasionally opposite, petiolate or sessile.
Flowers small to medium-sized, solitary (rarely
a few together in leaf axils), terminal. Corolla
lilac or white, funnelform; lobes slightly shorter
than tube. Filaments dilated at base, free; anthers
distinct. Ovary (2–)3-locular; style swollen at base
40
T.G. Lammers
or surrounded by a disc. Fruit capsular, dehiscent
by a terminal operculum formed from the enlarged
style base (or disc) and apex of ovary. Five species,
South Africa.
15. Roella L.
Roella L., Sp. Pl. 170 (1753); Adamson, J. S. African Bot. 17:
93–166 (1952), rev.
Subshrubs (rarely herbs). Leaves somewhat ericoid, crowded, often in axillary fascicles. Flowers
small to medium-sized, sessile, terminal, solitary
or in 2–5-flowered involucrate heads; bracts
sometimes pinnately spinose-lobed. Calyx lobes
4–5. Corolla campanulate; lobes 4–5. Stamens
4–5, included or slightly exserted; filaments
dilated basally, free from corolla; pollen 3-porate,
spinulose. Ovary 2-locular, crowned by an annular
or tumid nectary; ovules numerous; style sometimes with 2 glands below stigma. Fruit capsular,
dehiscent by an apical pore or operculum. Twenty
species, South Africa, divided by Adamson (1952)
into five series: ser. Roella (6 spp.), ser. Prostratae
Adamson (6 spp.), ser. Spicatae Adamson (4
spp.), ser. Squarrosae Adamson (3 spp.) and ser.
Muscosae Adamson (1 sp.).
16. Gunillaea Thulin
Gunillaea Thulin, Bot. Notiser 127: 166 (1974).
Annuals. Leaves sessile. Flowers small, pedicellate,
in more or less frondose monochasia. Calyx lobes
(3–)4–5. Corolla campanulate; lobes (3–)4–5. Stamens (3–)4–5, free. Ovary 2-locular; style included.
Fruit capsular, dehiscing irregularly between the
veins. 2n = 18. Two species, tropical Africa.
17. Prismatocarpus L’Hér.
Prismatocarpus L’Hér., Sert. Angl. 1 (1789), nom. cons.;
Adamson, J. S. African Bot. 17: 93–166 (1952), rev.
Subshrubs or perennial herbs. Leaves small, often
ericoid. Flowers small, terminal or axillary, solitary
or in clusters or sympodial panicles. Hypanthium
elongate. Corolla campanulate, funnelform, subcylindric, or rarely hypocrateriform, violet, blue,
pink or white; tube increasing in diameter towards
mouth. Stamens included or slightly exserted; filaments long and filiform or short and basally dilated,
free from corolla; pollen 3-porate, spinulose or verrucose. Ovary 2-locular, often topped by a glandular or tumid nectar disc; style included or exserted;
stigma rarely cylindric, sometimes subtended by
2 glands. Fruit capsular, splitting basipetally into
5 antipetalous segments. Twenty-nine species,
South Africa, divided by Adamson (1952) into two
subgenera: subg. Prismatocarpus (25 spp.) with
ser. Prismatocarpus (9 spp.), ser. Stricti Adamson
(7 spp.) and ser. Nitidi Adamson (9 spp.), and subg.
Afrotrachelium Adamson (4 spp.).
18. Treichelia Vatke
Treichelia Vatke, Linnaea 38: 700 (1874).
Dwarf annual. Flowers in dense terminal heads,
subtended by long linear bracts. Corolla cylindric
above a globose base, white or pale blue fading
yellow. Stamens included; filaments dilated basally,
free from corolla. Ovary 2-locular; ovules several;
style included, with a swollen subglobose base.
Fruit capsular, laterally dehiscent by medial
slits, covered by the style base. One species, T.
longibracteata (H. Buek) Vatke, South Africa.
19. Microcodon A. DC.
Microcodon A. DC., Monogr. Campan. 127 (1830).
Small annuals. Leaves sometimes subopposite.
Flowers small, in terminal corymbs or heads. Calyx
lobes foliose, enlarging in fruit. Corolla cylindric;
lobes shorter than tube. Stamens included; filaments linear, adnate to base of corolla; anthers
longer than filaments; pollen 3-porate, spinulose.
Ovary 5-locular, the locules antipetalous; style
included. Fruit capsular, loculicidal, apically
dehiscent by 5 valves. Four species, South Africa.
20. Merciera A. DC.
Merciera A. DC., Monogr. Campan. 369 (1830); Adamson, J.
S. African Bot. 20: 157–163 (1955), rev.; Cupido, Adansonia
III, 25: 33–44 (2003), rev.
Dwarf subshrubs or suffruticose perennials.
Leaves somewhat ericoid, dense, often fascicled.
Flowers small to medium-sized, solitary, axillary,
short-pedicellate or sessile, subtended by a pair
of bracteoles. Calyx lobes 4–5. Corolla tube long,
slender at base, gradually increasing in diameter
towards mouth; lobes 4–5, shorter than tube, sometimes unequal. Stamens 4–5, included; filaments
filiform, free from corolla; pollen 3-porate, spinulose. Ovary 1-locular or imperfectly 2-locular;
ovules 4, basal; style exserted; stigma bilobed.
Fruit dry, indehiscent, 1–2-seeded, crowned by the
persistent calyx lobes. Six species, South Africa.
Campanulaceae
21. Siphocodon Turcz.
Siphocodon Turcz., Bull. Soc. Imp. Naturalistes Moscou 25:
175 (1852).
Subshrubs. Leaves subulate, scale-like, sparse.
Flowers small, in few-flowered racemes or racemose panicles. Corolla cylindric; lobes shorter
than the tube. Stamens 5, epipetalous, included;
anthers almost equalling filaments. Ovary 2–3locular; ovules 2–5 in each locule, apical. Fruit
capsular, circumscissile, operculum crowned by
calyx lobes. Two species, South Africa.
22. Rhigiophyllum Hochst.
Rhigiophyllum Hochst., Flora 25: 232 (1842).
Subshrub. Leaves small, squarrose, densely imbricate in 4 rows. Flowers small, in terminal heads,
subtended by rigid leaf-life bracts. Corolla cylindric. Stamens 5, epipetalous, included; anthers
longer than filaments. Ovary 3-locular; ovules
several, apical; style exserted. Fruit capsular, circumscissile, operculum crowned by the persistent
style base. One species, R. squarrosum Hochst.,
South Africa.
23. Edraianthus A. DC.
Edraianthus A. DC. in Meisn., Pl. Vasc. Gen. 2: 149 (1839),
nom. cons.; Kuzmanov in Tutin et al., Fl. Eur. 4: 99–100
(1976), reg. rev.
Halacsyella Janch. (1910).
Protoedraianthus Lakušić (1988).
Caespitose perennial herbs. Flowers mediumsized, solitary or in capitula, terminal. Corolla
campanulate or infundibular, blue; lobes shorter
than tube. Stamens free; filament bases dilated;
pollen 3-porate. Ovary (2)3-locular; style included;
stigma lobes linear. Fruit capsular, dehiscing irregularly at apex. 2n = 32. Thirteen species, southeastern Europe.
24. Feeria Buser
Feeria Buser, Bull. Herb. Boissier 2: 517 (1894).
Suffruticose perennial. Leaves cauline, subsessile.
Flowers small, in a terminal flat-topped corymbose
cyme. Corolla infundibular, white with blue limb;
lobes shorter than corolla. Stamens included;
filaments slender; anthers shorter than filaments,
apiculate on connective, the inner pollen sacs
shorter than outer. Ovary 3-locular, crowned by
a narrow annular nectary; style exserted, swollen
41
towards apex. Fruit capsular, dehiscing apically
by 3 valves; seeds ca. 10. 2n = 34. One species, F.
angustifolia (Schousb.) Buser, Morocco.
25. Jasione L.
Jasione L., Sp. Pl. 928 (1753); Tutin in Tutin et al., Fl. Eur.
4: 100–102 (1976), reg. rev.; Parnell, Watsonia 16: 249–267
(1987), part. rev.
Jasionella Stoj. & Stef. (1933).
Annual, biennial or perennial herbs. Flowers
small, sessile or short-pedicellate, in terminal
involucrate heads. Corolla rotate, blue (rarely
white), split nearly to base. Filaments slender;
anthers connate at base, longer than filaments;
pollen 3-porate, spinulose. Ovary 2-locular. Fruit
capsular, loculicidal, dehiscing apically by 2 valves.
2n = 12, 14, 18, 24, 36, 48, 60. Sixteen species,
Mediterranean, except one in central Europe.
26. Musschia Dumort.
Musschia Dumort., Comment. Bot.: 28 (1822).
Long-lived monocarpic rosette shrub or polycarpic
suffruticose perennial. Flowers large, pedicellate,
in many-flowered pyramidal panicle. Calyx lobes
flushed with same colour as corolla. Corolla rotate,
bright yellow or reddish brown; lobes equalling
or longer than tube. Pollen 3–4(–8)-porate. Fruit
capsular, laterally dehiscent by numerous slits.
2n = 32. Two species, Madeira.
27. Azorina Feer
Azorina Feer, Bot. Jahrb. Syst. 12: 611 (1890).
Viscid rosette shrub. Leaves sessile. Flowers large,
pendulous, pedicellate, in lax panicles which arise
below the apical rosette. Corolla white or rose,
campanulate or urceolate; lobes shorter than the
tube. Stamens included; filaments dilated basally,
forming a nectar chamber; anthers shorter than
filaments; pollen 3–4-porate. Ovary (2–)3-locular,
topped by a green nectar disc with a thickened
orange rim; style included. Fruit capsular, laterally
dehiscent by (2)3 valves. 2n = 56. One sp., A.
vidalii (H. C. Watson) Feer, Azores.
28. Campanula L.
Fig. 6
Campanula L., Sp. Pl. 163 (1753); Fedorov, Fl. S.S.S.R.
24: 133–331 (1957), reg. rev.; Phitos, Oesterr. Bot. Z. 111:
208–230 (1964), part. rev.; Phitos, Oesterr. Bot. Z. 112:
449–498 (1965), part. rev.; Rechinger & Schimann-Czeika,
Fl. Iran. 13: 7–38 (1965), reg. rev.; Charadze, Zametki Sist.
Geogr. Rast. 32: 46–56 (1976), reg. rev.; Fedorov & Kovanda
42
T.G. Lammers
in Tutin et al., Fl. Eur. 4: 74–93 (1976), reg. rev.; Damboldt,
Fl. Turk. 6: 2–64 (1978), reg. rev.; Hong, Fl. Reip. Pop. Sin.
73: 78–92 (1983), reg. rev.; Carlström, Willdenowia 15:
375–387 (1986), part. rev.; Oganesian, Bot. Zhurn. (Moscow
& Leningrad) 78, 3: 145–157 (1993), sect. rev.; Sáez &
Aldasoro, Bot. J. Linn. Soc. 141: 215–241 (2003), subg. rev.
Roucela Dumort. (1822).
Symphyandra A. DC. (1830).
Diosphaera Buser (1894).
Tracheliopsis Buser (1894).
Campanulastrum Small (1903).
Brachycodon Fed. (1957), nom. illegit.
Astrocodon Fed. (1957).
Popoviocodonia Fed. (1957).
Brachycodonia Fed. (1961).
Annaea Kolak. (1979).
Gadellia Shulkina (1979).
Pseudocampanula Kolak. (1980).
Mzymtella Kolak. (1981).
Hyssaria Kolak. (1981).
Neocodon Kolak. & Serdyuk. (1982).
Hemisphaera Kolak. (1984).
Echinocodon Kolak. (1986), nom. illegit.
Echinocodonia Kolak. (1994).
Annual, biennial or perennial herbs, often with
thickened rootstocks or rhizomes. Leaves cauline
and often basally rosulate. Flowers small to large,
pendulous, erect, or horizontal, pedicillate, solitary or in various racemose, spicate, paniculate or
rarely capitulate inflorescences, rarely involucrate,
rarely cleistogamous. Calyx lobes 5, sometimes
with reflexed appendages in the sinuses. Corolla
campanulate, tubular, funnelform, bowl-shaped,
or rotate, violet, blue or white (rarely yellow, red
or pink); lobes shorter than tube to much longer
than tube. Stamens commonly included; filaments
dilated basally; anthers rarely connate; pollen
3(–6)-zonoporate or rarely 6–18-pantoporate.
Ovary (2–)3–5-locular; style straight or, if curved
towards apex, then exserted; nectar disc absent.
Fruit capsular, erect or pendent, laterally dehiscent
by (2–)3–5 basal, medial, or subapical pores or
valves; seeds small, numerous. 2n = 14, 16, 18, 20,
22, 24, 26, 28, 30, 32, 34, 36, 40, 46, 48, 50, 52, 54,
56, 58, 68, 70, 72, 80, 84, 90, 102. Four hundred and
twenty-one species, circumboreal in distribution,
extending as far south as eastern Africa, southern
Asia and northern Mexico; many species cultivated
and some naturalized outside their indigenous
range. Though many infrageneric taxa have been
proposed, there is no modern classification which
accounts for this large genus in its entirety.
Fig. 6.
Campanulaceae-Campanuloideae. Campanula
edulis. A Habit. B Flower (two petals removed). C Stamen.
D Flower (corolla and stamens removed). E Open capsule.
F Seed. (Thulin 1976)
29. Favratia Feer
Favratia Feer, Bot. Jahrb. Syst. 12: 610 (1890).
Perennial herb. Leaves cauline and basally rosulate. Flowers small, erect or horizontal, pedicillate, solitary, terminal or axillary. Calyx lobes 5,
unappendaged. Corolla urceolate, pale blue; lobes
much shorter than tube, each sinus with a large
conspicuous saccate appendage. Stamens included;
filaments dilated basally. Ovary 3-locular; style included, curved towards apex; nectar disc absent.
Fruit capsular, suberect, laterally dehiscent by 3
subapical pores; seeds small, numerous. 2n = 34.
One sp., F. zoysii (Jacq.) Feer, south-eastern Alps.
30. Zeugandra P.H. Davis
Zeugandra P.H. Davis, Hooker’s Icon. Pl. 35: pl. 3497 (1950).
Perennial herbs. Flowers medium-sized, shortpedicellate, pendent in lax panicles. Calyx lobes
Campanulaceae
alternating with recurved liguliform appendages.
Corolla narrowly infundibular, blue; lobes shorter
than tube. Filaments connate for half their length
or more, dilated basally; anthers connate. Ovary
3-locular; style exserted. Fruit capsular, laterally
dehiscent by 3 basal valves. Two species, Iran.
31. Adenophora Fisch.
Adenophora Fisch., Mém. Soc. Imp. Naturalistes Moscou 6:
165 (1823); Hong, Fl. Reip. Pop. Sin. 73, 2: 92–139 (1983),
reg. rev.
Perennial herbs, often with large tuberous roots.
Leaves cauline, rarely opposite or 3–6-verticillate.
Flowers small to medium-sized, pendulous, shortpedicillate, in loose racemes or panicles. Corolla
purple or blue (rarely white), campanulate, tubular, infundibular, or rarely urceolate; lobes shorter
than the tube. Stamens commonly included; filaments dilated basally, forming a nectar chamber;
anthers equalling or shorter than filaments; pollen
3–5-porate, echinate. Ovary 3-locular, crowned by
a large, thick, annular cylindric or cupulate nectary; style exserted. Fruit capsular, laterally dehiscent by 3 pores or valves. 2n = 34, 36, 68, 72, 102.
Sixty-five species, eastern Asia, one in western Europe and one in Crimea. Though many infrageneric
taxa have been proposed, no modern classification
accounts for this large genus in its entirety.
32. Hanabusaya Nakai
Hanabusaya Nakai, J. Coll. Sci. Imp. Univ. Tokyo 31: 62
(1911).
Keumkangsania Kim (1976), nom. illegit.
Perennial herb with a branched rhizome. Leaves
cauline, petiolate, 4–6 at middle of stem. Flowers large, pedicellate, pendent, solitary or in fewflowered racemes, terminal. Corolla campanulate,
light purple; lobes only 1/20 as long as tube. Stamens included; filaments dilated basally; anthers
about as long as filaments, connate; pollen 4–7porate, echinate. Ovary 3-locular, topped by a large
conic or hemispheric nectary. 2n = 34. One sp., H.
asiatica (Nakai) Nakai, Korea.
33. Theodorovia Kolak.
Theodorovia Kolak., Bot. Zhurn. (Moscow & Leningrad)
70: 7 (1985).
Fedorovia Kolak. (1980), nom. illegit.
Perennial herb with a thickened rootstocks. Leaves
apically rosulate. Flowers small, pedicillate, solitary, terminal. Calyx lobes 5, with small reflexed
43
appendages in the sinuses. Corolla tubular, dark
blue; lobes much shorter than tube. Stamens included; filaments dilated basally. Ovary 3-locular.
Fruit schizocarpous, the locules forming 3 mericarps which separate from the hypanthium; the 3
mericarps remain connected by the style which
splits into 3 basally. One sp., T. karakuschensis
(Grossh.) Kolak., Caucasus.
34. Muehlbergella Feer
Muehlbergella Feer, Bot. Jahrb. Syst. 12: 615 (1890).
Perennial herb from a thick rhizome. Leaves small,
sessile, dense. Flowers small, sessile, solitary, terminal. Hypanthium narrowly obconic. Calyx lobes
recurved and appressed to hypanthium after anthesis. Corolla tubular, blue, becoming scarious and
hyaline, persistent; lobes equalling or shorter than
tube. Filaments dilated basally. Ovary 3-locular;
style included. Fruit capsular, splitting from tip
to base into irregular fragments. One species, M.
oweriniana (Rupr.) Feer, Caucasus.
35. Sachokiella Kolak.
Sachokiella Kolak., Soob. Akad. Nauk Gruzinsk. S.S.R. 118:
595 (1985).
Biennial herb with thick fusiform root. Leaves
cauline. Flowers small, pedicillate, in an involucrate head. Corolla tubular, mauve or blue; lobes
shorter than tube. Filaments somewhat dilated
basally. Ovary 3 locular. Fruit capsular, splitting
laterally from base into 5 elongate segments; seeds
very large, few, spongy verrucose, erose winged.
One sp., S. macrochlamys (Boiss. & A. Huet) Kolak.,
Caucasus.
36. Peracarpa Hook. f. & Thomson
Peracarpa Hook. f. & Thomson, J. Proc. Linn. Soc., Bot. 2:
26 (1858); Barnesky & Lammers, Bot. Bull. Acad. Sin. 38:
49–56 (1997), rev.
Delicate stoloniferous herb. Leaves cauline, petiolate. Flowers very small, pedicellate, terminal or less
often axillary, solitary (rarely 2–17 in a fascicle).
Corolla funnelform, pale blue or white; lobes about
equalling tube. Stamens distinct, included; anthers
half as long as filaments; pollen 4–6-porate, spinulose. Ovary (2–)3-locular, topped by a fleshy trisulcate nectar disc; ovules 4–5 pairs per placenta; style
included; stigmas filiform. Fruit capsular, pendent,
rupturing irregularly at the base; seeds 10–16, large.
2n = 28, 30. One species, P. carnosa (Wall.) Hook.
f. & Thomson, eastern Asia.
44
T.G. Lammers
37. Homocodon D.Y. Hong
Homocodon D.Y. Hong, Acta Phytotax. Sin. 18: 473 (1980).
Annuals. Stems 3-winged. Leaves cauline, petiolate.
Flowers very small, sessile, solitary or paired, axillary. Calyx lobes spinose-toothed. Corolla campanulate, white or pale bluish; lobes equalling tube. Stamens included, distinct; filaments dilated at base;
anthers shorter than filaments; pollen 3–4-porate,
spinulose. Ovary 3-locular; style slightly exserted;
stigmatic lobes linear. Fruit capsular, dehiscent by
pores or by irregular tears. Two species, southern
China.
38. Legousia Durande
Legousia Durande, Fl. Bourgogne 1: 37 (1782); Tutin in
Tutin et al., Fl. Eur. 4: 94 (1976), reg. rev.
Specularia Heist. ex A. DC. (1830), nom. illegit.
Annuals. Leaves cauline, sessile or petiolate.
Flowers small to medium-sized, sessile or
short-pedicellate, in spike-like inflorescences.
Hypanthium elongate. Corolla rotate or broadly
campanulate, violet, blue, pinkish or white.
Stamens 5, distinct; filaments slender; anthers
much longer than filaments; pollen 3–5-porate,
spinulose. Ovary 3-locular with axile placentae
(rarely 1-locular with parietal placentae); stigmatic
lobes filiform. Fruit capsular, dehiscing laterally
by 3 medial or subapical valves. 2n = 20, 26. Seven
species, Mediterranean.
39. Triodanis Raf.
Triodanis Raf., New Fl. N. Am. 4: 67 (1838); McVaugh,
Wrightia 1: 13–52 (1945), rev.
Annuals. Leaves sessile or short-petiolate. Flowers medium-sized, sessile, 1–3(–8) in upper axils, forming a spike-like inflorescence, lower ones
smaller and cleistogamous. Hypanthium elongate.
Calyx lobes 3 in cleistogamous flowers. Corolla
lavender blue, rotate; lobes longer than tube. Stamens 5, distinct, free from corolla; filaments dilated basally; anthers longer than filaments. Ovary
inferior, 3-locular with axile placentae (rarely 1locular with parietal placentae). Fruit capsular, laterally dehiscent by 3 apical, medial or basal pores.
2n = 28, 56, 60. Six species, North America.
40. Heterocodon Nutt.
Heterocodon Nutt., Trans. Amer. Philos. Soc. n.s. 8: 255
(1842).
Annual. Leaves cauline, sessile. Flowers very small,
subsessile, solitary, opposite the upper leaves, the
lower cleistogamous. Corolla cylindric, white with
blue limb; lobes shorter than tube. Ovary 3-locular.
Fruit capsular, dehiscent by irregular basal pores.
One species, H. rariflorum Nutt., western North
America.
41. Githopsis Nutt.
Githopsis Nutt., Trans. Amer. Philos. Soc. n.s. 8: 258 (1842);
Morin, Syst. Bot. 8: 436–468 (1983), rev.
Annuals. Leaves cauline, sessile. Flowers small
to medium-sized, terminal, sessile or shortpedicellate, sometimes cleistogamous. Corolla
cylindric, campanulate, or funnelform, white
with purple, blue or white lobes; lobes equalling
or longer than tube. Stamens included; pollen
6–8-porate, spinulose. Ovary (2–)3-locular; placentation axile; style 2–3-lobed. Fruit capsular,
dehiscing by 1 irregular apical pore. 2n = 18,
20, 36, 38, 40. Four species, western North
America.
42. Asyneuma Griseb. & Schenk.
Asyneuma Griseb. & Schenk., Arch. Naturgesch. 18: 335
(1852); Damboldt, Boissiera 17: 1–128 (1970), rev.
Asyneumopsis Contandr., Quézel & Pamukç. (1972).
Annual, biennial or perennial herbs, with a thin
rhizome or fusiform root. Leaves cauline and/or
rosulate, subsessile. Flowers small to mediumsized, sessile, in a simple or branched spike or
dense cylindric heads. Corolla rotate, purple,
blue or white, divided nearly to base. Filaments
dilated basally; pollen 4–5-porate, spinulose.
Ovary (2–)3(–4)-locular; stigma lobes linear. Fruit
capsular, dehiscent laterally by (2–)3 medial to
subapical pores. 2n = 20, 22, 24, 28, 30, 32, 34, 48,
56, 68. Thirty-three species, Asia, eastern Europe,
northern Africa.
43. Cryptocodon Fed.
Cryptocodon Fed. in Kom., Fl. S.S.S.R. 24: 474 (1957).
Perennial herb with large tuberous roots. Flowers medium-sized, subsessile, in a terminal involucrate capitulum. Calyx lobes with small recurved
appendages in sinuses. Corolla blue, tubular, split
nearly its entire length. Ovary 3-locular; style included. Fruit capsular, laterally dehiscent by 3 medial pores. One sp., C. monocephalus (Trautv.) Fed.,
central Asia.
Campanulaceae
45
44. Petromarula Vent. ex R. Hedw.
48. Physoplexis (Endl.) Schur
Petromarula Vent. ex R. Hedw., Gen. Pl. 139 (1806).
Physoplexis (Endl.) Schur, Sert. Fl. Transsilv. 47 (1853).
Synotoma (G. Don) Rich. Schulz (1904), nom. illegit.
Perennial herb. Leaves petiolate, pinnate or pinnatisect, segments dentate or lobed. Flowers small,
in panicles. Corolla pale blue, infundibular, split
nearly to base. Filaments free, distinct, dilated
basally. Ovary 3-locular; style exserted; stigma
large, capitate. Fruit capsular, laterally dehiscent by
3 medial pores. 2n = 30. One species, Petromarula
pinnata (L.) A. DC., Crete.
45. Cylindrocarpa Regel
Cylindrocarpa Regel, Trudy Imp. S.-Peterburgsk. Bot. Sada
5: 258 (1877).
Fig. 7
Perennial herb. Leaves cauline and basal, petiolate. Flowers medium-sized, pedicellate, in an 8–20flowered terminal umbel. Corolla tubular, pinkish
lilac with dark violet apex; lobes much longer than
tube but connate in the apical third and bowed
out medially, forming five prominent fenestrations.
Pollen 4-porate, spinulose. Ovary 2-locular. Fruit
capsular, laterally dehiscent by 2 medial pores. 2n =
34, 68. One species, Physoplexis comosa (L.) Schur,
south-eastern Europe.
Perennial herb. Flowers small to medium-sized,
sessile, solitary and terminal. Hypanthium cylindric. Corolla blue, split nearly its entire length.
Stamens included; anthers much longer than filaments. Ovary 2–3-locular; style included; stigmatic
lobes filiform. Fruit capsular, laterally dehiscent
by 2–3 subapical slits. One species, C. sewerzowii
(Regel & Herder) Regel, central Asia.
46. Sergia Fed.
Sergia Fed. in Kom., Fl. S.S.S.R. 24: 474 (1957).
Perennial herbs with an enlarged woody caudex.
Leaves cauline, sessile or petiolate. Flowers small,
pedicellate, terminal. Corolla blue, campanulate or
funnelform, split for most its length. Stamens 4;
anthers much longer than filaments, appearing sessile. Ovary 3-locular. Fruit capsular, laterally dehiscent by 3 basal valves. Two species, central Asia.
47. Phyteuma L.
Phyteuma L., Sp. Pl. 170 (1753); Damboldt in Tutin et al.,
Fl. Eur. 4: 95–98 (1976), reg. rev.
Perennial herbs from a fleshy rootstock. Flowers
small to medium-sized, sessile or subsessile, in
terminal spicate or subglobular involucrate heads.
Calyx lobes 4–5. Corolla tubular, blue or lilac
(rarely yellow); lobes 4–5, much longer than tube
but coherent in the apical third and bowed out
medially, forming five prominent fenestrations,
eventually separating and spreading. Filaments
dilated basally; pollen 4-porate, spinulose. Ovary
2–3-locular. Fruit capsular, laterally dehiscent by
2–3 medial pores. 2n = 16, 18, 20, 22, 24, 26, 28.
Twenty-two species, Europe.
Fig. 7.
Campanulaceae-Campanuloideae. Physoplexis
comosa. A Habit. B Flower (corolla removed). C Flower
(corolla removed, male phase). D Flower (corolla removed,
female phase). (Schönland 1889)
46
T.G. Lammers
49. Michauxia L’Hér.
Michauxia L’Hér., Michauxia (1788), nom. cons.; Rechinger
& Schimann-Czeika, Fl. Iran. 13: 47–49 (1965), reg. rev.;
Damboldt, Fl. Turk. 6: 81–83 (1978), reg. rev.
Robust biennials. Leaves cauline and basally
rosulate, pinnately lobed or cleft. Flowers large,
subsessile or short-pedicellate, in spicate racemes
or panicles. Calyx lobes (6–)8–10, the sinuses
appendaged. Corolla white or pale pink, rotate,
split nearly to the base; lobes (6–)8–10, recurved.
Stamens (6–)8–10; filaments basally dilated;
anthers 3 times longer than filaments, apically mucronate or cuspidate; pollen 3-porate, spinulose.
Ovary (6–)8–10-locular; style exserted; stigmatic
lobes filiform. Fruit capsular, pendent, dehiscent
laterally by (6–)8–10 basal valves. 2n = 28, 30, 34.
Seven species, south-western Asia.
This genus has been called Mindium, though
the type of that name belongs to Canarina.
50. Trachelium L.
Trachelium L., Sp. Pl. 171 (1753).
Perennial herb. Leaves petiolate. Flowers small,
pedicellate, in a pedunculate corymb. Corolla
tubular, blue (rarely white); lobes much shorter
than tube. Style long-exserted. Fruit capsular,
laterally dehiscent by basal pores. 2n = 32, 34. Two
species, Mediterranean and often cultivated.
Nemacladoideae comprise 3 genera (2 monotypic) and 15 species endemic to the south-western
United States and adjacent Mexico.
51. Pseudonemacladus McVaugh
Pseudonemacladus McVaugh, N. Amer. Fl. 32A: 3 (1943).
Perennial herb. Leaves cauline, opposite or the
upper alternate, petiolate. Flowers very small, pedicellate, in a terminal pedunculate 5–30-flowered
raceme; pedicels subapically bibracteolate. Corolla
white or bluish; lobes dimorphic, the ventral larger.
Filaments connate in upper half, tube strongly recurved at apex, lacking any special glands or
appendages. Ovary half-inferior, 2-locular. Capsule loculicidal, dehiscent by 2 apical valves. One
species, P. oppositifolius (B.L. Rob.) McVaugh,
Mexico.
52. Nemacladus Nutt.
Fig. 8
Nemacladus Nutt., Trans. Amer. Philos. Soc. n.s. 8: 254
(1842); McVaugh, Amer. Midl. Nat. 22: 521–550 (1939), rev.;
Wimmer, Pflanzenr. IV.276c: 923–935 (1968), rev.; Morin &
Milburn in Hickman, Jepson Man.: 465–468 (1993), reg. rev.
II. Subfam. Nemacladoideae Lammers (1998).
Annual herbs (one species perennial). Inflorescence a terminal raceme (often comprising much
of the plant body) or rarely a series of 1–4 capitate
clusters. Flowers not resupinate. Calyx with the odd
(unpaired) lobe in a ventral (anterior) position.
Corolla bilaterally symmetric, with 3 dorsal and
2 ventral lobes. Stamens epigynous, the dorsal 2
commonly alternating with three flattened, round
glands; filaments connate above (rarely distinct),
the dorsal 2 (sometimes 4) commonly bearing
one or more transparent rod-like cells on a stipe
(or rarely sessile); anthers distinct, commonly
radiating at right angles to the filaments. Ovary
half-inferior (rarely nearly superior), 2-locular
with axile placentation (rarely 1-locular with
parietal placentation); stigma 2-lobed. Fruit
a loculicidal capsule, dehiscent by 2 apical valves
(these sometimes splitting longitudinally at apex)
or circumscissile. Seeds subglobose, ellipsoid, or
cylindric, smooth or variously pitted or striate.
Fig. 8.
Campanulaceae-Namacladoideae. Nemacladus
rigidus. A Habit. B Flower. C Capsule. D Seed. (Hickman
1993, with permission from the Jepson Herbarium.
© Regents of the University of California)
Campanulaceae
47
Annuals. Leaves rosulate, petiolate or sessile. Flowers very small, pedicellate, in a terminal raceme;
pedicels ebracteolate. Corolla white or bluish; lobes
monomorphic. Filaments distinct or connate in upper half, dorsal pair bearing one or more transparent rod-like cells on a stipe. Ovary half-inferior,
2-locular (rarely 1-locular). Capsule loculicidal, dehiscent by 2 apical valves, these sometimes splitting
longitudinally. 2n = 18. Thirteen species, western
North America.
54. Dialypetalum Benth.
53. Parishella A. Gray
55. Lobelia L.
Parishella A. Gray, Bot. Gaz. (Crawfordsville) 7: 94 (1882).
Lobelia L., Sp. Pl.: 929 (1753); McVaugh, N. Amer. Fl. 32A:
35–99 (1943), reg. rev.; Braga, Trib. Farm. 32: 1–8, 41–54
(1964), 33: 3–24 (1965), reg. rev.; Mabberley, Kew Bull. 29:
535–583 (1974), reg. sect. rev.; Ayers, Syst. Bot. 15: 296–327
(1990), part. rev.; Lammers in Wagner et al., Man. Fl. Pl.
Hawai’i: 473–480 (1990), sect. rev.; Wilbur, Sida 14: 555–567
(1991), reg. sect. rev.; Hong & Zhang, Acta Phytotax. Sin. 30:
146–162 (1992), reg. sect. rev.; Lammers, Sida 19: 87–110
(2000), sect. rev.; Lammers, Sida 21:591–623 (2004), sect.
rev.
Pratia Gaudich. (1825).
Hypsela C. Presl (1836).
Trimeris C. Presl (1836).
Galeatella (E. Wimm.) O. Deg. & I. Deg. (1962).
Neowimmeria O. Deg. & I. Deg. (1965).
Calcaratolobelia Wilbur (1997).
Annual. Leaves rosulate, petiolate. Flowers very
small, pedicellate, in a series of 1–3 terminal
capitula; pedicels ebracteolate. Corolla white;
lobes monomorphic. Filaments connate in upper
half, ventral 4 bearing sessile clusters of a few
transparent rod-like cells. Ovary half-inferior,
imperfectly 2-locular. Capsule circumscissile. One
species, P. californica A. Gray, western North
America.
III. Subfam. Lobelioideae Burnett (1835).
Herbaceous perennials (less often annual or
biennial), woody lianas (sometimes twining),
pachycaul rosette plants, subshrubs, shrubs,
treelets, or trees to 15 m tall, typically terrestrial,
rarely aquatic or epiphytic. Inflorescences commonly racemose, or flowers solitary in an axillary
(rarely terminal) position; flowers commonly
resupinate by torsion of the pedicel. Calyx with the
odd (unpaired) lobe in a ventral (anterior) position
prior to resupination. Corolla bilaterally symmetric, with two dorsal and three ventral lobes, or all
five lobes ventral. Stamens epigynous; filaments
connate, tube sometimes adnate to corolla tube;
anthers connate, forming an oblique tube. Ovary
inferior (rarely nearly superior), 2-locular with
axile placentation (rarely 1-locular with parietal or
basal placentation); stigma 2-lobed. Fruit a capsule
(commonly loculicidal) or berry.
Lobelioideae comprise 29 genera and 1,195 species. Though the subfamily is represented on all six
continents, two-thirds of the species are endemic to
the New World: South America harbours nearly half
and North America 17%. Another 14% are African,
12% Polynesian, 5% Australasian, 3% Asian and
less than 1% European.
Dialypetalum Benth. in Benth. & Hook. f., Gen. Pl. 2: 553
(1876).
Herbs and shrubs. Flowers small, in many-flowered
terminal panicles. Corolla actinomorphic, rotate,
split nearly to base, white, yellowish, greenish or
purplish. Filaments distinct or connate only at
apex; ventral 2 anthers with a bristle at apex. Fruit
capsular, dehiscent by 2 apical valves. Five species,
Madagascar.
Fig. 9
Annual or perennial herbs, shrubs, trees or giant
rosette plants (often pliestesial). Flowers small to
large, pedicellate, solitary and axillary or in terminal racemes (rarely secund or corymbose) or panicles. Calyx lobes rarely auriculate. Corolla unilabiate or bilabiate, blue, purple, red, rose, green, yellow or white, rarely with a narrow nectar spur; tube
dorsally cleft to base (rarely entire), sometimes fenestrate; lobes monomorphic (apices sometimes coherent) or dimorphic, the ventral larger. Filament
tube rarely adnate to corolla; ventral or all five anthers with apical tufts of stiff hairs, ventral sometimes with minute apical appendages, rarely all
nude. Ovary rarely almost superior. Plants rarely
dioecious. Fruit capsular, dehiscent apically by 2
valves, or a berry. 2n = 12, 14, 18, 24, 26, 28,
38, 42, 70, 140. Over 400 species, cosmopolitan,
divided by Murata (1995) into three subgenera:
subg. Lobelia (231 spp.), subg. Isolobus (A. DC.)
Y.S. Lian (48 spp.) and subg. Tupa (G. Don) E.
Wimm. (127 spp.). Subgenus Lobelia comprises
four sections: sect. Lobelia (22 spp.), sect. Heyneana
J. Murata (143 spp.), sect. Cryptostemon (E. Wimm.)
48
T.G. Lammers
J. Murata (9 spp.) and sect. Delostemon (E. Wimm.)
J. Murata (57 spp.). Isolobus comprises four sections: sect. Dioica (E. Wimm.) J. Murata (8 spp.),
sect. Pratia (Gaudich.) J. Murata (37 spp.), sect.
Paramezleria E. Wimm. (1 sp.) and sect. Isolobus
(A. DC.) C.B. Clarke (2 spp.). Tupa comprises seven
sections: sect. Tupa (G. Don) Benth. (4 spp.), sect.
Colensoa (Hook. f.) J. Murata (47 spp.), sect. Rhynchopetalum (Fresen.) Benth. (20 spp.), sect. Homochilus A. DC. (6 spp.), sect. Tylomium (C. Presl)
Benth. (37 spp.), sect. Revolutella E. Wimm. (9 spp.)
and sect. Galeatella E. Wimm. (4 spp.)
56. Solenopsis C. Presl
Solenopsis C. Presl, Prodr. Monogr. Lobel. 32 (1836); Crespo
et al., Pl. Syst. Evol. 210: 211–219 (1998), rev.
Annual or perennial herbs, sometimes scapose.
Leaves petiolate, sometimes basally rosulate.
Flowers pedicellate, solitary, axillary (appearing
terminal in scapose species); pedicels medially
1–3-bracteolate (rarely ebracteolate). Corolla
bilabiate, blue, sometimes white or yellow on lower
lip; tube entire; lobes subdimorphic, the ventral
slightly larger. Filament tube free from corolla;
ventral anthers aristate at apex, often also with tufts
of stiff hairs. Fruit capsular, dehiscent apically by
2 valves. Seeds ellipsoid to fusiform, strophiolate,
with weak longitudinal ridges, sulcate with keeled
walls. 2n = 22, 28. Six species, Mediterranean.
57. Wimmerella L. Serra, M.B. Crespo & Lammers
Wimmerella L. Serra, M.B. Crespo & Lammers, Novon 9:
415 (1999).
Annual or perennial herbs. Leaves petiolate or
sessile. Flowers pedicellate, solitary, axillary, or
in lax, often secund racemes; pedicels basally
bibracteolate or ebracteolate. Corolla bilabiate,
blue or white; tube entire; lobes dimorphic, the
ventral slightly larger, or monomorphic. Filament
tube free from corolla; ventral anthers aristate
at apex, often also with tufts of stiff hairs. Fruit
capsular, dehiscent apically by 2 valves. Seeds
ellipsoid to subglobose, estrophiolate, sulcate with
flattened walls. Ten species, South Africa.
58. Grammatotheca C. Presl
Grammatotheca C. Presl, Prodr. Monogr. Lobel. 43 (1836).
Repent herb. Flowers small, solitary, axillary, sessile, with a pair of bracteoles at base. Hypanthium
pedicelliform, twisted 180°. Corolla bilabiate, blueviolet with white eye; tube dorsally cleft for 2/3 its
length; lobes dimorphic, the ventral larger. All 5 anthers with apical tufts of stiff hairs. Fruit capsular,
dehiscent apically by 2 valves and also laterally by
irregular tears. One species, G. bergiana (Cham.)
C. Presl, South Africa and Australia.
59. Dielsantha E. Wimm.
Fig. 9. Campanulaceae-Lobelioideae. Lobelia welwitschii.
A Habit. B Flower. C Opened corolla. D Stamens and anther
tube. E Ovary, longitudinal section. F Stigma at different
developmental stages. G Capsule. H Seed. (Thulin 1984)
Dielsantha E. Wimm., Ann. Naturhist. Mus. Wien 56: 372
(1948).
Perennial herb. Flowers small, sessile, in fewflowered axillary fascicles. Hypanthium pedi-
Campanulaceae
49
celliform, twisted 180°. Corolla bilabiate, pale
violet; tube cleft to base; lobes dimorphic, the
ventral larger. Filament tube free from corolla; all
5 anthers with apical tufts of stiff hairs. Stigma
lobes filiform, 1/3 as long as style. Fruit capsular,
dehiscing by 5 or 6 irregular lateral ruptures. One
species, D. galeopsoides (Engl. & Diels) E. Wimm.,
western Africa.
hairs. Fruit capsular, dehiscent apically by 2 valves,
or a berry. 2n = 14, 28. Fourteen species, Australia,
New Zealand.
60. Monopsis Salisb.
Annuals. Flowers small, solitary; pedicels adnate
to the midrib of the subtending leaf. Hypanthium
dorsally oblique. Corolla bilabiate, green or yellow; tube dorsally cleft to base; lobes dimorphic,
the dorsal 1.5–3 times as long as the ventral but
more slender. Ventral anthers with minute apical
appendages and apical tufts of stiff hairs, sometimes all nude. Fruit capsular, dehiscent apically by
2 valves. Four species, New Guinea.
Monopsis Salisb., Trans. London Hort. Soc. 2: 37 (1817).
Annual or perennial herbs. Leaves sometimes opposite or whorled. Flowers small, solitary, axillary,
pedicellate or sessile and aggregated into a congested terminal raceme, commonly not resupinate;
pedicels ebracteolate or basally bibracteolate.
Corolla bilabiate with dimorphic lobes (the ventral
larger) or subrotate with monomorphic lobes,
blue, violet, yellow or orange; tube cleft dorsally to
base. Filament tube sometimes adnate to corolla
at base; all 5 anthers with apical tufts of stiff hairs.
Stigma lobes long, filiform. Fruit capsular, dehiscent apically by 2 valves. 2n = 28. Fifteen species,
Africa. Divided by Wimmer (1953) into three
sections: sect. Monopsis (1 sp.), sect. Dobrowskya
(C. Presl) Urb. (11 spp.) and sect. Parastranthus
(G. Don) E. Wimm. (3 spp.).
61. Unigenes E. Wimm.
Unigenes E. Wimm., Ann. Naturhist. Mus. Wien 56: 373
(1948).
Herb. Flowers very small, pedicellate, solitary,
axillary; pedicels ebracteolate. Corolla bilabiate,
white; tube dorsally cleft to base; lobes subdimorphic, the ventral slightly larger. Ventral anthers
with minute appendages and tufts of stiff hairs at
apex. Ovary nearly superior, 1-locular; placenta
basal. Fruit capsular, dehiscent from apex to base
by 2 valves; seed 1(2), white, large. One species, U.
humifusa (A. DC.) E. Wimm., South Africa.
62. Isotoma (R. Br.) Lindl.
Isotoma (R. Br.) Lindl., Edwards’ Bot. Reg. 12: pl. 964 (1826).
Annuals, sometimes dioecious. Flowers small to
medium-sized, solitary, axillary, or aggregated into
secund racemes; pedicels ebracteolate. Corolla bilabiate or rarely salverform, blue, rarely rose or
white; tube entire; lobes monomorphic. Filament
tube adnate to corolla above the middle; ventral
anthers with long bristle at apex plus tufts of stiff
63. Ruthiella Steenis
Ruthiella Steenis, Blumea 13: 127 (1965); Tuyn, Fl. Males.
ser. I, 6: 137–139 (1960), rev.
Phyllocharis Diels (1919), nom. illegit.
64. Diastatea Scheidw.
Diastatea Scheidw., Allg. Gartenz. 9: 396 (1841); McVaugh,
Bull. Torrey Bot. Club 67: 784–794 (1940), rev.
Annuals. Flowers small, in terminal, sometimes secund 5–30-flowered racemes (rarely solitary, axillary); pedicels ebracteolate. Corolla bilabiate, purplish, blue or white; tube entire, persistent on the
developing fruit and becoming scarious and hyaline; lobes dimorphic, the ventral larger. Filament
tube loosely adnate to the corolla at base; ventral 2 anthers with apical tufts of stiff hairs. Ovary
nearly superior, free from hypanthium for 4/5 of its
length. Fruit capsular, dehiscing to middle by 2 apical valves. Five species, Central and South America.
65. Palmerella A. Gray
Palmerella A. Gray, Proc. Amer. Acad. Arts 11: 80 (1876).
Perennial herb. Leaves sessile. Flowers mediumsized, in a 5–23-flowered subcapitate terminal
racemes; pedicels ebracteolate. Corolla bilabiate,
blue or purple; tube dorsally cleft from base for
half its length at maturity; lobes dimorphic, the
ventral larger. Filament tube dorsally loosely adnate to corolla; ventral anthers with a short apical
appendage and apical tufts of stiff hairs. Fruit capsular, dehiscent apically by 2 valves. One species,
Palmerella debilis A. Gray, western North America.
66. Porterella Torr.
Porterella Torr. in F.V. Hayden, Prelim. Rep. Geol. Surv. Montana: 488 (1872).
50
T.G. Lammers
Annual. Leaves much smaller than bracts, sessile.
Flowers small, fragrant, pedicellate, solitary, axillary. Corolla bilabiate, blue (rarely white), often
marked with yellow or white on ventral lip; tube
entire; lobes dimorphic, the ventral larger. Filament tube free from the corolla. Ventral anthers
with a horn-like bristle, all 5 with apical tufts of
stiff hairs. Ovary 2-locular with axile placentation.
Fruit capsular, dehiscent apically by 2 valves; seeds
fusiform, brown with dark apiculate tips. 2n = 22,
24. One species, P. carnosula (Hook. & Arn.) Torr.,
western North America.
67. Legenere McVaugh
Legenere McVaugh, N. Amer. Fl. 32A: 13 (1943); Ruiz de
Ciolfi, Bol. Soc. Argent. Bot. 17: 176–178 (1976), rev.
Emergent aquatic annuals. Leaves sessile. Flowers small, both chasmogamous and cleistogamous,
pedicellate, in a terminal raceme; pedicels ebracteolate. Corolla bilabiate, yellow; tube dorsally cleft
to base; lobes dimorphic, the ventral larger. Staminal column included; ventral anthers with minute
apical appendages. Ovary 1-locular with 2 parietal
placentae. Fruit capsular, dehiscent apically by 2
valves; seeds 20 or less. One species, L. valdiviana
(Phil.) E. Wimm., western North America, southern South America.
68. Howellia A. Gray
Howellia A. Gray, Proc. Amer. Acad. Arts 15: 43 (1879).
Submersed aquatic annuals. Stems branched,
flaccid, floating. Leaves sessile. Flowers small,
chasmogamous and cleistogamous, pedicellate,
solitary, axillary; pedicels ebracteolate. Corolla
bilabiate, white; tube dorsally cleft nearly to base;
lobes dimorphic, the ventral larger. Staminal
column included; ventral anthers with minute
apical appendages. Ovary 1-locular with 2 parietal
placentae. Capsule dehiscent laterally by irregular
ruptures; seeds 1–5, large. 2n = 22. One species,
H. aquatilis A. Gray, western North America.
Corolla bilabiate, blue (rarely pinkish or white), often marked with yellow or white on ventral lip; tube
entire; lobes dimorphic, the ventral larger. Ventral
anthers with a horn-like bristle plus apical tufts of
stiff hairs. Ovary 2-locular with axile placentation
or 1-locular with 2 parietal placentae. Capsule dehiscent by 3–5 longitudinal slits. 2n = 12, 16, 18, 20,
22, 24. Thirteen species, western North America.
70. Lysipomia Kunth
Lysipomia Kunth in Humb., Bonpl. & Kunth, Nov. Gen. Sp.
3: 318 (1819); McVaugh, Brittonia 8: 69–105 (1955), rev.;
Jeppesen, Fl. Ecuador 14: 136–151 (1981), reg. rev.
Rhizocephalum Wedd. (1858).
Dominella E. Wimm. (1953).
Dwarf perennial herbs, often forming cushions,
sometimes scapose. Leaves sessile or petiolate,
commonly rosulate. Flowers small to mediumsized, solitary, axillary, often not resupinate,
sometimes sessile; pedicels (when present)
ebracteolate. Corolla bilabiate, yellow or white;
tube entire (very rarely cleft dorsally to base); lobes
monomorphic or subdimorphic. Filament tube
adnate to corolla, at least at base; ventral anthers
commonly with long bristle at apex, sometimes
also with tufts of stiff hairs. Ovary 1-locular with
2 parietal placentae. Fruit a pyxis, operculum
umbonate. 2n = 20. Thirty species, Andean South
America.
71. Hippobroma G. Don
Hippobroma G. Don, Gen. Hist. 3: 717 (1834); McVaugh,
Bull. Torrey Bot. Club 67: 782–784 (1940), rev.
Perennial herbs. Leaves repand-dentate. Flowers
large, fragrant, solitary, axillary; pedicels short,
bibracteolate at base. Corolla salverform, white;
tube entire; lobes monomorphic. Filament tube adnate to corolla; all anthers with apical tufts of stiff
hairs. Fruit capsular, apically dehiscent by 2 valves.
2n = 28. One species, H. longiflora (L.) G. Don,
originally native to the West Indies but now naturalized throughout the tropics.
69. Downingia Torr.
Downingia Torr., Rep. Explor. RR Pac. Ocean 4: 116 (1857),
nom. cons.; McVaugh, Mem. Torrey Bot. Club 19, 4: 1–57
(1941), rev.; Ayers in Hickman, Jepson Man.: 460–462 (1993),
reg. rev.
Annuals. Leaves much smaller than bracts, sessile.
Flowers small, sessile, in a terminal raceme. Hypanthium pedicelliform, twisted 180° (rarely not).
72. Heterotoma Zucc.
Fig. 10
Heterotoma Zucc., Flora 15 (2, Beibl.): 100 (1832); Ayers,
Syst. Bot. 15: 296–327 (1990), rev.
Suffrutescent perennial herb. Flowers large,
spurred, in a pedunculate terminal raceme.
Hypanthium asymmetric, ventrally distended by
the nectar spur. Ventral calyx lobes displaced to
Campanulaceae
51
larger. Filament tube adnate to corolla basally or
rarely free; ventral anthers (rarely all 5) with apical
tufts of stiff white hair (rarely nude). Fruit capsular,
apically dehiscent by 2 valves; seeds slightly longer
than wide, testa reticulate. 2n = 28. Over 230
species, Central and South America, Greater Antilles. Wimmer (1953, 1968) divided the genus into
two sections: sect. Siphocampylus (203 spp.) with
four subsections: subsect. Hemisiphocampylus (A.
DC.) E. Wimm. (10 spp.), subsect. Siphocampylus
(180 spp.), subsect. Byrsanthes (C. Presl) E. Wimm.
(7 spp.) and subsect. Isochilus E. Wimm. (6 spp.);
sect. Brachysiphon E. Wimm. (31 spp.) also with
four subsections: subsect. Secundiflori E. Wimm.
(10 spp.), subsect. Altofissi E. Wimm. (6 spp.),
subsect. Megastomi E. Wimm. (11 spp.) and
subsect. Megalandri E. Wimm. (4 spp.).
74. Centropogon C. Presl
Centropogon C. Presl, Prodr. Monogr. Lobel. 48 (1836);
Jeppesen, Fl. Ecuador 14: 48–122 (1981), reg. rev.
Fig. 10. Campanulaceae-Lobelioideae. Heterotoma lobelioides. A Flowering branch and leaves. B Flower, partly
opened. C Flower, partly opened. D Anthers. E Stigma.
(Nash 1976)
tip of spur. Corolla unilabiate, orange or red with
yellow lobes, much of its length in the form of
a broad calcarate nectar spur; lobes monomorphic.
Ventral anthers with apical tufts of stiff hairs.
Fruit capsular, dehiscent by two apical valves.
2n = 14. One species, H. lobelioides Zucc., Central
America.
73. Siphocampylus Pohl
Siphocampylus Pohl, Pl. Bras. Icon. Descr. 2: 104 (1831);
Jeppesen, Fl. Ecuador 14: 151–170 (1981), reg. rev.
Suffrutescent herbs or shrubs, sometimes scandent or twining lianas. Leaves rarely opposite or
whorled. Flowers medium-sized to large, solitary,
axillary, rarely forming a terminal corymb or
raceme. Corolla bilabiate or tubular, red, orange,
pink, purple, yellow, green or white; tube entire or
very rarely fenestrate; lobes dimorphic, the dorsal
Suffrutescent herbs or shrubs, sometimes scandent lianas. Flowers medium-sized to large, solitary, axillary, rarely forming a terminal corymb or
raceme; pedicels commonly bibracteolate. Corolla
bilabiate or tubular, red, orange, pink, purple, yellow, green or white; tube entire or very rarely fenestrate, often constricted at base and/or inflated
at mouth; lobes dimorphic, the dorsal larger and
sometimes falcate. Filament tube adnate to corolla
basally; ventral anthers with apical tufts of stiff or
weak hairs (sometimes concrescent into a triangular scale). Fruit a berry, sometimes inflated; seeds
slightly longer than wide, reticulate. 2n = 28. Two
hundred and thirteen species, Central and South
America, Lesser Antilles. Lammers (1998b, 2002)
divided the genus into five sections: sect. Centropogon (49 spp.), sect. Siphocampyloides Benth. (100
spp.), sect. Wimmeriopsis McVaugh (40 spp.), sect.
Burmeisteroides Gleason (21 spp.) and sect. Niveopsis Lammers (1 sp.). Siphocampyloides was divided
into subsect. Brevilimbati E. Wimm. (89 spp.) and
subsect. Peruviani McVaugh (11 spp.); Wimmeriopsis into subsect. Falcati McVaugh and subsect.
Colombiani McVaugh (20 spp. each).
75. Burmeistera Triana
Fig. 11
Burmeistera Triana, Nuev. Gen. Esp. 13 (1855); Jeppesen, Fl.
Ecuador 14: 12–48 (1981), reg. rev.
Suffrutescent herbs or shrubs, sometimes scandent lianas. Flowers medium-sized to large, soli-
52
T.G. Lammers
cal tufts of stiff hairs. Ovary 2-locular, placentae
axile. Fruit capsular, dehiscent by two apical pores.
Six species, Polynesia.
77. Apetahia Baill.
Apetahia Baill., Bull. Mens. Soc. Linn. Paris 1: 310 (1882);
Florence, Allertonia 7: 248–253 (1997), rev.
Shrubs or small trees. Flowers large, solitary, axillary; pedicels bibracteolate at or above middle.
Corolla unilabiate, green, white, pink or violet; tube
cleft 1/4–1/5 of its length, curved slightly; lobes
monomorphic, spreading, forming 1/3–2/5 of the
corolla. Ovary 1-locular, placentae parietal (or appearing 2-locular/axile through intrusive growth of
placentae). Fruit capsular, dehiscent by two apical
valves which are often bifid. 2n = 28. Four species,
Polynesia.
78. Trematolobelia Zahlbr.
Fig. 11. Campanulaceae-Lobelioideae. Burmeistera virescens. A Flowering branch. B Leaves. C Flower. D Flower,
opened. E Anther tube and stigma. F Berry and seeds. (Nash
1976)
tary, axillary, rarely forming a terminal corymb
or raceme; pedicels ebracteolate. Corolla bilabiate,
green, yellow, purple or maroon; tube entire, often inflated at base and/or mouth; lobes dimorphic, the dorsal larger and often falcate. Filament
tube free or adnate to corolla basally; orifice of anther tube wide open, glabrous or with soft weak
hairs on ventral anther or all anthers. Fruit a berry,
sometimes much inflated; seeds much longer than
wide, reticulate. Over 100 species, Central America and Andean South America. Lammers (1998b,
2002) divided the genus into two sections: sect.
Burmeistera (54 spp.) and sect. Barbatae E. Wimm.
(48 spp.).
76. Sclerotheca A. DC.
Sclerotheca A. DC. in DC., Prodr. 7: 356 (1839).
Shrubs or small trees. Flowers large, solitary, axillary; pedicels bibracteolate at or below middle.
Corolla bilabiate, magenta, purple, green or yellow; tube entire or dorsally cleft to base; lobes
monomorphic. Ventral anthers or all five with api-
Trematolobelia Zahlbr. in Rock, Coll. Hawaii Publ. Bull. 2:
45 (1913); Lammers in Wagner et al., Man. Fl. Pl. Hawai’i:
485–488 (1990), rev.
Pliestesial rosette treelets. Leaves sessile or petiolate. Flowers large, pedicellate, in a terminal
50–400-flowered pedunculate panicle composed
of 5–20 horizontally radiating secund racemes;
pedicels bibracteolate. Corolla unilabiate or
bilabiate, red, rose, pink or white; tube dorsally
cleft to base; lobes monomorphic. Ventral anthers
with apical tufts of stiff hairs. Fruit capsular,
dehiscent laterally by 5–15 irregular pores after
decomposition of the fleshy exocarp; seeds winged.
2n = 28. Four species, Hawaiian islands.
79. Brighamia A. Gray
Brighamia A. Gray, Proc. Amer. Acad. Arts 7: 185 (1867);
Lammers, Syst. Bot. 14: 133–138 (1989), rev.; Lammers in
Wagner et al., Man. Fl. Pl. Hawai’i: 422–423 (1990), rev.
Polycarpic rosette treelets; stems unbranched, caudiciform, succulent. Leaves fleshy. Flowers large,
fragrant, in axillary 3–8-flowered racemes. Corolla
salverform, yellow or white; tube slender, straight,
entire at anthesis but eventually cleft for c. 1/3 its
length. Staminal column included; filament tube
adnate to corolla tube below middle; all 5 anthers
with apical tufts of stiff hairs. Capsule dehiscent
by two lateral longitudinal slits per locule; seeds
white, rugose. 2n = 28. Two species, Hawaiian
islands.
Campanulaceae
80. Delissea Gaudich.
Delissea Gaudich., Voy. Uranie: pl. 78 (1826); Lammers in
Wagner et al., Man. Fl. Pl. Hawai’i: 467–472 (1990), rev.;
Lammers, Syst. Bot. Monogr. 73: 1–75 (2005), rev.
Shrubs, treelets or trees; stems unbranched or sparingly branched. Flowers medium-sized to large, in
5–20-flowered axillary racemes. Corolla bilabiate
or unilabiate, white or greenish (rarely lilac); tube
dorsally cleft to middle, with a knob at the base
of the cleft (sometimes also with a lateral pair of
knobs); lobes monomorphic. Filament tube free
from corolla; ventral anthers with apical tufts of
stiff hairs. Fruit a purple berry; seeds white, transversely rugose. 2n = 28. Fifteen species, Hawaiian
islands. Lammers (2005) divided the genus into
three sections: sect. Delissea (4 spp.), sect. Rhytidospermae Lammers (4 spp.) and sect. Macranthae
(Hillebr.) Rock (7 spp.).
81. Cyanea Gaudich.
Cyanea Gaudich., Voy. Uranie: pl. 75 (1828); Lammers in
Wagner et al., Man. Fl. Pl. Hawai’i: 437–467 (1990), rev.
Shrubs, treelets or trees, rarely lianas, rarely
epiphytic; stems unbranched or sparingly
branched, sometimes muricate or aculeate (more
so in juveniles); latex sometimes yellow or tan.
Leaves sometimes muricate or aculeate (more
so in juveniles); margin sometimes pinnately
lobed, cleft, parted, or divided (often more so
in juveniles). Flowers medium-sized to large,
in (3–)5–25(–40)-flowered axillary racemes,
sometimes subumbellate. Corolla bilabiate or
unilabiate, white, magenta, purple, pink, greenish
or yellowish, sometimes longitudinally striped,
rarely muricate; tube dorsally cleft to middle;
lobes monomorphic. Filament tube free from
corolla or dorsally adnate for 1/3–1/2 of its length;
ventral anthers (rarely all 5) with apical tufts of
stiff hairs. Fruit a yellow, orange or purple berry;
seeds reticulate. 2n = 28. Seventy-eight species,
Hawaiian islands. Based on the phylogeny of
Givnish et al. (1995), the genus can be divided
into two sections: sect. Cyanea (61 spp.) and sect.
Delisseoideae (Hillebr.) Rock (17 spp.); the former
will eventually prove divisible into subordinate
taxa.
82. Clermontia Gaudich.
Clermontia Gaudich., Voy. Uranie: pl. 459 (1829); Lammers
in Wagner et al., Man. Fl. Pl. Hawai’i: 423–437 (1990), rev.;
Lammers, Syst. Bot. Monogr. 32: 1–94 (1991), rev.
53
Shrubs or small trees, sometimes epiphytic. Stems
branched repeatedly. Flowers medium-sized to
large, in axillary, 2(–10)-flowered, subumbellate
racemes. Calyx lobes frequently similar to corolla
in shape, texture and colour. Corolla bilabiate,
unilabiate, tubular, or rotate, white, green, purple
or rose; tube dorsally cleft to near the base; lobes
monomorphic. Fruit an orange or yellow berry;
seeds reticulate. 2n = 28. Twenty-two species, Hawaiian islands. Two sections were recognized by
Lammers (1991): sect. Clermontioideae (Hillebr.)
Rock (7 spp.) and sect. Clermontia (15 spp.).
Each was divided into three series: the former
into ser. Clermontioideae (Hillebr.) Lammers
(3 spp.), ser. Sarcanthae Lammers (2 spp.) and
ser. Unilabiatae Lammers (2 spp.), the latter
into ser. Kakeanae Lammers (7 spp.), ser. Parviflorae Lammers (4 spp.) and ser. Clermontia
(4 spp.).
IV. Subfam. Cyphocarpoideae Miers (1848).
Annual and perennial herbs. Leaves pinnatifid.
Flowers small to medium-sized, sessile, bibracteolate at base, in a 3–15-flowered terminal spike.
Hypanthium clavate or linear. Calyx with the odd
(unpaired) lobe in a ventral (anterior) position;
lobes pinnatifid. Corolla blue, lavender or white,
bilaterally symmetric, with a single cucullate
dorsal lobe bearing an apical appendage plus
a 4-lobed ventral lip with a gibbous palate; tube
entire. Stamens epipetalous, distinct, included.
Ovary inferior, bilocular with axile placentation;
stigma 2-lobed. Fruit a capsule, dehiscent via
irregular rupture of the lateral walls.
83. Cyphocarpus Miers
Fig. 12
Cyphocarpus Miers, London J. Bot. 7: 62 (1848); Wimmer,
Pflanzenr. IV.276c: 922–923 (1968), rev.
See subfamily description. Three species, Chile.
V. Subfam. Cyphioideae (A. DC.) Walp. (1852).
Perennial herbs with a subglobose or elongate
root tuber; stems erect or twining. Inflorescence
a terminal raceme. Calyx with the odd (unpaired)
lobe in a ventral (anterior) position. Corolla
bilaterally symmetric, bilabiate with 3 dorsal and
2 nearly distinct ventral lobes, or tubular with
5 subequal lobes. Stamens epigynous; filaments
distinct or connate; anthers distinct. Ovary inferior
(rarely almost superior), 2-locular with apical
54
T.G. Lammers
Fig. 12. Campanulaceae-Cyphocarpoideae. Cyphocarpus
psammophilus. A Habit. B Flower, opened. C Capsule.
D Seed. (Ricardi 1959)
placentation; style tipped by a fluid-filled stigmatic
cavity with a lateral pore. Fruit a loculicidal
capsule, dehiscent with 2 apical valves, these sometimes splitting longitudinally at apex so that the
capsule appears 4-valved; seeds circumferentially
winged and smooth, or 3-angled and coarsely
reticulate.
Cyphioideae has sometimes been circumscribed to include Cyphocarpus, Nemacladus,
Parishella and Pseudonemacladus (Schönland
1889; Wagenitz 1964; Wimmer 1968). Note that
with this expanded circumscription, the name
Cyphocarpoideae has priority.
84. Cyphia P.J. Bergius
Fig. 13
Cyphia P.J. Bergius, Descr. Pl. Cap. 172 (1767); Wimmer,
Pflanzenr. IV.276c: 935–1014 (1968), rev.
See subfamily description. Sixty-four species,
Africa, divided by Wimmer (1968) into two
sections: sect. Cyphia (51 spp.) and sect. Cyphiella
C. Presl (13 spp.).
Fig. 13.
Campanulaceae-Cyphioideae. Cyphia erecta.
A Habit. B Leaves. C Flower. D Flower, perianth partly
removed. E Flower, longitudinal section. F Stigma.
G Capsule. H seed. (Thulin 1984)
Selected Bibliography
Adamson, R.S. 1952. A revision of the genera Prismatocarpus and Roella. J. S. African Bot. 17: 93–166.
Batterman, M.R.W., Lammers, T.G. 2004. Branched foliar
trichomes of Lobelioideae (Campanulaceae) and the
infrageneric classification of Centropogon. Syst. Bot.
29, 2: 448–458.
Bentham, G. 1875. Notes on the gamopetalous orders belonging to the campanulaceous and oleaceous groups.
J. Linn. Soc., Bot. 15: 1–16.
Bentham, G. 1876. Campanulaceae. In: Bentham, G.,
Hooker, J.D., Genera Plantarum, vol. II. London:
Reeve, pp. 541–564.
Bigazzi, M. 1986. Ultrastructural and cytochemical observations on fibrillar intranuclear inclusions in the family
Campanulaceae. Caryologia 39: 199–210.
Campanulaceae
Bremer, K., Gustafsson, M.H.G. 1997. East Gondwana ancestry of the sunflower alliance of families. Proc. Natl
Acad. Sci. U.S.A. 94: 9188–9190.
Candolle, A. de 1839. Lobeliaceae, Campanulaceae. In: Candolle, A.P. de, Prodromus systematis naturalis regni
vegetabilis, vol. 7. Paris: Treuttel & Würtz, pp. 339–496,
784–792.
Carlquist, S. 1962. Ontogeny and comparative anatomy of
thorns of Hawaiian Lobeliaceae. Amer. J. Bot. 49: 413–
419.
Carlquist, S. 1969. Wood anatomy of Lobelioideae (Campanulaceae). Biotropica 1: 47–72.
Carlquist, S. 1992. Wood anatomy of sympetalous dicotyledon families: a summary, with comments on systematic
relationships and evolution of the woody habit. Ann.
Missouri Bot. Gard. 79: 303–332.
Carolin, R.C. 1960. The structures involved in the presentation of pollen to visiting insects in the order Campanales [sic]. Proc. Linn. Soc. New South Wales 85:
197–207.
Carolin, R.C. 1967. The concept of the inflorescence in the
order Campanulales. Proc. Linn. Soc. New South Wales
92: 7–26.
Chapman, J.L. 1966. Comparative palynology in Campanulaceae. Trans. Kansas Acad. Sci. 69: 197–200.
Chase, M.W, Soltis, D.E., Olmstead, R.G., Morgan, D., Les, D.,
Mishler, B.D., Duvall, M.R., Price, R.A., Hills, H.G.,
Qiu, Y., Kron, K.A., Rettig, J.H., Conti, E., Palmer, J.D.,
Manhart, J.R., Sytsma, K.J., Michaels, H.J., Kress, W.J.,
Donoghue, M.J., Clark, W.D., Hedrén, M., Gaut, B.S.,
Jansen, R.K., Kim, K.-J., Wimpee, C.F., Smith, J.F.,
Furnier, G.R., Strauss, S., Xiang, Q., Plunkett, G.M.,
Soltis, P.S., Swensen, S., Eguiarte, L.E., Learn, G.H.
Jr., Barrett, S.C.H., Graham, S., Dayananadan, S.,
Albert, V.A. 1993. Phylogenetics of seed plants: an
analysis of nucleotide sequences from the plastid gene
rbcL. Ann. Missouri Bot. Gard. 80: 528–580.
Cosner, M.E., Jansen, R.K., Lammers, T.G. 1994. Phylogenetic relationships in the Campanulales based on rbcL
sequences. Pl. Syst. Evol. 190: 79–95.
Cronquist, A. 1981. An integrated system of classification
of flowering plants. New York: Columbia University
Press.
Dunbar, A. 1975. On pollen of Campanulaceae and related
families with special reference to the surface ultrastructure. Bot. Notiser 128: 73–118.
Dunbar, A. 1984. Pollen morphology in Campanulaceae, IV.
Nordic J. Bot. 4: 1–19.
Erbar, C., Leins, P. 1989. On the early floral development
and the mechanisms of secondary pollen presentation
in Campanula, Jasione and Lobelia. Bot. Jahrb. Syst.
111: 29–55.
Erbar, C., Leins, P. 1996. Distribution of the character states
“early sympetaly” and “late sympetaly” within the
“Sympetalae Tetracyclicae” and presumably allied
groups. Bot. Acta 109: 427–440.
Fedorov, A.A. 1957. Campanulaceae. In: Shishkin, B.K. (ed.)
Flora URSS, vol. 24. Moscow: Akademia Nauk, pp. 126–
450, 459–475.
Gadella, T.W.J. 1966. Some notes on the delimitation of genera in the Campanulaceae. Proc. Konink. Ned. Akad.
Wetensch. ser. C 69: 502–521.
Givnish, T.J., Sytsma, K.J., Smith, J.F., Hahn, W.J. 1995.
Molecular evolution, adaptive radiation, and ge-
55
ographic speciation in Cyanea (Campanulaceae,
Lobelioideae). In: Wagner, W.L., Funk, V.A. (eds)
Hawaiian biogeography: evolution on a hot spot
archipelago. Washington: Smithsonian Institution
Press, pp. 288–337.
Gustafsson, M.H.G. 1995. Petal venation in the Asterales
and related orders. Bot. J. Linn. Soc. 118: 1–18.
Gustafsson, M.H.G., Bremer, K. 1995. Morphology and phylogenetic interrelationships of the Asteraceae, Calyceraceae, Campanulaceae, Goodeniaceae, and related
families (Asterales). Amer. J. Bot. 82: 250–265.
Gustafsson, M.H.G., Backlund, A., Bremer, B. 1996. Phylogeny of the Asterales sensu lato based on rbcL sequences with particular reference to the Goodeniaceae.
Pl. Syst. Evol. 199: 217–242.
Hickey, L.J., Wolfe, J.A. 1975. The bases of angiosperm
phylogeny: vegetative morphology. Ann. Missouri Bot.
Gard. 62: 538–589.
Hickman, J. 1993. The Jepson Manual. Higher Plants of
California. University of California Press.
Hong, D.-Y. 1995. The geography of the Campanulaceae: on
the distribution centres. Acta Phytotax. Sin. 33: 521–
536.
Hong, D.-Y., Lian, Y.S., Shen, L.D. 1983. Campanulaceae. In:
Flora Reipublicae Popularis Sinicae, vol. 73, 2. Beijing:
Science Press, pp. 1–177.
Kaplan, D.R. 1967. Floral morphology, organogenesis and
interpretation of the inferior ovary in Downingia bacigalupii. Amer. J. Bot. 54: 1274–1290.
Kårehed, J., Lundberg, J., Bremer, B., Bremer, K. 1999. Evolution of the Australasian families Alseuosmiaceae, Argophyllaceae, and Phellinaceae. Syst. Bot. 24: 660–682.
Kolakovskii, A.A. 1987. System of the Campanulaceae family
from the Old World (in Russian). Bot. Zhurn. (Moscow
& Leningrad) 72: 1572–1579.
Kolakovskii, A.A. 1994. The conspectus of the system of the
Old World Campanulaceae (in Russian). Bot. Zhurn.
(Moscow & Leningrad) 79: 109–124.
Lammers, T.G. 1991. Systematics of Clermontia (Campanulaceae: Lobelioideae). Syst. Bot. Monogr. 32: 1–94.
Lammers, T.G. 1992. Circumscription and phylogeny of the
Campanulales. Ann. Missouri Bot. Gard. 79: 388–413.
Lammers, T.G. 1993. Chromosome numbers of Campanulaceae. III. Review and integration of data for subfamily
Lobelioideae. Amer. J. Bot. 80: 660–675.
Lammers, T.G. 1998a. Nemacladoideae, a new subfamily of
Campanulaceae. Novon 8: 36–37.
Lammers, T.G. 1998b. Review of the Neotropical endemics
Burmeistera, Centropogon, and Siphocampylus (Campanulaceae: Lobelioideae), with description of 18 new
species and a new section. Brittonia 50: 233–262.
Lammers, T.G. 2002. Seventeen new species of Lobelioideae
(Campanulaceae) from South America. Novon 12: 206–
233.
Lammers, T.G. 2005. Revision of Delissea (Campanulaceae:
Lobelioideae). Syst. Bot. Monogr. 73: 1–75.
Lancucka-Srodoniowa, M. 1977. New herbs described from
the Tertiary of Poland. Acta Palaeobot. 18: 37–44.
Lancucka-Srodoniowa, M. 1979. Macroscopic plant remains
from the freshwater Miocene of the Nowy-Sacz Basin
(West Carpathians, Poland). Acta Palaeobot. 20: 3–117.
Leins, P., Erbar, C. 1990. On the mechanisms of secondary
pollen presentation in the Campanulales-Asteralescomplex. Bot. Acta 103: 87–92.
56
T.G. Lammers
Leins, P., Erbar, C. 2003. The pollen box in Cyphiaceae (Campanulales). Intl J. Pl. Sci. 164 suppl. 5: S321–S328.
Lundberg, J., Bremer, K. 2003. A phylogenetic study of the
order Asterales using one morphological and three
molecular data sets. Intl J. Pl. Sci. 164: 553–578.
Michaels, H.J., Scott, K.M., Olmstead, R.G., Szaro, T.,
Jansen, R.K., Palmer, J.D. 1993. Interfamilial relationships of the Asteraceae: insights from rbcL sequence
variation. Ann. Missouri Bot. Gard. 80: 742–751.
Murata, J. 1995. A revision of infrageneric classification
of Lobelia (Campanulaceae-Lobelioideae) with special
reference to seed coat morphology. J. Fac. Sci. Univ.
Tokyo sect. 3 15: 349–371.
Nash, D.L. 1976. Campanulaceae. In: Nash, D.L., Dieterle, J.V.A. (eds) Flora of Guatemala. Fieldiana Bot. 24
(XI, 4): 396–431.
Olmstead, R.G., Michaels, H.J., Scott, K.M., Palmer, J.D.
1992. Monophyly of the Asteridae and identification
of their major lineages inferred from DNA sequences
of rbcL. Ann. Missouri Bot. Gard. 79: 249–265.
Olmstead, R.G., Bremer, B., Scott, K.M., Palmer, J.D. 1993.
A parsimony analysis of the Asteridae sensu lato based
on rbcL sequences. Ann. Missouri Bot. Gard. 80: 700–
722.
Philipson, W.R. 1948. Studies in the development of the
inflorescence, V. The raceme of Lobelia dortmanna L.,
and other campanulaceous inflorescences. Ann. Bot. II
12: 147–156, pl. 4.
Presl, C.B. 1836. Prodromus monographiae Lobeliacearum.
Prague: Theophilus Haase.
Ricardi, M. 1959. Un Cyphocarpus nuevo para Chile. Bol.
Soc. Argent. Bot. 7: 247–250.
Rosén, W. 1932. Zur Embryologie der Campanulaceen und
Lobeliaceen. Acta Horti Gothob. 7: 31–42.
Rosén, W. 1949. Endosperm development in Campanulaceae and closely related families. Bot. Notiser 1949:
137–147.
Schönland, S. 1889. Campanulaceae. In: Engler, A.,
Prantl, K., Die natürlichen Pflanzenfamilien, IV, 5.
Leipzig: W. Engelmann, pp. 40–70.
Shen, L.D., Hong, D.Y. 1983. Codonopsis. In: Hong, D.Y. (ed.)
Flora Reipublicae Popularis Sinicae, vol. 73, 2. Beijing:
Science Press, pp. 32–69.
Shetler, S.G. 1979. Pollen-collecting hairs of Campanula
(Campanulaceae). I. Historical review. Taxon 28: 205–
215.
Shetler, S.G., Morin, N.R. 1986. Seed morphology in North
American Campanulaceae. Ann. Missouri Bot. Gard.
73: 653–688.
Shrestha, K.K. 1997. Taxonomic revision of the SinoHimalayan genus Cyananthus (Campanulaceae). Acta
Phytotax. Sin. 35: 396–433.
Shulkina, T.V. 1980. The significance of life-form characters
for systematics, with special reference to the family
Campanulaceae. Pl. Syst. Evol. 136: 233–246.
Takhtajan, A. 1980. Outline of the classification of flowering
plants (Magnoliophyta). Bot. Rev. 46: 225–359.
Takhtajan, A. 1997. Diversity and classification of flowering
plants. New York: Columbia University Press.
Thulin, M. 1976. Campanulaceae. In: Polhill, R.M. (ed.)
Flora of tropical East Africa. Rotterdam: Balkema.
Thulin, M. 1984. Lobeliaceae. In: Polhill, R.M. (ed.) Flora of
tropical East Africa. Rotterdamm: Balkema.
Tjon Sie Fat, L. 1978. Contribution to the knowledge of
cyanogenesis in angiosperms. 2. Cyanogenesis in Campanulaceae. Proc. Koninkl. Ned. Akad. Wetensch. C 81:
126–131.
Tobe, H., Morin, N.R. 1996. Embryology and circumscription of Campanulaceae and Campanulales: a review of
literature. J. Pl. Res. 109: 425–435.
Wagenitz, G. 1964. Reihe Campanulales. In: Melchior, H.
(ed.) A. Engler’s Syllabus der Pflanzenfamilien,
ed. 12, Band 2. Berlin: Gebrüder-Bornträger, pp.
478–497.
Wimmer, F.E. 1943. Campanulaceae-Lobelioideae, I. Teil. In:
Mansfeld, R. (ed.) Das Pflanzenreich, IV.276b. Leipzig:
W. Engelmann, pp. 1–260.
Wimmer, F.E. 1953. Campanulaceae-Lobelioideae, II Teil.
In: Stubbe, H., Noack, K. (eds) Das Pflanzenreich,
IV.276b. Berlin: Akademie Verlag, pp. 261–813.
Wimmer, F.E. 1968. Campanulaceae-Cyphioideae. In:
Stubbe, H. (ed.) Das Pflanzenreich, IV.276c. Berlin:
Akademie-Verlag, pp. 917–1014.
Yeo, P.F. 1993. Platycodoneae [sic], a new tribe in Campanulaceae. Taxon 42: 109.
Carpodetaceae
Carpodetaceae Fenzl, Denkschr. Königl.-Baier. Bot. Gesell. Regensburg 3: 155 (1841).
Abrophyllaceae Nakai (1943).
M.H.G. Gustafsson
Evergreen shrubs or trees with unicellular hairs.
Leaves alternate, estipulate, petiolate, serrate. Inflorescences terminal and/or axillary, paniculate,
bracts minute. Flowers bisexual or functionally
female, actinomorphic. Sepals (4)5(7), small,
free or basally fused. Petals (4)5(7), free, valvate
in bud. Stamens (4)5(7), alternipetalous, free,
anthers tetrasporangiate, introrse. Intrastaminal
nectar-disc present or absent. Ovary superior or
inferior, (3)5-locular, placentation axile. Ovules
many, anatropous, unitegmic. Style simple, distally
branched, or virtually absent. Stigma capitate
or lobed. Fruit a berry or a loculicidal capsule.
Seeds numerous, small, alveolate, with hard testa,
endosperm abundant, embryo minute.
Three genera with five species in eastern Australia, New Guinea, New Zealand and the Solomon
Islands.
Vegetative Morphology Juvenile plants of
Carpodetus serratus often show a characteristic
growth pattern, with slender, zigzag, divaricate
branches. This type of branching is typical for
a number of other New Zealand plants. Leaves are
uniformly serrate (very slightly in Abrophyllum
microcarpum), with glandular teeth. The venation
is prominent on the abaxial side of the leaf, and can
be described as semicraspedodromous. Domatia
sometimes develop in the abaxial axils of midrib
and lateral veins in Carpodetus.
Vegetative Anatomy. The characteristic indumentum of the three genera of Carpodetaceae was
described by Al-Shammary and Gornall (1994).
Hairs on young stems, leaves and inflorescences are
all of one type, almost identical in the three genera: unicellular, eglandular and curved (Fig. 14E).
The base of these hairs is often sunken, and they
have a very thick wall and a warty cuticle. The leaf
mesophyll and stem pericycle of Abrophyllum and
Cuttsia contain idioblasts with granular contents,
possibly some kind of latex. Carpodetus lacks these
structures but has crystalliferous idioblasts in leaf
mesophyll and floral parts (Gustafsson and Bremer
1997). The wood of Carpodetus serratus has both
uniseriate and very large multiseriate rays. Vessel
perforation plates are scalariform, with numerous
(average 80) cross bars. Large rhomboidal crystals
occur in the ray cells (Patel 1973).
Floral Morphology. Flowers are predominantly bisexual but Carpodetus is sometimes
gynodioecious, i.e. some plants bear functionally female flowers with rudimentary stamens.
Merosity is variable, and 4-, 5- and 6-merous
flowers may occur on the same plant. The abaxial
side of sepals and petals have the same kind of
indumentum as is found on vegetative parts.
There may be a slight basal fusion in the sepal
whorl, and in Carpodetus serratus the calyx leaves
a circular scar after abscission (Fig. 14C). At least
in Carpodetus, petals are basally fused early in
ontogeny (Tobe and Raven 1999). Filaments are
short in Abrophyllum, not exceeding the anthers
in length, while in the other genera filaments are
three to several times longer, protruding from the
widely open, star-like flowers (Fig. 14B, F, G). The
ovary is inferior in Carpodetus but, after anthesis,
the apical portion enlarges disproportionately, so
that the locules and placentae reach above the line
of perianth insertion. The ovary of Carpodetus is
crowned by a pulvinate disc that may be radially
furrowed. The stigma is deeply (3)5-lobed in
Abrophyllum and Cuttsia, and in the latter genus
the style is distally branched, particularly so after
anthesis. In Carpodetus, the stigma is capitate but
may have a few central pits and numerous, slight
indentations along the margin.
Pollen Morphology. Pollen grains of Carpodetaceae are of two distinctive types, tetrahedral
tetrads in Carpodetus and tricolporate monads in
Abrophyllum and Cuttsia (Hideux and Ferguson
1976; Praglowski and Grafström 1985). The surface
58
M.H.G. Gustafsson
pattern is striate to rugulate or, in Carpodetus,
smooth or with irregular, flattened elements.
The infratectum consists of wide columellae in
Abrophyllum and Cuttsia, and is very thin in
Carpodetus, consisting of scattered, small irregular
elements.
Karyology. For Carpodetus, diploid chromosome numbers of 2n = 28 and 30 have been
reported (Hair and Beuzenberg 1960).
Pollination and Reproductive Systems.
The flowers are widely open at anthesis, and
the nectar (produced at least in Cuttsia and
Carpodetus) is easily accessible. Cuttsia flowers
have been referred to as an example of a generalist
entomophilous syndrome (Williams and Adam
1994), and flowers of Carpodetus arboreus are
reported to have an unpleasant odour (van Royen
1983), which could indicate adaptation to insect
pollination, perhaps by flies.
Fruit and Seed. Abrophyllum has fleshy berries,
whilst those of Carpodetus have a thin, leathery
wall. The fruit of Cuttsia is a dry capsule with loculicidal dehiscence. The seeds of Carpodetus are
much larger than those of the other genera, and
tend to become angular due to dense packing in
the fruit (Fig. 14D). All genera have seeds with
a highly characteristic alveolate surface pattern, resulting from the presence of strong thickenings on
the proximal and radial walls of the testa epidermis cells (Krach 1976; Fig. 14D). The embryo is
very small, 1/6 to less than 1/10 of the length of the
seed. Endosperm is abundant, and contains fatty
substances and aleuron grains.
Dispersal. The dark-coloured berries of Abrophyllum and Carpodetus are presumably birddispersed, whilst the small seeds of Cuttsia lack an
obvious adaptation for dispersal.
Phytochemistry. The polyphenols leucodelphinidin, quercetin and kaempferol have been
detected in Carpodetus serratus, and a triterpene,
lupeol, occurs in the bark of the same species
(Hegnauer 1973).
Relationships Within the Family. Cuttsia
and Abrophyllum are morphologically highly similar, and they constitute tribe Cuttsieae of Saxifragaceae-Escallonioideae in the system of Engler
(1930). Molecular data confirm a close relationship
between these genera (Gustafsson and Bremer
1997).
Affinities. The genera here included in Carpodetaceae have traditionally been placed in
the highly heterogeneous Saxifragaceae sensu
lato. Praglowski and Grafström (1985) found
palynological similarities between Carpodetus
and Ericaceae, and pointed to the possibility of
a close relationship between these taxa. Findings
from studies of DNA variation in the last few years
(Gustafsson and Bremer 1997; Lundberg 2001)
have confirmed the monophyly of Carpodetaceae
as here delimited, and indicated a position distant
from both Saxifragaceae and Ericaceae, viz. in
Asterales sensu lato. This ordinal placement
is supported by morphological characters, e.g.
valvate corolla aestivation, and general characters of Asteridae such as unitegmic ovules and
alternipetalous stamens. Carpodetaceae are but
one of several small, woody, southern hemisphere
families which have recently been found to belong
in Asterales. One of these families, the monotypic
Rousseaceae from Mauritius, is sister to Carpodetaceae, according to recent phylogenetic studies
based on molecular data and morphology (Lundberg 2001; Lundberg and Bremer 2003). Lundberg
(2001) reduced Carpodetaceae to a subfamily of
Rousseaceae, and this has often been followed in
later studies and classification schemes, e.g. APG
II (2003). The idea of an affinity between these two
taxa is not new; already Fenzl (1841) considered,
on morphological grounds, Carpodetaceae to
stand between Saxifragaceae and Rousseaceae.
Most of the morphological characters shared by
Roussea and Carpodetaceae are widespread in
Asterales. One exception is a carpel number of
five, which is otherwise seen only in some genera
of Campanulaceae; most Asterales have two or
three. Carpodetaceae (together with Roussea) are
sister to Pentaphragmataceae plus Campanulaceae
in the most recent, combined molecular and
morphological analysis of Asterales, but this
relationship is only weakly supported (Lundberg
and Bremer 2003).
Distribution and Habitats. Abrophyllum
and Cuttsia occur in rainforests in New South
Wales and southern Queensland, particularly
along watercourses. Carpodetus arboreus is found
in montane and subalpine forests of New Guinea
and the Solomon Islands, often as part of the
undergrowth, and reaches altitudes above 3,500 m.
Carpodetaceae
59
Carpodetus serratus occurs in various kinds of
forest up to 1,000 m altitude in most regions of New
Zealand.
Economic Importance. The wood of Carpodetus serratus is reported to be strong and tough, and
has been used for, e.g. tool handles. Frequent damage due to wood-boring insects reduces, however,
its economic importance. Abrophyllum ornans is
occasionally cultivated as an ornamental tree in
Australia.
Key to the Genera
1. Flowers epigynous, stigma capitate, seeds angular
1. Carpodetus
– Flowers hypogynous, stigma (3)5-lobed, seeds ovoid
2
2. Filament shorter than anther, stigma sessile, fruit baccate
2. Abrophyllum
– Filament longer than anther, style well-developed,
fruit capsular
3. Cuttsia
Genera of Carpodetaceae
1. Carpodetus J.R. Forst. & G. Forst.
Fig. 14A–E
Carpodetus J.R. Forst. & G. Forst., Char. Gen. Pl. t. 17: 33
(1776).
Argyrocalymma K. Schumann ex K. Sch. & Ltb. (1900).
Shrubs or trees to 20 m tall. Inflorescences often
equalling leaves in length. Petals white, yellowish or greenish. Filaments much longer than anthers. Ovary inferior or almost so, crowned by conspicuous, intrastaminal nectar-disc. Style present,
stigma capitate. Fruit a black or greyish, leathery berry. Seeds angular. Pollen grains in tetrahedral tetrads. 2n = 28, 30. Two species, C. arboreus
(K. Schum. & Lauterb.) Schltr. in New Guinea and
the Solomon Islands, and C. serratus J.R. Forst. &
G. Forst. in New Zealand. C. arboreus is very variable in size, shape and texture of the leaves, and
a number of different species have been described
from its area, but van Royen (1983) considered
these to belong to a single, polymorphic species.
2. Abrophyllum Hook. f.
Fig. 14F
Fig. 14. Carpodetaceae. A–E Carpodetus serratus. A Flowering branch. B Flower. C Young fruit. D Seed. E Hair.
F Abrophyllum ornans. Flower. G Cuttsia viburnea. Flower
Two species, A. microcarpum Domin in southern
Queensland and A. ornans Hook f. in New South
Wales and southern Queensland.
3. Cuttsia F. Muell.
Fig. 14G
Cuttsia F. Muell., Fragm. V, t. 70: 47 (1865).
Shrub or tree to 15 m tall. Inflorescences often
equalling leaves in length. Petals white. Filaments
much longer than anthers. Ovary superior, with
nectariferous base. Stigma 5-lobed or style distally
5-branched. Fruit a loculicidal capsule. Seeds
minute, ovoid. Pollen in monads, tricolporate.
A single species, C. viburnea F. Muell. in New
South Wales and southern Queensland.
Abrophyllum Hook. f. in Benth. Fl. Austral. II: 437 (1864).
Shrubs or small trees to 8 tall. Inflorescences much
shorter than leaves. Petals yellowish. Filaments
shorter than anthers. Ovary superior. Stigma (3)5lobed, sessile. Fruit a blackish, fleshy berry. Seeds
minute, ovoid. Pollen in monads, tricolporate.
Selected Bibliography
Al-Shammary, K.I.A., Gornall, R.J. 1994. Trichome anatomy
of the Saxifragaceae s.l. from the southern hemisphere.
Bot. J. Linn. Soc. 114: 99–131.
60
M.H.G. Gustafsson
APG II (2003) An update of the Angiosperm Phylogeny
Group classification for the orders and families of
flowering plants: APG II. The Angiosperm Phylogeny
Group. Bot. J. Linn. Soc. 141: 399–436.
Engler, A. 1930. Saxifragaceae. In: Engler, A., Prantl, K., Die
natürlichen Pflanzenfamilien, ed. 2, vol. 18a. Leipzig:
Engelmann, pp. 74–226.
Fenzl, E. 1841. Carpodetus Forster. Denkschr. Königl.-Baier.
Bot. Gesell. Regensburg 3: 155–173.
Gustafsson, M.H.G., Bremer, K. 1997. The circumscription
and systematic position of Carpodetaceae. Austral.
Syst. Bot. 10: 855–862.
Hair, J.B., Beuzenberg, E.J. 1960. Contributions to a chromosome atlas of the New Zealand flora. 4. Miscellaneous
families. N. Z. J. Sci. 3: 432–440.
Hegnauer, R. 1973. Chemotaxonomie der Pflanzen, Band 6.
Basel: Birkhäuser.
Hideux, M.J., Ferguson, I.K. 1976. The stereostructure of the
exine and its evolutionary significance in the Saxifragaceae sensu lato. In: Ferguson, I.K., Muller, J. (eds)
The evolutionary significance of the exine. London:
Academic Press, pp. 327–377.
Krach, J.E. 1976. Samenanatomie der Rosifloren, 1. Die
Samen der Saxifragaceae. Bot. Jahrb. Syst. 97: 1–60.
Lundberg, J. 2001. The asteralean affinity of the Mauritian Roussea (Rousseaceae). Bot. J. Linn. Soc. 137: 267–
276.
Lundberg, J., Bremer, K. 2003. A phylogenetic study of the
order Asterales using one morphological and three
molecular data sets. Intl J. Pl. Sci. 164: 553–578.
Patel, R.N. 1973. Wood anatomy of the dicotyledons indigenous to New Zealand. 2. Escalloniaceae. N. Z. J. Bot. 11:
421–434.
Praglowski, J., Grafström, E. 1985. The genus Carpodetus (Escalloniaceae): a pollen morphological enigma.
Grana 24: 11–21.
Royen, P. van 1983. The alpine flora of New Guinea, vol. 4.
Vaduz: Cramer.
Tobe, H., Raven, P.H. 1999. Floral structures of Alseuosmiaceae, Argophyllaceae, Carpodetaceae, Phellinaceae
and Rousseaceae: additional members in Asterales.
In: Abstract Volume XVI International Botanical
Congress, St. Louis, Missouri Botanical Garden,
Abstract 19.13.3., p. 241.
Williams, G., Adam, P. 1994. A review of rainforest pollination and plant-pollinator interactions with particular
reference to Australian subtropical rainforests. Austral.
Zool. 29: 177–212.
Compositae
Compositae Adans., Fam. Pl. 2: 103 (1763), nom. alt. et cons.
Asteraceae Martynov, Tekhno-Bot. Slovar: 55 (1820), nom. cons.
A.A. Anderberg, B.G. Baldwin, R.G. Bayer, J. Breitwieser, C. Jeffrey, M.O. Dillon,
P. Eldenäs, V. Funk, N. Garcia-Jacas, D.J.N. Hind, P.O. Karis, H.W. Lack, G. Nesom,
B. Nordenstam, Ch. Oberprieler, J.L. Panero, C. Puttock, H. Robinson, T.F. Stuessy,
A. Susanna, E. Urtubey, R. Vogt, J. Ward and L.E. Watson
Introduction with Key to Tribes
C. Jeffrey
Herbs, annual or biennial and monocarpic or
perennial and polycarpic or sometimes monocarpic, subshrubs, shrubs or less often trees,
leptocaul or sometimes pachycaul, often especially
when herbaceous or suffruticose with tuberous
roots or rhizomes or lignotuberous rootstock,
sometimes lianas (vines), usually terrestrial, rarely
epiphytic or aquatic, sometimes succulent, usually
with one or more of various types of glandular
and eglandular hairs, commonly the glandular
biseriate and the eglandular uniseriate; tissues
with schizogenous secretory canals (resin-ducts)
and/or with articulated lacticifers; nodes (1-)3- to
multilacunar. Leaves alternate or opposite, rarely
whorled, usually simple but often lobed or divided,
exstipulate; stomata anisocytic or anomocytic.
Unit inflorescence a capitulum (head), with rare
exceptions surrounded by an involucre of one
to several series of protective bracts (phyllaries),
capitula sometimes solitary at the apices of more
or less leafless stems (scapes) but usually few
to very many in often corymbiform cymose
synflorescences (inflorescences, capitulescences)
of various types, sometimes aggregated into often
involucrate capituliform syncephalia (glomerules)
of the second or even third order. Receptacle
paleate with persistent or caducous vascularized
scales (paleae, pales, chaff) subtending some or all
of the florets, fimbrilliferous with non-vascularized
fimbrils or scale-like processes surrounding the
bases of the florets, setulose, hairy or naked and
then smooth, areolate with polygonal areoles or
alveolate with depressions in which the florets are
inserted. Flowers (florets) small, 1–500 or more per
capitulum, sessile or subsessile; ovary inferior, of 2
(rarely 3) united carpels, unilocular, with 1 erect,
basal ovule; ovule anatropous, tenuinucellate,
unitegmic; calyx represented by a pappus formed
of (1–)2 to many awns, scales (squamae, squamulae), setae or hairs (rays) in 1 or more series,
homomorphic or heteromorphic, or by a more
or less coroniform or auriculiform structure, or
completely absent, never green and herbaceous;
corolla gamopetalous, of (3–)5(–6) united petals,
more or less regular (actinomorphic) and equally
or unequally (3–)5(–6)-lobed or -toothed with the
lobes or teeth valvate, or filiform with the lobes
reduced or absent or with a minute ray, or variously
zygomorphic, bilabiate with a 2-lobed internal
(adaxial) lip and a 3-lobed external (abaxial)
lip, pseudobilabiate with an unlobed internal
(adaxial) lip and a 4-lobed external (abaxial) lip,
ligulate with an apically 5-dentate abaxial ligule,
or radiate with an abaxial 0–3(–4)-dentate ray1 ,
the different types variously arranged within the
capitulum, the florets either all alike (homomorphic, isomorphic, capitulum homogamous) and
all regular (capitulum discoid), all ligulate (ligulate
capitulum) or all bilabiate, or of more than one
type (heteromorphic, anisomorphic, capitulum
heterogamous) with the inner (disc florets) regular (or rarely bilabiate) and perfect (bisexual,
hermaphrodite) or functionally staminate (male)
and the outer (ray florets) radiate, often pistillate
(female) or sometimes sterile (neuter), in one
or more series (capitulum radiate), or the outer
filiform pistillate, usually in several series, and the
inner regular, perfect or functionally staminate
(capitulum disciform), rarely the corolla absent
from the pistillate florets, occasionally all the
florets pistillate or staminate and the plants dioecious or monoecious, rarely the florets variously
otherwise arranged. Stamens with the filaments
inserted on the corolla-tube, equal in number to
and alternating with the corolla-lobes; filaments
1
In descriptions, the disposition of the corolla-lobes is sometimes
indicated by a formula, giving the number of lobes forming the
adaxial and abaxial parts of the limb respectively, e.g. ligulate
(0/5), pseudobilabiate (1/4), radiate (0/3 or 0/4), bilabiate (2/3),
regular (5/0).
62
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
usually free, rarely connate, the upper part of
the filament usually with cells with thickened
walls, forming a split cylindrical or balusterform
anther-collar (or filament-collar); anthers united
into a tube surrounding the style, very rarely free,
dithecal, tetrasporangiate or rarely bisporangiate,
introrse, dehiscent by longitudinal slits, usually
with an apical appendage, rounded, sagittate or
tailed at the base; tapetum integumentary; pollen
mostly tricolporate, usually echinate (spiny),
sometimes echinolophate or lophate (with a pattern of raised ridges) or spinulate (microechinate,
spinulose), often caveate; nectary a thickened scale
or cup surrounding the style base; style solitary,
elongating through the anther-tube and extruding
the pollen at its summit, apically divided (except
sometimes in functionally staminate florets) into
2 (or rarely 3) short to long branches (style arms)
with stigmatic areas on their inner (adaxial) surfaces, the apices of the style arms acute to rounded,
truncate or variously appendaged; stigmas dry,
papillose. Fruit unilocular, 1-seeded, indehiscent,
usually an achene (cypsela), very rarely a drupe,
crowned by the persistent pappus or the pappus
caducous or absent; abscission scar surrounded by
a carpopodium, distinguished by the form of its
cells and the texture of its surface, of 1 to many
rows of cells, indistinct to prominent, sometimes
apparently absent; embryo straight; endosperm
scant, forming a thin layer around the embryo.
The largest family of flowering plants, cosmopolitan except for Antarctica, with over 1,600
genera and 23,000 species (excluding apomictic
microspecies), especially well represented in grassland, wooded grassland and montane vegetation,
comparatively few in humid tropical lowland
forests.
Vegetative Morphology. The majority of
Compositae are subshrubs, shrubs or perennial
herbs, adapted to moderately xeric conditions, but
most life forms are represented, from ephemeral
desert annuals, pyrophytes and hemicryptophytes
to trees of 30 m or more, including especially in
Senecioneae (Fioretto and Alfani 1988) leaf and
stem succulents, also halophytes, marsh plants,
lianas, epiphytes and aquatics, though the latter
two life forms are comparatively infrequent and
submerged aquatics rare.
Herbaceous members of the non-asteroid
tribes growing in areas where grazing pressure
is significant are often thistleoid, i.e. with spiny
leaves and/or involucral bracts; this syndrome is
mostly found in the tribes Cynareae, Gundelieae
and parts of Arctotideae, and to a lesser extent in
Cichorieae. Its absence from the asteroid tribes
probably reflects the generally greater development
of chemical defences in the latter. Subshrubby
forms are usually evergreen, shrubs and trees
usually also evergreen but sometimes deciduous.
Pachycaul, megaphytic, polycarpic or sometimes
monocarpic trees, shrubs or herbs occur widely
on tropical mountains and oceanic islands, e.g.
Centaurodendron (Cynareae, Juan Fernández
Islands), Sonchus (Cichorieae, Macaronesia),
Dendroseris (Cichorieae, Juan Fernández Islands),
Espeletia (Millerieae/Heliantheae s.l., Andes), Argyroxiphium (Madieae/Heliantheae s.l., Hawaiian
islands), Lachanodes (Senecioneae, St. Helena)
and Dendrosenecio (Senecioneae, tropical east
African mountains).
Underground storage organs are common in
perennial herbaceous and shrubby forms and may
be represented by thickened taproots, root tubers,
tuberous rhizomes (or rarely a corm) or a lignotuber, the last especially common in pyrophtyic
and other geoxylic forms of tropical grasslands and
wooded grasslands.
The leaves of Compositae are exstipulate and
commonly alternate, but opposite leaves are characteristic of most Heliantheae s.l., the Liabeae and
some Vernonieae, and occur sporadically in several other tribes, e.g. Astereae and Senecioneae.
The leaves may be petiolate or sessile; the lower
leaves in many herbaceous perennials frequently
have an attenuate petioloid base. All the leaves may
be of more or less equal size or, as often in perennial herbs, the lower and basal may be larger than
the upper and may form a rosette; caulirosulate
forms also occur. The lamina is usually simple, although often lobed or divided, sometimes repeatedly and very deeply so, rarely truly compound
with separate leaflets. In Mutisia (Mutisieae), the
leaf-tip is modified into a tendril, in Cissampelopsis
(Senecioneae) and a few related genera, the petioles
are prehensile. The leaf venation may be parallelopinnate, pinnate, palmato-pinnate or palmate. The
presence of domatia has been demonstrated (Cerana and Ariza Espinar 1995) in Mikania (Eupatorieae/Heliantheae s.l.). Hydathodes are common.
Vegetative Anatomy. The anatomical features
of Compositae are subject to considerable homoplasy and are therefore only rarely group-definitive
at or above the generic level. Like that of the leaves,
the stem anatomy of Compositae (Carlquist 1966)
Compositae
reflects the fact that Compositae in general are
adapted to moderately dry conditions. The pith is
sometimes hollow, sometimes septate and often
lignified. Herbaceous stems have generally a single
ring of collateral bundles which may be closely
spaced or even form a complete cylinder in the
more woody members; medullary bundles are
frequently recorded and cortical bundles may
also occur. Secondary growth is usually present,
sometimes anomalous, especially in scandent
(e.g. Mikania, Eupatorieae/Heliantheae s.l.) or
secondarily woody forms. In general, the wood of
Compositae is more or less indistinguishable from
that of other higher asterid families. Most woody
Compositae have normal wood, some (e.g. Dendroseris, Cichorieae, Dendrosenecio, Senecioneae,
Chrysanthemoides, Calenduleae) have features
indicative of secondary woodiness, derived from
herbaceous ancestors. The vessels occur singly or
in groups, commonly in radial chains or tangential
bands. The vessel elements usually have simple
perforation plates, occasionally some of them are
scalariform or reticulate; the pits in the vessel
element walls are alternate, often somewhat larger
on the walls facing parenchyma. Helical sculpturing is often present on the vessel walls. Libriform
fibres are present, they are wide to narrow, thickor thin-walled and occasionally septate; tracheids
and fibre-tracheids are absent. Wood parenchyma
is scant and mostly paratracheal, usually only
one layer thick. Multiseriate and uniseriate rays
commonly co-occur, the former almost always the
more abundant, 4–10(–18) cells wide; in a few
Mutisioideae only uniseriate rays occur; a few
genera have rayless wood. The multiseriate rays
are heterocellular with both vertically and radially
elongated cells. The ray cells are wide to narrow,
with thin, unlignified or more commonly thick,
lignified walls, often with conspicuous pits. Storied
wood structure is rather common; it may take
the form of patches of storied fibres or vessels,
or may extend to all the elements of the axial
xylem; in Brachylaena (Tarchonantheae/Mutisieae
s.l.), Gochnatia (Gochnatieae/Mutisieae s.l.) and
Olearia (Astereae), the rays are also storied.
Growth rings are common, usually expressed as
wider vessel elements at the start of a growth ring.
Prismatic crystals are present in a few genera of
Mutisioideae, Astereae and Anthemideae, usually
in the ray parenchyma. Resin-like deposits are
common, usually as minute droplets in the cell
lumina. The nodes are usually 3–multi-lacunar,
rarely unilacunar.
63
The leaves of Compositae are occasionally
glabrous but both glandular and non-glandular
hairs (trichomes) are frequently present, the basic
types being biseriate glandular and uniseriate
non-glandular. Glandular hairs may be superficial or depressed in grooves or pits; they have
a 1–2-multiseriate stalk and an unicellular or
multicellular head. The following types of nonglandular hairs have been documented (Metcalfe
and Chalk 1950): unicellular to uniseriate base and
a long, flagelliform terminal cell (a very widespread
type); likewise but with an incipiently to unequally
or equally 2-armed terminal cell with the arms
ascending or parallel to the epidermal surface (also
very widespread); stellate with uniseriate stalk
and branched terminal cell; candelabriform with
several tiers of stelliform branches (e.g. Scorzonera,
Cichorieae); bladder-like, with inflated terminal
cell; multiseriate, shaggy (e.g. Vernonia, Vernonieae); and peltate, scale-like (e.g. Olearia, Astereae).
Many of these trichome types commonly occur
also on the stems and various floral parts. Although
subject to much homoplasy, indumentum types
can provide taxonomically useful data. In many
Heliantheae s.l., the hair bases are surrounded by
a rosette of silicified cells, making the leaf surface
rough to the touch. Stomata are mesogenous and
usually anisocytic or anomocytic. The hydathodes
of Compositae (Lersten and Curtis 1985) are
always marginal, usually on the teeth, and are of
a rather unspecialized type; the water-stomata
are similar in size to the ordinary stomata, but
are usually sunken and are permanently open.
The mesophyll is usually dorsiventral, but may be
isolateral or radial (e.g. in Werneria, Senecioneae);
ecologically specialized features, such as an aqueous or lignified hypodermis and bundle sheath
extensions, are commonly found. Kranz anatomy
is found in Heliantheae s.l. in a monophyletic
group (subtribe Chrysanthellinae) of seven genera
in Coreopsidodineae, and also in Pectis and some
species of Flaveria. The petiole usually has a single
arc of separate vascular bundles.
Two internal secretory systems are widespread
in Compositae (Carlquist 1966, 1976) – articulated
lacticifers (or lacticiferous cells) with triterpenerich latex, and schizogenous secretory canals (resin
ducts) usually lined with an epithelium. Lacticifers
are found in the phloem (or pericycle), secretory
canals in the cortex or the region of the endodermis, usually associated with the vascular bundles
(opposite or between them), occasionally also in
the rays. In the asteroid tribes, secretory canals
64
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
are usually present in the roots, rhizomes, aerial
stems and leaves (absent in Gnaphalieae), often
in the pericarp and sometimes in the cotyledons
(in Senecioneae), and lacticiferous cells are rare.
In the non-asteroid tribes, various expressions of
both lacticifers (or lacticiferous cells) and secretory canals are found. Lacticifers are characteristic of Cichorieae, where secretory canals are rare
(found in the roots of Scolymus and Scorzonera),
and Gundelieae; they are also found (or are replaced
by lacticiferous cells) in some genera of Mutisieae
(e.g. Gongylolepis), Cynareae (e.g. Cardopatium,
Atractylis, Carlina [Carlininae] and many genera
of Carduinae [Xeranthemum group]), Arctotideae
(e.g. Gazania) and Vernonieae (e.g. Vernonia) and
in most Liabeae. In a few genera of Barnadesioideae
and Mutisioideae, both secretory systems are apparently absent. Secretory cavities, often forming
translucent dots or streaks on the leaves and sometimes other organs (e.g. phyllaries), may supplement or replace the secretory canals.
Floral structure and Development. In the
majority of Compositae, the capitulum (Harris
1999) is the functional flower and it usually acts
as a single attraction unit. Its structure reflects
a balance of various functions concerned with pollination, the breeding system, dispersal, seed germination and defence (Stuessy and Garver 1996).
However, reduction and aggregation of capitula
into syncephalia which function as attraction units
of the second or even third order (Claßen-Bockhoff
1992, 1996) is not infrequent. Like primary capitula, these may be provided with floral (ray
floret) or extrafloral (coloured phyllary) pseudocorollas. Second-order syncephalia are common
in Gnaphalieae and the subtribe Nassauviinae
of Mutisieae and also occur, e.g. in CynareaeEchinopsinae, Inuleae and Heliantheae. Thirdorder syncephalia are exemplified by Platycarpha
(? Arctotideae), Gundelia (Gundelieae), Triplocephalum (Inuleae) and Lagascea (Heliantheae). In
Gundelia, for example, the one-flowered primary
capitula are grouped into numerous secondary
capitula of 5–7 primary capitula which in turn are
grouped into a large syncephalium of the third
order (Claßen-Bockhoff et al. 1989).
The onset of capitulum development (Harris
1995; Palmer 1996) is marked by a broadening of
the apical meristem. Phyllaries, florets and receptacular bracts (when present) are initiated by subsurface periclinal cell divisions. When receptacular bracts are present, they are frequently initiated
as part of a common primordium with the corresponding floret (e.g. in Tithonia, Heliantheae and
Xeranthemum, Cynareae); they may also be initiated after the appearance of the floral primordium
(e.g. in Madia, Madieae/Heliantheae s.l.); rarely,
the receptacular bract is initiated first, in the axil
of which the floret primordium then appears (e.g.
in Rudbeckia, Heliantheae). Receptacular bracts
may also be formed late in development, as enations from the receptacular surface (e.g. in Chrysopsis, Astereae); such bracts are probably not homologous with those of the other developmental
types. The receptacular outgrowths characteristic
of most Cynareae are likewise initiated in middevelopment, and are not homologous with receptacular bracts (paleae) in terms of initiation time.
Receptacular bracts (Stuessy and Spooner 1988)
are found in some Mutisieae, some Cynareae (Carduinae), some Vernonieae, some Inuleae, many Heliantheae s.l. and occasionally in other tribes (e.g.
Cichorieae, Liabeae, Astereae and Anthemideae).
The order of floret initiation and differentiation on the receptacle is closely associated with
capitulum type (Harris 1995). In homogamous capitula, the order is almost always strictly acropetal.
In heterogamous capitula, the order is often mixed,
partly acropetal and partly basipetal. On the basis
of primordium type, three groups of corolla types
may be recognized:
1. bilabiate, ligulate and regular (disc) perfect florets – primordia radially symmetrical, usually
pentagonal-circular; early stages of development similar; initiation acropetal;
2. ray and radiant florets – primordia triangular
or bilaterally symmetrical; delayed in development and sometimes also in initiation with
respect to the disc florets;
3. filiform florets, apically truncate, lobed or
with a minute ray – primordia consistently
circular and small; corollas abbreviated; all
traces of stamens completely absent; initiation
and/or development acropetal, basipetal,
bidirectional or synchronous.
Ray-floret primordia are similar in the early stages
in both the asteroid and the non-asteroid tribes in
which rays occur. The development of the Mutisia
type of ray floret differs least from that of the perfect
types of floret.
In floral development (Leins and Erbar 1987;
Harris 1995), the organogenetic order is 1, corolla,
2, androecium, 3, gynoecium. The pappus may be
initiated at any stage but it is seldom the first organ
Compositae
to be initiated. When pappus initiation is integral
with development, three basic types of initiation
are observed: sequential, in 5 (or more) sites alternate with the corolla-lobes; random, in available
space; and as a ring meristem, from which many
individual pappus primordia subsequently develop
(a more derived state found, e.g. in Astereae and Inuleae). Alternatively, the pappus may be initiated
late as enations around the summit of the ovary,
during the stage of floral expansion and differentiation. Such pappi are probably not homologous
with those of the other developmental types.
The corolla is initiated as a ring meristem
(meristematic rim or ring wall) which develops
before the more or less simultaneous initiation
of the corolla-lobe primordia upon it (Leins and
Erbar 2000), although in the non-asteroid tribes
an irregular successional development of the
corolla-lobe primordia has been reported in the
development of bilabiate and ligulate corollas (Harris 1995). After initiation of the stamen primordia
alternate with the corolla-lobes, intercalary growth
(which occurs in the ring meristem zone when it
is present) carries the stamens upwards, forming
the corolla-stamen tube. The true corolla-tube
(above the point of insertion of the stamens) is
formed independently earlier in development. In
the asteroid tribes, stamen primordia initiation
may be simultaneous or in a spiral sequence; in
the non-asteroid tribes, an irregular helical type of
stamen initiation is commonly found.
The vascular structure of the flower (floret) of
Compositae is relatively simple, and in perfect florets usually conforms to the following pattern. At
the base of the ovary, the single floret vascular bundle divides into six, one of which goes to the ovule,
the other five, in the ovary wall, to the summit of
the ovary, where two of them each send a branch
to the style and style arms. Each of the five traces
then splits, sending one branch to a stamen and
one to the top of the corolla to a sinus between adjacent corolla-lobes; here the corolla bundles fork
to unite in pairs at the apices of the lobes. These
corolla bundles represent the fused lateral bundles
of adjacent petals; median petal traces are absent.
Simplification of this pattern may be observed,
with reduction of the ovary wall bundles from 5 to 4
(as in Bidens, Coreopsideae/Heliantheae s.l.) or 2
(as in Lactuca, Cichorieae). More complex patterns
are also found, with more bundles in the ovary
wall and median corolla-lobe bundles present.
Sometimes a ring of 10 bundles (5 fused laterals
and 5 median bundles) is found in the ovary
65
wall, inside of which 4 carpellary bundles extend
into the style. Such and similar more complex
patterns are exhibited by some Barnadesioideae
(e.g. Schlechtendalia), Stifftieae (e.g. Stenopadus,
Wunderlichia) and Vernonieae (e.g. Proteopsis),
and here represent probable plesiomorphic states
(the 10-bundle wall condition is also found in
Calyceraceae). More complex venation patterns
sometimes occur in other tribes (e.g. Heliantheae),
are here frequently associated with an increase in
size of the achene and/or corolla, and probably
represent derived conditions. However, increase in
size with retention of the simple 5-bundle pattern
is also found (e.g. as in Tragopogon, Cichorieae).
The adaxial corolla epidermal cells exhibit
a number of microstructural surface ornamentation patterns which, especially in ray and ligulate
corollas, are broadly correlated with systematic
position and functionally with the pollinator
attraction mechanism – the reflexion of visible and
ultraviolet light, and the nature and distribution
of visible and ultraviolet light-absorbing pigments
(Baagøe 1980; Hansen 1991b).
In Compositae, the stamens are typically syngenesious, with connate anthers (rarely free, in
some wind-pollinated forms) and free filaments
(very rarely partly or wholly connate). The upper part of the filament forms an anther-collar (or
filament-collar), usually distinguished by the size,
shape and wall thickness of its cells, in the form
of an adaxially split cylinder or baluster (Meiri and
Dulberger 1986). The anthers usually bear an apical
appendage, variable in size and proportions and
also consistency; it may be lignified (as in Barnadesioideae, Stifftieae, Mutisieae and Cynareae)
or soft (as in Lactuceae, Arctotideae and the asteroid tribes); occasionally it is absent. At the base, the
anthers may be calcarate (with a considerable portion of the thecae protruding below the filament
insertion point, i.e. below the top of the anthercollar) or ecalcarate (with basal insertion of the
filament). Calcarate anthers are typical of the nonasteroid tribes, ecalcarate of the asteroid tribes,
although they also occur, e.g. in a few Mutisieae
s.l. (Mutisieae, Gochnatieae and Dicomeae). The
base of the anther may be rounded, acute, sagittate or prolonged on each side into a sterile tail,
which varies from fine and unbranched to stout and
branched. The endothecial cells of the pollen-sacs
have walls which are variously thickened; generally
the thickenings are polarized or lateral, with special
states occurring in Catananche (Cichorieae), other
Cichorieae and some Cynareae (Carduinae). Ra-
66
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
dially disposed thickenings are the most common
in Eupatoriodinae (Eupatorieae), but also occur in
some other Heliantheae s.l. (e.g. Dahlia, Coreopsideae/Heliantheae s.l.).
The nectary (when present) is commonly
a thickened scale or cup alongside or surrounding
the style base at the top of the ovary; it is provided
with numerous stomata, through which the nectar
is secreted (Galetto 1995).
The style in Compositae is single, and in perfect florets consists of a shaft divided at the apex
into 2 (rarely 3) style arms. The shaft is usually glabrous, rarely hairy and may be uniform in diameter or swollen at the base (stylar node). The inner
(adaxial) faces of the style arms bear the stigmatic
surfaces. The stigmatic area is dry and papillose
and may cover the entire inner surface of each style
arm or form two marginal bands which vary from
widely separated to contiguous, from broad to narrow and from free to united towards the apex of the
style arm. Entire stigmatic areas are general in the
non-asteroid tribes, marginal characteristic of the
asteroid tribes, although this feature is subject to
reversal. The style arms themselves vary in many
characters, including: orientation (tangential or radial to the capitulum); disposition (erect to connivent or reflexed); length; degree of fusion; shape
of apex; presence or absence of a sterile appendage
above the stigmatic areas; form of apical appendage
(when present); presence or absence of sweeping
hairs on the outer (abaxial) surface; presence or absence of an articulation (swelling) at the base of the
style arms; distribution and form of the sweeping
hairs on, and character of the outer surface (papillose or smooth) of the style arms and/or upper
part of the shaft. In pistillate or functionally pistillate florets, the stylar structure is generally much
simpler and in staminate or functionally staminate florets, the style is commonly undivided at
the apex and devoid of stigmatic areas; such styles
may exhibit various elaborations of the sweepinghair mechanism. Particular style and style arm features are characteristic of many subtribes, tribes or
groups of tribes, and their variation is a reflexion
of the evolutionary development and diversification of the pollen-presentation mechanism (Thiele
1988). Appendaged style arms are especially characteristic of Astereae (papillose and broadly conical
to subulate) and Eupatoriodinae of Heliantheae s.l.
(highly developed and conspicuous).
1987; Ahlstrand 1988, 1992) is fairly uniform, although variation in detail occurs which may be of
taxonomic significance as, e.g. in the infratribal
classification of Anthemideae and Arctotideae and
in the tribal classification of Carduoideae. The
ovule is anatropous, unitegmic and tenuinucellate;
usually a solitary vascular bundle enters the
chalaza and penetrates the integument to a greater
or lesser extent. The micropyle is usually short.
In some tribes (e.g. Cynareae, Cichorieae, Calenduleae, Heliantheae), the inner epidermis of the
integument differentiates into an integumentary
tapetum. The archesporium is usually unicellular
but sometimes multicellular or rarely 2-cellular.
The megaspore tetrad is usually linear, rarely
T-shaped; most often the chalazal megaspore
develops, in a few species the micropylar. The
monosporic 8-nucleate Polygonum type of embryo
development is most frequently found, the bisporic
Allium type less frequently and almost exclusively
in the asteroid tribes (with the exception of Heterolepis, ? Arctotideae); various tetrasporic types
have also been recorded. Variation may be found
even within one and the same species. In apomicts,
development may be bisporic (Taraxacum type),
tetrasporic (e.g. Antennaria type) or aposporic
(Hieracium type). Fertilization is porogamous,
double and premitotic. Endosperm development is
usually cellular, less often (as in Cynareae and some
representatives of some other tribes) nuclear, usually without haustoria. Both types may sometimes
occur in one and the same genus. Embryogenesis
is of the asteroid type with a few variations, the
basic variant being the Senecio variant.
The stamens of Compositae are usually
4-locular, rarely 2-locular. The walls of the loculi
develop centrifugally, and consist of epidermis,
endothecium (with cell wall thickenings), a middle
layer which is ephemeral, and the tapetum. The
tapetum is initially cellular and multinucleate,
then partial dissolution of the cell walls occurs;
in the post-meiotic stage, it becomes amoeboid,
sometimes forming a false periplasmodium;
eventually it forms the pollenkit and is absent
from the mature anther. The archesporium may be
uniseriate or multiseriate, tetrad formation is of
the simultaneous type and the tetrads tetrahedral,
isobilateral or cruciform. The mature pollen grains
are solitary and 3-celled. Abnormal meiosis is
often observed in apomictic species.
Embryology and Pollen development. Embryogenesis in Compositae (Batygina and Yakovlev
Palynology. The pollen in Compositae (Stix
1960; Skvarla et al. 1977) is tricolporate or some-
Compositae
times triporate, usually echinate or spinulate,
generally sphaeroidal with a transverse furrow in
the endexine perpendicular to each colpus and
a tectate ektexine. A cavea (space), where the
columellae are detached from the foot layer in the
regions between the apertures or colpi, is often
present. The ektexine generally consists of a foot
layer, columellae (bacula) and a tectum. There
is considerable and systematically significant
variation on this basic pattern, especially in
surface ornamentation and exine stratification.
General evolutionary trends are an increase in the
surface relief and in the porosity of the ektexine,
both a reflection of an evolutionary increase in the
capacity of the pollen grain wall to hold substances
important in pollen presentation and pollination.
Spinulate pollen grains are plesiomorphic in
Compositae, and are shared with Calyceraceae and
some other related families (Hansen 1991a). They
occur in Barnadesioideae (DeVore and Stuessy
1995; Urtubey and Telleria 1998), in Mutisioideae
and some Carduoideae. The intracolpar concavities
found in some Calyceraceae and Barnadesioideae
may represent a synapomorphy for the two families. The apomorphic echinate condition appears
to have arisen independently at least three times
– in Dicomeae (Mutisieae s.l.) in the Pleiotaxis
group of genera, in Cynareae (Serratula type), and
in the vernonioid group of tribes (Cichorioideae
and Asteroideae). Lophate pollen, in which the
surface is ridged in a polygonal pattern, is found –
either in the echinolophate (with echinate ridges)
or the psilolophate (with smooth ridges) variant –
in some Barnadesioideae and in some members of
the tribes Arctotideae, Vernonieae and Cichorieae
(Blackmore 1986). Its occurrences in these taxa
are also examples of parallel or convergent evolution; the lophate condition must be considered
derivative, as it depends upon the acquisition of
a mechanism for the differential deposition of callose which is absent in the spinulate and echinate
forms. It is of functional significance in, e.g. the
storage and release of exine-held substances and
in its mechanical attributes. Its absence from the
asteroid tribes may possibly be correlated with the
development of the double tectum in the latter.
Two main types of exine stratification can be
recognized, lactucoid (found in the non-asteroid
tribes) and asteroid. The lactucoid type exhibits
one or more of the following features (Vezey
et al. 1994): branched infratectal columellae (e.g.
Schlechtendalia, Barnadesioideae; Onoseris, Mutisieae); infratectal columellae that appear to be
67
continuations or branches of the basal columellae
(e.g. Gerbera, Mutisieae) and/or more than one
layer of internal tectum (e.g. Carthamus, Cynareae;
Arctotis, Arctotideae); slender, widely separated
basal columellae (e.g. Tragopogon, Cichorieae)
or thick but widely separated basal columellae
(e.g. Vernonia, Vernonieae; Liabum, Liabeae).
Lactucoid pollen is generally ecaveate, having
basal columellae which arise from the foot layer
and one to several levels of slender columellae
and internal tecta above the basal columellae, but
caveate pollen is found in Arctotideae and in some
Liabeae. Asteroid pollen is generally caveate, lacking basal columellae and having instead a cavity,
the cavea, and is characterized by a double tectum.
The double tectum is a synapomorphy of the asteroid tribes and does not occur in the non-asteroid
tribes (except perhaps in Liabeae as a parallelism).
In some asteroid tribes (e.g. Calenduleae, parts
of Inuleae and Heliantheae), more complex tectal
structural elements occur, which appear to be
transformations of the double tectum. In most Anthemideae (except, e.g. Ursinia), the pollen grains
are ecaveate and have a thick foot layer with basal
columellae; however, these structures are not homologues of similar structures in lactucoid pollen,
though in the past the lactucoid forms were also
often referred to as anthemoid. Lactucoid pollen
is generally larger than asteroid pollen, and has
a smaller diameter/exine thickness ratio, 6.2–7.5 as
opposed to 8.0–10.0, except for 6.4 in Anthemideae.
Independent of this larger-scale variation in
exine structure, in some tribes of Asteroideae (and
rarely also in the non-asteroid tribes) minute internal foramina are present in the structural elements of the ektexine; pollen with such foramina
is commonly found in members of Heliantheae s.l.
and has become known as the helianthoid type; it
is also found in a few genera of Senecioneae, e.g.
Doronicum, Pericallis, Packera (Bain et al. 1997),
but the majority of Senecioneae have pollen devoid
of such internal foramina, and such asteroid pollen
is referred to as the senecionoid type.
The great variation in pollen form and
structure in Compositae provides much taxonomically valuable information, and various
pollen-type groups have been proposed by various authors. In general, the following groups
are characterized by particular palynological features (or transformations of them):
(1) Barnadesioideae, (2) Mutisieae s.l. (StifftieaeMutisieae s.s.-Dicomeae-Tarchonantheae-Pertyeae)-Echinopsinae, (3) other Cynareae, (4) Arc-
68
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
totideae, (5) Moquinieae-Gundelieae-VernonieaeLiabeae-Cichorieae, (6) Inuleae, (7) Gnaphalieae,
(8) Senecioneae-Calenduleae-Astereae-Heliantheae
s.l., and (9) Anthemideae. On a more detailed
level, particular pollen types may characterize one
or more subtribes, groups of genera or genera;
only rarely is more than one type found within
a particular genus, e.g. Munnozia (Liabeae).
Pollination. Compositae are characterized by
secondary pollen presentation (Thiele 1988; Leins
and Erbar 1990; Erbar and Leins 1995; DeVore and
Stuessy 1995) in which the pollen is shed into the
anther-tube by introrse dehiscence of the syngenesious anthers, extruded by action of the style and
presented on the outer surface of the style shaft
and style arms, which then open to expose the stigmatic areas on their upper (adaxial) surfaces. In
some, the filaments are insensitive and the pollen
is presented merely by upgrowth of the style; in
others, the filaments are sensitive (thigmotropic)
and contract on being touched, so that the anthers
withdraw and the pollen is exposed for transport.
In all, the anther-collars, the abaxial and lateral epidermal cells of which have lignified walls, support
and guide the style. Three variants of the presentation mechanism may be recognized; the pollen may
be dragged out by adhesion to papillae-like microhairs on the style exterior, brushed out by sweeping
hairs on the outside of the style arms and style shaft
or on an appendage to the style arms, or pumped
out by spreading sweeping hairs on the apices of
the style arms. The drag variant is characteristic
of Barnadesioideae, Stifftieae and some Mutisieae
and Carduoideae, the brush variant of most Carduoideae, all Cichorioideae and some Asteroideae,
and the pump variant of most Asteroideae.
The pollinator rewards are mainly nectar and
pollen. The nectary is situated at the style base and
the nectar is secreted through stomata. The nectar
is sugar-rich (hexose or sucrose); in some, it contains amino acids which attract flies, or sometimes
pyrrolizidine alkaloids which attract Lepidoptera
(Brown 1984). The pollen is lipid-rich. The tapetum forms pollenkit which plays an essential role
in the presentation and transfer of pollen. It keeps
the pollen grains attached to the style, holds them
together in clumps, facilitates adhesion to the pollinator, acts as an attractant to the pollinator, and
facilitates adhesion to the stigma, the surfaces of
which are papillose and dry, lacking exudate.
The majority of Compositae are adapted
to specific pollinators and are not generalists,
as sometimes previously thought. They exhibit
intricate and precise mechanisms of pollinator
attraction, species identification, pollinator rewards and reward presentation (Lane 1996). Some
Barnadesioideae, Mutisioideae, Dicomeae and
Cynareae with elongated corollas are pollinated
by hummingbirds or sunbirds, others by longtongued insects such as sphingids. Pollination
by flies is characteristic of Compositae of some
montane areas, such as the Andes and the Southern
Alps of New Zealand. However, the most important pollinators of Compositae are solitary bees,
with which a special relationship has developed,
including learned recognition of species and orientation cues and learned manipulation for reward
extraction. A few Compositae are wind-pollinated,
for example, the genera Artemisia (Anthemideae)
and Ambrosia (Heliantheae) and some species of
Espeletia (Millerieae/Heliantheae s.l.; Berry and
Calvo 1989).
Karyology. Haploid chromosome numbers in
Compositae range from 2 to 120, although very
high polyploids and very low numbers are infrequent, and species with n = 9, 10, 12, 17 or 18
form 50% of all Compositae for which chromosome
numbers are known. The most common number is
9, and x = 9 is possibly the basic (plesiomorphic)
number for the family as a whole. Variation in chromosome number has been brought about mainly
by the processes of allopolyploidy (including amphidiploidy), aneuploid change and chromosome
loss (Solbrig 1977). Dysploid reduction is most
characteristic of Compositae (especially annuals)
of more arid and/or disturbed areas, and appears
to be correlated with speciation under such conditions. Polyploidy is more characteristic of mesic,
perennial herbaceous and woody species occupying relatively stable habitats.
Breeding Systems. The majority of Compositae are sporophytically self-incompatible; the incompatibility is strong but not absolute and selfcompatibility is not uncommon; a few cases of capitular cleistogamy are known. The perfect florets
are protandrous but the capitulum as a whole is protogynous when the outer florets (ray or filiform) are
pistillate or functionally so. Autogamy may occur
with floret age. The following capitular conditions
are met with: hermaphroditism (monocliny); gynomonoecy, monoecy, androdioecy, gynodioecy,
dioecy and andromonoecy. Monoclinous capitula
are homogamous (monomorphic) with all the flo-
Compositae
rets (disc, bilabiate or ligulate) perfect; outbreeding
is facilitated by the floral protandry and the centripetal maturation of the florets. Dioecious capitula are also monomorphic but either entirely pistillate or entirely staminate; outbreeding is achieved
by spatial separation of the capitula, which occur on
different plants. The remaining capitular types are
heteromorphic (heterogamous), most commonly
with the inner florets perfect or functionally staminate and the outer (filiform or ray) florets pistillate. The gynomonoecious condition, in which
there are one or more outer rows of pistillate florets and central perfect florets, is very common;
outbreeding is facilitated by the capitular protogyny of the outer florets. The monoecious condition is similar, but here the central florets are functionally staminate. The androdioecious, gynodioecious, andromonoecious and various intermediate
conditions are less frequently observed. Apomixis
is common in a number of genera, such as Hieracium, Pilosella and Taraxacum (all Cichorieae;
den Nijs and Menken 1996), Blumea (Inuleae) and
Antennaria (Gnaphalieae).
Fruit and Seed. Variation in fruit and seed characters provides much taxonomically useful information (Reitbrecht 1974; Grau 1980; Sáenz 1981;
Velez 1981; Reese 1989; Short et al. 1989; Eriksson
1991; Dittrich 1996; Werker 1997). The fruit is oneseeded, normally an achene (sometimes but superfluously termed a cypsela), very rarely a drupe,
usually regarded as indehiscent but often with preformed dehiscence lines opening by pressure of the
germinating embryo. Achenes may be terete, angled or variously dorsiventrally obcompressed or
laterally compressed, unribbed or with 2–many ribs
of various degrees of prominence, apically truncate
to variously attenuate or beaked, and vary in the
form of the apical part of the fruit (where the pappus, when present, is inserted) and in the form of
the carpopodium. Within a capitulum, the achenes
may be all alike (homomorphic) or heteromorphic
(of 2 or 3 kinds); such heterocarpy is quite frequent
and has been recorded, for example, in 39 genera
of the tribe Cichorieae (Voytenko 1989a; Voytenko
and Oparina 1990). The main structural elements
of the fruit are pappus (when present), pericarp,
testa, endosperm and embryo.
The pericarp epidermis (epicarp) is uniseriate,
sometimes papillate or otherwise surface-ornamented, sometimes lignified and commonly bears
hairs of various types – uniseriate barnadesioid
in Barnadesioideae, multicellular in Echinopsinae,
69
one-celled in most other Cynareae, in other
tribes twin hairs which are sometimes myxogenic;
various glandular hairs may also be present. The
mesocarp may consist of various elements. A hypodermis of one to several layers of sclerified or
unsclerified, parenchymatous to prosenchymatous
cells is often developed. Commonly, sclerenchyma
is present to a greater or lesser extent; it may occur in discrete bundles or bands or form a continuous ring; often it surrounds the vascular
bundles. In Heliantheae s.l., a phytomelanin layer
is usually found between the hypodermis and the
sclerenchymatous tissue. Schizogenous secretory
canals (resin ducts) may be present, usually
associated with the vascular bundles. The inner
part of the mesocarp and the inner epidermis of
the pericarp (endocarp) may be present at achene
maturity and the endocarp even lignified, but
commonly one or both become obliterated.
The testa may also become disorganized at
maturity but in a number of tribes, its cellular
structure is maintained and then the form of the
cells of the testa epidermis and the type of thickening of their walls provide taxonomically useful
variation as, for example, in the tribes Mutisieae
s.l. (Stifftieae, Gochnatieae, Mutisieae, Dicomeae),
Cynareae, Cichorieae and, to a lesser extent, Anthemideae and Inuleae. Endosperm is constantly
present; it forms a layer 1–3 cells thick enveloping the embryo, and consists of living cells rich in
oil and protein but devoid of starch. Its cell walls
are thickened and composed mainly of mannose
(Grau and Hopf 1985). The endosperm apparently
serves as an additional protective layer for the embryo. The embryo is large, straight, dicotyledonous
(rarely syncotyledonous) and with hypocotyl and
radicle developed. The seed is rich in protein and
oil but poor in oligosaccharides and starch; they
form respectively 10–48, 4–70, 2–4 and 0–2% of the
dry mass.
The pappus in its manifold forms has a protective function and may also act as a dispersal mechanism when the achene is mature. Other specialized
structures associated with dispersal, such as wings,
hooks and the elaiosomes of some Cynareae, may
also be developed.
Dispersal. The disseminules of Compositae are
dispersed mainly by wind, sometimes externally by
mammals, less often by ants or other agents. The
disseminule is usually the achene, sometimes enclosed by a phyllary or receptacular bract. In other
cases, the capitulum is the unit of dispersal, the
70
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
achenes (one or more) being retained within the involucre. Such synaptospermy is most characteristic
of Compositae of semi-arid and arid areas (Gutterman and Ginott 1994). It may be partial (as in Crepis
aculeata, where the inner achenes are dispersed
singly and the outer together, or Didelta spinosa
where the outer are dispersed singly and the central
remain in the capitulum) or complete. In Didelta
carnosa, several-fruited capitula fragments act as
disseminules, while in Gundelia tournefortii the
disseminules are one-fruited capitula of the second
order (Claßen-Bockhoff 1996); the lignified involucre is composed of the phyllaries of the 4–6 sterile
lateral capitula. The capitula of Compositae suffer
strongly from phytophagous insects, most of which
attack the fruits. In synaptospermy, the protection
afforded by the testa and pericarp is reinforced by
(or transferred to) more peripheral organs in which
the achenes are enclosed. Wind dispersal may be
facilitated by various forms of capillary pappus, by
compression of the achene, by the development of
wings or (in synaptospermous species) by the development of chaffy involucral bracts. Corky ribs
characterize some water-dispersed achenes, elaiosomes those dispersed by ants. The development
of barbs or hooks on the pappus or other parts
of the achenes or, in synaptospermous species, on
the involucre (such as in Arctium, Cynareae) is frequent in those Compositae usually dispersed by
mammals. In heterocarpic Compositae, the different achene morphs may have different dispersal
strategies as well as different dormancy and germination requirements, and may thus enable the
species to take advantage of a wider range of temporal and environmental conditions.
Phytochemistry. Compositae exhibit a rich
and varied assortment of secondary metabolites,
which serve as storage compounds or as chemical defence mechanisms, the development of
which has undoubtedly been important in the
evolutionary success of the family. The combined
occurrence of inulin-type fructans, acetylenic fatty
acid derivatives transformed into cyclic (aromatic
or heterocyclic) compounds and sesquiterpene
lactones is almost as characteristic of the family as
is the capitulum (Hegnauer 1977).
Inulins, synthesized as a homologous series of
β-2-1-linked polymers and stored in solution in
the vacuole, are the main storage carbohydrates of
Compositae.
The acetylenes (formerly referred to as polyacetylenes on account of their multiple acetylenic
bonds; that term is now restricted by chemists
to denote polymers of acetylene) of Compositae
(Zdero and Bohlmann 1990) and their derivatives
are nemacidal, antibiotic and toxic, and occur
mainly in the resin ducts. The most common
acetylene is the C13 pentaynene, but this is absent
in Cichorieae, Astereae and Anthemideae; in the
latter two tribes (except for Ursinia and a few
related genera of Anthemideae), it is replaced
by particular C10 and C17 acetylenes. The C17
acetylenes are also found in Cichorieae and
Senecioneae, in the former another C10 acetylene
also occurs. Thiophene derivatives of the C13 pentaynene occur in some Heliantheae s.l. (including
Tagetodinae [Tageteae]), Inuleae, Arctotideae,
Gochnatieae, Cynareae (including Echinops)
and Mutisieae, while spiroketalenol ethers (also
present as thiophene derivatives) are characteristic
of Anthemideae. In their acetylene chemistry,
Mutisieae and Gochnatieae resemble Cynareae,
while acetylenes with pyran or furan attachments
are found in Cynareae, Anthemideae, Inuleae and
Heliantheae s.l. and a special type is present in
Gnaphalieae. The presence of phytomelanin in
the pericarp is characteristic of Heliantheae s.l.,
although secondarily absent in some members and
occurring as a parallelism in a few Vernonieae.
Sesquiterpene lactones (Seaman 1982) are
bitter and toxic and occur in the latex, in the subcuticular cavities of glandular hairs or extruded in
glandular hairs. The two most significant aspects
of their chemistry are the patterns of skeletal and
substitutional diversity; superimposed upon each
skeleton is a set of substituents which collectively
define a specific compound (Seaman and Funk
1983). Three levels of biogenetic complexity may be
recognized. The sesquiterpene lactone chemistry
of the non-asteroid tribes is generally less complex
than that of Asteroideae; the main structural types
are guaianolides and germacranolides; eudesmanolides are relatively infrequent and only the
bourbonolides of Vernonieae represent the highest
level of biogenetic complexity. The most commonly occurring types are as follows: Stifftieae:
cis-lactonized germacranolides and eudesmanolides; Mutisieae (in Nassauviinae): lactonic and
non-lactonic isocedrene derivatives; Gochnatieae,
Dicomeae, Pertyeae, Tarchonantheae (all Mutisieae
s.l.): germacranolides, cis-cis germacradienolides,
trans-lactonized eudesmanolides (in Gochnatieae
and Dicomeae) and guaianolides; Cynareae:
guaianolides and germacranolides; Cichorioideae: guaianolides (uncommon in Arctotideae),
Compositae
germacranolides (in Moquinieae, Vernonieae,
Cichorieae and Liabeae), trans-lactonized eudesmanolides (in Moquinieae) and heliangiolides
(in Vernonieae). In Asteroideae, sesquiterpene
lactones are absent from Heliantheae-Tagetodinae
(Tageteae) and Calenduleae and rare in Astereae;
they are elaborated in Inuleae, many Heliantheae
s.l. (including Eupatoriodinae [Eupatorieae]),
Gnaphalieae and Anthemideae. Senecioneae are
very distinct in their development of furanoeremophilanes (also oxidized to lactones). The
commonly occurring types are as follows: Astereae
(rarely): eudesmanolides and eremophilanolides;
Anthemideae: eudesmanolides, guaianolides and
germacranolides; Inuleae (including Gnaphalieae):
eudesmanolides, germacranolides, xantholides
and guaianolides, and less frequently ambrosanolides, helenanolides, pseudoguaianolides,
seco-pseudoguaianolides and seco-helenanolides;
Heliantheae s.l.: ambrosanolides, helenanolides,
heliangiolides, germacranolides, melampolides,
eudesmanolides, guaianolides, xantholides and
seco-ambrosanolides; and Senecioneae: furanoeremophilanes and aromatic furanoeremophilanes.
An 8-α-oxygen function is characteristic of the
germacranolides of Mutisioideae, Carduoideae,
Vernonieae, Cichorieae and other non-asteroid
tribes. In Heliantheae s.l., there is an 8-β function, often (in Eupatoriodinae [Eupatorieae])
an oxygenated C5 ester. In Anthemideae, the
C8 position can be free or functionalized. The
guaianolides of Cynareae are mainly 6,12-olides
with an 8-α-oxygen function whereas in Inuleae
and Heliantheae s.l., 8,12-lactones are widespread
and again the β orientation is favoured. The presence of eudesmanolides (santanolides), especially
6,12-olides and often as 11,13-dihydro derivatives,
is characteristic of Anthemideae whereas 8,12eudesmanolides predominate in Heliantheae s.l.
and Inuleae. Sesquiterpene lactones are apparently
absent from Barnadesioideae.
Other Compositae chemical constituents of
note include triterpenes (which occur free in the
latex or in the lipid fraction of tissues), steroids,
essential oils, carotenoids, flavonoids, chromenes
(benzopyrans) and benzofurans (prenylated
acetophenones, Proksch and Kunze 1996), lignans
and other phenolics, labdanes and kauranes
(chemical defences), coumarins, prenylated
coumarins, thymol derivatives, diterpenoids, alkaloids, cyanogenetic glycosides (both valine- and
phenylalanine-derived), inositol esters, isobutyl
amides (insecticidal) and seed oils. The isomeric
71
cyclitols L-inositol and scyllitol are accumulated
together. The terpenoid essential oils of Compositae are complex mixtures of steam-volatile
constituents which occur in secretory glandular
hairs and the resin ducts; components include
acetylenes, phenylpropanols and chromenes,
thymol derivatives, monoterpenoids, sesquiterpenes (widespread) and (in Ursinia and related
genera of Anthemideae) furanosesquiterpenes.
Among the flavonoids (Bohm and Stuessy 2001),
quercetagetin-based yellow flavonols are characteristic of Compositae; gossypetin-based flavonols
are rare. Yellow anthochlor pigments (chalcones
and aurones), resorcinol-based, occur in the
Heliantheae-Coreopsidodinae (Coreopsideae) and
some other Heliantheae s.l., and phloroglucinolbased in some Cynareae (e.g. Carthamus) and
Gnaphalieae (e.g. Helichrysum and Gnaphalium).
Anthocyanins are important contributors to flower
colour, e.g. in such ornamentals as Chrysanthemum, Dahlia and Callistephus. Cyanidin is
the major anthocyanin, delphinidin occurs only
occasionally. The anthocyanin pattern is relatively
simple, acylation is rare and methylation absent.
The phytochemistry of Compositae is a rich
source of potentially valuable taxonomic information; the co-occurrence (or co-absence)
of specific chemical compounds (or groups of
compounds) is often indicative of taxonomic
affinity at the subfamily and lower levels (Zdero
and Bohlmann 1990). For example, 5-methyl
coumarins and onoseriolides are present in both
Mutisieae-Mutisiinae and Mutisieae-Nassauviinae
(Zdero et al. 1986). Mutisieae as a whole are of
interest in showing some chemical similarities to
the asteroid, rather than to the non-asteroid tribes.
For instance, p-hydroxyacetophenones, prenyl or
geranyl coumarins and nerolidol derivatives occur
in Mutisieae and Asteroideae but are absent from
Cichorioideae (except for the presence of nerolidol
derivatives in Vernonieae) and Carduoideae;
however, the presence of 5-alkyl coumarins and
chromenes is shared with Vernonieae, whereas
isocedrene derivatives are unique to Mutisieae.
Labdanes, clerodanes and their derivatives occur in Astereae, Inuleae, Heliantheae s.l. and
Senecioneae, abietanes in Inuleae and Heliantheae,
beyeranes and their derivatives in all Asteroideae
except Astereae, thymol derivatives in all Asteroideae except Senecioneae and most Astereae,
prenyl phloroglucinols in Inuleae and Heliantheae,
furanosesquiterpenes in Cynareae and Vernonieae, and (possibly as a parallelism) in Ursinia
72
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
and related genera of Anthemideae. Inuleae and
Heliantheae are also linked by their acetylene
and sesquiterpene lactone chemistry, as are
Gochnatieae, Tarchonantheae, Dicomeae, Pertyeae
(all Mutisieae s.l.) and Cynareae; Anthemideae
and Astereae are linked by their acetylene chemistry and the co-occurrence of umbelliferone
derivatives. Calenduleae relate to Inuleae and
Heliantheae by the co-occurrence of special
diterpenes and prenylated p-oxyacetophenones.
In the rich chemistry of Compositae lies the basis
of their very widespread use as medicinal plants.
Subdivisions and Relationships Within the
Family. Compositae can be divided into two
monophyletic groups of markedly unequal size –
the small monotribal South American subfamily
Barnadesioideae and the rest (Jansen and Palmer
1987, 1988; Jansen et al. 1991; Bremer and Jansen
1992; Kim et al. 1992; Kim and Jansen 1995;
Panero and Funk 2002). Within the rest (here
called non-barnadesioid Compositae), a number
of putatively monophyletic groups have been
identified, the largest of which, the asteroid group
of tribes (subfamily Asteroideae) appears to be
the most derived (Bremer 1987, 1994; Kadereit
1989; Kim et al. 1992); it consists of the tribes
Inuleae (including Plucheeae), Heliantheae s.l.
(including Eupatorieae), Gnaphalieae, Astereae,
Anthemideae, Calenduleae, Corymbieae (Panero
and Funk 2002) and Senecioneae. Putatively sister
to this group of tribes is the cichorioid group
of tribes (subfamily Cichorioideae sensu stricto)
in which may be included the tribes Gymnarrheneae (Panero and Funk 2002), Moquinieae
(Robinson 1994), Vernonieae, Liabeae, Cichorieae
(syn. Lactuceae), Gundelieae (Karis et al. 2001)
and Arctotideae. Together, Asteroideae and Cichorioideae constitute the vernonioid group of
tribes, for the monophyly of which there is strong
molecular as well as considerable morphological
support. Tentatively, two further subfamilies may
be recognized, these being Carduoideae with the
tribes Gochnatieae, Hecastocleideae, Dicomeae,
Cynareae and Pertyeae, and a basal Mutisioideae.
Possibly, Mutisioideae and Carduoideae together
form the sister group of the vernonioid group of
tribes (i.e. Cichorioideae plus Asteroideae) (Bayer
and Starr 1998) but it is probable that they are
paraphyletic with respect to the vernonioid group
(Jansen and Palmer 1988; Jansen et al. 1991; Kim
et al. 1992, 2002; Karis et al. 1992; Panero and Funk
2002).
Barnadesioideae (Gustafsson et al. 2001;
Urtubey and Stuessy 2001) are distinguished by
a number of unique morphological features, such
as the abundant 3-celled hairs – each with a short,
cup-shaped basal cell, a rounded intermediate
cell (Erbar and Leins 2000) and a longer, straight
terminal cell – on all the floral parts, and unique
types of testa-epidermis (Grau 1980) and pollen
grains (Urtubey and Telleria 1998; Zhao et al. 2000).
Non-barnadesioid Compositae are distinguished
by the presence of a cpDNA inversion absent
from Barnadesioideae and all other angiosperms
(Jansen and Palmer 1987), and by the presence of
twin hairs on the achenes and a bristly pappus,
although these morphological features are subject
to reversal, transformation and loss. Mutisioideae
share with Barnadesioideae a number of plesiomorphies, such as the mutisioid and crested,
smooth types of petal adaxial epidermis patterns
and spinulate pollen grains (Hansen 1991a, b). The
subfamily Mutisioideae can be divided into two
main groups – a basal, perhaps paraphyletic group
(Stifftieae) with short (or rarely long, Hyaloseris
type), rounded, essentially glabrous style arms
of the Wunderlichia, Stifftia and Quelchia types,
and a second tribe Mutisieae s.s. (subtribes
Mutisiinae, Gerberinae and Nassauviinae) with
short to long, spreading-hairy style arms of
the Mutisia, Trixis and Nassauvia types (Jeffrey
1967). Carduoideae, perhaps characterized by the
constancy of the Gochnatia type of testa epidermal
cell wall thickening (Grau 1980) or derivative
types thereof (Dittrich 1996; Häffner 2000), share
a number of plesiomorphies with Mutisioideae;
the more plesiomorphic members (Gochnatieae,
Hecastocleideae, Tarchonantheae) are marked by
short to long or lipped style arms of the Gochnatia
and Seris (Richterago) types, the more advanced
by short to long, finely to coarsely hairy style arms,
often with the upper part of the stylar shaft slightly
expanded (Pertya, Dicoma and Pleiotaxis types).
The more plesiomorphic members (tribe Pertyeae)
of the latter group retain the mutisioid types of
petal adaxial epidermis patterns, the rest (including the tribes Dicomeae and Tarchonantheae
and the uncertainly placed genus Adenocaulon,
see Katinas 2000) exhibit tabular, senecionoid or
rugose patterns (Hansen 1991b), but all retain the
plesiomorphic spinulate pollen, except for a few
tropical African genera (Pleiotaxis and its allies) of
Dicomeae which have apomorphic echinate pollen.
The vernonioid group of tribes (i.e. Cichorioideae and Asteroideae as here circumscribed) is
Compositae
marked by a 9-base pair deletion in the ndhF gene
(Kim and Jansen 1995), and is strongly supported
as a monophyletic group also by other molecular
(e.g. Jansen and Kim 1996) and by morphological
data (Karis et al. 1992). Within this group, Cichorioideae, although fairly well supported by molecular data, are poorly characterized morphologically
(the presence of long, acute style arms is perhaps
a synapomorphy but, if so, it must be considered as
subject to reversal). Asteroideae are strongly supported as monophyletic not only by molecular data,
including a duplication in the rbcL gene (Kim et al.
1992), but also morphologically, e.g. stigmatic areas in two marginal bands and disc floret corollas
regular, with short lobes, although again these features are subject to reversal.
The basal branches of non-barnadesioid
Compositae, of the tribal groups within them, and
of many of the subfamilies, tribes and subtribes
themselves tend to be poorly resolved in both
morphological and molecular cladistic analyses.
This is indicative of rapid evolution in the early
stages of diversification of the taxa concerned,
no doubt consequent upon the attainment of new
arogenetic levels providing the basis for rapid
adaptive radiation and cladogenesis. Improvement
in the efficiency of the pollen presentation and
pollination mechanisms as well as innovations
in chemical defences appear to have played
particularly important roles in this respect. The
sister-group relationships of the tribes within Carduoideae, Cichorioideae and Asteroideae had been
largely unresolved (Bremer 1996; Jansen and Kim
1996; Bayer and Starr 1998) but Panero and Funk
(2002) have recently clarified tribal relationships
within the subfamilies, based on a yet to be fully
published study of 13,000 base pairs of chloroplast
DNA from 120 taxa across the family. There is
some evidence (molecular, morphological and
palynological) that, in Cichorioideae, Moquinieae
are close to Vernonieae, and that Vernonieae
and Liabeae may be sister groups; they are also
linked to Cichorieae by the common presence of
similar (vernonioid) style arms and the tangential
orientation of the gynoecium (Robinson 1984).
Cichorieae and Gundelieae are also sister groups
(Karis et al. 2001; Panero and Funk 2002). The
sister-group relationships of Arctotideae are unclear, although the tribe is undoubtedly a member
of Cichorioideae.
In Asteroideae (Karis 1993b), there is fairly
good evidence (chemical, molecular and morphological) that Anthemideae and Astereae may
73
be sister groups. Inuleae (including Plucheeae)
may be sister to Heliantheae s.l., a relationship
more supported by molecular and chemical than
by morphological data. That Anthemideae and
Astereae together form a clade with Calenduleae
and Gnaphalieae is fairly well supported by some
molecular evidence; the sister-group relationships
within this clade have been clarified by Panero
and Funk (2002). Senecioneae stand apart from
the other tribes of Asteroideae on both molecular
and chemical grounds, and their sister-group
relationships remain unresolved.
Subtribal delimitation within Compositae is
currently in a state of flux and, in some tribes,
satisfactory and generally accepted subtribal taxonomy is yet to be established. Currently, it would
appear more prudent to recognize at subtribal level
only the more well-established major clades within
each tribe, to treat smaller, clearly delimited putatively monophyletic groups within them as informal, infrasubtribal groups of genera, and to leave
unclassified below the tribal level those genera the
affinities of which have to date not been established. In Cynareae, recent analyses (Kim et al.
1992; Susanna et al. 1995; Wagenitz and Hellwig
1996; Petit 1997; Häffner and Hellwig 1999; Häffner
2000; Petit et al. 2000; Garcia-Jacas et al. 2002), although giving somewhat conflicting results, have
demonstrated the paraphyly of Carduinae, as previously constituted, with respect to Centaureinae as
previously recognized; thus, only Carlininae, Cardopatiinae, Echinopsinae and the Carduinae in the
broad sense are at present well-established taxa.
The three subtribes of Liabeae are based largely
on the results of morphological cladistic analysis
but molecular data are as yet lacking. Apparently
basal in Vernonieae is the yellow-flowered genus
Distephanus but otherwise, owing mainly to the
paucity of our understanding of the Old World
representatives, the subtribal situation remains unclear (Keeley and Jansen 1995; Keeley and Chan
2003), in spite of recent studies (Robinson 1996,
1999a, b). In Cichorieae, well-defined at the moment, both morphologically and molecularly, are
only the subtribes Scolyminae, Crepidinae sensu
lato and Scorzonerinae; possibly, the large subtribe
Crepidinae sensu lato (i.e. including all the other
subtribes often recognized) may prove to be subdivisible into a number of smaller monophyletic
subtribes; especially the New World representatives
(Microseridinae sensu lato) and Hyoseridinae appear to have some evidence in favour of their recognition at subtribal level (Whitton et al. 1995; Koop-
74
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
man et al. 1998; Kim et al. 1999a; Lee et al. 2003). Of
the other cichorioid tribes, Liabeae and Gundelieae
are small enough to be considered monosubtribal,
while the subtribes Gorteriinae and Arctotidinae
of Arctotideae are more or less supported by both
molecular and morphological (including developmental) data (Karis et al. 2001; Funk and Chan
2003).
Within Inuleae sensu lato, various groups at
subtribal level have been recognized (Anderberg
1991b, c) but, as in other tribes, it is probable
they will need considerable reassessment when
the results of more detailed morphological and
molecular cladistic analyses become available.
In recent years, the plucheoid genera of Inuleae
sensu lato have been treated by some authorities
as constituting a distinct tribe, Plucheeae. In
molecular analyses, however, Inuleae sensu stricto
and Plucheeae always appear as sister groups (Kim
and Jansen 1995; Eldenäs et al. 1998, 1999), and
thus the monophyly of Inuleae is not violated by the
inclusion in it of the plucheoid genera (Anderberg
et al. 2005). Many small entities have been recognized at subtribal level in Heliantheae s.l. (Stuessy
1977; Turner and Powell 1977; Robinson 1981),
including Eupatoriodinae as treated at tribal rank
(King and Robinson 1987), but further molecular
data have shown that these schemes were in need
of modification, as some of the entities recognized
appeared to be paraphyletic or even polyphyletic
(Karis 1993a; Jansen and Kim 1996; Panero et al.
1999; Schilling et al. 1999; Schmidt and Schilling
2000; Baldwin and Wessa 2000a, b; Ito et al.
2000a, b; Baldwin et al. 2002, 2003). Recognition
of putatively monophyletic taxa such as the
Blepharispermum group (Eriksson 1991) and the
subtribes Heleniinae, Coreopsidinae, Eupatoriinae
and Tagetinae (including Flaveriinae) left what was
an admittedly paraphyletic residue – Helianthinae
(including Ecliptinae) – which required further
analysis, now reflected in the treatments of the
helianthoid tribes in this volume. Subtribal delimitation in Asteroideae is proceeding. Molecular
studies in the Heliantheae alliance have provided
clear evidence of subtribal relationships. With the
exception of Tageteae (Tagetodinae), Eupatorieae
(Eupatoriodinae) and Coreopsideae (Coreopsidodinae), subtribal delimitation is robustly
established. In Gnaphalieae, the six subtribes (Anderberg 1991a) recently recognized appear to need
considerable reassessment in the light of evidence
from recent morphological and molecular studies
(Bayer and Puttock 1999; Bayer et al. 2000, 2002,
2003). Astereae have been variously divided into
subtribes on morphological grounds (Zhang and
Bremer 1993; Nesom 1994c), but neither scheme
is wholly supported by molecular studies (Lane
et al. 1996; Noyes and Rieseberg 1999; Noyes
2000); austral genera appear to be basal, indicating
a possible Gondwanan origin for the tribe, as is
also the case for their putative sister tribe, Anthemideae; subtribes recently recognized (Bremer
and Humphries 1993) appear in several cases not
to represent monophyletic groups; again, austral
(African) genera appear to be basal (among them
the chemically distinctive Ursinia and its relatives)
whereas the northern-hemisphere genera form
a derived, monophyletic group (Anthemidinae
sensu lato; Francisco-Ortega et al. 1997; Watson
and Evans 1999; Oberprieler and Vogt 2000;
Watson et al. 2000). Calenduleae are small enough
to be considered monosubtribal, whereas further
molecular and morphological analyses are needed
to clarify the subtribal delimitation and relationships, and the affinities of problematical genera
such as Packera, within Senecioneae (Swenson
and Bremer 1999). The reference of Corymbium
to a distinct monogeneric tribe, Corymbieae, has
recently been established (Panero and Funk 2002).
The tribal and infratribal systematics adopted
in this work is that accepted by the respective
authors of the tribes concerned, and at the tribal
rank departs from the system presented in this
introduction (Jeffrey 2004) in that D.J.N. Hind
treats the tribes Stifftieae, Mutisieae, Gochnatieae,
Hecastocleideae, Tarchonantheae, Dicomeae and
Pertyeae within a broadly circumscribed Mutisieae
sensu lato, and J.L. Panero the supersubtribes
i–xii and D.J.N. Hind and H. Robinson the supersubtribe xiii, of Heliantheae s.l., respectively
as the tribes Athroismeae, Helenieae, Coreopsideae, Neurolaeneae, Tageteae, Chaenactideae,
Bahieae, Polymnieae, Heliantheae (sensu stricto),
Millerieae, Madieae, Perityleae and Eupatorieae,
i.e. the tribes accepted in the subfamilial and tribal
classification of Compositae given by Panero and
Funk (2002), in which 11 subfamilies and 35 tribes
are accepted. Here, it is considered preferable
to retain the familiar, broad circumscription of
Heliantheae of Cronquist (1955) and Robinson
(1981), but with the addition of Eupatorieae, shown
to be an ingroup of such a broadly circumscribed
Heliantheae by recent molecular studies (Kim and
Jansen 1995; Baldwin et al. 2002; Panero and Funk
2002). A few other genera are placed otherwise by
the authors of the tribal accounts.
Compositae
Family Affinities. The affinities of Compositae have been clarified in recent years by molecular taxonomic studies (Lammers 1992; Olmstead
et al. 1993; Chase et al. 1993; Michaels et al. 1993;
Gustafsson et al. 1996; Jansen and Kim 1996; Wagenitz 1997; Albach et al. 2001; see also introduction to this volume), supplemented by morphological cladistic analyses (Gustafsson and Bremer
1995; Hansen 1997), and there is now strong evidence that Calyceraceae, Goodeniaceae (including
Brunoniaceae) and Menyanthaceae are successive
sister families of Compositae. These four families
together are closely related to Stylidiaceae (incl.
Donatiaceae) and a clade of Alseuosmiaceae, Phellinaceae and Argophyllaceae, and more distantly
related to Rousseaceae/Carpodetaceae, Campanulaceae (including Lobeliaceae) and Pentaphragmataceae (Lundberg and Bremer 2003).
The Menyanthaceae-Compositae group is
characterized by the presence of scalariform
perforation plates, the frequent occurrence of
sclerified idioblasts, the lack of endosperm haustoria, the presence of binucleate pollen grains
and binucleate tapetal cells, the production of
herbivore defences by the mevalonate pathway,
and the fusion of the lateral veins of adjacent
petals for some part of their length, before they
separate to join the mid-vein of each petal at
the apex of the corolla-lobe. Compositae, Calyceraceae and Goodeniaceae are united by the
presence of bifurcating columellae in the pollen
grain wall and the presence of anther-collars in
at least some members. With Calyceraceae, the
Compositae-Barnadesioideae share a number of
possible synapomorphies, such as pollen wall
structure, the aggregation of pollen by pollenkit,
some anther microcharacters, the unilocular ovary
and the type of pollen presentation mechanism.
History and Palaeontology. The fossil record
of Compositae is sparse and consists mostly of dispersed pollen grains. The earliest records are of
both spinulate (mutisioid) and echinate (probably
asteroid) pollen grains from the Palaeocene to midEocene of the west coast of South America, of spinulate grains from the Eocene of Chile and of echinate
grains from the Eocene of Brazil. Unfortunately, uncertainties of identification and/or dating attend all
these pre-Oligocene records, and the earliest, definitely identified and dated records of Compositae
pollen are from the early Oligocene of the western
USA, the mid-Oligocene of Chile and central Europe, and the late Oligocene of North America. The
75
late Oligocene also sees the first records of macrofossils – leaves, involucres and achenes of unknown
tribal and generic affinities. From the Miocene onwards, fossils become more abundant, diverse and
widespread (Graham 1996).
Because the Oligocene pollen records are of
comparatively advanced types and are found on
several different continents, it is reasonable to assume that the family dates back at least to the
Eocene-Oligocene boundary, i.e. 38 Ma b.p. This
accords well with the evidence for the age of the
split between Goodeniaceae and Calyceraceae plus
Compositae, which was probably connected with
the isolation of South America from Antarctica
and has been dated to 48 ± 5 Ma b.p. (DeVore
and Stuessy 1995). Substitution rate calculations
of chloroplast DNA rbcL gene sequences also give
at least 38 Ma b.p. as the age of Compositae (Bremer and Gustafsson 1997). Late Eocene tectonic
and climatic changes undoubtedly played a significant role in the early evolution of Compositae and
the geographical spread of Compositae during the
Miocene (10–20 Ma b.p.) was correlated with the
development of extensive dry to semi-arid ecosystems with seasonal precipitation. In this respect,
the storage of carbohydrate in the form of highly
soluble fructans in the vacuole, giving the plants the
ability rapidly to adjust vacuole size and therefore
osmotic potential, enabled Compositae to exploit
conditions of limited water availability under climates with seasonal rainfall or seasonally low soil
water temperatures (Hendry 1996). Coupled with
the development of effective dispersal mechanisms
and effective chemical defences against herbivores,
this enabled Compositae to flourish and spread
in the grassland and wooded grassland vegetation
types which became widespread during the late
Miocene, and pre-adapted them to the cooler, xeric
conditions which developed during the Pliocene.
Economic Importance. Compositae are prominent among the plants utilized by peoples in native
cultures in all parts of the world, especially for
medicinal purposes. For example, in China 500
(Huang and Ling 1996) and in Mexico 180 (Heinrich 1996) species of wild-growing Compositae
are currently employed in traditional medicine.
Preparations of Compositae are variously valued
for their antibiotic, antifungal, antihelmintic,
antiplasmodial, expectorant, sedative, diuretic,
spasmolytic, haemostatic, immunostimulatory or
anti-inflammatory properties. Wild Compositae
species are also used as foods, fish poisons, fodder,
76
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
and sources of oil and nectar. Industrial products
for which Compositae (cultivated or wild) provide
raw materials include insecticides, medicines,
soaps, detergents, varnishes, paints, cosmetics,
rubber, perfumes, food products, and flavourings
and colourants for foods and drinks (Garg and
Sastry 1996; Viswanathan and Singh 1996). Many
Compositae are also highly valued as ornamentals
in both commercial and recreational horticulture.
Among cultivated Compositae, the following
are among the more important of over 260 species
currently in cultivation for other than ornamental
purposes: Cynara cardunculus, cardoon and globe
artichoke, edible leaves and capitula; Carthamus
tinctorius, safflower, edible and industrial oil;
Cichorium intybus, chicory, leaf vegetable, roots
provide a coffee substitute and also yield fructans;
Cichorium endivia, endive, salad vegetable; Pterocypsela indica, Indian lettuce, salad vegetable;
Lactuca sativa, lettuce, salad vegetable; Scorzonera hispanica, Spanish salsify, scorzonera, root
vegetable; Acmella oleracea, Para cress, culinary
herb; Echinacea purpurea, purple coneflower,
medicinal, immunostimulatory; Helianthus annuus, sunflower, edible and industrial oil and
edible seeds, the most important crop plant of
the family; Helianthus tuberosus, Jerusalem artichoke, root vegetable; Smallanthus sonchifolius,
yacon, root vegetable; Artemisia dracunculus,
tarragon, flavouring and oil used in perfumery;
Tanacetum cinerariifolium, pyrethrum, yields
the insecticidal monoterpenes called pyrethrins;
Glebionis coronarium, crown daisy, green vegetable; and Petasites japonicus, butterbur, green
vegetable. Among the most important of the very
numerous genera cultivated as ornamentals are
Gerbera (Mutisieae, Barberton daisy); Dahlia
(Coreopsideae/Heliantheae s.l., dahlia); Tagetes
(Tageteae/Heliantheae s.l., French and African
marigolds), Xerochrysum (Gnaphalieae, everlasting), Callistephus (Astereae, China aster), Symphyotrichum (Astereae, Michaelmas daisy), Chrysanthemum (Anthemideae, florist’s chrysanthemum)
and Pericallis (Senecioneae, florist’s cineraria).
A number of Compositae are of negative economic significance, inasmuch as they are more
or less noxious weeds of gardens, cultivated
fields, pastures and plantations. Among the
more important weedy genera are Acroptilon,
Carduus, Cirsium, Centaurea and Onopordum (all
Cynareae), Chondrilla (Cichorieae), Parthenium,
Ambrosia, Chromolaena and Mikania (all Heliantheae s.l.), Chrysanthemoides (Calendulae) and
Senecio (Senecioneae). Parthenium hysterophorus
causes a contact dermatitis; the pollen of Ambrosia
species is a cause of hay fever, and many Senecio
species are highly hepatotoxic to grazing livestock.
Classification of Compositae
(Asteraceae)
I. Subfamily Barnadesioideae (D. Don) Bremer & Jansen
(1992).
1. Tribe Barnadesieae D. Don (1830).
Genera 1–9
Non-Barnadesioid Compositae
II. Subfamily Mutisioideae (Cass.) Lindl. (1829).
1. Tribe Stifftieae D. Don (1830).
Genera 10–12, 19–22, 77–78
2. Tribe Mutisieae Cass. (1819) s.s.
Genera 13–18, 24–47, 50–55, 57–59, 61–64, 66–72
Genera of uncertain placement
Genera 23, 48–49, 65
III. Subfamily Carduoideae Cass. ex Sweet (1829).
1. Tribe Gochnatieae (Benth.) Panero & V.A. Funk (2002).
Genera 56, 60, 73–75
2. Tribe Hecastocleideae Panero & V.A. Funk (2002).
Genus 76
3. Tribe Tarchonantheae Kostel. (1833).
Genera 80–82
4. Tribe Dicomeae Panero & V.A. Funk (2002).
Genera 83–87
5. Tribe Cynareae Lam. et DC. (1806) syn. Tribe Cardueae
Cass. (1819).
Genera 92–164
Genera of uncertain placement
Genera 165–166, 395
6. Tribe Pertyeae Panero & V.A. Funk (2002).
Genera 79, 88–91
Vernonioid group of tribes
IV. Subfamily Cichorioideae (Juss.) Chev. (1828).
1. Tribe Gymnarrheneae Panero & V.A. Funk (2002).
Genus 167
2. Tribe Moquinieae H. Rob. (1994).
Genera 168–169
3. Tribe Vernonieae Cass. (1819).
Genera 170–287
4. Tribe Liabeae (Cass. ex Dumort.) Rydb. (1927).
Genera 288–303
5. Tribe Cichorieae Lam. et DC. (1806).
Genera 304–389
6. Tribe Gundelieae DC. ex Lecoq et Juillet (1831).
Genera 390–391
7. Tribe Arctotideae Cass. (1819).
Genera 392–394, 396–408
V. Subfamily Asteroideae (Cass.) Lindl. (1829).
1. Tribe Corymbieae Panero & V.A. Funk (2002).
Genus 409
2. Tribe Senecioneae Cass. (1819).
Genera 410–559
3. Tribe Calenduleae Cass. (1819).
Genera 560–570
4. Tribe Gnaphalieae (Cass.) Lecoq et Juillet (1831).
Compositae
Genera 571–755
5. Tribe Astereae Cass. (1819).
Genera 756–960
6. Tribe Anthemideae Cass. (1819).
Genera 961–1071
7. Tribe Inuleae Cass. (1819).
Genera 1072–1137
8. Tribe Heliantheae Cass. (1819) s.l.
Genera 1138–1619
i. Supersubtribe Athroismodinae (Panero & V.A. Funk)
C. Jeffrey (2004) (Tribe Athroismeae)
Genera 1138–1143
ii. Supersubtribe Heleniodinae (Raf.) C. Jeffrey (2004)
(Tribe Helenieae)
Genera 1144–1156
iii. Supersubtribe Coreopsidodinae (Lindl.) C. Jeffrey
(2004) (Tribe Coreopsideae)
Genera 1157–1186
iv. Supersubtribe Neurolaenodinae (Rydb.) C. Jeffrey
(2004) (Tribe Neurolaeneae)
Genera 1187–1191
v. Supersubtribe Tagetodinae (Cass.) C. Jeffrey (2004)
(Tribe Tageteae)
Genera 1192–1223
vi. Supersubtribe Chaenactidodinae (Rydb.) C. Jeffrey
(2004) (Tribe Chaenactideae)
Genera 1224–1226
vii. Supersubtribe Bahiodinae (Rydb.) C. Jeffrey (2004)
(Tribe Bahieae)
Genera 1227–1246
viii. Supersubtribe Polymniodinae (H. Rob.) C. Jeffrey
(2004) (Tribe Polymnieae)
Genus 1247
ix. Supersubtribe Helianthodinae (Cass.) C. Jeffrey (2004)
(Tribe Heliantheae)
Genera 1248–1360
x. Supersubtribe Milleriodinae (Lindl.) C. Jeffrey (2004)
(Tribe Millerieae)
Genera 1361–1394
xi. Supersubtribe Madiodinae (Benth.) C. Jeffrey (2004)
(Tribe Madieae)
Genera 1395–1430
xii. Supersubtribe Peritylodinae (Rydb.) C. Jeffrey (2004)
(Tribe Perityleae)
Genera 1431–1437
xiii. Supersubtribe Eupatoriodinae (Cass.) C. Jeffrey (2002)
(Tribe Eupatorieae)
Genera 1438–1619
Asteroid genus of uncertain tribal position
Genus 1620
Descriptions of the Subfamilies
of Compositae
77
pseudoradiate or ligulate, sessile or pedunculate;
synflorescences variable. Receptacle pilose, rarely
with pales or glabrous. Involucre cylindrical to
hemispherical, phyllaries in several series. Florets 1 to numerous, isomorphic or anisomorphic,
generally hermaphrodite; corollas tubular, pseudobilabiate, bilabiate, subpseudobilabiate, ligulate
or subligulate, often villous. Stamens 5 (3–5 in
the central flowers); filaments free or rarely fused,
inserted at different levels; anthers ecaudate to
tailed, with apical appendage entire, emarginate
or bilobed. Pollen with or without depressions,
lophate, microechinate, scabrate-microechinate or
smooth. Style shortly bilobed or bifid, glabrous
or papillose below bifurcation. Achenes densely
villous, with straight simple hairs, rarely glabrous.
Pappus uniseriate or rarely absent.
Barnadesioideae are the smallest subfamily of
Asteraceae, with 91 species in 9 genera, endemic
to South America. This subfamily, earlier treated
at the subtribal level within Mutisieae, was newly
recognized by Bremer and Jansen (1992) based primarily on molecular restriction site (and subsequent sequence, e.g. Bayer and Starr 1998) analyses.
All Asteraceae (except for Barnadesioideae) have
a 22-kb inversion in cpDNA (Jansen and Palmer
1987, 1988). This not only helps define the subfamily, but also places it in a basal phylogenetic
position within the family. Diagnostic morphological features include the presence of axillary
spines, unbranched “barnadesioid” hairs (with an
epidermal cell very slightly differentiated and two
other cells, one short with thick walls and the other
longer with thin walls) on corollas, achenes, pappus and vegetative structures (Cabrera 1959, 1961;
Bremer 1987; Bremer and Jansen 1992), and frequent occurrence of pseudobilabiate corollas (Bremer 1987, 1994). Flavonoid profiles are quite simple (Bohm and Stuessy 1995). Phylogenetic analyses of intergeneric relationships based on morphology (Urtubey and Stuessy 2001) and DNA sequences (Gustafsson et al. 2001) have given incongruent results; more study is evidently needed. Biogeographic hypotheses for the subfamily suggest
a southern South American origin (Stuessy et al.
1996). Comprising only the tribe Barnadesieae.
I. Subfam. Barnadesioideae
(D. Don) Bremer & Jansen (1992).
Shrubs, trees, or perennial or annual herbs, usually
with fascicled nodal spines. Leaves variable, entire,
mostly pinnatinerved, many xeromorphic. Capitula homogamous or heterogamous, discoid or
II. Subfam. Mutisioideae
(Cass.) Lindl. (1829).
Shrubs, trees or perennial or rarely annual
herbs, rarely scandent. Leaves usually alternate,
78
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
usually simple, entire, serrate, denticulate or
lobulate. Capitula homogamous or heterogamous,
bilabiato-radiate, bilabiate or discoid, rarely ligulate. Involucre cylindrical to urceolate-subglobose,
phyllaries few- to many-seriate, imbricate, gradate.
Receptacle usually epaleaceous. Florets 1–many,
5-merous, marginal usually bilabiate and often
radiate, inner bilabiate or sometimes regular and
corolla deeply 5-lobed, rarely all florets regular or
ligulate; corolla-lobes long, with an apical tuft of
minute hairs, those of inner lip of bilabiate florets
coiled. Anthers calcarate and caudate, tails usually
long, often variously branched or fimbriate.
Pollen ecaveate, microechinate. Style shaft usually
glabrous, arms mostly short, obtuse to rounded
and glabrous, sometimes truncate and penicillate,
with stigmatic papillae covering all inner surface.
Achenes usually with twin hairs. Pappus usually
present, commonly of bristles, often uniseriate.
Mutisioideae comprise c. 60 genera and 720
species, mostly South American. They here are circumscribed in the sense of Hansen (1991b), except
that certain genera included by him in the subfamily (as tribe Mutisieae) are here transferred to
Carduoideae.
III. Subfam. Carduoideae
Cass. ex Sweet (1826).
Perennial, biennial or less often annual herbs,
shrubs or rarely trees, rarely scandent. Leaves
alternate, usually simple, entire, serrate, denticulate or lobulate, especially in herbaceous
members often spiny. Capitula homogamous or
heterogamous, discoid or discoid with marginal
florets sterile and radiant, rarely bilabiato-radiate,
radiate or ligulate. Involucre narrowly cylindrical
to urceolate-subglobose, phyllaries 3- to manyseriate, imbricate, often spiniferous. Receptacle
epaleaceous and very often setulose, rarely paleaceous. Florets 1 to many, 5-merous, all or inner
regular or subregular, outer sometimes radiant,
rarely bilabiato-radiate or radiate, very rarely all
ligulate; corolla-lobes long, those of inner lip of
bilabiate florets straight or with incurved apex,
very rarely coiled. Anthers calcarate and caudate,
tails usually long, sometimes pilose or fringed.
Pollen usually ecaveate, spiny or microechinate.
Style arms short to long, obtuse to rarely acute,
glabrous or with dorsal hairs, sometimes not
divergent for most of their length, with stigmatic
papillae covering all inner surface; style shaft often
with an articulation at or below the branching
point, marked by a ring of hairs and/or an increase
in diameter, glabrous below the articulation,
usually hairy above it. Achenes with twin hairs,
simple hairs or glabrous. Pappus usually present,
of bristles or scales, isomorphic or heteromorphic.
Carduoideae comprise at least 93 genera and
2,600 species, mostly in the Old World. The subfamily as here circumscribed includes all mutisioid
genera which in molecular studies come out above
Mutisieae and below the cichorioid-asteroid clade,
as well as Cynareae sensu lato. The constancy of the
Gochnatia type of testa epidermal cell wall thickening (Grau 1980) or its derivatives define the subfamily. Although this type also occurs, possibly as
a parallelism, in some Mutisieae, it is there associated with different style and/or pollen types.
IV. Subfam. Cichorioideae
(Juss.) Chev. (1828).
Perennial, biennial or annual herbs, shrubs or trees,
rarely scandent, very rarely aquatic. Leaves alternate or opposite, usually simple, entire to deeply
lobed, in herbaceous members sometimes spiny.
Capitula homogamous to heterogamous, discoid,
ligulate or radiate. Involucre narrowly to broadly
cylindrical or campanulate to subglobose, phyllaries (1-)2- to many-seriate, usually imbricate. Receptacle epaleaceous, often alveolate, or paleaceous.
Florets 1–many, usually 5-merous, all regular, all
ligulate or very rarely bilabiate, or inner regular
and outer bilabiate or radiate; corolla-lobes usually
long, not coiled. Anthers calcarate, caudate or ecaudate, tails usually simple. Pollen ecaveate or sometimes appearing caveate, spiny, sometimes echinolophate, psilolophate or lophate, globose. Style
arms commonly long, tapered, acute, hairy dorsally, sometimes shaft thickened apically and arms
short to long; stigmatic papillae covering all inner surface. Achenes with twin hairs. Pappus usually present, usually of bristles or scales, sometimes
heteromorphic.
Cichorioideae are here accepted in the sense
of Bremer (1994), comprise 241 genera and about
2,900 species, and are well represented in both the
Old and the New Worlds.
V. Subfam. Asteroideae
(Cass.) Lindl. (1829).
Perennial to annual herbs or shrubs, less often
trees, sometimes scandent, epiphytic or aquatic,
sometimes succulent. Leaves alternate or opposite,
Compositae
simple but not infrequently highly dissected, not
spiny. Capitula heterogamous or homogamous,
radiate, disciform or discoid, very rarely ligulate.
Involucres cylindrical to subglobose, phyllaries 1to few-, less often many-seriate, imbricate or eximbricate. Receptacle epaleaceous or paleaceous.
Florets 1 to many, (3–4–)5(–6)-merous, all regular,
or inner regular and outer radiate or filiform,
narrowly tubular and subtruncate or microradiate,
very rarely outer bilabiate or all ligulate; corollalobes short, deltoid, rarely long. Anthers almost
always ecalcarate, caudate or ecaudate, tails simple,
usually slender. Pollen almost always caveate,
spiny, with double tectum. Style arms short to
long, tapered and acute to obtuse or truncate, often
appendaged; stigmatic papillae mostly confined
to two marginal, distant to contiguous, apically
confluent or separate lines, rarely covering entire
inner surface of style arms. Achenes with twin
hairs, in many helianthoid members with a phytomelanin layer in the pericarp (“carbonized”).
Pappus present or absent, usually of bristles or
scales or coroniform, sometimes auriculiform or
of awns, sometimes heteromorphic.
Asteroideae, here accepted in the sense of
Bremer (1994), are the largest subfamily of
Compositae, comprising 1,210 genera and about
17,000 species, of which Heliantheae sensu lato
form 480 and c. 5,600 respectively. They are widely
distributed on all continents, except Antarctica.
Key to the Tribes and Tribally
Unplaced Genera2
1. Corolla inside and outside and all other floral parts
bearing simple, uniseriate, eglandular, 3-celled hairs,
the basal cell of which has a centrally depressed, distal
transverse wall
I. Barnadesieae (p. 87)
– Corolla inside and outside and all other floral parts
without such hairs, though often with hairs of various
other kinds
2
2. Plant a heterocapitular amphicarpic rosulate herb,
forming both subterranean cleistogamous and
subaerial chasmogamous capitula
IV. Gymnarrheneae (p. 147)
– Plant otherwise, if heterocapitular, then all capitular
forms subaerial
3
2
The tribes in this key (I–XXX) are those accepted by the
authors of the tribal accounts. Where a final lead ends in alternative tribes, the user should try the unknown in the keys
to genera of each tribe. If a plant to be identified is known
or thought to be a member of the Heliantheae alliance (tribes
XVIII–XXX), the user may go directly to the key to the corresponding tribes on p. 391. Stylar characters, unless otherwise
stated, relate to the styles of perfect florets.
79
3. Capitula heteromorphic, homogamous, with only pistillate (female) florets, or only functionally staminate
(male) or perfect (hermaphrodite) florets, rarely pistillate capitula with a few functionally staminate florets,
or staminate capitula with some marginal pistillate
florets
4
– Capitula homomorphic, heterogamous (radiate or disciform) or if homogamous (discoid), then all florets
perfect or rarely the outer enlarged and sterile (radiant)
16
4. Staminate and pistillate capitula borne on the same
plant (plants monoecious)
5
– Staminate (or perfect) and pistillate capitula borne on
different plants (plants dioecious or gynodioecious) 7
5. Herbs; achenes with a phytomelanin layer in pericarp
XXVI. Heliantheae (p. 440)
– Shrubs; achenes without a phytomelanin layer in pericarp
6
6. Capitula 1-flowered
V. Moquinieae (p. 148)
– Capitula many-flowered
XV. Astereae (p. 284)
7. Leaves opposite, achenes with a phytomelanin layer in
pericarp
8
– Leaves alternate, achenes without a phytomelanin
layer in pericarp
10
8. Plant scandent, capitula with 4 phyllaries and 4 florets
XXX. Eupatorieae (p. 510)
– Plant a shrub or small tree, phyllaries and florets > 4 9
9. Phyllaries dimorphic, pappus of 2–3 awns
XX. Coreopsideae (p. 406)
– Phyllaries subequal, pappus of a few minute squamellae or absent
XIX. Helenieae (p. 400)
10. Leaves spiny
III. Cynareae (Cardueae) (p. 123)
– Leaves not spiny
11
11. Phyllaries in one series, with or without a few much
smaller outer bracts (calyculus); pappus of bristles
present
XII. Senecioneae (p. 208)
– Phyllaries 2- or > 2-seriate, if uniseriate, then pappus
absent
12
12. Phyllaries papery, brownish or coloured, not green
XIV. Gnaphalieae (p. 246)
– Phyllaries not papery, often green, herbaceous
13
13. Anthers ecaudate, without basal tails
127
– Anthers caudate, with basal tails, often long and
prominent
14
14. Capitula solitary; corollas white, anthers ecalcarate;
Australia
XVII. Inuleae (p. 374)
– Capitula few to many, variously arranged or if solitary,
then corollas purple; anthers calcarate; not Australia
15
15. Style shaft slightly broadened and scabrid in upper
part; style arms dorsally scabrid; South America
V. Moquinieae (p. 148)
– Style shaft not broadened and glabrous (South
America) or short-pilose (E Asia) in upper part;
style arms dorsally glabrous (South America),
short-papillose (Africa/Madagascar) or short-pilose
(E Asia)
II. Mutisieae (p. 90)
16. Capitula with some or all florets ligulate (0/5), with
strap-shaped apically 5-toothed or 5-lobed abaxial
limb
17
– Capitula without ligulate florets, florets variously regular (5/0), pseudobilabiate (1/4), bilabiate (2/3), or radiate (0/3, 0/4)
21
17. Latex (milky juice) abundantly present in all vegetative
parts; all florets ligulate VIII. Cichorieae (p. 180)
80
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
– Latex (milky juice) absent; all or some florets ligulate
18
18. Leaves opposite; achenes with phytomelanin layer in
pericarp
XX. Coreopsideae (p. 406)
– Leaves alternate; achenes without phytomelanin layer
in pericarp
19
19. Outer phyllaries and leaves spiny
III. Cynareae (Cardueae) (p. 123)
– Outer phyllaries and leaves not spiny
20
20. Herbs; style shaft with sweeping hairs in upper part
VI. Vernonieae (p. 149)
– Shrubs or small trees; style shaft glabrous
II. Mutisieae (p. 90)
21. Capitula with some or all florets bilabiate (2/3) or
rarely pseudobilabiate (1/4)
22
– Capitula entirely without bilabiate florets
31
22. All florets of capitulum bilabiate, although outer sometimes with enlarged abaxial lip and appearing radiate
23
– Only marginal or only inner florets bilabiate or pseudobilabiate
24
23. Achenes with large, elongated crystals in epidermal
cells; style arms with sweeping hairs extending to below the bifurcation; annual herbs; India
XVII. Inuleae 1136. Nanothamnus (p. 390)
– Achenes without or with other kinds of crystals in
epidermal cells; trees, shrubs or perennial herbs, if
annual herbs, then New World and/or style shaft glabrous
II. Mutisieae (p. 90)
24. Peripheral florets radiate, inner florets bilabiate; S
Africa
XVI. Anthemideae (p. 342)
– Peripheral florets bilabiate, inner florets regular 25
25. Herbs or shrubs; achenes with phytomelanin layer in
pericarp
91
– Herbs, shrubs or small trees, if herbs or shrubs, then
without phytomelanin layer in pericarp
26
26. Filaments connate; achenes dorsally spiny, outcurved
166. Dipterocome (p. 147)
– Filaments free; achenes not dorsally spiny nor outcurved
27
27. Phyllaries uniseriate
28
– Phyllaries 2- or > 2-seriate
29
28. Scandent shrub; achenes without stipitate glands
XII. Senecioneae (p. 208)
– Erect perennial herb; achenes with conspicuous stipitate glands
II. Mutisieae (p. 90)
29. Pappus of a single plumose bristle; Africa
XIV. Gnaphalieae (p. 246)
– Pappus otherwise, usually of numerous barbellate
bristles or scarious scales
30
30. Anthers ecalcarate, ecaudate; disc florets functionally
staminate
XV. Astereae (p. 284)
– Anthers calcarate, caudate, often conspicuously so;
disc florets perfect, rarely functionally staminate or
pistillate
128
31. Capitula 1-flowered, or 1- to several-flowered and
aggregated into secondary compound heads (syncephalia)
32
– Capitula 2- to many-flowered and not aggregated into
secondary compound heads (syncephalia), although
inflorescence sometimes congested
50
32. Leaves and/or phyllaries spiniferous; plant caulescent
or if acaulescent, then corollas white or pale blue 33
– Leaves and/or phyllaries not spiniferous or if leaves
spiniferous, then plant acaulescent and corollas purple
35
33. Syncephalia with one central perfect floret and 4–6
peripheral functionally staminate florets, further aggregated into a somewhat elongated terminal tertiary
head; plant lacticiferous
IX. Gundelieae (p. 199)
– Syncephalia with all florets perfect, not further aggregated into a somewhat elongated terminal tertiary
head; plants not or only weakly lacticiferous
34
34. Individual capitula surrounded by a tuft of bristles
or white plumose hairs; syncalathia globose or hemispherical, with a few small or leaf-like basal bracts;
Old World
III. Cynareae (Cardueae) (p. 123)
– Individual capitula not surrounded by a tuft of bristles or white hairs; syncalathia surrounded by ovate,
marginally spiny bracts; North America
II. Mutisieae (p. 90)
35. Achenes without a phytomelanin layer in the pericarp,
usually not black
36
– Achenes with a phytomelanin layer in the pericarp,
often therefore appearing black or streaked with black
42
36. Style shaft and style arms both glabrous
II. Mutisieae (p. 123)
– Style shaft glabrous or with hairs or papillae in upper
part below the bifurcation, style arms with hairs or
papillae dorsally or at least apically
37
37. Phyllaries papery or cartilaginous, hyaline, brownish
or coloured; style arms truncate, with apical hairs;
capitula homogamous, rarely heterogamous
XIV. Gnaphalieae (p. 246)
– Phyllary and style characters otherwise; capitula heterogamous or homogamous
38
38. Style shaft with hairs on upper part, style arms narrow, acute, unappendaged, hairy dorsally and with
stigmatic papillae covering whole of inner (ventral)
surface; capitula homogamous
39
– Style characters all or some otherwise; capitula heterogamous or homogamous
40
39. Plant an acaulescent rosulate herb; capitula crowded
in a large syncalathium sessile in the centre of the leaf
rosette; S Africa
X. Arctotideae (p. 200)
– Plant a tree, shrub or caulescent herb; capitula distinct
or if aggregated into syncephalia, then these not sessile in the centre of a leaf rosette; New World, if Old
World, then capitula 4-flowered and phyllaries decussate
VI. Vernonieae (p. 149)
40. Capitula 1-flowered, not aggregated into secondary
compound heads (syncephalia), although sometimes
in congested inflorescences
41
– Capitula 1- to several-flowered, aggregated into
sphaerical to spiciform secondary heads (syncephalia)
XVII. Inuleae (p. 374)
41. Plant a caulirosulate herb; leaves pinnately veined;
phyllaries 5–7-seriate, imbricate; anthers with prominent tails; E Asia
II. Mutisieae (p. 90)
– Plant a rosulate scapigerous herb; leaves parallelveined; phyllaries 2-seriate; anthers ecaudate; S Africa
XI. Corymbieae (p. 207)
42. Style arm appendages longer than the stigmatic lines,
spathulate, or if not spathulate, then phyllaries and
florets 4; syncalathia spherical clusters or glomerules;
corollas not yellow
XXX. Eupatorieae (p. 510)
Compositae
– Style arm appendages otherwise, shorter than the stigmatic lines, or style arms unappendaged; corollas yellow or not
43
43. Leaves and/or phyllaries with pellucid glands or pustules, aromatic
XXII. Tageteae (p. 420)
– Leaves and phyllaries without pellucid glands or pustules, usually not aromatic
44
44. Plant a prostrate, rosulate, annual herb; pappus of
many scales, fused at the base and deciduous as a unit
XXXIII. Chaenactideae (p. 431)
– Plant an annual or perennial herb or shrubby; pappus
absent or otherwise
45
45. Capitula crowded in glomerules surrounded by leafy
bracts; phyllaries uniseriate, connate; leaves opposite
XXVI. Heliantheae (p. 440)
– Capitula otherwise; leaves opposite or alternate
46
46. Plant a shrub with brittle whitish stems; leaves opposite, succulent, glaucous
XXII. Tageteae (p. 420)
– Plant otherwise; leaves opposite or alternate, and most
subsucculent
47
47. Leaves alternate; capitula disciform or discoid; achenes tangentially flattened, laterally ciliate; anthers
tailed; Africa, Madagascar, India
XVIII. Anthroismeae (p. 395)
– Leaves opposite or at least some capitula radiate; achenes otherwise; anthers not tailed; New World, Australia, adventive elsewhere
48
48. Leaves alternate; shrubs or small trees
XXVI. Heliantheae (p. 440)
– Leaves opposite; herbs
49
49. Capitula with peripheral tubular florets enclosed in
a perigynium; styles of disc florets undivided
XXVII. Millerieae (p. 477)
– Capitula without peripheral florets enclosed in a perigynium; styles of disc florets divided, style arms truncate
XXII. Tageteae (p. 420)
50. Phyllaries uniseriate or varyingly uniseriate-biseriate;
receptacle epaleaceous, without scales subtending florets, although sometimes with bristles or projections
51
– Phyllaries 2- to many-seriate, if uniseriate, then receptacle paleaceous, with scales subtending all or some
florets, each scale subtending a single floret
72
51. Achenes without a phytomelanin layer in the pericarp,
usually not black
52
– Achenes with a phytomelanin layer in the pericarp,
therefore often appearing black or streaked with black
62
52. Phyllaries cartilaginous, hyaline, greenish or greenishbrown; style arm apices conical
XIV. Gnaphalieae (p. 246)
– Phyllaries and style arm apices either or both otherwise
53
53. All or at least the disc florets with tetramerous corollas
54
– All or at least the disc florets with pentamerous corollas
56
54. Pappus of conspicuous bristles
XII. Senecioneae (p. 208)
– Pappus coroniform or absent, not of conspicuous bristles
55
55. Leaf blades obovate, usually toothed, rarely entire
XVIII. Athroismeae (p. 395)
81
– Leaf blades linear, entire, with sheathing base
XII. Senecioneae (p. 208)
56. Pappus of bristles present, at least on some achenes 61
– Pappus absent, coroniform or of a single scale
57
57. Pappus of a single scale XII. Senecioneae (p. 208)
– Pappus coroniform or absent
58
58. Phyllaries with narrow hyaline margins or all or at least
the inner with broad, pale membranous margins; style
arms truncate, penicillateXVI. Anthemideae (p. 342)
– Phyllaries and style arms either or both otherwise 59
59. Achenes large, terete, triquetrous or flattened, winged
or wingless, straight or curved, glabrous, smooth or
aculeate, sometimes fenestrate, or fruits drupaceous;
achenes homomorphic or heteromorphic; pappus absent
XIII. Calenduleae (p. 241)
– Achenes small, columnar, angled, terete or compressed, never fenestrate, not drupaceous, unwinged,
homomorphic; pappus coroniform or absent
60
60. Style arms with an ovate-lanceolate or linear appendage
XV. Astereae (p. 284)
– Style arms without an apical appendage, truncate to
obtuse, or disc floret styles undivided
XII. Senecioneae (p. 208)
61. Involucre saucer-shaped to subglobose; marginal filiform florets 2- to many-seriate, tubular, with or without a short, narrow ray; Old World tropics
XV. Astereae (p. 284)
– Involucre cylindrical, sometimes calyculate; marginal
pistillate florets absent or uniseriate and radiate, rarely
tubular; widespread
XII. Senecioneae (p. 208)
62. Style arm appendages longer than the stigmatic lines,
prominent; capitula discoid; corollas never yellow
XXX. Eupatorieae (p. 510)
– Style arm appendages shorter than stigmatic lines;
capitula radiate, disciform or discoid; corollas often
yellow
63
63. Leaves and/or phyllaries with pellucid glands or pustules, aromatic
XXII. Tageteae (p. 420)
– Leaves and phyllaries without pellucid glands or pustules
64
64. Disc corollas all 4-lobed
65
– Disc corollas 5-lobed, sometimes some also 4- or 6lobed
67
65. Capitula radiate
XXIX. Perityleae (p. 507)
– Capitula discoid or disciform
66
66. Achenes flattened, with wings or usually a corky, often
ciliate margin
XXIX. Perityleae (p. 507)
– Achenes 4-sided, obpyramidal, if flattened, then capitula disciform
XXIV. Bahieae (p. 433)
67. Leaves all or some opposite or whorled, or proximal
rosulate
68
– Leaves alternate
69
68. Ray corollas white; pappus of about 8 erose scales
XXIV. Bahieae (p. 433)
– Ray corollas some shade of yellow; pappus of persistent bristles, fragile awns or awns and scales, or absent,
or capitula discoid
XXVIII. Madieae (p. 492)
69. Plant a shrub; capitula discoid
XXIV. Bahieae (p. 433)
– Plant an annual or perennial herb, if a shrub, then
capitula radiate
70
70. Plant an annual herb or shrub
XXVIII. Madieae (p. 492)
82
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
– Plant a perennial herb
71
71. Disc florets functionally staminate; Cuba
XXI. Neurolaeneae (p. 417)
– Disc florets perfect; W North America
XXVIII. Madieae (p. 492)
72. Receptacle paleaceous, with a scale subtending each
floret or each of some of the florets, scales sometimes
spiniform
73
– Receptacle epaleaceous, without scales subtending the
florets, although sometimes with bristles or receptacular or alveolar projections between or around the
florets
129
73. Leaves all alternate
74
– Leaves some or all opposite or whorled
92
74. Style shaft with an articulation below the bifurcation,
marked by a ring of hairs and/or an increase in diameter; style arms short and shortly hairy dorsally, or
long and not divergent for most of their length, velvety
dorsally
III. Cynareae (Cardueae) (p. 123)
– Style shaft without an articulation below the bifurcation; style arms various
75
75. Style arms tangentially spreading, narrow, acute, unappendaged, hairy dorsally, with stigmatic papillae
over whole of ventral surface; corollas not yellow; capitula homogamous; receptacular scales sometimes
spiniform
VI. Vernonieae (p. 149)
– Style arm characters otherwise; capitula homogamous
or heterogamous; corollas often yellow; receptacular
scales never spiniform
76
76. Plant an annual herb with heterogamous capitula sessile 2–4 together in the stem bifurcations, achenes adnate to 1 or 2 of the receptacular scales, and shortly
tailed anthers; NE Asia
1620. Symphyllocarpus (p. 574)
– Plant otherwise
77
77. Achenes without a phytomelanin layer in pericarp,
usually not black
78
– Achenes with a phytomelanin layer in pericarp, thus
often black or streaked with black
93
78. Style arms with apical appendages above the stigmatic
lines; anthers usually ecaudate
79
– Style arms unappendaged, rounded or truncate; anthers usually caudate
80
79. Ray florets neuter, sterile XIX. Helenieae (p. 400)
– Ray florets pistillate and fertile or absent
XV. Astereae (p. 284)
80. Style arms apically truncate, penicillate, with apical
hairs
81
– Style arms apically rounded, with hairs or papillae
dorsally, or glabrous
86
81. Phyllaries herbaceous, without scarious margins and
apex
82
– Phyllaries papery or cartilaginous, hyaline, brownish
or coloured, or with distinct brownish to pallid scarious membranous margins and apex
83
82. Plant a herb; North America
XIX. Helenieae (p. 400)
– Plant a shrub; S Africa XVI. Anthemideae (p. 342)
83. Pappus absent, a rim or a cartilaginous corona or auricle
84
– Pappus of bristles, scales or spines present
85
84. Phyllaries papery, brownish or white; pappus absent
XIV. Gnaphalieae (p. 246)
– Phyllaries with distinct brownish to pallid membranous scarious margins and apex; pappus absent, coroniform or auriculiform
XVI. Anthemideae (p. 342)
85. Phyllaries papery, hyaline, whitish to coloured; pappus
of bristles, scales or 2 spines; Australia, Africa
XIV. Gnaphalieae (p. 246)
– Phyllaries not papery, but with scarious margins and
apex; pappus of scales; Africa
XVI. Anthemideae (p. 342)
86. Capitula homogamous, discoid
87
– Capitula heterogamous, radiate or disciform
89
87. Style arms glabrous (South America) or coronatepapillose around apices (tropical Africa)
II. Mutisieae (p. 90)
– Style arms with sweeping hairs and papillae dorsally
88
88. Corollas yellow; tropical Africa
XVIII. Athroismeae (p. 395)
– Corollas white, cream, purplish or lavender; North
America
XIX. Helenieae (p. 400)
89. Rays purple, lilac or white
XIV. Gnaphalieae (p. 246)
– Rays yellow or absent and capitula disciform
90
90. Achenes with one large calcium oxalate crystal in each
epidermal cell, or if without crystals, then capitula
disciform and corolla mauve, or stems winged and
receptacular paleae flattened
XVII. Inuleae (p. 374)
– Achene epidermal cells without crystals; receptacular
paleae enfolding achenes
XVIII. Athroismeae (p. 395)
91. Pappus of 8–10 scales
XXIV. Bahieae (p. 433)
– Pappus absent, of 2 scales and sometimes 2 fragile
awns, or of numerous plumose bristles
107
92. Achenes without a phytomelanin layer in the pericarp
123
– Achenes with a phytomelanin layer in the pericarp,
thus often black or streaked with black
93
93. Receptacular scales in one series between ray florets
and disc florets
XXVIII. Madieae (p. 492)
– Receptacular scales subtending some or all of the nonperipheral florets, but not in a single series between
the ray and disc florets
94
94. Capitula discoid; corollas never yellow; style arm appendages much longer than the stigmatic surfaces,
prominent, lanceolate to clavate; phyllaries all similar
or gradate; pappus never of barbed awns
XXX. Eupatorieae (p. 510)
– Capitula radiate, disciform or discoid; corolla often
yellow; style arm appendages usually shorter than the
stigmatic surfaces, if longer, then phyllaries dimorphic
and pappus of a few barbed awns
95
95. Pappus absent, or of a few teeth or awns, or of scales
and minute bristles or awns, or of barbed awns, or
cupuliform or coroniform, but not of 3 or more similar,
conspicuous, often plumose scales or bristles
96
– Pappus of all or disc florets of 3 or more similar, conspicuous, often plumose scales or bristles
108
96. Styles of disc florets undivided, or apically very shortly
bifid; disc florets functionally staminate
97
– Styles of disc florets with free style arms; disc florets
perfect or functionally staminate
111
Compositae
83
97. Plants caulirosulate herbs, shrubs or small trees, or
megaphytic rosulate herbs, sometimes monocarpic;
Ecuador, Colombia, Venezuela (Paramos)
XXVII. Millerieae (p. 477)
– Plant of different habit and usually also habitat
98
98. Achenes each enfolded by a phyllary or wholly enclosed in a conceptacle formed by an adnate phyllary
XXVII. Millerieae (p. 477)
– Achenes not enfolded by phyllaries nor enclosed in
conceptacles
99
99. Capitula disciform, or if radiate, then achenes each
associated with a phyllary and adjacent two disc florets
and scales
XXVI. Heliantheae (p. 440)
– Capitula radiate; achenes not so associated
100
100. Achenes obpyriform or broadly obpyramidal, not flattened parallel to the involucre
XXVII. Millerieae (p. 477)
– Achenes more or less flattened tangentially, parallel to
the involucre
101
101. Achenes winged
102
– Achenes not winged, although sometimes strongly
flattened
103
102. Plant scandent
XX. Coreopsideae (p. 406)
– Plant an annual or perennial herb or shrub, not scandent
XXVI. Heliantheae (p. 440)
103. Pappus of 2–3 awns
XX. Coreopsideae (p. 406)
– Pappus of a few scales, minutely coroniform or absent
104
104. Leaves alternate
XXVI. Heliantheae (p. 440)
– Leaves opposite
105
105. Pappus a shallow corona XXV. Polymnieae (p. 439)
– Pappus absent
106
106. Leaf blades trilobed; S Asia
XX. Coreopsideae (p. 406)
– Leaf blades elliptic to lanceolate, unlobed; South
America
XXVII. Millerieae (p. 477)
107. Phyllaries 2- to 5-seriate; pappus of plumose bristles,
rarely absent
XXVII. Millerieae (p. 477)
– Phyllaries 1-seriate; pappus absent or of 2 scales with
or without 2 fragile awns XXVIII. Madieae (p. 492)
108. Leaves all or mostly alternate
109
– Leaves opposite
XXI. Neurolaeneae (p. 417)
and
XXVII. Millerieae (p. 477)3
109. Leaves bipinnate, segments filiform to linear, leaves all
alternate
XXIV. Bahieae (p. 433)
– Leaves with entire to pinnatilobed margins
110
110. Proximal leaves rosulate or opposite, distal alternate
XXVIII. Madieae (p. 492)
– Proximal and distal leaves all alternate
XXI. Neurolaeneae (p. 417)
and
XXVII. Millerieae (p. 477)3
111. Leaves mostly alternate, proximal rosulate or opposite; phyllaries each enfolding the subtended ray floret
proximally
XXVIII. Madieae (p. 492)
– Leaves all alternate or all opposite, or opposite and
the upper alternate; phyllaries not so enfolding the
subtended ray floret, or if so, then leaves opposite 112
112. Achenes strongly 9–11-ribbed; shrubs; SW North
America, N Mexico
XXII. Tageteae (p. 420)
– Achenes otherwise; herbs, shrubs or trees; widespread
113
113. Plant a shrub with succulent, deeply dissected leaves,
achenes fusiform, 4-angled; San Felix and San Ambrosio Islands, N Chile
XXIX. Perityleae (p. 507)
– Plant and achenes otherwise; widespread
114
114. Plant aquatic or of very moist places; stem fistulose,
rooting at nodes; receptacular scales wrapping tightly
around florets; capitula minutely radiate or disciform,
rays minute or absent; pantropical
XXI. Neurolaeneae (p. 417)
– Plant without above combination of features;
widespread
115
115. Achenes of the ray and of the disc (when present) tangentially flattened (flattened parallel to the involucre,
obcompressed); receptacular scales more or less flat to
navicular and sometimes cucullate
116
– Achenes of the disc radially flattened (compressed),
although those of the ray sometimes tangentially flattened, or achenes of the disc not flattened; receptacular
scales conduplicate, rarely more or less flat to navicular, cucullate or bristle-like
121
116. Leaves alternate, with expanded leaf-bases
XXVII. Millerieae (p. 477)
– Leaves opposite, or if alternate, then without expanded
leaf-bases
117
117. Leaves alternate
118
– Leaves all or mostly opposite
119
118. Plant with Kranz anatomy; style arm appendages as
long as or longer than stigmatic surfaces
XX. Coreopsideae (p. 406)
– Plant not with Kranz anatomy; style arm appendages
much shorter than stigmatic surfaces or absent
XXVI. Heliantheae (p. 440)
119. Style arm appendages usually vascularized, penicillate
and papillose; receptacular scales more or less flat
XX. Coreopsideae (p. 406)
– Style arm appendages non-vascularized, acute to
cylindrical, papillose, or absent and represented by
an apical tuft of papillae; receptacular scales more or
less flat, navicular or cucullate
120
120. Capitula discoid
XXVI. Heliantheae (p. 440)
– Capitula radiate
XXVI. Heliantheae (p. 440)
and
XXVII. Millerieae (p. 477)4
121. Plant with outer phyllaries herbaceous, shortly connate and inner membranous or scarious, fusiform to
linear or narrowly oblanceolate achenes with conspicuously attenuate apices, and pubescent filaments
XX. Coreopsideae (p. 406)
– Plant without above combination of features
122
122. Leaves alternate, at least in part
XXVI. Heliantheae (p. 440)
– Leaves all opposite
XXVI. Heliantheae (p. 440)
and
XXVII. Millerieae (p. 477)4
123. Style arms with a lanceolate to linear, papillose appendage above the stigmatic areas
XV. Astereae (p. 284)
– Style arms without appendages
126
124. Stigmatic papillae covering entire inner surface of
style arms; style arms dorsally and upper part of shaft
with long sweeping hairs
125
– Stigmatic papillae confined to two marginal or submarginal lines on inner surface of style arms; style
3
4
See also key to the Heliantheae alliance tribes, p. 391.
See also key to the Heliantheae alliance tribes, p. 391.
84
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
125.
–
126.
–
127.
–
128.
–
129.
–
130.
–
131.
–
132.
–
133.
–
134.
–
135.
–
136.
arms truncate, with hairs apically, or rounded with
hairs dorsally
126
Capitula radiate; disc florets yellow; Americas
VII. Liabeae (p. 175)
Capitula discoid; florets purplish; tropical Africa
VI. Vernonieae (p. 149)
Pappus absent or an entire or denticulate rim; capitula
radiate, disciform or discoid
XVI. Anthemideae (p. 342)
Pappus of scales or awns, sometimes connate; capitula
radiate, rays yellow with purple bands dorsally, or blue
or white
XIV. Gnaphalieae (p. 246)
Pappus bristles, if present, whitish to tawny; New
World
XV. Astereae (p. 284)
Pappus bristles purple; Sinohimalaya
165. Cavea (p. 146)
Inner phyllaries whitish to pinkish-violet or purple, or
ending in a long, pungent, often pink or purple apex
III. Cynareae (Cardueae) (p. 123)
Inner phyllaries otherwise
II. Mutisieae (p. 90)
Pappus of at least some achenes at least in part of one
or more conspicuous scabrid, barbellate, plumose or
smooth capillary bristles as long as or longer than the
corolla
171
Pappus absent, rim-like, coroniform, auriculiform or
of short to long and conspicuous free or variously connate scales (sometimes dissected distally into bristles),
aristae or awns, devoid of conspicuous bristles at least
as long as the corolla, or if bristles present, then these
shorter than the corolla, inconspicuous and often caducous
130
Achenes with a phytomelanin layer in the pericarp,
thus often black or streaked with black
131
Achenes without a phytomelanin layer in pericarp,
usually but not always not black
154
Leaves and/or phyllaries with pellucid glands or pustules, if without them, then plant a perennial herb with
3-seriate involucres with the outer phyllaries broad
and herbaceous (Mexico, SE North America), or phyllaries succulent or achenes strongly 9–11-ribbed
XXII. Tageteae (p. 420)
Leaves and phyllaries without pellucid glands or pustules, plant and/or phyllaries and achenes otherwise,
achenes mostly with 5 or fewer ribs
170
Pappus absent, or of a few teeth or awns, or of small
scales and minute bristles and awns, or cupuliform or
coroniform, but not of 3 or more conspicuous, often
erose, plumose or lanciniate scales nor of numerous,
sometimes erose awns
133
Pappus of at least the disc florets of 3 or more conspicuous, often erose, plumose or lanceolate scales, sometimes dissected into bristles distally, or of numerous,
sometimes erose awns
144
Pappus present
138
Pappus absent
134
Leaves all or mostly alternate
135
Leaves all opposite, or if alternate, then viscid, or glandular and achenes 4-angled
137
Capitula discoid
XXIII. Chaenactideae (p. 431)
Capitula radiate
136
Rays orange-yellow; receptacle flat to shallowly convex
XXIV. Bahieae (p. 433)
– Rays yellow or white; receptacle convex to conical
XXVIII. Madieae (p. 492)
137. Leaves sessile, clasping or shallowly to strongly perfoliate
XXVIII. Madieae (p. 492)
– Leaves petiolate or if sessile, then not clasping nor
perfoliate
142
138. Pappus of numerous scales, erose and fused at the base,
ciliate or erose and free or lanceolate and laciniate and
free
140
– Pappus of a few awns or scales or coroniform
139
139. Capitula disciform, or if radiate or discoid, then pappus of 2–3 awns fused to the wings of the achenes, or
outer phyllaries 1/10th of the length of the inner, and
inner enclosing the achenes
XXVI. Heliantheae (p. 440)
– Capitula radiate, if discoid, then pappus and phyllaries
otherwise; pappus of 1–2 pairs of opposite scales, of
2–4 bristles or awns and small scales, or a small corona
of fimbriate scales
143
140. Pappus scales united at base, pappus shed as a unit
XXIII. Chaenactideae (p. 431)
– Pappus scales free, persistent
141
141. Achenes obconical or obpyramidal, black or brown;
disc florets usually numerous
XXI. Neurolaeneae (p. 417)
– Achenes obovoid, terete and slightly flattened to quadrangular, black; disc florets 1–6
XXVI. Heliantheae (p. 440)
142. Leaves viscid, or glandular and achenes 4-angled
XXIX. Perityleae (p. 507)
– Leaves not viscid, or if glandular, then achenes not
4-angled
XXVI. Heliantheae (p. 440)
143. Capitula radiate; pappus of 1–2 pairs of opposite scales
or of 2 (rarely 3–4) awns and small scales
XXVIII. Madieae (p. 492)
– Capitula radiate or discoid; pappus of 4 scales alternating with bristles or a small corona of fimbriate scales
XXIX. Perityleae (p. 507)
144. Pappus of 5–10 or many erose scales, fused at the base
and deciduous as a unit
XXIII. Chaenactideae (p. 431)
– Pappus of (2–)4 to many scales or awns, free and persistent, rarely fused at the base and persistent
145
145. Scales of pappus with thickened, often excurrent
midrib
XXIV. Bahieae (p. 433)
– Scales of pappus without prominent thickened midrib,
or pappus of awns or of awns and scales
146
146. Scales and/or awns several (usually 4–10)
147
– Scales and/or awns numerous
150
147. Pappus of 2–4 opposite, erose scales or awns
XXVIII. Madieae (p. 492)
– Pappus otherwise
148
148. Plant a rosulate perennial herb; disc florets functionally staminate; Cuba XXI. Neurolaeneae (p. 417)
– Plant an annual or non-rosulate perennial herb or
shrub; disc florets perfect, rarely functionally staminate, or capitula discoid; New World
149
149. Leaves mostly alternate, entire to pinnately or deeply
lobed
XXVIII. Madieae (p. 492)
– Leaves opposite, if alternate, then leaves 1–3-pinnate
with linear segments
XXIV. Bahieae (p. 433)
150. Capitula discoid
151
Compositae
– Capitula radiate or disciform
152
151. Leaves alternate
XXI. Neurolaeneae (p. 417)
– Leaves opposite
XXI. Neurolaeneae (p. 417)
or
XXIV. Bahieae (p. 433)
or
XXVI. Heliantheae (p. 440)5
152. Leaves opposite
XXI. Neurolaeneae (p. 417)
or
XXVI. Heliantheae (p. 440)5
– Leaves all or mostly alternate
153
153. Achenes with large glands
XXVIII. Madieae (p. 492)
– Achenes without large glands
XXI. Neurolaeneae (p. 417)
154. Style arms with entire inner surface covered with stigmatic papillae
155
– Style arms with stigmatic papillae confined to two
submarginal, distant to almost contiguous, distinct or
apically confluent bands
159
155. Style shaft with an articulation below the bifurcation,
marked by a ring of hairs and/or an increase in diameter, hairy above the articulation; style arms hairy or
velvety dorsally, short to long and, when long, often
not divergent for much of their length; leaves and/or
phyllaries often spiny
158
– Style shaft without an articulation below the bifurcation, of uniform diameter, glabrous or hairy in upper
part; leaves and phyllaries not spiny
156
156. Arms of style short, with hairs or papillae dorsally,
style shaft glabrous; capitula disciform, few-flowered
(temperate South America) or discoid, 1–3-flowered
(E Asia)
II. Mutisieae (p. 90)
– Arms of style usually long, with long sweeping hairs
dorsally extending down shaft to below bifurcation;
widespread
157
157. Capitula radiate; corollas usually yellow; leaves opposite; latex usually present; New World
VII. Liabeae (p. 175)
– Capitula discoid; corollas never yellow; leaves alternate or rarely (Old World) opposite; latex usually absent; widespread
VI. Vernonieae (p. 149)
158. Capitula radiate, rarely discoid; corollas usually yellow; tropical and S Africa X. Arctotideae (p. 200)
– Capitula discoid or radiant; if corollas yellow, then
plant not African
III. Cynareae (Cardueae) (p. 123)
159. Achenes large, terete, triquetrous or flattened, winged
or wingless, straight to curved, glabrous, smooth or aculeate, sometimes fenestrate, homomorphic or heteromorphic, or fruits drupaceous; pappus absent; phyllaries herbaceous
XIII. Calenduleae (p. 241)
– Achenes usually small, columnar, angled, terete or
compressed, never fenestrate, usually but not always
homomorphic, never drupaceous; pappus present or
absent; phyllaries various
160
160. Style arms with an apical papillose appendage
above the stigmatic bands, hairy dorsally below the
appendage
161
– Style arms usually obtuse to truncate, rarely acute or
with a much prolonged apex, hairy or papillose apically and/or dorsally, not appendaged
162
161. Receptacle strongly alveolate, alveolae forming aristate baskets or cups around achenes, or densely se5
See also key to the Heliantheae alliance tribes, p. 391
–
162.
–
163.
–
164.
–
165.
–
166.
–
167.
–
168.
–
169.
–
170.
–
85
tose; achene epidermal cells with large, quadrate or
isodiametric crystals
XIX. Helenieae (p. 400)
Receptacle alveolate or not, if alveolate, then alveolae not forming baskets or cups around achenes, nor
densely setose; achene epidermal cells usually with
raphides
XV. Astereae (p. 284)
Disc floret corollas 4-lobed
163
Disc floret corollas 5-lobed
164
Phyllaries rather dry and scarcely herbaceous, often
with pale or brownish scarious margins and/or apices;
style arms truncate, penicillate
XVI. Anthemidae (p. 342)
Phyllaries herbaceous or membranous; style arms
short, acute to obtuse, apically papillose
XVIII. Athroismeae (p. 395)
Capitula radiate
166
Capitula disciform or discoid
165
Plant a perennial herb with alternate leaves, herbaceous 4–5-seriate phyllaries, zygomorphic peripheral
florets, achenes with 4 strongly thickened ribs and
pappus of basally fused scales; SW tropical Africa
XVIII. Athroismeae (p. 395)
Plant not with above combination of features
167
Phyllaries papery or cartilaginous; style arms truncate with apical hairs or obtuse with dorsal hairs; rays
yellow or yellow with purple bands abaxially; pappus
absent or of free scales; S Africa
XIV. Gnaphalieae (p. 246)
Phyllaries, style arms, rays and pappus not providing
the above combination of features; widespread 167
Style arms more or less acute, with stigmatic papillae
in two submarginal, rarely almost contiguous, apically
confluent bands; style arms with acute sweeping hairs
ending above the bifurcation or obtuse sweeping hairs
extending down to below the bifurcation; phyllaries
herbaceous
XVII. Inuleae (p. 374)
Style arms and phyllaries not with above combination
of characters
168
Phyllaries papery or cartilaginous at least in part,
hyaline, whitish, brownish or variously coloured, very
rarely herbaceous; capitula discoid or disciform; style
arms truncate, hairy apically, or rounded or with
a much prolonged apex and hairy dorsally
XIV. Gnaphalieae (p. 246)
Phyllaries herbaceous, or rather dry and usually with
scarious margins and/or apices; capitula radiate, discoid or disciform; style arms truncate, apically penicillate or with a tuft of papillae
169
Phyllaries herbaceous; cells of apical anther appendages heavily sclerified, appendages carinate;
style arms with apical tuft of papillae
XIX. Helenieae (p. 400)
Phyllaries rather dry and scarcely herbaceous, usually
with scarious margins and/or apices; cells of anther
appendages not sclerified, appendages soft, flat; style
arms penicillate
XVI. Anthemideae (p. 342)
Capitula discoid, although rarely abaxial lobes
of marginal floret corollas enlarged and ray-like;
corollas never yellow; style arm appendages usually
much longer than the stigmatic bands, prominent,
lanceolate to clavate
XXX. Eupatorieae (p. 510)
Capitula radiate, disciform or discoid; corollas often
yellow; style arm appendages usually shorter than stigmatic bands or style arms unappendaged
132
86
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
171. Achenes with a phytomelanin layer in the pericarp,
therefore often black or streaked with black
189
– Achenes without a phytomelanin layer in the pericarp,
not black or if black, then for some other reason 172
172. Style shaft with an articulation below the bifurcation,
marked by a ring of hairs and/or an increase in diameter, usually hairy or papillose above the articulation; style arms with hairs or papillae dorsally, with
stigmatic papillae covering the whole inner surface,
sometimes long and united for most of their length;
capitula homogamous, radiant or radiate
177
– Style shaft uniform in diameter below the bifurcation
or rarely somewhat narrowed in upper part, without
an articulation, glabrous or with sweeping hairs in upper part below the bifurcation; capitula never radiant
173
173. Style arms usually long, with long sweeping hairs dorsally extending down shaft below the bifurcation, unappendaged; stigmatic papillae covering entire inner
surface of style arms
174
– Style arm and shaft characters otherwise; style arms
unappendaged or appendaged, with stigmatic papillae
over entire inner surface or more usually confined to
two submarginal bands
178
174. Leaves opposite; South America
VII. Liabeae (p. 175)
– Leaves alternate; Old and New Worlds
175
175. Corollas not yellow, or if of some shade of yellow, then
leaves triplinerved; capitula discoid; widespread
VI. Vernonieae (p. 149)
– Corollas yellow; leaves pinnately veined; capitula discoid or radiate; Old World
176
176. Capitula radiate or discoid, sessile; leaves dentatespinulate, spine-tipped; S Africa
X. Arctotideae (p. 200)
– Capitula discoid, pendunculate; leaves dentatelobulate, unarmed; NW Africa
IX. Gundelieae (p. 199)
177. Capitula radiate; S Africa X. Arctotideae (p. 200)
– Capitula discoid or radiant
182
178. Stigmatic papillae covering whole of inner surface of
style arms
184
– Stigmatic papillae confined to two free or apically
confluent, distant to almost contiguous submarginal
bands
179
179. Style arms ending in a sterile appendage above the
stigmatic bands
183
– Style arms unappendaged, although sometimes ending in a more or less conical tuft of papillae or hairs,
apex acute, obtuse or truncate
180
180. Anther appendages carinate, with cells strongly sclerified; North America
XIX. Helenieae (p. 400)
– Anther appendages flat, soft, cells not sclerified;
widespread
181
181. Phyllaries with broad, pale to dark brown or black
scarious margins; C AsiaXVI. Anthemideae (p. 342)
– Phyllaries without broad scarious margins, although
sometimes wholly or partly papery or cartilaginous,
often herbaceous
185
182. Capitula numerous, in inflorescences with densely
racemose branches; shrub; leaves not spiny; Brazil
V. Moquinieae (p. 148)
– Capitula variously arranged, but not in inflorescences
with densely racemose branches; habit various, usu-
183.
–
184.
–
185.
–
186.
–
187.
–
188.
–
189.
–
190.
–
ally herbaceous, rarely shrubby; leaves spiny or unarmed; Old World, if not Old World, then not Brazil
III. Cynareae (Cardueae) (p. 123)
Plant a broom-like shrub or treelet with small, yellow,
discoid capitula and sagittate or caudate anther bases;
SW North America, Mexico
XII. Senecioneae (p. 208)
Plant of various habit; capitula discoid, radiate or disciform, yellow or not; anther bases commonly rounded
or truncate, rarely caudate; worldwide
XV. Astereae (p. 284)
Capitula discoid, or if radiate, then rays red; corollalobes much longer than wide; anther bases strongly
tailed
II. Mutisieae (p. 90)
Capitula radiate, with yellow rays, if discoid, corollas
yellow, lobes about as long as wide; anther bases not
tailed
XII. Senecioneae (p. 208)
Phyllaries papery or cartilaginous at least in part, hyaline, white, brownish or variously coloured, often yellow or pink; style arms truncate and hairy apically,
or rounded or with a much prolonged apex and hairy
dorsally and apically; stigmatic bands not confluent
apically
XIV. Gnaphalieae (p. 246)
Phyllaries usually herbaceous and green, style arms
apically conical, rounded, obtuse, acute or truncate,
with hairs or papillae apically or dorsally and sometimes extending downwards on to shaft below the bifurcation; stigmatic bands confluent or not confluent
apically
186
Stigmatic bands apically confluent; style arms more
or less acute, with usually acute sweeping hairs dorsally ending above the bifurcation, or usually obtuse
sweeping hairs extending to below the bifurcation 187
Stigmatic bands distant, not confluent apically; style
arms apically conical, obtuse or truncate; sweeping
hairs otherwise distributedXII. Senecioneae (p. 208)
Plant a scandent shrub; Cuba
XVII. Inuleae 1137. Feddea (p. 390)
Plant a herb, subshrub, non-scandent shrub or tree
188
Pappus purple, of numerous uniseriate barbellate bristles, bristles in staminate florets shorter and apically
slightly dilated; corolla externally with robust, acute,
multicellular hairs; Sinohimalaya 165. Cavea (p. 146)
Pappus otherwise, not purple; corollas without such
hairs; widespread
XVII. Inuleae (p. 374)
Capitula discoid, although rarely abaxial lobes of peripheral floret corollas enlarged and ray-like; corollas never yellow; style arm appendages usually much
longer than the stigmatic bands, prominent, lanceolate
to clavate
XXX. Eupatorieae (p. 510)
Capitula radiate, disciform or discoid; corollas often
yellow; style arm appendages usually shorter than stigmatic bands or style arms unappendaged
190
Leaves and/or phyllaries with pellucid glands or pustules, if without them, then plant a perennial herb with
2–3-seriate involucres with the outer phyllaries broad
and herbaceous (Mexico, SE North America) or achenes 9–10-ribbed and leaves succulent, opposite, or if
alternate, then receptacle setose
XXII. Tageteae (p. 420)
Leaves and phyllaries without pellucid glands or pustules; plant and/or phyllary and achene features otherwise, achenes usually with 5 or fewer ribs
191
Compositae
191. Achenes 4–5-angled, cylindrical to obconical
193
– Achenes tangentially or radially flattened
192
192. Plant an annual herb; achenes tangentially flattened,
without wings or corky ciliate margin; disc floret corollas 5-lobed
XXIV. Bahieae (p. 433)
– Plant a shrub or perennial or annual herb; achenes
tangentially or radially flattened, with wings or a welldeveloped corky, ciliate margin; disc floret corollas 4or 5-lobed
XXIX. Perityleae (p. 507)
193. Pappus of 4 scales alternating with 4 bristles
XXIX. Perityleae (p. 507)
– Pappus of numerous bristles
194
194. Phyllaries 2–3-seriate and subequal or 3–5-seriate and
gradate
XXIV. Bahieae (p. 433)
– Phyllaries (1–)2-seriate, outer herbaceous and each
opposite a ray floret, inner membranous or absent
XXVIII. Madieae (p. 492)
I. Tribe Barnadesieae D. Don (1830).
T.F. Stuessy and E. Urtubey
Shrubs, trees, or perennial or annual herbs. Stems
erect or decumbent, with or without axillary spines.
Basal leaves sometimes rosulate. Cauline leaves alternate, fasciculate, opposite or whorled, often with
spinescent apex, sessile to petiolate, entire, often
xeromorphic. Synflorescences solitary or aggregated (cymose-racemose or cymose-corymbose
heads). Involucre campanulate, cylindrical,
turbinate or hemispherical; phyllaries in 4–14series, frequently apically spinulose, velutinous to
glabrous. Receptacle pilose, rarely with pales or
glabrous. Capitula homogamous or heterogamous,
discoid, pseudoradiate or ligulate. Florets 1–c. 135,
white, yellow, orange, pink, purple to violet, isomorphic, generally hermaphrodite or sometimes
unisexual or with only marginal florets unisexual;
corolla 5-merous, actinomorphic or zygomorphic
(pseudobilabiate, bilabiate, subpseudobilabiate,
ligulate or subligulate), or anisomorphic, with
marginal flowers hermaphrodite (with pseudobilabiate 1/4 corollas), and with central flowers
hermaphrodite or unisexual (with actinomorphic
or zygomorphic, i.e. pseudobilabiate, bilabiate,
ligulate or subligulate, corollas), often villous.
Anthers ecaudate to tailed; apical appendage
entire, emarginate or bilobed; filaments free or
connate, inserted at apex or base of corolla (rarely
in between), Style shortly bilobed or bifid, glabrous
or papillose below bifurcation. Achenes villous
(“barnadesioid hairs”), rarely only at the apex.
Pappus often plumose, sometimes scaly, barbellate
or setaceous. Pollen with or without depressions,
microechinate, scabrate-microechinate, granulate
87
(sparsely microechinate), spinulate, smooth, or
rarely lophate. 2n = 16, 48, 50, 54, 100, 108.
Exclusively South American, in the Andes from
Venezuela to Patagonia in Argentina, in central
Chile and eastwards into Brazil. Nine genera and
91 species.
Key to the Genera
1. Herbs
2
– Subshrubs, shrubs or trees
4
2. Plants erect; leaves long and strap-shaped, hairy
1. Schlechtendalia
– Plants spreading; leaves various
3
3. Pappus plumose
2. Doniophyton
– Pappus scaly, ciliate
3. Duseniella
4. Capitula one-flowered
4. Fulcaldea
– Capitula with more than one flower
5
5. Capitula heteromorphic; pollen lophate
6
– Capitula isomorphic; pollen various
7
6. Shrubs or trees, 0.6–20 m tall; spiny
5. Barnadesia
– Subshrubs to 5 cm tall; unarmed
6. Huarpea
7. Apical appendage emarginate or bilobed
7. Dasyphyllum
– Apical appendage entire
8
8. Corollas pseudobilabiate (1/4); stamens inserted on
throat of corolla tube
8. Arnaldoa
– Corollas tubular, rarely pseudobilabiate or pseudoligulate; stamens inserted at base of corolla tube
9. Chuquiraga
Genera of Barnadesieae
1. Schlechtendalia Less.
Fig. 15A–E
Schlechtendalia Less., Linnaea 5: 242 (1830), nom. cons.
Perennial herbs, unarmed, to 1 m. Leaves basally
rosulate, on stems opposite, linear, sessile, amplexicaul, at apex mucronate, entire, parallel-nerved
with numerous nerves. Capitula homogamous, discoid, pedunculate, in racemose or cymose clusters. Involucre hemispherical, c. 5-seriate. Receptacle flat, pilose. Florets numerous, yellow, isomorphic, hermaphrodite; corollas 5-merous, pseudobilabiate (1/4), pilose. Stamens inserted near base
of corolla; filaments free; anthers shortly sagittate, with apical appendage entire. Styles bifid, papillose below bifurcation. Achenes turbinate, villous. Pappus plumose. Pollen with 1 depression per
mesocolpus, microgranulate, sparsely microechinate. 2n = 16. One species, S. luzulaefolia Less.,
endemic to Brazil, Uruguay and Argentina.
2. Doniophyton Wedd.
Fig. 15F–J
Doniophyton Wedd., Chloris Andina 1: 7, 8 (1855); Katinas
& Stuessy, Pl. Syst. Evol. 206: 33–45 (1997), rev.
88
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Chuquiraga Juss. “anomale” DC. (1838).
Chuquiraga Juss. sect. Gymnophoranta Remy (1848).
Herbs with some secondary growth, 2.5–8 cm.
Stems ascendent or decumbent, fasciculate spines
present or absent. Leaves alternate, sessile, linear
to linear-lanceolate, at apex spiny, entire, 1-nerved,
involute or plicate. Capitula heterogamous, discoid, sessile, solitary. Involucre hemispherical
to campanulate, 4–5-seriate. Receptacle flat or
convex, pilose. Florets 40–135, yellow, isomorphic;
corollas 5-merous, tubular, pilose. Marginal florets
female. Central florets hermaphrodite. Stamens
inserted at base of corolla tube; filaments free;
anthers long-sagittate, with apical appendage
entire. Style shortly bifid, papillose below bifurcation. Achenes turbinate, densely villous.
Pappus plumose. Pollen without depressions,
scabrate-microechinate. 2n = 48, 50. Two species
in northern Chile and Patagonian Argentina.
3. Duseniella K. Schum.
Fig. 15K–O
Duseniella K. Schum., Just’s Bot. Jahresber. 28, 1: 475
(1902).
Annual herbs, unarmed, to 10 cm. Leaves basally
opposite, becoming alternate further up, sessile,
linear to linear-lanceolate, at apex mucronate,
entire, 3-nerved. Capitula heterogamous, discoid, sessile, solitary. Involucre campanulate, c.
4-seriate. Receptacle convex, naked. Florets 30–35,
yellow, isomorphic, corollas 5-merous, tubular,
pilose. Marginal flowers female. Central flowers
hermaphrodite. Stamens inserted near base of
corolla; filaments free; anthers long-sagittate, with
apical appendage entire. Style bifid, papillose
below bifurcation. Achenes cylindrical, villous.
Pappus scaly, ciliate. Pollen without depressions,
microechinate. One species, D. patagonica K.
Schum., endemic to Patagonian Argentina.
4. Fulcaldea Poir. ex Lam.
Fulcaldea Poir. ex Lam., Encycl. Meth. Bot. suppl. 5: 375
(1817).
Shrubs or small trees, 3–4 m. Stems erect, spiny.
Leaves alternate, petiolate, elliptic, at apex spiny,
entire, glabrous, 3-nerved. Capitula one-flowered,
sessile, in corymbose cymes. Involucre cylindrical,
5–7-seriate. Receptacle flat, pilose. Florets violet
or white, hermaphrodite; corolla 5-merous, tubular, pilose. Stamens 5, inserted between throat and
base of corolla; filaments free; anthers obtuse, with
Fig. 15. Compositae-Barnadesieae. A–E Schlechtendalia
luzulaefolia. A Habit. B Pseudobilabiate corolla. C Style.
D Shortly sagittate anther. E Lanceolate pappus trichome.
F–J Doniophyton anomalum. F Habit. G Tubular corolla.
H Style. I Long sagittate anther. J Plumose pappus trichome.
K–O Duseniella patagonica. K Habit. L Tubular corolla.
M Style. N Long sagittate anther. O Lanceolate pappus
trichome
Compositae
89
apical appendage entire. Style shortly bifid, papillose and distinctly swollen below lobes. Achenes
cylindrical, villous. Pappus plumose. Pollen without depressions, spinulose. One species, F. laurifolia Poir. ex Lam., endemic to southern Ecuador and
north-western Peru.
5. Barnadesia Mutis
Fig. 16A–K
Barnadesia Mutis in L. f., Suppl. Pl. 55 (1782); Urtubey, Ann.
Missouri Bot. Gard. 86: 57–117 (1999), rev.
Bacasia Ruiz & Pav. (1794).
Diacantha Less. (1830), non Lag. (1811).
Penthea (D. Don) Spach (1841), non Lindl. (1838).
Shrubs or trees, 0.6–20 m. Stems erect, spiny. Leaves
alternate or fascicled, sessile to petiolate, ovate, elliptic to obovate, at apex mucronate or spinescent,
at base usually obtuse, entire, 1- or 3-nerved. Capitula heterogamous or homogamous, pseudoradiate,
sessile or pedunculate, solitary or in cymose aggregates. Involucre campanulate, turbinate or cylindrical, 6–14-seriate. Receptacle flat, pilose. Florets
9 or 16, pink, red, purple, rarely white, iso- or 2–
3-morphic, hermaphrodite or unisexual. Stamens
inserted on throat (or rarely at base of corolla tube
or between throat and base of corolla tube); filaments free or fused; anthers obtuse or bulging, with
apical appendage entire. Style bilobed, smooth below bifurcation. Achenes turbinate or cylindrical,
villous. Marginal florets 8 or 13, hermaphrodite;
corollas pseudobilabiate (1/4). Stamens 5. Pappus
plumose. Central florets 1 or 3, iso- or anisomorphic, hermaphrodite or unisexual; corollas tubular (3, 5-merous), pseudobilabiate (1/4), bilabiate
(1/3 or rarely 2/3), subligulate or ligulate (0/5). Stamens 3–5. Pappus plumose, barbellate or setaceous.
Pollen lophate, radially symmetric or asymmetric,
smooth. 2n = 50, 100. Nineteen species distributed
from the eastern Andes in Colombia southwards
into north-western Argentina and south-eastern
Brazil.
6. Huarpea Cabrera
Huarpea Cabrera, Bol. Soc. Argent. Bot. 4, 1/2: 129 (1951).
Subshrubs, unarmed, c. 5 cm. Leaves alternate,
subrosulate, sessile, amplexicaul, linear, at apex
spiny, entire, 1-nerved, revolute, xeromorphic.
Capitula heterogamous, sessile, solitary. Involucre
cylindric-campanulate, c. 4-seriate. Receptacle
flat, pilose. Florets 6, white, dimorphic, 5-merous.
Marginal florets 5, hermaphrodite; corollas pseudobilabiate (1/4). Central flower 1, male; corolla
Fig. 16. Compositae-Barnadesieae. A–K Barnadesia odorata. A Habit. B Tubular corolla. C Subligulate corolla. D Ligulate corolla. E Pseudobilabiate corolla. F Style. G Anther.
H Shortly sagittate anther. I Simple pappus trichome. J Barbellate pappus trichome. K Plumose pappus trichome. L–
S Chuquiraga erinacea var. erinacea. L Habit. M Tubular
corolla. N Subligulate corolla. O Subpseudobilabiate corolla.
P Pseudobilabiate corolla. Q Style. R Long sagittate anther.
S Plumose pappus trichome
90
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
tubular. Filaments of stamens free; anthers obtuse
or shortly sagittate, with entire apical appendage.
Style bilobed, smooth below bifurcation. Achenes
turbinate or cylindrical. Pappus plumose (central
florets with a single villous bristle or without
pappus). Pollen lophate, radially symmetrical,
smooth. One species, H. andina Cabrera, endemic
to San Juan Province of Argentina.
7. Dasyphyllum Kunth
Dasyphyllum Kunth in Humb., Bonpl. &. Kunth, Nova Gen.
Sp. ed. folio 4: 13, tab. 308 (1818); Cabrera, Revista Mus. La
Plata, secc. Bot. 9: 21–100 (1959), rev.
Flotovia Spreng. (1826).
Chuquiraga sect. Erinesa D. Don (1830).
Piptocarpha Hook. & Arn. (1835).
Flotowia Endlicher (1838).
Shrubs or trees, 0.5–20 m. Stems erect or decumbent, with or without spines. Leaves alternate,
fasciculate or whorled, sessile to petiolate, ovate,
elliptic to obovate, at apex mucronate or spiny,
entire, 1- or 3-nerved. Capitula homogamous
or heterogamous, discoid, sessile or pedunculate, solitary, glomerate, cymose or racemose.
Involucre campanulate, turbinate or cylindrical,
6–14-seriate. Receptacle flat, pilose, sometimes
with paleae. Florets 6–90, white to yellowish,
isomorphic, hermaphrodite or unisexual; corollas
5-merous, tubular, pseudobilabiate (1/4), subpseudobilabiate, rarely bilabiate (2/3), subligulate or
ligulate, pilose, rarely glabrous. Stamens inserted
on throat or near base of corolla tube; filaments
free; anthers sagittate, with apical appendage
emarginate or bilobed. Style bifid, papillose (rarely
hairy) below bifurcation. Achenes turbinate
or cylindrical, villous, rarely glabrous. Pappus
plumose. Pollen with 3–23 depressions (rarely
lacking), sparsely microechinate. Forty species,
distributed from Venezuela south to Chile and
Argentina and eastwards to Brazil.
ple, isomorphic, hermaphrodite; corollas pseudobilabiate (1/4). Stamens inserted on the throat of
the corolla tube; filaments free, with or without
barnadesioid hairs; anthers sagittate, with apical
appendage entire. Style bifid, papillate bellow bifurcation. Achenes turbinate or cylindric, densely
hirsute. Pappus plumose. Pollen with 4 paraporal
depressions for mesocolpi, microechinate. 2n = 48,
54. Three species endemic to northern Peru and
southern Ecuador, in xerophytic habitats.
9. Chuquiraga Juss.
Fig. 16L–S
Chuquiraga Juss., Gen. Pl.: 178 (1789); Ezcurra, Darwiniana
26: 219–284 (1985), rev.
Johannia Willd. (1803).
Joannesia Pers. (1807), orth. var.
Joannea Spreng. (1818), orth. var.
Intricately branched shrubs, 0.25–2 m. Stems
erect or compressed into cushions, often spiny.
Leaves alternate, opposite, whorled or fasciculate,
sessile, ovate, linear to obovate, at apex spiny; 1- or
3-nerved, flat or revolute, xeromorphic. Capitula
homogamous, discoid, sessile, solitary. Involucre
campanulate, turbinate or cylindrical, 4–8(–12)seriate. Receptacle alveolate, pilose. Florets 5–100,
yellow to orange, isomorphic, hermaphrodite;
corollas 5-merous, tubular, rarely pseudobilabiate
(1/4), subpseudobilabiate or subligulate, pilose.
Stamens inserted at base of corolla tube; filaments
free; anthers sagittate, with apical appendage
entire. Style bifid, papillate bellow bifurcation.
Achenes turbinate, villous. Pappus plumose. Pollen
without depressions, microechinate. 2n = 54, 108.
Twenty-three species from the Andes of Colombia
south into Argentina and Chile, frequently in
xeromorphic habitats.
II. Tribe Mutisieae Cass. (1819).
D.J.N. Hind
8. Arnaldoa Cabrera
Arnaldoa Cabrera, Bol. Soc. Argent. Bot. 10: 21–45 (1962);
Sagástegui Alva & Stuessy, Arnaldoa 1: 9–21 (1993), rev.
Shrubs, 1–4 m. Stems erect, with axillary spines.
Leaves alternate, shortly petiolate, ovate, elliptic or
obovate, at apex mucronate or spiny, at base attenuate or obtuse, entire, 3-nerved. Capitula homogamous, discoid, sessile, solitary. Involucre campanulate, 8–15-seriate. Receptacle flat, pilose. Florets
30–95, cream-white, light orange to orange or pur-
Herbs, subshrubs, shrubs, trees, rarely climbers
or ramblers, glabrous or with simple, glandular,
malpighiaceous or stellate hairs, often glabrescent.
Leaves usually evenly spaced, sometimes rosulate
or densely spiralled, usually alternate, rarely opposite, lamina simple, variously shaped, usually
herbaceous, venation trinervate, pinnate, sometimes parallel or very rarely palmate, margins entire or lobed, serrate or denticulate, rarely spiny
or pinnatisect, lamina rarely pinnate or impar-
Compositae
ipinnate with a simple or branched tendril. Inflorescences scapose or scapiform, cymose or of
corymbose or paniculate, axillary or terminal clusters, sometimes of glomerules, very rarely of true
syncalathia. Capitula small to very large, usually
chasmogamous, very rarely cleistomagous, usually
monoecious, homogamous or heterogamous, radiate, very rarely ligulate, rarely disciform or discoid, one- to many-flowered; involucres cylindrical
to globular or urceolate; phyllaries imbricate, fewto many-seriate, sometimes uniseriate, rarely calyculate, rarely distant, usually gradate, often chartaceous or herbaceous, usually homomorphic; receptacles flat, convex or rarely conical, scrobiculate,
foveolate, fimbrillate or alveolate, glabrous or variously pubescent, usually epaleaceous. Florets rarely
all actinomorphic, usually marginal and disc florets
distinct; marginal florets usually variously bilabiate
(2/3) or pseudobilabiate (1/4), and often distinctly
rayed, sometimes ligulate (0/5), hermaphrodite,
female or neuter, corollas glabrous or variously
pubescent; staminodes rarely present; disc florets
usually bilabiate (2/3), sometimes actinomorphic
(5/0), usually fertile; corollas glabrous or variously
pubescent, lobes short or long. Stamens usually
conspicuously exserted from corolla; filaments glabrous or rarely pubescent or papillate, collars inconspicuous or sometimes distinct and variously
enlarged or flattened; anther appendages usually
acuminate or apiculate and several times as long
as wide, sometimes thickened at apex and knoblike, sometimes truncate or rounded; anthers calcarate and caudate, rarely ecalcarate, tails usually
long-acute, entire or variously laciniate, sometimes
conspicuously branched or pilose. Pollen tricolporate, exine tectate, perforate, psilate, spinulose
or echinate, tectum poorly differentiated or with
distinct columellae. Styles usually well exserted
from corolla and anther cylinder; style base sometimes with conspicuous nectary, with or without
distinctive basal node, glabrous, style shaft usually glabrous, rarely papillose in upper part, style
arms usually relatively short, apices acute, obtuse
to rounded, or (Nassauviinae) truncate and usually penicillate, often with a lip around inner margins, usually apically pilose, hairs acute or obtuse
to rounded. Achenes fusiform or sometimes distinctly beaked, terete, ribbed or angled or very
rarely flattened, glabrous or variously setuliferous,
commonly with twin-hairs, or rarely tomentose
with long tortuous hairs, or appearing papillate,
very rarely with stalked and sticky glands (Adenocaulon); carpopodium sometimes absent, more
91
usually a narrow annulus, sometimes cylindrical;
pappus sometimes absent but usually of uniseriate,
sometimes biseriate to multiseriate, simple, barbellate, subplumose or plumose bristles, sometimes
flattened or scale-like, persistent or deciduous, separate or sometimes basally connate.
Eighty-two genera with more than 950 species,
most in South America but also in North America, Africa/Madagascar and Asia; one genus in Australia.
Panero and Funk (2002) placed several genera
of Mutisieae into separate tribes and subfamilies.
This treatment is not recognized here.
Classification of Mutisieae s. l.
1. Stifftia group
Genera 10–12
2. Stenopadus group
Genera 13–23
3. Subtribe Nassauviinae
Genera 24–47
Problematic placement (probably within Nassauviinae)
Genera 48–49
4. Subtribe Mutisiinae
Genera 50–65
5. Subtribe Gerberinae
Genera 66–72
6. Subtribe Gochnatiinae
Genera 73–75
7. Hecastocleis group
Genus 76
8. Nouelia group
Genera 77–78
9. Catamixis group
Genus 79
10. Subtribe Tarchonanthinae
Genera 80–81
11. Dicoma group
Genera 82–87
12. Pertya group
Genera 88–91
Key to the Genera
1. All capitula ligulate, or rarely with mixtures of ligulate
and bilabiate florets in one capitulum as well as ligulate
capitula on one plant
2
– Capitula containing actinomorphic, bilabiate, or pseudobilabiate florets, or mixtures thereof
4
2. Corollas red; leaves buff-tomentose beneath; Guyana
Highlands
14. Glossarion (G. rhodanthum)
– Corollas cream or yellow; leaves glabrous, glabrescent
or arachnoid pubescent; Argentina and Bolivia or Himalayas
3
3. Achenes glabrous or with moderately long setulae and
stipitate-glandular; style arms long; capitula few to
several in terminal or axillary clusters or solitary and
terminal; leaves usually opposite (sometimes fasci-
92
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
–
4.
–
5.
–
6.
–
7.
–
8.
–
9.
–
10.
–
11.
–
12.
–
13.
–
14.
–
15.
–
16.
–
17.
culate on axillary brachyblasts); phyllaries yellow;
pappus setae few-seriate; Argentina and Bolivia
11. Hyaloseris
Achenes densely long-setuliferous and appearing
sericeous; style arms short; capitula numerous in
large terminal corymbs; leaves alternate; phyllaries
greenish often with purplish apices; pappus setae
uniseriate; western Himalayas, India 79. Catamixis
Style arm apices flattened (subtribe Tarchonanthinae)
5
Style arm apices cylindrical or swollen
6
Pappus absent; capitula usually hemispherical or globular
81. Tarchonanthus
Pappus present; capitula turbinate or campanulate
80. Brachylaena
Receptacles entirely (or partially) paleaceous
7
Receptacles epaleaceous
15
Pappus absent (or reduced to a few (3–5) caducous
flattened setae); achene setulae basally forked twinhairs
85. Erythrocephalum
Pappus of capillary, or flattened, barbellate or plumose
setae (rarely absent on central achenes); achene setulae, when present, twin-hairs (apices connate or
forked)
8
Florets all actinomorphic; style arm apices acute or
obtuse
9
Florets all bilabiate; style arm apices truncate (subtribe
Nassauviinae p.p.)
11
Corollas magenta, sometimes with golden lobes
21. Stenopadus
Corollas cream to yellowish
10
Corollas glabrous; pappus setae subpaleaceous with
margins densely plumose towards apices, strawcoloured; corolla lobes tightly coiled; Brazil (Bahia
and Goías southwards)
12. Wunderlichia
Corollas with dense tufts of hairs at bases of sinuses
of lobes; pappus setae flattened with barbellate margins, cream-coloured; corolla lobes erect or slightly
recurved; Guyana Highlands
20. Stomatochaeta
Leaves linear, margins strongly revolute, entire; corollas yellow; pappus biseriate
37. Pleocarphus
Leaves lanceolate, ovate or orbicular, margins flat, entire, lobed or pinnatisect, sometimes spiny; corollas
white, pink, purple, rarely yellow; pappus setae uniseriate
12
Leaves reduced to spiny segments
29. Oxyphyllum
Leaves with relatively broad, unarmed lamina
13
Leaves (at least lower) sessile
43. Marticorenia
Leaves usually distinctly petiolate
14
Leaf margins pinnatisect; florets 2 per capitulum; phyllaries biseriate, outer foliaceous; annual herbs
46. Moscharia
Leaf margins entire or lobed; florets few to many
per capitulum; phyllaries uniseriate, sometimes with
an outer calyculus, never foliaceous; perennial herbs,
shrubs or lianes
36. Jungia
Style arms truncate or rarely rounded; style hairs obtuse to rounded (subtribe Nassauviinae p.p.)
16
Style arms acute or obtuse; style hairs mostly acute 37
Achenes epappose
17
Achenes pappose
20
Achenes rostrate; inflorescences scapiform, capitula
usually solitary; receptacle pubescent
47. Cephalopappus
– Achenes erostrate; inflorescences lax cymes, corymbs,
panicles or capitula solitary, axillary; receptacle glabrous
18
18. Achenes (of female florets) conspicuously stipitateglandular (of male florets, glandular)
48. Adenocaulon
– Achenes glabrous, sparsely papillose or densely lanose
19
19. Corollas bilabiate; annual herbs; leaves petiolate, leaf
bases not amplexicaul; inflorescences lax few-headed
corymbs; all florets hermaphrodite and fertile
45. Pamphalea
– Corollas actinomorphic, 5-lobed; perennial rhizomatous herbs; leaves sessile and pseudopetiolate,
leaf bases amplexicaul; inflorescences solitary axillary capitula; marginal florets female, disc florets
hermaphrodite and functionally male
49. Eriachaenium
20. Inflorescence scapiform (or with few capitula) and
arising from a basal rosette of leaves
21
– Inflorescences glomerules, cymes, panicles or capitula
solitary, terminal on leafy stems
22
21. Leaf bases vaginate about scape; corollas white; pappus
setae plumose
30. Macrachaenium
– Leaf bases clasping stem but not vaginate; corollas
orange; pappus setae coarsely barbellate 41. Criscia
22. Inflorescences glomerules
23
– Inflorescences cymes, corymbs, panicles or capitula
solitary
25
23. Pappus of coarse setae; leaf lamina pinnately lobed or
pinnatisect, margins unarmed, arachnoid or tomentose; capitula 2-(rarely 3-)flowered 31. Polyachyrus
– Pappus of caducous scales; leaf lamina entire or pinnatisect with denticulate spiny margins, or with few
long spines; capitula 3–6-flowered
24
24. Pappus scales linear or with broad flattened rachis,
margins ciliate to plumose; corollas white or rarely
violet or yellowish
27. Nassauvia
– Pappus scales apically lacerate-plumose; corollas
white or blue, often both on same plant 28. Triptilion
25. Pappus of caducous scales
27. Nassauvia
– Pappus setae capillary or broadened at base
26
26. Receptacles always glabrous
27
– Receptacles usually pubescent (with hairs or papillae)
30
27. Annual or perennial rosulate herbs; corollas white,
pink, lilac or purple
28
– Low shrubs; corollas yellow
29
28. Pappus setae biseriate, capillary, barbellate; corollas
white
44. Holocheilus
– Pappus setae usually uniseriate, flattened and broadened at base, barbellate or plumose; corollas white,
pink, lilac or purple
35. Leucheria
29. Achenes with stipitate-glandular beak; leaves pinnatisect, sparsely to moderately stipitate-glandular; capitula large and on elongated pedicels 39. Dolichlasium
– Achenes cylindrical and lacking stipitate-glandular setulae; leaves coarsely dentate and glandular-punctate;
capitula relatively small and short-pedicellate
40. Ameghinoa
30. Stems armed either with pairs of recurved spines at
nodes or with spines terminating branches or axes of
inflorescences
31
– Stems and inflorescence axes completely unarmed 32
Compositae
31. Clambering woody vine-like plants; stems bearing
a pair of recurved persistent spines at each node; inflorescences axillary cymes; corollas yellow; Greater
Antilles
25. Berylsimpsonia
– Stems and side branches often terminating in a spine,
but otherwise unarmed; inflorescences racemes or
panicles; corollas pink or purple; Argentina, Bolivia,
Chile, Peru
24. Proustia
32. Densely caespitose dwarf shrubs; leaves acicular with
expanded bases, margins highly revolute; Argentina
(Patagonia)
34. Burkartia
– Herbs, subshrubs, shrubs or small trees; leaves with
relatively broad lamina, ovate, broadly ovate, oblanceolate, spathulate, cordate or lyrate-pinnatifid
33
33. Rosettiform sometimes caespitose herbs, rarely tall
and leafy; inflorescences solitary and sessile in leaf
rosette, scapiform with solitary or few capitula or fewheaded panicles; stem leaves, if present, sessile and
amplexicaul but never decurrent; corollas blue, purple,
violet, red, or crimson, sometimes yellow, but rarely
white or cream
32. Perezia
– Subshrubs, shrubs (sometimes scandent or trailing)
or small trees or, if herbaceous, then inflorescences
corymbose and stem winged or with decurrent based
leaves
34
34. Corollas typically pinkish light purple or purple,
rarely white; subshrubs (stems with basal rosette
of leaves, appearing scapiform) or densely leafy
scandent shrubs; style arms > 1 mm long with
rounded apices
33. Acourtia
– Corollas white, yellow or orangish; non-scapiform
subshrubs, laxly leafy scandent shrubs or often
densely leafy shrubs; style arms < 1 mm long and
rounded or > 1 mm long and truncate or if > 1 mm
long and rounded, then capitula solitary, large and
terminal
35
35. Low subshrubs; inflorescence large terminal solitary
capitula; leaves and phyllaries with excentrically
branched ‘T’-shaped and stipitate-glandular hairs;
phyllaries biseriate and almost distant; anther
cylinder exserted from corolla throat; pappus setae
broadened at base and often branched, off-white;
Chile
42. Leunisia
– Herbs, subshrubs, scandent or trailing shrubs or small
trees and, if subshrubs, then capitula never large nor
solitary; leaves and phyllaries glabrous or with flagellate eglandular and stipitate-glandular hairs; phyllaries (1–)2–5-seriate and usually imbricate; anther
cylinder only partially exserted from corolla throat;
pappus setae of ± uniform diameter and simple, white,
straw-coloured, yellowish or reddish
36
36. Inflorescences usually terminal cymes, corymbs or
panicles, sometimes pseudoglomerules; style arms
> 1 mm long, apices truncate; receptacle pilose
or densely long-pubescent; corollas yellow, orange
or rarely white, variously pubescent or sometimes
glabrous; pappus setae 2–3(–4)-seriate
38. Trixis
– Inflorescences terminal solitary or tightly grouped
clusters of small capitula; style arms < 1 mm long,
apices rounded; receptacle papillate; corollas white or
yellowish white, glabrous; pappus setae uniseriate
26. Lophopappus
37. Florets all actinomorphic
38
93
– Florets clearly bilabiate, pseudobilabiate or pseudoligulate, capitula sometimes heterogamous and
radiate
55
38. Plants dioecious, capitula with only staminate or only
pistillate flowers
39
– Plants not dioecious, capitula otherwise
40
39. Corollas whitish or cream; style shaft glabrous; inflorescences many-headed, usually leafy panicles or
corymbs, capitula sessile or short-pedicellate
73. Gochnatia (sect. Moquiniastrum)
– Corollas purple; style shaft short-pilose beneath style
arms; inflorescence of solitary sessile or subsessile
capitula on brachyblasts subtended by bud and leafscales in 2–3 series
91. Myripnois
40. Plants soon appearing leafless (leafy only on young
shoots); stems terminating in spine
60. Cyclolepis
– Plants conspicuously leafy; stems unarmed
41
41. Basal half of stem with only scale leaves; normal leaves
(large and obovate or trilobed) rosulate or alternate at
base of inflorescence; pappus setae usually somewhat
flattened and dilated at apices
89. Macroclinidium
– Leaves rosulate at base or apex of stem or alternate
and descrescent upwards or leaves similar throughout,
sometimes with axillary brachyblasts, leaves small or
medium-sized; pappus setae of uniform width, if flattened, then narrowing towards apex or reduced to
a corona
42
42. Pappus a scale-like corona; capitula clustered in centre
of secondary spiny foliaceous bracts; USA (Nevada and
California)
76. Hecastocleis
– Pappus of capillary or paleaceous setae or absent; capitula free or in aggregations lacking outer secondary
foliaceous bracts; outside USA
43
43. Corolla lobes stiff and erect and sinuses of lobes
densely pubescent; inflorescences solitary terminal
capitula embedded in densely tomentose, densely
leafy stem apices
19. Chimantaea
– Corolla lobes usually recurved or tightly coiled and sinuses of lobes glabrous; inflorescences multi-headed
or if one-headed, then stem apices not densely tomentose and capitulum not embedded in apical leaves 44
44. Capitula solitary on relatively long, slender multibracteolate axillary pedicels, bracteoles scale-like;
corolla lobes tomentose; receptacle with alveolae
margins ciliate or fimbriate between achenes; corollas
bright yellow
65. Chucoa
– Capitula in corymbs or panicles and sessile or shortpedicellate, if solitary, then sessile or on short pedicels
and terminal or axillary and bracteoles scale-like or
leaf-like; corolla lobes glabrous, glandular-punctate or
pilose; receptacle with alveolae margins scarcely discernible or appearing honeycombed; corollas white,
cream red, violet, purple, brownish, rarely yellow and
then often pale
45
45. Pappus uniseriate and plumose (rarely absent); plants
with basal rosettes of leaves, or with short naked
basal portion of stem beneath leaf rosette or rarely
leafy throughout; inflorescences usually spiciform,
racemose or narrow-paniculate; achene setulae, when
present, short or long twisted twin-hairs
90. Ainsliaea
– Pappus usually 2- to 3-seriate, barbellate or plumose
or combinations of barbellate and plumose setae or
94
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
46.
–
47.
–
48.
–
49.
–
50.
–
51.
–
52.
–
53.
–
54.
if uniseriate, then barbellate and setae united at base;
inflorescences of solitary axillary or terminal capitula,
or of many-headed panicles or compound corymbs;
achene setulae, when present, straight twin-hairs 46
Phyllary apices pungent, outer phyllaries narrow and
usually squarrose
84. Dicoma
Phyllary apices acute, obtuse or rounded, outer phyllaries usually erect and never pungent
47
Leaves congested and sometimes fasciculate on
brachyblasts, or evenly spaced on stems and
uniform; capitula usually solitary and sessile or
short-pedicellate on brachyblasts or stem apices or
in few-headed panicles; leaves small and subulate,
lanceolate or oblong to ovate; anther thecae drying
brownish or blackish
88. Pertya
Stems basally or apically rosulate or stems leafy
throughout and leaves often descrescent towards
inflorescence; inflorescences scapiform (single- or
few- to many-headed), cymose, pseudocorymbose or
pseudopaniculate with many capitula or capitula large
terminal and solitary; leaf laminas medium to large;
anther thecae not drying brownish or blackish
48
Capitula with 1 floret
49
Capitula with 2 or more florets
50
Pappus setae straw-coloured; plants poorly branched
shrubs or monopodial small trees
22. Quelchia
Pappus setae magenta; plants clambering shrubs or
vines
10. Stifftia (S. uniflora)
Apical anther appendages acuminate to long-caudate
and knob-like at very tip
86. Pleiotaxis
Apical anther appendages apiculate or lanceolate to
long-acute
51
Leaves usually sticky and distinctly pockmarked and
glandular-punctate; corollas lilac or white; pappus
setae broad and flattened at base; inflorescences dense
glomerules or pseudoracemes
74. Pentaphorus
Plants lacking glandular punctae; corollas yellow
to orange, cream or white; pappus setae capillary,
sometimes coarse; inflorescences corymbs, racemes,
sometimes capitula solitary, terminal or axillary 52
Inflorescences dense terminal corymbs or glomerules;
achene setulae long twisted twin-hairs; Old World
(Asia)
78. Leucomeris
Inflorescences scapiform, of solitary terminal capitula or of cymes or panicles; achenes glabrous
or setuliferous and setulae straight twin-hairs,
stipitate-glandular, or uniseriate; New World (Central
and South America)
53
Style arms markedly divergent, relatively long; apical
anther appendages lanceolate to long-acute; achenes
cylindrical, glabrous or very sparsely setuliferous,
setulae short stipitate glands or eglandular uniseriate
and multicellular; pappus setae 4–5-seriate, usually
spreading, of plants with dense axillary or terminal
cymes straw-coloured, of those with solitary capitula
highly coloured and orange, reddish, purple or
magenta
10. Stifftia
Style arms scarcely bifid and very short; apical
anther appendages usually apiculate; achenes usually
turbinate, usually densely setuliferous, setulae
twin-hairs and sometimes stipitate glands; pappus
setae 1–2-seriate, erect, straw-coloured
54
Shrubs or trees; inflorescences solitary or clustered
terminal capitula or capitula in many-headed cymes,
–
55.
–
56.
–
57.
–
58.
–
59.
–
60.
–
61.
–
62.
–
63.
–
64.
–
65.
–
66.
–
pseudocorymbs or pseudopanicles; capitula usually
sessile, subsessile or short-pedicellate; pappus setae
free to base
73. Gochnatia
Small subshrubs; inflorescences scapiform with
a long, distinct peduncle above leaves; capitula
usually long-pedicellate; pappus setae basally connate
into distinct annulus
75. Richterago
Plants appearing leafless, minute linear-spathulate
leaves falling rapidly
58. Aphylloclados
Plants conspicuously leafy, stems leafy throughout or
with apical or basal rosettes of leaves
56
Plants dioecious, capitula with only pistillate or only
staminate flowers
55. Lycoseris
Plants not dioecious, capitula otherwise
57
Pappus setae dimorphic, outer series multiseriate,
capillary and barbellate, inner series of awn-like
scales with laciniate margins
51. Urmenetia
Pappus setae monomorphic, all barbellate or plumose
58
Achenes obcompressed with 2 or 4 ribs
64. Lulia
Achenes cylindrical, fusiform or turbinate
59
Leaves with simple or branched tendrils at apices,
leaves often pinnate, sometimes simple 50. Mutisia
Leaves lacking tendrils at apices
60
Capitula radiate and marginal (ray) florets with conspicuous ray limb, either with marginal floret corollas
bilabiate and disc floret corollas actinomorphic, or
marginal and disc floret corollas bilabiate
61
Capitula discoid and all floret corollas bilabiate and
hermaphrodite
82
Marginal (ray) floret and disc floret corollas all
bilabiate
62
Marginal (ray) floret bilabiate and disc floret corollas
actinomorphic
72
Pappus setae plumose
63
Pappus setae barbellate
64
Prostrate rhizomatous rosettiform perennial herbs or
subshrubs; leaves fleshy
52. Pachylaena
Erect, ascending or clambering subshrubs or shrubs;
leaves usually herbaceous
50. Mutisia (sect. Holophyllum, sect. Fruticosa)
Leafy-stemmed erect or prostrate annual or perennial
herbs or dense caespitose or lax subshrubs or shrubs;
inflorescences of solitary sessile or subsessile terminal
capitula or a short few-headed cyme; ray florets female
and lacking staminodes, neuter or hermaphrodite 65
Acaulescent rosulate scapigerous herbs; scapes usually
very long and with few to several scale-like bracteoles,
scapes often markedly elongating in fruit; ray florets
usually with staminodes (subtribe Gerberinae)
66
Densely caespitose shrubs with sessile capitula; outer phyllaries not foliaceous; corolla tube
arachnoid-pubescent or glabrescent, usually sparsely
glandular-punctate
53. Brachyclados
Herbs or very rarely small shrubs with more or less
pedicellate capitula; outer phyllaries often foliaceous;
corolla tube glabrous
54. Chaetanthera
Plants with stout rhizomes; involucres mostly broad
and hemispherical; achenes short and ovoid; anther
filaments papillate; achene setulae flattened and
spathulate; ray limbs pubescent and sometimes
glandular-punctate
66. Trichocline
Plants usually with slender rhizomes and often wiry
or fibrous roots; involucres mostly turbinate; achenes
Compositae
67.
–
68.
–
69.
–
70.
–
71.
–
72.
–
73.
–
74.
–
75.
–
76.
usually long, cylindrical, sometimes beaked; anther
filaments glabrous; achene setulae inflated or short
or very long twin-hairs or absent; ray limbs usually
glabrous
67
Receptacles fimbriate
68
Receptacles alveolate
69
Achenes densely silky setuliferous with abundant, very
long, white pointed setulae; capitula conspicuously
radiate and limb of ray florets long and conspicuous,
exceeding pappus, pink; length of scape at least twice
leaf length; Turkish Armenia and central Asia
69. Uechtritzia
Achenes glabrous or densely papillate; capitula
appearing discoid with limb of ray florets very short,
slender, strap-shaped, white; length of scape scarcely
exceeding leaf length; South Africa
71. Perdicium
Capitula with only ray and disc florets
70
Capitula with ray, submarginal and disc florets
71
Plants with vernal (radiate and chasmogamous) and
autumnal (cleistogamous) generations of capitula;
achene setulae fine with acute apices, apices scarcely
divided or rarely divided to base; ray limbs concolorous above and beneath but sometimes with limb
(pink or purple) differently coloured from tube (white
or cream); Central America and Asia 68. Leibnitzia
Plants with only one generation of capitula; achene
setulae ‘sausage-shaped’ with rounded apices, apices
never divided; ray limbs discolorous, white above and
pink or purple beneath; Australia 70. Amblysperma
Pappus setae usually uniseriate and often united at
base, setae fine; ray limbs shorter than involucre;
corollas usually white, rarely purplish; capitula either
nodding or erect in bud and flower becoming erect in
fruit; plants often with vernal (radiate and chasmogamous) and autumnal (cleistogamous) generations of
capitula; Central and South America 67. Chaptalia
Pappus setae usually biseriate, setae relatively coarse;
ray limbs usually considerably longer than involucre;
corollas white, yellow, pink, red, purple; capitula always erect; plants with only chasmogamous capitula;
East and South Africa, Madagascar, southern Asia
72. Gerbera
Corollas intense orange to orange-red; phyllaries with scarious apical appendages or linear to
linear-lanceolate
56. Cnicothamnus
Corollas white, yellow, pink, reddish or purple;
phyllaries lacking apical appendages
73
Leaves filiform; ray florets purple; Chile
59. Gypothamnium
Leaves round, elliptic, lanceolate, hastate or lyrate;
ray florets usually white or pink (if violet, then leaves
broad)
74
Pappus setae purple
62. Ianthopappus
Pappus setae whitish, straw-coloured or fawn
75
Pachycaul shrubs or small trees with very densely
tomentose, poorly branched stems; South Africa
82. Oldenburgia
Annual to perennial herbs or well-branched shrubs or
trees, sometimes simple-stemmed or poorly branched
subshrubs and inflorescences appearing scapiform 76
Achene base with dense tuft of setulae, otherwise
setuliferous between ribs; ray corollas deep red (rarely
white); ray florets neuter
83. Passacardoa
95
– Achenes glabrous, glandular-punctate or uniformly
setuliferous; ray corollas white, pink, purple or violet;
ray florets hermaphrodite or female
77
77. Florets few (5–6); leaf pubescence of malpighiaceous
hairs; leaves linear-lanceolate to oblong-lanceolate
93. Hyalis
– Florets numerous (> 10); leaf pubescence absent or of
simple hairs; leaves round, ovate, hastate or lyrate 78
78. Ray or marginal floret corollas pink, purple or violet
61. Onoseris
– Ray or marginal floret corollas white
79
79. Tall shrubs or small trees; capitula large; Old World
(China)
77. Nouelia
– Small shrubs or subshrubs; capitula medium; Old
World (Madagascar) or New World (Argentina,
Bolivia, Brazil, Chile, Peru)
80
80. Inflorescences scapiform with solitary capitula or
few capitula in panicles; achenes setuliferous, setulae
twin-hairs; New World (Brazil)
75. Richterago
– Inflorescences solitary, terminal and sessile and
surrounded by apical leaves; achenes glabrous or
glandular-punctate; Old World (Madagascar) or New
World (Argentina, Bolivia, Chile, Peru)
81
81. Stems and leaves densely white- to grey-tomentose,
leaves concolorous; achenes glandular-punctate; Old
World (Madagascar)
87. Gladiopappus
– Stems resinous and short-pubescent, leaves sticky
and glabrous above and densely pubescent beneath;
achenes glabrous; New World (Argentina, Bolivia,
Chile, Peru)
57. Plazia
82. Pappus setae plumose; corollas yellow
50. Mutisia (sect. Isantha)
– Pappus setae barbellate; corollas white, red or purple,
rarely pale yellow
83
83. Phyllaries subequal and pubescent throughout;
pappus setae caducous; capitula solitary on long, very
sparsely bracteolate peduncles
16. Eurydochus
– Phyllaries gradate and glabrous or pubescent only
at apices; capitula, if solitary, on relatively short
peduncles with leaf-like bracts, otherwise few to many
capitula in corymbs or cymes
84
84. Florets few (2–5, rarely 6)
85
– Florets several to numerous (8–60)
87
85. Inflorescences terminal and many-headed; corollas
reddish; achenes setuliferous; Dominican Rep.
23. Salcedoa
– Inflorescences axillary; corollas white; achenes
glabrous; Guyana Highlands
86
86. Outer phyllaries with densely pubescent apices;
sinuses of corolla lobes sparsely to moderately
pubescent; phyllaries c. 4-seriate; adjacent basal
anther appendages distinct; inflorescences axillary,
of solitary capitula or few-headed cymes, often
obscured by subtending leaves or in terminal clusters
on ascending, leafy subterminal flowering stems,
capitula usually erect
17. Achnopogon
– Outer phyllaries pubescent throughout; sinuses of
corolla lobes glabrous; phyllaries 6–8-seriate; adjacent
basal anther appendages connate; inflorescences
axillary of few (1–)2–9 capitula, capitula usually
pendulous
18. Neblinaea
87. Leaves linear or linear-lanceolate and conspicuously
1-veined; corollas white, reddish or purple; achenes
glabrous or setuliferous
13. Duidaea
96
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
– Leaves broadly lanceolate, oblanceolate or obovate,
with reticulate, pinnate or subparallel venation;
corollas white, pale yellow or red; achenes glabrous 88
88. Corollas red; inflorescence of subsessile or shortpedunculate solitary terminal capitula; outer phyllaries floccose or glabrescent; leaves buff-tomentose
beneath; florets 12–22
14. Glossarion
– Corollas white or pale yellow; inflorescence of
long-pedunculate large solitary terminal capitula or
few- to many-headed and corymbiform or umbelliform; outer phyllaries glabrous; leaves glabrous or
pubescent beneath; florets 6–60
15. Gongylolepis
II.1. Stifftia Group
Tribe Stifftieae D. Don (1830).
Shrubs, vines or trees. Leaves alternate or opposite, coriaceous. Inflorescence solitary terminal
capitula or paniculate or racemose; capitula
discoid; phyllaries few- to many-seriate; involucres
cylindrical, campanulate or globose; receptacle
epaleaceous or paleaceous. Florets few to many,
rarely solitary; corollas bilabiate, actinomorphic
or rarely ligulate. Achenes cylindrical or fusiform,
glabrous or sparsely setuliferous; carpopodium
annular to short-cylindrical; pappus setae 3–5seriate, capillary and barbellate or subpaleaceous
and densely plumose.
10. Stifftia J.C. Mikan
Stifftia J.C. Mikan, Delec. Fl. Faun. Brasil. I 1, tab. 1 (1820),
nom. cons., Robinson, Syst. Bot. 16: 685–692 (1991), key;
Hind & Semir, Kew Bull. 53: 617–622 (1998), part. rev.
Shrubs, vines or trees. Leaves persistent, alternate,
simple, lamina herbaceous or coriaceous. Capitula
in dense axillary cymes, few-headed apical clusters,
or solitary, 1- to many-flowered, homogamous, discoid; involucres narrowly cylindrical to turbinate;
phyllaries few- to many-seriate, subimbricate
to imbricate, usually with many smaller bracts
grading down pedicels; receptacle flat to slightly
convex, epaleaceous. Florets actinomorphic,
hermaphrodite; corollas deeply 5-lobed, whitish
to yellow or orangish yellow, lobes linear, tightly
coiled; basal anther appendages long-caudate,
short-papillate or laciniate; style base lacking
basal node, glabrous, shaft glabrous, arms short,
divergent, rounded to acute, glabrous. Achenes
cylindrical, glabrous or sparsely setuliferous;
carpopodium annular to short-cylindrical; pappus
setae 4–5-seriate, persistent, prominent, often
brightly coloured. Eight species, Brazil, French
Guyana. One species is relatively widely cultivated
in the Tropics.
11. Hyaloseris Griseb.
Hyaloseris Griseb., Symb. Fl. Argent.: 212 (1879); Ariza
Espinar, Kurtziana 7: 195–211 (1973), rev.
Dinoseris Griseb. (1879).
Shrubs or small trees. Leaves opposite, simple,
lamina lanceolate, narrowly elliptic to elliptic,
denticulate or entire, apices acute or obtuse. Capitula two to several subsessile in dense terminal
or axillary clusters or terminal, homogamous,
few- or many-flowered, usually appearing ligulate;
involucre cylindrical (in Hyloseris s.str.) or campanulate (in Dinoseris), usually surrounded by
dense scale-like bracteoles; phyllaries 6–7-seriate,
imbricate, gradate, all straw-coloured; receptacle
small, epaleaceous, glabrous. Corollas off-white
or cream to yellow, glabrous, ligulate or bilabiate
and then with 3- to 4-toothed outer lip; basal
anther appendages extremely long, long-caudate,
retrorsely long-pilose towards base; style base
lacking basal node, glabrous, shaft glabrous,
arms long, divergent and often recurved, acute
and short-papillose. Achenes fusiform to longfusiform, 5- or 10-ribbed; carpopodium large,
procurrent in upper part with body; pappus setae
few-seriate, persistent, coarsely barbellate, offwhite to straw-coloured. Seven species, Argentina,
Bolivia.
12. Wunderlichia Riedel ex Benth. & Hook. f.
Fig. 17
Wunderlichia Riedel ex Benth. & Hook. f., Gen. Pl. 2, 1: 489
(1873); Barroso & Maguire, Revista Brasil. Bot. 33: 379–406
(1973), rev.
Shrubs or trees. Leaves alternate, deciduous, lamina elliptic, oblong-elliptic or broadly obovate to
orbicular, coriaceous. Capitula solitary, terminal,
or few to several in dense panicles or lax racemes
to scorpioid cymes, usually erect, medium or large,
homogamous, discoid; involucre campanulate,
globose or infundibuliform; phyllaries 4–10seriate, gradate, imbricate, persistent; receptacle
flat to convex, paleaceous. Florets actinomorphic,
many (to 300+), yellowish to cream; corollas
glabrous, lobes linear, coiled throughout or only
at apex; filaments long, often ‘swan-necked’ at
anthesis; basal anther appendages caudate, usually
entire or appearing somewhat contorted; style
base with enlarged basal node, or node absent
Compositae
97
13. Duidaea S.F. Blake
Duidaea S.F. Blake, Bull. Torrey Bot. Club 58: 496 (1931);
Pruski, Fl. Venez. Guayana 3: 261–263 (1997), reg. rev.
Fig. 17. Compositae-Mutisieae. Wunderlichia cruelsiana.
A Leaf. B Inflorescence. C Floret. (Drawings by Margaret
Tebbs)
and distinctive nectary present, glabrous, style
glabrous, long-exserted from anther cylinder,
arms short, scarcely divided, or appearing connate/adnate. Achenes 10-ribbed; carpopodium
very narrow, pale; pappus setae 3–4-seriate, falling
as a unit, subpaleaceous, sometimes barbellate
below and densely plumose above, straw-coloured.
Five species, endemic to Brazil (Bahia, Espírito
Santo, Goiás, Minas Gerais, Rio de Janeiro).
II.2. Stenopadus Group
Small to large shrubs or trees. Leaves alternate,
coriaceous. Inflorescence solitary axillary or terminal capitula or cymes, corymbs or glomerules;
capitula homogamous, discoid or ligulate; involucres campanulate or cylindrical; phyllaries
3–8-seriate; receptacles usually paleaceous; florets
hermaphrodite, few to many, rarely solitary; corollas actinomorphic and deeply 5-lobed or ligulate or
bilabiate. Achenes cylindrical, ribbed, glabrous or
setuliferous; pappus setae capillary and barbellate
or flattened with barbellate to subplumose margins.
Shrubs (with extremely woody bases) or dwarf
trees. Leaves alternate or densely spiralled, simple,
lamina linear, linear-lanceolate or oblanceolate.
Capitula usually solitary, axillary or subterminal,
or on medium-length pedicels, homogamous;
involucre hemispherical or cylindrical to campanulate; phyllaries c. 3-seriate, imbricate,
gradate; receptacle scarcely convex, epaleaceous,
pubescent. Florets few to many (8–24), bilabiate,
hermaphrodite, fertile; corollas white, red or
reddish purple, outer lip short 3-toothed, scarcely
curved or apparently strongly coiled, inner of two
long coiled lobes; basal anther appendages caudate
(sometimes very long), rough or pilose, obtuse
or acute; style base lacking obvious basal node,
glabrous, shaft glabrous, arms moderately long,
usually recurved/coiled at maturity, apices obtuse
to truncate. Achenes c. 10-ribbed, cylindrical;
carpopodium annular; pappus setae biseriate,
flattened at base, margins almost subplumose,
coarsely barbellate above, straw-coloured. Four
species, Venezuela (Venezuelan Guyana).
14. Glossarion Maguire & Wurdack
Glossarion Maguire & Wurdack, Mem. New York Bot. Gard.
9: 390 (1957); Pruski, Fl. Venez. Guayana 3: 279–281 (1997),
reg. rev.
Guaicaia Maguire (1967).
Shrubs or small trees. Leaves alternate, shortpetiolate, lamina elliptic to narrowly lanceolate,
entire. Capitula solitary, axillary, homogamous,
discoid or ligulate; involucre turbinate to narrowly
campanulate or cylindrical; phyllaries imbricate,
gradate, persistent; receptacle slightly convex,
epaleaceous, scarcely alveolate, long-pilose. Florets bilabiate or ligulate, bilabiate corollas with
3-toothed outer lip and inner lip of 2 long linear
lobes, lobes somewhat coiled, glabrous, ligulate
corollas usually with limb tightly rolled in apical
portion; corollas rose-coloured to orange-red;
basal anther appendages appearing truncate,
densely short-pilose, often appearing retrorsely
so; style with small glabrous basal node, shaft
glabrous, arms moderately long or short, spreading
to coiled or erect. Achenes glabrous, 10-ribbed,
cylindrical; carpopodium narrow, pale; pappus
setae multiseriate, capillary, barbellate, cream or
98
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
rust-coloured. Two species, northern Brazil and
southern Venezuela in the Guyana Shield.
15. Gongylolepis R.H. Schomb.
Gongylolepis R.H. Schomb., Linnaea 20: 759 (1847); Pruski,
Bol. Mus. Para. Emilio Goeldi, ser. Bot. 7: 352–367 (1991),
rev.; Fl. Venez. Guayana 3: 284–293 (1997), reg. rev.
apices truncate to obtuse. Achene 10-ribbed;
carpopodium annular; pappus setae 2–3-seriate,
barbellate, fragile, caducous, shorter than corolla
tube, usually bronze-coloured. One species, E.
bracteatus Maguire & Wurdack, Brazil, Venezuela
(Guyana Highlands).
Small to large shrubs or trees. Leaves simple,
alternate or densely spiralled, lamina usually large,
broadly lanceolate to obovate, entire. Capitula
solitary, sessile or pedicellate, or few to many
corymbose to subumbellate, homogamous, fewto many-flowered (6–150), large; involucre hemispherical or campanulate; phyllaries imbricate,
gradate, 4–6-seriate; receptacle convex, epaleaceous. Florets hermaphrodite; corollas bilabiate,
white or pale yellow, sometimes yellowish or
reddish, outer lip tightly rolled outwards, apex
with three short teeth, glabrous, inner of two long,
linear, tightly rolled lobes; basal anther appendages
long-caudate, adjacent pairs fused, usually smooth;
style base with large nectary, glabrous, shaft glabrous throughout, cream or purple, arms bifid,
short and scarcely bifid, or of moderate length
and usually recurved, glabrous, apices truncate
or obtuse, usually purplish. Achenes cylindrical,
usually 10-ribbed; carpopodium annular; pappus
setae biseriate to few-seriate, setae about as long
as corolla tube, finely barbellate, cream, pale
yellow or rust-coloured. Fourteen species, Brazil,
Colombia, Guyana (Guyana Highlands) Venezuela
(Venezuelan Guyana).
17. Achnopogon Maguire, Steyermark & Wurdack
16. Eurydochus Maguire & Wurdack
Neblinaea Maguire & Wurdack, Mem. New York Bot. Gard.
9: 391 (1957).
Achnopogon Maguire, Steyermark & Wurdack, Mem. New
York Bot. Gard. 9: 437 (1957); Pruski, Fl. Venez. Guayana 3:
197–199 (1997), reg. rev.
Shrubs or small trees. Leaves densely spiralled or
rosulate, lamina coriaceous, elliptic, oblanceolate
or obovate, entire. Capitula solitary, sessile, axillary or in few-headed cymes or in axillary cymes at
apices of long flowering branches, homogamous,
bilabiate; involucres narrow-cylindrical; phyllaries c. 4-seriate, imbricate, gradate; receptacle
small, epaleaceous, naked. Florets few (2–6),
hermaphrodite; corollas white, zygomorphic,
outer lip of three long lobes, inner of two long,
often tightly rolled lobes; basal anther appendages
caudate, retrorsely pilose; style base with distinct
glabrous basal node, shaft glabrous throughout,
arms long, eventually coiled, truncate, mammillose
outside at apices. Achenes cylindrical, obscurely
10-ribbed; carpopodium distinct, annular; pappus
setae 3–5-seriate, flattened at base, margins
finely barbellate throughout, straw-coloured. Two
species, Venezuela (Guyana Highlands).
18. Neblinaea Maguire & Wurdack
Eurydochus Maguire & Wurdack, Bol. Soc. Venez. Ci. Nat.
20: 57 (1958).
Trees or treelets. Leaves simple, alternate, usually in terminal clusters at stem apices, lamina
large, elliptic or oblanceolate. Capitula solitary,
subterminal on long sparsely bracteolate pedicels,
homogamous; involucre hemispherical to campanulate; phyllaries imbricate, few-seriate (c. 6–8),
subequal; receptacle broad, naked, convex. Florets numerous (40–50), hermaphrodite; corollas
bilabiate, red, outer lip three-toothed at apex,
usually partially to wholly tightly coiled, inner
of two tightly rolled linear lobes; basal anther
appendages long-caudate, free (Pruski 1997) or
connate (Maguire and Wurdack 1958), puberulous; style base with pronounced basal nectary,
shaft glabrous, arms relatively short, recurved,
Poorly branched shrubs or trees/treelets. Leaves
alternate to ± densely spiralled, pseudopetiolate,
oblanceolate, entire. Capitula solitary or few to several in cymes, homogamous, bilabiate; involucre
narrow-cylindrical; phyllaries c. 6-seriate, imbricate, gradate; receptacle small, epaleaceous. Florets
few (2–5), hermaphrodite, fertile; corollas white,
glabrous; outer lip short 3-toothed, inner of two
long rolled lobes; basal anther appendages caudate,
pilose; style base with glabrous node, arms short,
slightly recurved, obtuse to rounded. Achenes
cylindrical, obscurely ribbed; carpopodium large,
merging with and procurrent on to base of achene;
pappus setae c. 2–3-seriate, somewhat flattened at
base, coarsely barbellate, straw-coloured, vaguely
pinkish. One species, N. promontorium Maguire
Compositae
& Wurdack, Venezuela (Guyana Highlands) and
neighbouring Brazil.
19. Chimantea Maguire, Steyerm. & Wurdack
Chimantea Maguire, Steyerm. & Wurdack, Mem. New York
Bot. Gard. 9: 428 (1957); Maguire et al., Mem. New York Bot.
Gard. 9: 428–434 (1957), rev.; Pruski, Fl. Venez. Guayana 3:
239–245 (1997), reg. rev.
Small shrubs or rather low trees/treelets (to c. 9 m).
Leaves sessile or pseudopetiolate, spiralled, lamina linear, oblanceolate, broadly elliptic or obovate,
entire. Capitula solitary, sessile, terminal, discoid,
homogamous; involucre campanulate; phyllaries
multiseriate, persistent; receptacle flat or slightly
concave, epaleaceous or with a few outer paleae,
alveolate. Florets hermaphrodite, few to many (7–
35, rarely to 100); corollas actinomorphic, 5-lobed,
yellowish or yellowish-green, lobes stiff, very long,
erect; basal anther appendages caudate, entire or
scarcely ‘erose’/pilose; style base lacking basal node
but immersed in large lobed nectary, glabrous; style
glabrous throughout, arms short to medium, acute
or obtuse. Achene 10-ribbed; carpopodium absent;
pappus setae 3-seriate, flattened at base, barbellate,
straw-coloured. Nine species, Venezuela (Guyana
Highlands).
20. Stomatochaeta (S.F. Blake)
Maguire & Wurdack
Stomatochaeta (S.F. Blake) Maguire & Wurdack, Mem. New
York Bot. Gard. 9, 3: 388 (1957); Pruski, Brittonia 41: 35–40
(1989), rev.; Pruski, Fl. Venez. Guayana 3: 370–374 (1997),
reg. rev.
Stenopadus S.F. Blake subg. Stomatochaeta S.F. Blake
(1931).
Trees, treelets (or possibly shrubs). Leaves alternate
or sometimes pseudowhorled, simple, sessile or
pseudopetiolate, lamina oblanceolate, obovate, entire. Capitula solitary, terminal, often surrounded
by a pseudowhorl of leaves, homogamous, discoid;
involucres cylindrical to hemispherical; phyllaries few-seriate, imbricate, gradate; receptacle
sparsely paleaceous or epaleaceous, glabrous,
flat to slightly concave. Florets few to many,
hermaphrodite, all fertile; corollas actinomorphic,
cream-coloured, corolla lobes stiff, erect, long;
basal anther appendages caudate, long, apices
coarsely and irregularly short-papillose; style base
lacking basal node, glabrous, shaft glabrous, arms
relatively short, ascending, scarcely divided, acute
or possibly subobtuse. Achenes often 4-ribbed;
99
carpopodium absent; pappus setae 3-seriate,
unequal, persistent, flattened throughout, coarsely
barbellate, straw-coloured. Six species, Brazil,
Guyana, Venezuela.
21. Stenopadus S.F. Blake
Stenopadus S.F. Blake, Bull. Torrey Bot. Club 58: 489 (1931);
Maguire & Wurdack, Mem. New York Bot. Gard. 9: 366–392
(1957), reg. rev.; Pruski, Bol. Mus. Para. Emlio Goeldi, ser.
Bot. 7: 372–384 (1993), reg. rev.; Fl. Venez. Guayana 3:
364–370 (1997), reg. rev.
Trees or shrubs. Leaves alternate or loosely
spiralled, simple, oblanceolate or round, entire.
Capitula solitary, terminal or rarely in few-headed
cymes, homogamous, discoid; involucre campanulate, sometimes subtended by a pseudowhorl of
reduced leaf-like bracts; phyllaries multiseriate
(c. 8-seriate), imbricate, gradate; receptacle flat to
slightly concave or slightly convex, epaleaceous
or paleaceous with narrowly lanceolate paleae.
Florets few to many (5–100), actinomorphic,
hermaphrodite; corolla magenta, lobes straight,
partially coiled or strongly coiled; apical anther
appendages long-acute; basal anther appendages
caudate, short or long, entire, irregular or
antrorsely short-papillose, sometimes connate
with adjacent anthers; style base lacking basal
node, glabrous, shaft glabrous, arms relatively
short or of medium length, scarcely separated or
obviously bifid and coiled, short-papillose outside,
obtuse, usually with marginal lip. Achenes c.
10-ribbed (sometimes obscurely); carpopodium
absent; pappus setae 3-seriate, flattened throughout, barbellate, apices somewhat broadened,
straw-coloured. Fifteen species, Brazil, Colombia,
Ecuador, Venezuela (Guyana Highlands).
22. Quelchia N.E. Br.
Quelchia N.E. Br., Trans. Linn. Soc. London, Bot. 6: 41
(1901); Maguire & Steyermark, Mem. New York Bot. Gard.
9: 428–434 (1957), reg. rev.; Pruski, Bol. Mus. Para. Emílio
Goeldi, ser. Bot. 7: 370–372 (1993), reg. rev.; Fl. Venez.
Guayana 3: 353–355 (1997), reg. rev.
Poorly branched shrubs or small trees. Leaves
simple, alternate, sometimes densely clustered towards branch apices, lamina elliptic, oblanceolate
or obovate, entire. Capitula in dense terminal or
subterminal cymes of glomerules, single-flowered,
homogamous; involucres usually cylindrical,
sometimes slightly constricted at apex; phyllaries
c. 3–4-seriate, imbricate; receptacle small, epalea-
100
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
ceous, naked. Florets hermaphrodite, usually
actinomorphic; corollas red or white to cream,
lobes 5, long, spreading to slightly recurved; basal
anther appendages sagittate, acute or obtuse,
somewhat papillose; style base with basal node,
glabrous, shaft purplish, glabrous throughout,
arms short, recurved. Achenes c. 10-ribbed;
carpopodium a narrow annulus; pappus setae
multiseriate, usually united at base into prominent
callus, persistent, barbellate, straw-coloured or
sometimes reddish. Four species, endemic to
Guyana Shield in southern Venezuela.
or paleaceous, paleae enclosing accompanying
floret. Florets 3–70, rarely 2, heteromorphic, or
homomorphic and all with bilabiate corollas,
hermaphrodite and fertile; style arm apices
rounded, obtuse or truncate, short-papillate,
sometimes penicillate. Achenes fusiform or cylindrical, very rarely compressed, ribbed or terete,
very rarely rostrate, glabrous or setuliferous; carpopodium a narrow annulus or short-cylindrical,
or absent; pappus rarely absent or setae 2–4-seriate,
sometimes uniseriate, barbellate throughout or
apically subplumose, or entirely plumose.
23. Salcedoa F. Jiménez R. & Katinas
24. Proustia Lag.
Salcedoa F. Jiménez R. & Katinas, Syst. Bot. 29: 991 (2004).
Proustia Lag., Amen. Nat. Españ. 1, 1: 33 (1811); Fabris,
Revista Mus. La Plata n.s., Secc. Bot. 11: 23–49 (1968), rev.
Moderately branched treelet. Leaves alternate,
lamina simple, margins entire, obtuse. Inflorescence terminal, pseudocorymbose, many-headed
(20–30), capitula pedicellate, erect, discoid, homogamous, medium to large; involucre cylindrical;
phyllaries 4–5-seriate, imbricate, gradate; receptacle glabrous, alveolate, epaleaceous. Florets few
(4–5), hermaphrodite, fertile; corollas reddish or
cream-coloured, bilabiate or rarely actinomorphic
and 5-lobed, glabrous; anther cylinder exserted;
apical anther appendages oblong-lanceolate, apices
apiculate; basal appendages caudate, tails pilose;
style base glabrous; style shaft glabrous; style
arms short and short-bifid. Achenes cylindrical
to turbinate, setuliferous, setulae long twin-hairs,
apices unequal and undivided; carpopodium
annular(?); pappus setae biseriate, barbellate,
outer series capillary, inner flattened, reddish. One
species, S. mirabaliarum F. Jiménez R. & Katinas,
Dominican Republic.
II.3. Subtribe Nassauviinae
(Cass.) Dumort. (1829).
Annual or perennial herbs, subshrubs, shrubs,
sometimes clambering, or trees. Leaves alternate,
rarely in a basal rosette or densely clustered on
brachyblasts. Inflorescences short leafy axillary
cymes, dense, few- to many-headed terminal
corymbs, or appearing glomerulate with capitula
densely aggregated, or sometimes scapiform;
capitula homogamous and discoid or sometimes
appearing almost radiate; involucres turbinate,
cylindrical, campanulate or hemispherical, very
rarely acetabuliform or cochleariform; phyllaries (1–)3–5-seriate; receptacles epaleaceous
Scandent or erect shrubs or rarely small trees,
stems unarmed or spiny. Leaves simple, alternate,
lamina elliptic, ovate or oblong, entire, denticulate or dentate-spinose. Capitula racemose or
paniculate, erect or pendent, pedunculate or
sessile, homogamous, few-flowered; involucre
campanulate; phyllaries few-seriate, imbricate,
gradate; receptacle flat. Florets few, hermaphrodite,
sweet-smelling; corollas bilabiate, outer lip with
expanded 3-toothed limb, inner 2-dentate, pink
or purple; basal anther appendages caudate; style
with basal node, glabrous, arms truncate, papillose. Achenes fusiform, 4-ribbed; carpopodium
annular; pappus biseriate, barbellate, apically
subplumose, straw-coloured, yellow, pink, or
purplish. n = 26, 27. Three species, Peru, Bolivia,
Chile, Argentina.
25. Berylsimpsonia B.L. Turner
Berylsimpsonia B.L. Turner, Phytologia 74: 351 (1993);
Turner, Phytologia 74: 349–355 (1993), key.
Clambering woody shrubs, 1–5 m tall, stems with
recurved spines at each node. Leaves alternate,
simple, very short-petiolate, lamina entire to serrulate, scarcely spinulose. Capitula in short leafy
axillary cymes, sessile or very short-pedicellate; involucres turbinate; phyllaries 3–4-seriate, gradate;
receptacle pubescent. Florets 3–6 per capitulum;
corollas yellow, glabrous, bilabiate, outer lip short
three-toothed, inner deeply 2-lobed, lobes strongly
coiled; basal anther appendages long-tailed, bases
irregularly ‘bearded’; style base slightly expanded
but lacking basal node, shaft glabrous, arms
glabrous, rounded or obtuse, short-papillate.
Compositae
Achenes fusiform to narrowly oblanceolate,
ribbed; carpopodium cylindrical, procurrent on
base of ribs; pappus 2–3-seriate, setae numerous,
barbellate, tawny. Two species, Cuba, Dominican
Republic, Haiti, Puerto Rico, Santo Domingo.
26. Lophopappus Rusby
Lophopappus Rusby, Bull. Torrey Bot. Club 21: 487 (1894);
Cabrera, Bol. Soc. Argent. Bot. 5: 37–50 (1953), reg. rev.;
Faúndez & Macaya, Not. Mens. Mus. Nac. Hist. Nat. 332:
3–6 (2000), reg. rev.
Well-branched shrubs. Stems often viscous. Leaves
alternate or in dense axillary clusters, simple, often
viscous, lamina often viscous and shiny when
covered in exudate, entire or dentate, sometimes
minutely so. Capitula solitary or in tight clusters
at apices of branches, sessile or short-pedicellate,
homogamous; involucres cylindrical to narrowly
campanulate; phyllaries few-(3-)seriate, with apical spine; receptacles small, flat to slightly convex,
papillate. Florets usually few, yellowish white to
white, usually sweet-smelling, hermaphrodite,
fertile; corollas distinctly two-lipped, either outer
lip short to moderately 3-toothed or deeply
3-lobed, inner of two long, usually coiled lobes or
2-toothed with teeth moderately long, or corolla
actinomorphic and deeply 5-lobed; basal anther
appendages caudate, entire; style base sometimes
with distinct node, glabrous, shaft gradually
thickening upwards and then contracting beneath
branching point of arms or uniformly cylindrical,
arms moderately long, divergent, often coiled at
maturity, acute, mammillose outside. Achenes
5-ribbed, usually narrowed to almost attenuate at
base; carpopodium cylindrical; pappus uniseriate,
setae barbellate at base, becoming subplumose
towards apices, usually straw-coloured. Six species,
Argentina, Bolivia, Chile, Peru.
Lophopappus is rather similar to Proustia but
differs in its solitary or few, grouped capitula, the
lack of terminal spines on short branches, to some
degree corolla colour (white in Lophopappus, pink
or purple in Proustia). The setulae on the achenes of Proustia, when present, are long, twisted or
spiralled twin-hairs. Preliminary molecular work
(Funk et al., pers. comm.) supports their separation, although they are clearly closely related. The
recent treatment of genera for the Flora of Peru
(Ferreyra 1995) has suggested that Lophopappus
be treated as congeneric with Proustia. They are
treated as separate genera here.
27. Nassauvia Comm. ex Juss.
101
Fig. 18
Nassauvia Comm. ex Juss., Gen. Pl.: 175 (1789); Cabrera,
Darwiniana 24: 283–379 (1982), rev.
Calopappus Meyen (1834).
Perennial herbs, subshrubs or shrubs, often
compact and caespitose. Leaves alternate, sessile,
usually densely crowded, lamina ovate, lanceolate,
rarely spathulate, entire, dentate, denticulatespiny, or with few long spines. Capitula generally
in complex, terminal, often dense, sometimes
globular synflorescences, rarely solitary or in
few-headed dichasia, rarely with solitary shortpedicellate axillary capitula forming a terminal
‘spike’, capitula sessile or subsessile, homoga-
Fig. 18. Compositae-Mutisieae. Nassauvia dentata. A Flowering shoot. B Capitulum. C Floret. (Drawings by Margaret
Tebbs)
102
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
mous; involucre cylindrical; phyllaries biseriate;
receptacle epaleaceous, glabrous. Florets few,
(2–4–)5, hermaphrodite; corollas white, rarely
violet-pinkish or yellowish, bilabiate, outer lip
with a conspicuous 3-toothed limb, inner bilobed,
lobes recurved; basal anther appendages caudate,
usually acute with scarcely finely laciniate margins;
anther collar thickened and markedly constricted
at base; style base with distinct node, glabrous,
shaft glabrous, arms linear, truncate, penicillate.
Achenes turbinate or obovate, ribbed or sometimes
conspicuously compressed with two lateral ribs;
carpopodium usually an inconspicuous, narrow
white annulus; pappus uniseriate, of few (5)
caducous linear scales or several setae with broad
flattened rachis and margins ciliate to plumose,
white to straw-coloured. n = 11, 22, c. 44. Circa 40
species, Argentina, Bolivia, Chile, Falkland Islands.
The genus has been divided into two subgenera
and the type subgenus into four sections by Cabrera
(1982).
28. Triptilion Ruiz & Pav.
Triptilion Ruiz & Pav., Fl. Peruv. Prodr.: 102, t. 22 (1794).
Annual or perennial herbs or subshrubs. Leaves
alternate, in basal rosettes in some herbaceous
plants, lamina entire or pinnatisect with denticulate spiny margins. Capitula densely aggregated
into paniculate glomerules, sessile to shortpedicellate, few-flowered, discoid, homogamous;
involucre biseriate, ovoid to cylindrical; phyllaries
with pungent apices; receptacle convex, longciliate or rarely naked. Florets hermaphrodite;
corollas glabrous, white or blue, bilabiate, outer
lip 3-toothed, inner 2-toothed, strongly coiled;
basal anther appendages sagittate, entire; style
shaft glabrous, apically bifid, arms truncate,
penicillate. Achenes cylindrical, attenuate towards
base; carpopodium absent; pappus of few (3–5)
apically lacerate-plumose caducous scales. Circa
12 species, Argentina and Chile.
29. Oxyphyllum Phil.
Oxyphyllum Phil., Fl. Atacam.: 28, tab. 4 (1860).
Erect shrub, each leaf axil with a dense cluster
of simple, linear, spine-tipped immature leaves.
Stem leaves pinnatifid, rarely entire, segments and
apices spiny. Capitula in a terminal, dense, few- to
many-headed corymb, homogamous, discoid; involucre cylindrical to narrowly campanulate; phyllaries 3-seriate, imbricate, gradate, with distinct
apical spine; receptacle glabrous, small, scarcely
convex in centre portion, paleaceous, paleae few
(3), broad. Florets few, dimorphic, outer florets apparently sterile and each subtended by an internal
palea, inner florets fertile; corollas pinkish-white;
outer florets bilabiate, outer lip a large limb, short
3-toothed at apex, inner of one coiled lobe; basal
anther appendages very long-caudate, entire; anther collar distinctly broadened; style base swollen,
glabrous, shaft glabrous, arms divergent, truncate,
usually with a slight corona of short papillae; inner florets similar to outer but inner lip of two
long, tightly coiled lobes. Achenes of outer florets apparently abortive, those of inner florets dark
brown; carpopodium absent; pappus uniseriate, setae plumose, white. One species, O. ulicinum Phil.,
Chile (Atacama Desert).
30. Macrachaenium Hook. f.
Macrachaenium Hook. f., Fl. Antarctica 2: 321 (1847).
Perennial herb. Leaves mostly in loose basal rosette,
alternate, lamina oblong, ovate or broadly ovate,
coarsely and deeply, often irregularly, lobed, sometimes pinnatifid to almost runcinate-pinnatifid,
often irregularly subdentate. Capitula solitary
on scapiform peduncles, homogamous, usually
discoid, sometimes conspicuously radiate, apparently nodding; involucre campanulate; phyllaries
biseriate, outer usually much shorter than inner;
receptacle flat to slightly convex, epaleaceous,
glabrous. Florets bilabiate, all hermaphrodite and
fertile; corollas white. Ray florets (when present)
two-lipped, outer lip conspicuous, short 3-toothed,
inner of two long coiled lobes; basal anther appendages absent, base of thecae rounded; style
base lacking basal node, glabrous, shaft glabrous,
arms short to moderate, acute with marginal
thickening, mammillose, glabrous outside. Disc
florets two-lipped, outer not conspicuous, short
3-toothed, inner of two long coiled lobes; basal
anther appendages caudate, entire; style base and
shaft as in ray florets, arms short, acute, mammillose. Achenes of ray and disc florets identical,
cylindrical; carpopodium evident or not; pappus
usually biseriate, setae plumose, off-white, strawcoloured or sometimes pale rust-coloured. One
species, M. gracile Hook. f., Argentina and Chile.
31. Polyachyrus Lag.
Polyachyrus Lag., Amen. Nat. Españ. 1, 1: 37 (1811); Ricardi
& Weldt, Gayana, Bot. 26: 1–41 (1974), rev.
Compositae
Decumbent, scandent or prostrate subshrubs or
shrubs, rarely herbs. Leaves alternate, lamina
pinnate-lobed or pinnatisect, entire or coarsely
dentate. Capitula in solitary apical glomerules or
glomerules in pseudocorymbs, each subtended
by one bract; capitula numerous, sessile, 2or rarely 3-flowered; involucres cochleariform;
phyllaries 5, outer enclosing outer floret, inner two
including inner floret; receptacle naked. Florets
hermaphrodite, fertile, sweet-smelling; corollas
bilabiate, outer lip expanded to a 3-toothed limb,
inner deeply 2-lobed, revolute/coiled, white or
pink; basal anther appendages caudate, margins
laciniate (glabrous?); style with distinct basal
nectary (possibly node in some species), glabrous,
arms divergent, truncate or obtuse, dorsally
papillose, appearing penicillate. Achenes terete;
carpopodium indiscernible or absent; pappus
uniseriate, often detached as unit, plumose, white.
n = 21. Seven species, Peru, Chile.
32. Perezia Lag.
Perezia Lag., Amen. Nat. Españ. 1, 1: 31 (1811); Vuilleumier,
Contr. Gray Herb. 199: 1–163 (1969), sect. rev.
Perennial, usually strongly rosettiform, sometimes
caespitose herbs, rarely tall leafy-stemmed herbs.
Leaves simple, radical or alternate, entire or lyrate,
lamina linear, narrowly lanceolate, spathulate
to broadly ovate, often ciliate to lacerate, entire,
coarsely serrate or dentate to biserrate, pinnatifid
or deeply lobed, often spinous. Capitula on
1–(2)-headed scapes arising from basal rosette,
or in few- to many-headed dense or lax and
spreading panicles; capitula appearing radiate but
being discoid, homogamous, usually erect, rarely
nodding; involucre broadly cylindrical, turbinate
or hemispherical; phyllaries few-seriate to multiseriate, gradate, imbricate, often with terminal
spine; receptacle convex, epaleaceous, usually
pubescent. Florets usually several to many (8–40);
corollas bilabiate, yellow, blue, purple, violet, red,
or crimson, rarely white or cream, occasionally
with outer lip of one colour and inner of another
(usually yellow); outer lip a 3-toothed limb, inner
usually tightly rolled, linear and short 2-toothed at
apex; basal anther appendages very long-sagittate,
often poorly laciniate; style base with distinct
glabrous basal node, shaft glabrous throughout,
arms relatively short, truncate, purplish or white.
Achenes cylindrical to fusiform; carpopodium
short-cylindrical or scarcely evident; pappus setae
numerous, persistent, slightly longer than corolla
103
tube, 2- or few-seriate, barbellate, brownish. n = 4,
8, 12. Circa 32 species, South America (Argentina,
Bolivia, Brazil, Chile, Paraguay, Peru, Uruguay).
33. Acourtia D. Don
Acourtia D. Don, Trans. Linn. Soc. London 16: 203 (1830);
Bacigalupi, Contr. Gray Herb. 97: 1–81 (1931), rev. (sub
Perezia); Turner, Phytologia 38: 456–468 (1978), part. rev.
Scandent shrubs or subshrubs. Leaves few in a basal
rosette, or many, cauline and alternate, simple,
lamina small to large, narrowly ovate, oblanceolate,
oblong, spathulate, cordiform, lyrate-pinnatifid or
broadly elliptic to broadly ovate, serrate, spinous,
coarsely dentate or rarely entire. Capitula 1–(2–3),
arising on scape from basal rosette, few to many
in dense terminal or axillary clusters, or many
in a dense thyrse or in broad thyrsoid panicle
well exceeding upper stem leaves, homogamous,
few- to many-flowered (4–60), rarely appearing
radiate when outer florets bilabiate and inner
florets actinomorphic; involucre turbinate to
campanulate; phyllaries imbricate, gradate, 3–8seriate; receptacle usually glabrous, sometimes
appearing fimbriate or sparingly short-pubescent,
epaleaceous, flat to slightly convex. Florets
hermaphrodite; corolla cream, white, pinkish,
light purple or purple, usually bilabiate, outer lip 3toothed, inner of two linear lobes, rarely few inner
florets with actinomorphic corollas (with 5 linear
lobes); basal anther appendages long-caudate,
entire or varyingly sparingly laciniate, long-acute;
style base somewhat expanded into node, glabrous,
shaft purplish, glabrous, arms purplish, recurved
(sometimes strongly so), truncate, papillate, occasionally appearing coronate. Achenes cylindrical
to fusiform, usually with narrowed apex and
apical callus, obscurely ribbed or with distinct
paler ribs/lines; carpopodium annular; pappus
setae 1–2-seriate, numerous, persistent, barbellate,
fawn, whitish or sometimes dark greyish brown.
n = 27, 28. Circa 80 species, USA, Mexico.
34. Burkartia Crisci
Burkartia Crisci, Bol. Soc. Argent. Bot. 17, 3/4: 242 (1976).
Dwarf caespitose shrub forming hemispherical
cushions. Leaves sessile, densely spiralled, acicular,
margins strongly revolute. Capitula solitary, terminal, usually appearing sessile although shortly
pedunculate, few-flowered, radiate, homogamous;
involucre turbinate to campanulate; phyllaries
2–3-seriate, imbricate; receptacle slightly convex,
104
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
epaleaceous, densely short-pubescent. Florets
hermaphrodite, fertile; corollas white, bilabiate,
outer lip glabrous, short 3-toothed, inner long
2-lobed (scarcely short 2-toothed in some florets),
lobes coiled; basal anther appendages caudate,
pilose; anther collar slightly thickened; style base
with basal node, shaft glabrous, arms markedly
divergent, truncate, short-papillate outside. Achenes cylindrical, obscurely ribbed; carpopodium
short-cylindrical or conical, straw-coloured; pappus usually 3-seriate, setae barbellate, off-white.
One species, B. lanigera (Hook. & Arn.) Crisci,
Argentina (Patagonia).
35. Leucheria Lag.
Leucheria Lag., Amen. Nat. Españ. 1, 1: 32 (1811); Crisci,
Darwiniana 20: 9–126 (1976), rev.
Annual or perennial herbs. Leaves alternate, rosulate, lamina linear, narrowly lanceolate, spathulate,
oblong or ovate, entire, dentate, coarsely lobed,
pinnatisect or pinnatifid. Capitula solitary or
in few- to many-headed corymbs or panicles,
pedicellate, appearing radiate; involucre campanulate; phyllaries usually biseriate; receptacle
convex, glabrous, epaleaceous. Florets several
to many, homomorphic, hermaphrodite, fertile,
outer lip white, pink, lilac or wine-coloured;
corollas 2-lipped, outer short 3-toothed, often
slightly more prominent, inner usually very shortly
2-toothed, often rolled; basal anther appendages
caudate, entire or somewhat erose; anther collar
cylindrical, usually markedly narrowed beneath
and then expanding to flattened filament; style
base with basal node, glabrous, shaft glabrous,
arms moderately long, truncate, short-papillate
outside. Achene terete; carpopodium annular,
straw-coloured; pappus setae usually uniseriate,
setae flattened and united at base, barbellate or
rarely plumose, white or whitish. n = 20. Forty-six
species, Argentina, Bolivia, Chile, Peru.
36. Jungia L. f.
florets; involucre narrowly cylindrical, turbinate,
campanulate or hemispherical; phyllaries uniseriate, subequal, each enclosing a marginal floret. Florets few to many, hermaphrodite, all fertile; corollas white, violet pink, purple or rarely yellow, bilabiate, outer lip enlarged into 3-toothed limb, inner usually deeply bifid, lobes revolute to coiled;
basal anther appendages caudate, entire; style base
with basal node, glabrous, shaft glabrous, arms
markedly bifid and recurved through upper part of
anther cylinder, truncate, penicillate. Achenes slender, fusiform to turbinate, 5-ribbed; carpopodium
a broad annulus with somewhat lobed upper margin; pappus uniseriate or biseriate, setae barbellate,
subplumose or plumose, usually white or strawcoloured, rarely grey or bright orange red, sometimes variable. n = 18, 21. Circa 32 species, Central
and South America.
Harling (1995) recognized four sections in the
genus.
Fig. 19
Jungia L. f., Suppl. Pl.: 58 (1782), nom. cons.; Harling, Acta
Regiae Soc. Sci. Litt. Gothob. Bot. 4: 1–133 (1995), rev.
Tostimontia Diaz Piedrahita (2001).
Perennial herbs, rarely rosulate, subshrubs, shrubs
or lianes. Leaves alternate, lamina cordate, usually
lobed, entire, serrate, dentate or crenate. Capitula
in usually terminal corymbs or panicles, sometimes with dense glomerules, homogamous, appearing radiate by enlarged outer lip of marginal
Fig. 19.
Compositae-Mutisieae. A–C Jungia woodii.
A Mature leaf. B Flowering shoot. C floret. D–F Eriachaenium magellanicum. D Plant with stolons. E Floret showing
pubescent achene. F Floret with naked achene. (Drawings
by Margaret Tebbs)
Compositae
37. Pleocarphus D. Don
Pleocarphus D. Don, Trans. Linn. Soc. London 16: 228
(1830).
Shrub. Leaves sessile, linear, entire, margins conspicuously revolute. Capitula many to numerous
in an elongated panicle, pedicellate, homogamous;
involucre turbinate to campanulate; phyllaries
2–(3)-seriate, imbricate; receptacle scarcely convex, pubescent, paleaceous, paleae few, usually
surrounding central florets. Florets homomorphic,
hermaphrodite, fertile; corollas yellow, essentially
two-lipped, outer lip of a 2- or 3-short-toothed
limb, inner of 3 or 2 long coiled lobes; basal anther
appendages extremely long-caudate, entire or
sparsely short-pilose/papillose; style base with
basal node, glabrous, shaft glabrous, arms long,
apically truncate with a corona of short papillae.
Achenes long, apically and basally attenuate,
5-ribbed; carpopodium cylindrical, upper margins
procurrent with ribs; pappus setae biseriate, setae
numerous, barbellate, straw-coloured. n = 26. One
species, P. revolutus D. Don, Chile.
38. Trixis P. Browne
105
Argentina, Bolivia, Brazil, Chile, Colombia, Costa
Rica, Cuba, Ecuador, El Salvador, Guatemala,
Honduras, Mexico, Nicaragua, Panama, Paraguay,
Peru, Uruguay, USA, Venezuela, West Indies.
39. Doliclasium Lag.
Doliclasium Lag., Amen. Nat. Españ. 1, 1: 33 (1811).
Small, moderately branched low shrub. Leaves
alternate, petiolate, lamina pinnatisect, pinnae
opposite, ovate. Capitula large, terminal, solitary, homogamous, erect; involucre narrowly
campanulate; phyllaries biseriate to 3–4-seriate
with outer two series often well separated from
inner subequal series; receptacle epaleaceous,
glabrous. Florets hermaphrodite, numerous,
homogamous, fertile; corollas yellow, bilabiate,
with outer distinct 3-toothed limb and inner lip
of two long rolled lobes; basal anther appendages
Fig. 20
Trixis P. Browne, Civ. Nat. Hist. Jamaica: 312 (1756);
Anderson, Mem. New York Bot. Gard. 22: 1–68 (1972), reg.
rev.; Katinas, Darwiniana 34: 27–108 (1996), reg. rev.
Perennial herbs, subshrubs, shrubs, scandent/
trailing shrubs or small trees. Leaves alternate,
lamina simple, narrowly lanceolate, elliptical,
oblanceolate or obovate to oblong, entire or often
denticulate. Capitula in usually terminal, sometimes axillary, lax cymes, corymbs or panicles,
occasionally aggregated into pseudoglomerules,
homogamous; involucre cylindrical, campanulate
or hemispherical, sometimes an outer calyculus
present; phyllaries imbricate, pubescent, (1–)2–
(3–5)-seriate; receptacle flat, alveolate, pilose
or densely long-pubescent. Florets few to many
(5–c. 70), hermaphrodite, all fertile; corollas
yellow to orange, rarely white, bilabiate, outer
lip 3-toothed, inner deeply bifid; basal anther
appendages caudate, glabrous or papillose; style
base with distinct basal node, glabrous, shaft
glabrous, arms truncate, penicillate. Achenes
5-ribbed, cylindrical to turbinate, ± beaked and
usually with distinct expanded apical callus above
beak; carpopodium annular to short-cylindrical;
pappus setae 2–3(–4)-seriate, markedly longer
than phyllaries, barbellate, persistent, white,
yellowish or reddish. n = 27. Circa 50–60 species,
Fig. 20. Compositae-Mutisieae. Trixis vauthieri. A Flowering shoot. B Floret. (Drawings by Margaret Tebbs)
106
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
long-caudate, usually acute, margins laciniate to
short-pilose; style base with basal node, glabrous,
shaft glabrous, arms long, dilated at very apices,
obtuse to truncate, papillate outside and appearing
coronate. Achenes fusiform, slender, rostrate; pappus biseriate to multiseriate, setae with thickened
and flattened rachis at base, subplumose to very
coarsely barbellate, more densely so towards base,
off-white to white. One species, D. lagascae D. Don,
Argentina (Patagonia).
40. Ameghinoa Speg.
Ameghinoa Speg., Revista Fac. Agron. La Plata 3: 539
(1897).
Low, densely branched shrub. Leaves often in dense
clusters on brachyblasts, simple, coarsely dentate.
Capitula few in terminal clusters, short-pedicellate,
homogamous; involucre narrow-campanulate;
phyllaries 2-seriate, gradate, imbricate; receptacle
flat, epaleaceous, alveolate, glabrous. Florets
hermaphrodite, isomorphic, 20–30; corollas
yellow, bilabiate, outer lip a scarcely enlarged
limb with 3 short apical teeth, inner of 2 long
linear lobes, often rolled; basal anther appendages
long-caudate, sparsely pilose/laciniate; style base
somewhat swollen, glabrous, shaft glabrous,
arms short, slightly bifid, truncate and papillate
around edge outside, appearing coronate. Achenes
cylindrical; carpopodium dark, procurrent with
base of achene; pappus setae uniseriate, somewhat
fragile and easily detached, coarsely barbellate,
off-white. n = 26. One species, A. patagonica Speg.,
endemic to Argentina (Patagonia).
41. Criscia Katinas
Criscia Katinas, Bol. Soc. Argent. Bot. 30: 60 (1994).
Perennial herb or subshrub. Leaves rosulate, lamina obovate to broadly obovate, entire. Capitula
solitary or 2–4 on scapes, large, bilabiate, homogamous; involucre hemispherical or broadly campanulate; phyllaries c. 4-seriate, imbricate, gradate; receptacle epaleaceous, glabrous. Florets all bilabiate,
hermaphrodite, fertile; corollas orange, marginal
florets with more pronounced, shortly 3-toothed
outer lip, inner divided into 2 often straight (at least
when young) or somewhat rolled long linear lobes;
inner florets similar to marginal but with shorter
outer lip; basal anther appendages caudate, entire
or somewhat contorted/irregular; style base with
distinct basal node, glabrous, shaft glabrous, arms
moderately long, recurved to coiled, rounded to
truncate, papillate and appearing almost ‘crowned’
by short papillae. Achene obscurely constricted
towards apex almost into a short rostrum; carpopodium short-cylindrical; pappus setae c. 3–4seriate, setae flattened towards base, coarsely barbellate, pinkish- to light rusty-brown. One species,
C. stricta (Spreng.) Katinas, Argentina, Brazil and
Uruguay.
42. Leunisia Phil.
Leunisia Phil., Linnaea 33: 120 (1864).
Viscid low ‘subshrub’ or perennial herb. Leaves
alternate to loosely spiralled, irregularly toothed
or sometimes entire. Capitula terminal, solitary,
homogamous, discoid; involucre turbinate; phyllaries biseriate, scarcely imbricate, subequal; receptacle epaleaceous, convex, densely pubescent.
Florets numerous, hermaphrodite, fertile; corollas yellow, bilabiate, outer lip short 3-toothed, inner long 2-lobed; basal anther appendages caudate, pilose; style base with node, glabrous, shaft
glabrous, arms short, connate, short-papillate outside. Achenes cylindrical; carpopodium concolorous and procurrent with base of achene; pappus
setae biseriate, broad, united at base, barbellate, often branched, off-white. One species, L. laeta Phil.,
Chile.
43. Marticorenia Crisci
Marticorenia Crisci, J. Arnold Arb. 55: 38 (1974).
Shrub with short woody caudex. Leaves alternate,
lamina ovate, lobulate, becoming lanceolate
above. Capitula in a lax many-headed corymb,
homogamous; involucre hemispherical; phyllaries biseriate; receptacle ± concave, slightly
pubescent, paleaceous, paleae conduplicate about
florets, apices laciniate. Florets hermaphrodite;
corollas violet-pink, bilabiate, outer lip 3-toothed,
inner bifid, of two revolute lobes; basal anther
appendages tailed; style shaft glabrous, arms
truncate, penicillate. Achenes cylindrical; pappus
uniseriate, of numerous white plumose setae.
n = 22. One species, M. foliosa (Phil.) Crisci, Chile.
44. Holocheilus Cass.
Holocheilus Cass., Bull. Sci. Soc. Philom. 1818: 73 (1818);
Cabrera, Revista Mus. La Plata, Secc. Bot. 11: 1–15 (1968),
rev.
Perennial rosulate herbs. Leaves usually few,
loosely rosulate, lamina medium and coarsely
Compositae
dentate, crenate or entire, or large and pinnatisect
or coarsely lobed. Capitula in large, few- to
many-headed terminal corymbs or cymes, rarely
scapose, homogamous, small to medium, erect;
involucre hemispherical; phyllaries uniseriate to
biseriate; receptacle epaleaceous, glabrous. Florets
hermaphrodite, bilabiate; corollas white, outer lip
an enlarged limb with three short apical teeth, inner of 2 medium to short, usually coiled lobes; basal
anther appendages long-caudate, entire, anther
collar prominent, distinctly narrowed compared
with rest of filament; style base enlarged (probably
into node, rather than nectary), glabrous, shaft
glabrous, arms short, truncate, papillae forming
corona. Achenes fusiform; carpopodium a distinct
annulus, of same colour as achene; pappus setae
biseriate, persistent, barbellate, spreading to ascending, white or straw-coloured. n = 11, 18. Seven
species, Argentina, Brazil, Paraguay, Uruguay.
45. Pamphalea Lag.
Pamphalea Lag. (orig. Panphalea), Amen. Nat. Españ. 1, 1:
34 (1811); Cabrera, Notas Mus. Eva Perón, Bot. 16: 225–237
(1953), rev.
107
pinnatisect. Capitula in lax, terminal corymbs, discoid, homogamous; involucre biseriate; phyllaries
dimorphic, outer foliaceous, inner convolute and
enclosing marginal florets; receptacle convex, glabrous, paleaceous, paleae narrowly lanceolate. Florets few, hermaphrodite, bilabiate; corollas pink or
violet, outer florets hermaphrodite with 3-toothed
outer limb, inner lip bifid, inner florets sterile; basal
anther appendages long-caudate, entire; style base
with distinct basal node/nectary, glabrous, shaft
glabrous, arms bifid, truncate, penicillate. Achenes
fusiform; carpopodium short-cylindrical; pappus
setae uniseriate, short, flattened, persistent, with
long-ciliate/subplumose margins, brownish, sometimes absent on central florets. n = 20. Two species,
Chile.
47. Cephalopappus Nees & Mart.
Fig. 21
Cephalopappus Nees & Mart., Nova Act. Phys.-Med. Acad.
Caes. Leop.-Carol. Nat. Cur. 12, 1: 5, tab. 1 (1824).
Perennial rosulate stoloniferous herbs. Leaves with
ovate to narrowly obovate lamina, coarsely dentate.
Capitula solitary, scapose, rarely scape branched
with two (or few) capitula, homogamous; involucre
Slender annual or perennial rhizomatous herbs.
Leaves mostly basally rosulate, alternate, entire
and linear-lanceolate to orbicular, coarsely lobed
or lyrate-pinnatifid. Capitula in lax, relatively
few-headed corymbs, small, appearing radiate
(although all florets identical), homogamous; involucre campanulate to hemispherical; phyllaries
few, 1–2-seriate, imbricate, subequal (uniseriate)
or gradate (biseriate); receptacle flat, epaleaceous, glabrous, sometimes fimbriate. Florets
hermaphrodite, many; corollas white, bilabiate,
outer lip with an expanded 3-toothed limb, inner
of two fairly wide linear rolled lobes; basal anther
appendages sagittate, entire, anther collar slightly
narrower than rest of filament and constricted at
base; style base with distinct basal node, glabrous,
shaft glabrous, arms truncate, penicillate. Achenes
often included within involucre, inflated, usually
with apical corona/callus; carpopodium a narrow
annulus; epappose. n = 8. Nine species, Argentina,
Brazil, Paraguay, Uruguay.
46. Moscharia Ruiz & Pav.
Moscharia Ruiz & Pav., Fl. Peruv. Prodr.: 91 (1794), nom.
cons.; Crisci, Contr. Gray Herb. 205: 163–173 (1974), rev.
Annual odiferous herbs. Leaves alternate, simple,
lamina elliptic, entire, coarsely dentate or lobed to
Fig. 21. Compositae-Mutisieae. Cephalopappus sonchifolius. A Flowering plant. B Floret. C Mature achene.
(Drawings by Margaret Tebbs)
108
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
broadly saucer-shaped; phyllaries 2–3-seriate; receptacle epaleaceous, glabrous, scarcely alveolate.
Florets many, hermaphrodite, all fertile; corollas
white, few opening at a time and lost very rapidly,
leaving green achene, bilabiate when open, lobes
spreading; basal appendages sagittate, entire (or
sometimes subentire); anther collars markedly
enlarged; style base lacking basal node, glabrous,
shaft glabrous, arms obtuse and rounded, apically
densely to moderately short-pilose. Achenes
slender, glabrous; carpopodium apparently absent;
pappus absent. One species, C. sonchifolius Nees &
Mart., endemic to Brazil.
Genera of Problematic Placement
(Probably Within Nassauviinae)
48. Adenocaulon Hook.
Adenocaulon Hook., Bot. Misc. 1: 19 (1830); Bittman,
Candollea 45: 389–420, 493–518 (1990), rev.
Perennial herbs. Leaves alternate or forming
basal rosette, lamina ovate, broadly triangular or
lyrate-pinnatifid, entire to coarsely lobed. Capitula
many in lax panicles, peduncles and pedicels with
persistent and moderately long stipitate-glandular
hairs; capitula small, few-flowered, disciform,
heterogamous, hemispherical, erect; involucre
distant; phyllaries uniseriate, in many species
a cupule with phyllaries ‘connate’ in bottom
half; receptacle convex to almost conical in some
species, epaleaceous, glabrous. Marginal florets
uniseriate, female, fertile; corollas actinomorphic
or slightly zygomorphic (bilabiate), 4- or 5-lobed,
whitish or yellowish-white, staminodes present,
style bifid; central florets few, hermaphrodite,
functionally male, usually lacking achene, corollas
actinomorphic, 5-lobed, style undivided; basal
anther appendages caudate but without distinct
tails; style arms scarcely divided or connate, papillate outside. Achenes of female florets obovoid,
with conspicuous stipitate glands densest in
upper half, achenes of male florets, when present,
usually glabrous; carpopodium on base of basal
constriction of achene, annular; pappus absent.
n = 23. Five species, Argentina, Chile, eastern
Asia, Nepal, western Canada and USA, Guatemala.
49. Eriachaenium Sch. Bip.
Fig. 19
Eriachaenium Sch. Bip., Flora 38: 120 (1855).
Perennial rhizomatous herbs. Leaves alternate,
lamina obovate to elliptic, undulate, sometimes
serrate. Capitula solitary, axillary, small, sessile or
on short-bracteolate side shoots; involucre campanulate; phyllaries 1–(2)-seriate, few, subequal,
not imbricate; receptacle very small, epaleate,
glabrous. Marginal florets female, few; corollas
white (pinkish in some herbarium material),
actinomorphic, usually 5-lobed; anthers absent;
style base lacking basal node, glabrous, shaft
glabrous, arms short, short-papillose outside.
Achenes inflated, conspicuously densely lanose;
carpopodium not evident; pappus absent. Central
florets very few, hermaphrodite, functionally male;
corollas white, actinomorphic, 5-lobed; basal
anther appendages scarcely caudate, short-pilose;
style base lacking basal node, glabrous, shaft
glabrous, arms short, markedly divergent, usually
with lip around margins, short-papillose outside.
Achenes sterile, pappus absent. n = 23. One
species, E. magellanicum Sch. Bip., Argentina and
Chile (Patagonia and Tierra del Fuego).
II.4. Subtribe Mutisiinae
(Cass.) Dumort. (1829).
Annual or perennial herbs, subshrubs, shrubs
or trees, sometimes climbers. Leaves alternate,
sometimes rosulate, simple or rarely pinnate or
imparipinnate with a simple or branched tendril.
Inflorescences solitary terminal or leaf-opposite
capitula, or few- to many-headed corymbs or
panicles; capitula radiate and homogamous or
heterogamous, discoid and homogamous or very
rarely disciform; involucres cylindrical, turbinate,
campanulate or hemispherical; phyllaries 2–8seriate; receptacle epaleaceous, alveolate; ray
florets, when present, female, rarely neuter, uniseriate, corollas bilabiate; disc florets hermaphrodite,
actinomorphic and corollas deeply 5-lobed, or
bilabiate; style arms short, glabrous. Achenes
cylindrical or fusiform, very rarely obcompressed,
glabrous or setuliferous; carpopodium absent,
annular or stopper-shaped; pappus setae uniseriate
to multiseriate, persistent, barbellate, subplumose
or plumose.
50. Mutisia L. f.
Fig. 22
Mutisia L. f., Suppl. Pl.: 57 (1781); Cabrera, Opera Lilloana
13: 1–227 (1965), rev.
Plants perennial subshrubs or shrubs, often
climbers. Leaves simple, subulate, narrowly lanceolate or ovate, entire or dentate, apices with or
without terminal tendril, rarely deeply pinnatisect
Compositae
or coarsely lobed or deeply partite, or pinnately
compound with few to several pairs of leaflets and
rachis always with a terminal simple, 3- or 5-fid tendril. Capitula small to large, solitary, leaf-opposed
or terminal, erect or pendulous, homomorphic
with all florets hermaphrodite or heteromorphic
with marginal florets female, ‘subligulate’ and disc
florets hermaphrodite and bilabiate; involucre
short- or long-cylindrical or campanulate, sometimes very broadly so; phyllaries multiseriate,
imbricate, gradate; receptacle epaleaceous, naked,
convex to almost flat. Ray florets absent or few to
many, female; corollas yellow, orange, pink, purple
or white, bilabiate, outer lip a conspicuous, short
3-toothed limb, inner of 2 shorter linear lobes,
sometimes strongly reduced; anthers rudimentary
or present only as filaments; style base lacking
basal node but with enlarged basal portion, arms
shortly bifid or apparently adnate. Disc florets
few to numerous, hermaphrodite; corollas usually
clearly bilabiate with outer three-toothed limb
and inner lip of two linear, usually long straight
lobes, rarely subligulate with all lobes appearing
on one limb but one lobe separated by deeper
sinus, usually yellow; anther collar sometimes
109
discernible as abrupt change in colour of filament
near connection with anther; basal anther appendages extremely long-caudate, usually entire,
sometimes sparsely laciniate, often somewhat
contorted; style base lacking distinct basal node
but with enlarged basal portion, shaft glabrous,
arms relatively short, short-papillose outside,
adnate, subacute. Achenes fusiform, indistinctly
ribbed; pappus setae uniseriate, with broad, flattened rachis, margins plumose, white or greyish,
usually about as long as corollas. n = 13, 23, 24, 26.
Circa 62 species, Argentina, Brazil, Bolivia, Chile,
Colombia, Ecuador, Paraguay, Peru, Uruguay.
Six sections have been recognized by Cabrera
(1965).
51. Urmenetia Phil.
Urmenetia Phil., Fl. Atacam.: 26, tab. 3 (1860).
Subshrub or perennial herb. Leaves alternate and
loosely rosulate, lamina broadly elliptic or ovate,
denticulate. Capitula terminal on scape, radiate,
homogamous; involucre turbinate to campanulate
and hemispherical; phyllaries few-seriate, gradate,
imbricate; receptacle epaleaceous, glabrous, flat to
slightly convex, alveolate. Ray florets uniseriate,
female; corollas bilabiate, white or pink, outer lip
short 3-toothed at apex, inner of two long twisted
filiform lobes; anther cylinder reduced to apparently free staminodes; style base with distinct basal
node, shaft glabrous, expanding considerably to
middle, upper half considerably thicker, purple,
arms short, short-pilose outside, with distinct
labia. Disc florets numerous, hermaphrodite, fertile; corollas yellow, actinomorphic, short 5-lobed,
lobes erect to slightly spreading; basal anther
appendage caudate, short-pilose; style base with
distinct basal node, surface very short-papillate,
shaft glabrous, expanding gradually upwards,
arms short, somewhat enlarged, acute, shortpilose. Achenes apparently 5-ribbed; carpopodium
indistinct; pappus heteromorphic, ferrugineous,
outer series capillary, multiseriate, unequal, of
numerous finely and sparsely barbellate setae,
inner series of few awn-like scales with laciniate
margins and long-attenuate apices. One species, U.
atacamensis Phil., Chile and Argentina.
52. Pachylaena D. Don ex Hook. & Arn.
Fig. 23
Pachylaena D. Don ex Hook. & Arn., Companion Bot. Mag.
1: 106 (1835).
Fig. 22. Compositae-Mutisieae. Mutisia subspinosa. A Flowering shoot. B Ray floret. C Disc floret. (Drawings by Margaret Tebbs)
Prostrate, rhizomatous rosulate herbs or subshrubs. Leaves simple, lamina spathulate, fleshy,
110
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
solitary, terminal, sessile or short-pedicellate,
radiate, heterogamous; involucre campanulate or
hemispherical; phyllaries 3–4-seriate, imbricate,
gradate; receptacle flat to slightly convex, naked,
alveolate. Florets numerous, marginal female, disc
hermaphrodite; corollas yellow, of marginal florets
bilabiate, outer lip an enlarged 3-toothed limb,
inner deeply bifid, lobes spiralled to coiled, corollas of disc florets bilabiate but lacking markedly
enlarged limb, outer lip 3-toothed, inner bifid,
lobes coiled or recurved; basal anther appendages
sagittate, tails pilose; style lacking basal node,
glabrous, shaft gradually thickening towards style
arms, arms scarcely separating, short, obtuse,
apices very short-papillose. Achenes cylindrical;
carpopodium obscure; pappus biseriate, barbellate, whitish or yellowish. n = 23 + 2B. Three
species, Chile, Argentina.
54. Chaetanthera Ruiz & Pav.
Fig. 23. Compositae-Mutisieae. Pachylaena atriplicifolia.
A Flowering plant. B Ray floret. C Disc floret. (Drawings by
Margaret Tebbs)
finely to coarsely and often irregularly denticulate,
apices rounded or obtuse. Capitula solitary, terminal, sessile to subsessile, radiate, large; involucre
broadly campanulate to hemispherical; phyllaries
3–5-seriate, imbricate, gradate; receptacle flat to
concave, epaleaceous, alveolate to reticulate, naked.
Florets heteromorphic, marginal florets female,
disc florets hermaphrodite; corollas of marginal
florets yellowish-red to pink, bilabiate, outer
lip an enlarged 3-toothed limb, corollas of disc
florets yellow, bilabiate, outer lip scarcely enlarged,
3-toothed, inner bifid, lobes coiled; basal anther appendages caudate, pilose; style lacking basal node,
glabrous, arms short, scarcely bifid, obtuse. Achenes cylindrical; pappus 2–3-seriate, setae broad
and flattened at base, margins plumose, somewhat
fragile, whitish. Two species, Argentina, Chile.
53. Brachyclados D. Don
Brachyclados D. Don, Philos. Mag. 11: 391 (1832); Cabrera,
Fl. Patagonica 7: 316–318 (1971), key.
Dwarf caespitose or lax shrubs. Leaves alternate,
simple, lamina linear-lanceolate, entire. Capitula
Chaetanthera Ruiz & Pav., Fl. Peruv. Prodr.: 106, tab. 23
(1794); Cabrera, Revista Mus. La Plata, ser. 2, 1: 87–210
(1937), rev.
Luciliopsis Wedd. (1856).
Erect or prostrate annual or perennial herbs
or subshrubs, monoecious, rarely dioecious.
Leaves opposite and decussate with connate bases,
subulate, or alternate and linear, oblanceolate
or spathulate, entire, finely serrate or dentate,
sometimes almost pectinate. Capitula terminal,
solitary, sessile, rarely 2 or 3 in a cyme, radiate
or disciform, heterogamous, rarely homogamous,
small to medium; involucre campanulate or cylindrical; phyllaries 2–4-seriate; receptacle usually
flat, epaleaceous, glabrous. Ray florets female;
corollas white, yellow, rarely orange or reddish,
bilabiate, outer lip an enlarged usually 3-toothed
limb, rarely lacking teeth, inner of two very short,
rarely long lobes, style as in disc florets. Disc
florets hermaphrodite or female; corollas yellow,
bilabiate, outer lip usually only slightly longer than
inner, 3-toothed at apex, inner of two short lobes;
basal anther appendages caudate, pilose; style base
glabrous and lacking basal node, shaft glabrous,
arms short, bifid, short-papillose outside. Achenes terete or sometimes compressed, marginal
sometimes filiform and sterile; carpopodium
annular; pappus 1–2-seriate, often flattened at
base, barbellate or subplumose or almost plumose
towards base, whitish to fawn. n = 11, 12, 14. Circa
42 species, Argentina, Bolivia, Chile, Peru.
Compositae
Seven subgenera were recognized by Cabrera
(1937).
55. Lycoseris Cass.
Lycoseris Cass., Dict. Sci. Nat. 33: 474 (1824); Egeröd &
Ståhl, Nordic J. Bot. 11: 549–574 (1991), rev.
Dioecious subshrubs or shrubs, usually scandent.
Leaves alternate, simple, lamina ovate, elliptic,
lanceolate, entire or serrulate. Capitula solitary,
terminal or few to several in corymbs or racemes,
large, many-flowered, female often considerably
larger than male; involucre hemispherical to
campanulate; phyllaries c. 6-seriate in male plants,
c. 8-seriate in female plants, in female capitula
usually longer; receptacle flat to convex, alveolate
from coalesced acicular bristles between achenes.
Florets usually numerous, heteromorphic; corollas
orange to orange-red, sometimes yellow or violet.
Ray florets uniseriate, sterile, bilabiate, outer
lip an expanded (1–)3(–5)-toothed limb, inner
apparently absent or a single linear lobe; disc
florets actinomorphic, relatively short 5-lobed,
lobes sometimes of varying lengths, glabrous;
functional anthers present only in disc florets of
male capitula; basal anther appendages caudate,
long-entire, sometimes with ‘erose’ margins; style
base glabrous, of female florets scarcely enlarged
but lacking basal node, of male florets with distinct
node, shaft of male and female florets glabrous,
arms of female florets spreading, flattened, margins papillose, of male florets scarcely divergent
or sometimes divergent in ray florets. Achenes
cylindrical, ±5-ribbed; carpopodium annular, narrow; pappus setae numerous (150–200) in female
florets, few to many (−50) in male florets, flattened,
margins barbellate, sometimes with ± dilated
apices, fragile, whitish. Eleven species, Central
America (Guatemala), northern and western South
America (Bolivia, Colombia, Brazil), Mexico, USA.
56. Cnicothamnus Griseb.
Cnicothamnus Griseb., Abhand. Königl. Gesell. Wissensch.
Göttingen 19: 196 (1874); Cabrera, Fl. Prov. Jujuy, Rep.
Argent. X: 576–579 (1978), key.
Shrubs or small trees. Leaves alternate, simple,
lamina broadly elliptic or ovate, dentate. Capitula
solitary, terminal, large, ± sessile; involucre
broadly campanulate to globose; phyllaries multiseriate, imbricate, gradate; receptacle glabrous,
convex, naked, scarcely reticulate. Florets numerous, hermaphrodite, heteromorphic; corollas
intense orange to orange-red, marginal bilabiate
111
with expanded 3-toothed outer lip, inner lip
profoundly bilobed, lobes spiralled or recurved
only at apex, disc corollas deeply divided, lobes
erect, recurved at apex; basal anther appendages
sagittate, tails long, linear, glabrous; style lacking
basal node, glabrous, arms scarcely divided,
short, obtuse, glabrous. Achenes cylindrical,
densely long-setuliferous; carpopodium annular,
indistinct; pappus setae 3-seriate, flattened in
lower part, barbellate, straw-coloured. n = 22. Two
species, Argentina, Bolivia.
57. Plazia Ruiz & Pav.
Plazia Ruiz & Pav., Fl. Peruv. Prodr.: 104 (1794); Cabrera,
Darwiniana 9: 363–386 (1951), rev.
Harthamnus H. Rob. (1980).
Shrubs, often resinous. Leaves spiralled, lamina
ovate or oblong, entire. Capitula solitary, terminal,
surrounded by leaves, radiate; involucre campanulate; phyllaries few-seriate, imbricate, gradate; receptacle flat to slightly convex, epaleaceous, naked.
Florets numerous, hermaphrodite, all fertile; corollas white to pink, glabrous, marginal florets bilabiate, outer lip distinctly 3-toothed, inner profoundly bifid, lobes coiled, inner florets actinomorphic, deeply divided; basal anther appendages caudate, short-pilose, base ± penicillate; style with
basal node, glabrous, shaft glabrous, arms scarcely
bifid, obtuse, glabrous or short-papillose outside.
Achenes glabrous, attenuate towards base and narrowed beneath apical callus; carpopodium almost
indiscernible from body of achene, upper margins
procurrent on base of achene; pappus multiseriate,
setae barbellate, tawny to straw-coloured. n = 27.
Three species, Peru, Bolivia, Argentina, Chile.
58. Aphylloclados Wedd.
Aphylloclados Wedd., Chloris And. 1: 11 (1855); Cabrera,
Darwiniana 9: 363–386 (1951), rev.
Almost leafless, well-branched odoriferous shrubs.
Leaves alternate, simple, sessile, minute, rapidly
falling, linear-spathulate, entire. Capitula solitary,
terminal rarely in few-headed scorpioid-like
cymes, radiate or disciform; involucre campanulate; phyllaries 3–5-seriate, imbricate, gradate;
receptacle flat, naked, alveolate, fimbriate. Florets
several to many (c. 10–40), heterogamous or
homogamous; corollas lilac to purple, outer florets
bilabiate, with an outer 3-toothed limb, inner
lip profoundly 2-lobed, lobes linear, recurved to
coiled, disc florets actinomorphic, profoundly
5-lobed, lobes recurved, pubescent at apices;
112
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
basal anther appendages sagittate, long-pilose;
style with basal node, glabrous, shaft glabrous,
purplish, arms short, scarcely divergent, apices
obtuse. Achenes turbinate; carpopodium shortcylindrical; pappus 2–3-seriate, setae barbellate to
subplumose especially near apices, straw-coloured.
Five species, Bolivia, Chile, Argentina.
biseriate, long-barbellate below, barbellate to subplumose towards apices, straw-coloured. n = 27.
One species, C. genistoides D. Don, Argentina, Bolivia, Chile(?), Paraguay.
59. Gypothamnium Phil.
Annual or perennial herbs, subshrubs or suffrutices. Leaves radical or alternate, lamina small to
large, linear to broadly ovate or subrhomboid,
sometimes hastate, sagittate or lyrate. Capitula
solitary or few to many in panicles, usually radiate,
rarely discoid, erect; involucre hemispherical,
campanulate or turbinate; phyllaries many-seriate,
imbricate, gradate, persistent and spreading
after loss of achenes; receptacle flat, glabrous
or long-pilose, sometimes fimbrillate. Ray florets, when present, uniseriate, female; corollas
violet or purple, bilabiate, outer lip enlarged,
3-toothed, inner inconspicuous, usually 2-lobed,
sometimes rudimentary or even lacking, anthers
rudimentary; style base with basal node, glabrous,
shaft glabrous, arms appearing adnate or scarcely
divergent at very apices, often distinctly clavate,
glabrous. Disc florets hermaphrodite, fertile,
numerous; corollas usually yellow, reddish or
purple, tubular, 5-lobed, lobes equal or unequal, or
sometimes subligulate with an enlarged 4-toothed
limb and inner lip a single long, linear lobe;
basal anther appendages long-caudate, often
sparsely retrorsely short-papillate, anther collar
with obvious constriction at base; style base with
distinct basal node, glabrous, shaft glabrous, arms
short, obtuse, papillate at least around margin.
Achenes cylindrical, 3–6-ribbed, sometimes very
conspicuously, narrowed beneath distinct apical
callus; carpopodium a distinct annulus with upper
margins procurrent on base of achene body,
sometimes eccentric; pappus setae 2-seriate to
multiseriate, barbellate, isomorphous and capillary or heteromorphous with inner series larger
and broader, more numerous and sometimes
darker than outer series, yellowish, straw-coloured
or off-white. n = 18. Circa 32 species, Central and
South America (Argentina, Bolivia, Colombia,
Costa Rica, Ecuador, Guatemala, Mexico, Peru).
Gypothamnium Phil., Fl. Atacam.: 27, t. 3C (1860).
Moderately branched glabrous shrubs. Leaves
spiralled, often ascending, simple, linear, somewhat fleshy, entire. Capitula terminal, solitary,
radiate, medium to large; involucres cup-shaped;
phyllaries glabrous; receptacle flat, naked. Florets
heteromorphic, heterogamous, marginal florets
uniseriate, female, spreading, ± reflexed; disc florets numerous, hermaphrodite; corollas glabrous,
purple or pinkish-purple, marginal bilabiate,
outer lip enlarged, narrow, 3-toothed, inner
profoundly bifid, lobes coiled, disc actinomorphic,
profoundly 5-fid, lobes coiled; basal anther appendages caudate, tails sparsely long-pilose; style
base scarcely enlarged but lacking basal node,
shaft glabrous, arms short, obtuse, scarcely bifid,
glabrous or sparsely short-pilose outside. Achenes
± turbinate; carpopodium scarcely discernible;
pappus 2–4-seriate, setae straw-coloured, outer
series capillary, barbellate, inner series flattened,
margins barbellate, apices ± inflated. n = 36. One
species, G. pinifolium Phil., Argentina, Chile.
60. Cyclolepis D. Don
Cyclolepis D. Don, Philos. Mag. 11: 392 (1832).
Gynodioecious spiny shrubs. Leaves alternate, simple, lamina small, coriaceous, entire. Capitula few
to several on short side shoots, sessile, spicate, discoid, heterogamous; involucre campanulate; phyllaries few-seriate, lowermost scale-like; receptacle
small, glabrous, flat to slightly convex. Female florets few, fertile; corollas yellowish, equally fivelobed, or sometimes with one much longer lobe;
anther cylinder lacking; style base without basal
node, glabrous, shaft glabrous, arms relatively long,
bifid, glabrous, edges prominent. Hermaphrodite
florets few, fertile; corolla yellowish, equally fivelong-lobed, lobes somewhat rolled; anther collar
inconspicuous; basal anther appendages caudate,
pilose; style base lacking basal node, glabrous, shaft
glabrous, arms long, glabrous. Achenes densely
setuliferous; carpopodium annular; pappus setae
61. Onoseris Willd.
Onoseris Willd., Sp. Pl. 3, 3: 1702 (1803); Ferreyra, J. Arnold
Arb. 25: 349–395 & Lam. I–IX (1944), rev.
62. Ianthopappus Roque & D.J.N. Hind
Ianthopappus Roque & D.J.N. Hind, Novon 11: 97 (2001).
Subshrub. Leaves alternate, lamina coriaceous,
elliptic to orbicular. Capitula few to many in
Compositae
lax corymbs, radiate, heterogamous; involucre
few-seriate; phyllaries densely sericeous outside,
apices long-acute; receptacle convex, glabrous,
naked. Ray florets female, fertile; corolla glabrous,
bilabiate, outer limb white, often purplish beneath, short three-toothed, inner lip of two long
rolled lobes; basal anther appendages caudate,
long-attenuate, margins pilose; style base with
basal node, glabrous, shaft glabrous, arms purple, short, with thickened margins. Disc florets
hermaphrodite, fertile, actinomorphic; corollas
purplish, lobes revolute; basal anther appendages
caudate, long-attenuate, margins pilose; style
base with basal node, glabrous, shaft glabrous,
arms purple, short-bifid, with thickened margins.
Achenes long-cylindrical, obscurely 10-ribbed;
carpopodium annular; pappus setae 3-seriate,
barbellate, apices slightly dilated, purplish to red
wine-coloured. One species, I. corymbosus (Less.)
Roque & D.J.N. Hind, northern Argentina, the
extreme south of Brazil, Uruguay.
63. Hyalis D. Don ex Hook. & Arn.
Hyalis D. Don ex Hook. & Arn., Companion Bot. Mag. 1:
108 (1835).
Rhizomatous shrubs. Leaves alternate, simple,
lamina linear-lanceolate to oblong-lanceolate,
entire. Capitula few in corymbs, radiate, pedicellate; involucre narrowly campanulate; phyllaries
3-seriate, imbricate, gradate; receptacle flat, naked.
Florets hermaphrodite, 5–6, fragrant; corollas glabrous, usually pink, sometimes white or purplish,
marginal florets 4–5, bilabiate, outer lip 3-toothed,
inner profoundly bilobed, lobes coiled, central floret solitary, actinomorphic, deeply divided, lobes
revolute to tightly coiled; basal anther appendages
sagittate, tails laciniate to long-pilose; style shaft
glabrous, arms scarcely bifid, obtuse, glabrous.
Achenes obovoid to turbinate, 10-ribbed; carpopodium a narrow annulus; pappus multiseriate,
setae numerous, barbellate, often slightly inflated
and ± subplumose towards apices, whitish. n = 27.
Two species, Bolivia, Paraguay, Argentina.
64. Lulia Zardini
113
alveolate. Ray florets uniseriate, female; corollas
yellow or orangish, glabrous, bilabiate, outer lip
a ray, 3-toothed at apex, considerably longer than
inner, inner strongly 2-lobed, lobes linear, often
twisted along their length; staminodes present;
style base lacking basal node, expanding gradually
towards upper part, shaft glabrous, arms short,
short-papillose outside, margins thickened. Disc
florets hermaphrodite; corollas yellow, bilabiate,
lips equal, outer 3-toothed at apex, inner 2-lobed,
lobes long-acute, erect, filaments glabrous; basal
anther appendages very long-caudate, pilose; style
base lacking basal node, glabrous, shaft glabrous,
arms short-pilose outside, margins somewhat
thickened. Achenes obcompressed with 2 or
often 4 ribs, rostrate; pappus setae 2–3-seriate,
flattened, barbellate, off-white, apices dilated and
conspicuously darker than main stipe, often fawn.
One species, L. nervosa (Less.) Zardini, Brazil.
65. Chucoa Cabrera
Chucoa Cabrera, Bol. Soc. Argent. Bot. 6, 1: 40 (Nov. 1955).
Weberbaueriella Ferreyra (1955), non Weberbauerella Ulbr.
(1906) [Leguminosae-Papilionoideae].
Poorly and laxly branched shrubs. Leaves alternate, lamina oblanceolate, spinose-dentate
with few spines, apices spiny. Capitula solitary,
long-pendunculate, axillary, or two capitula in divaricately branched axillary inflorescence, discoid,
homogamous, slightly nodding before anthesis
but erect in flower; involucre turbinate to narrowly
campanulate; phyllaries 4–5-seriate, gradate, imbricate, subulate, ending in a long spine; receptacle
flat to slightly convex, epaleaceous, alveolate,
margins of alveolae ciliate. Florets actinomorphic,
hermaphrodite, several; corollas 5-lobed, yellow;
basal anther appendages long-sagittate, usually
entire, sometimes appearing laciniate/rough;
style base with glabrous basal node, shaft glabrous throughout, arms short, usually adpressed,
short-papillate outside. Achenes cylindrical; carpopodium stopper-shaped with thickened upper
margin; pappus setae multiseriate (3+), persistent,
barbellate, more densely so towards apices. One
species, C. ilicifolia Cabrera, endemic to Peru.
Lulia Zardini, Bol. Soc. Argent. Bot. 19: 255 (1980).
Subshrub or perennial herb. Leaves alternate, simple, lamina coriaceous, entire. Capitula solitary,
terminal on scapes, radiate; involucre campanulate
or hemispherical; phyllaries 4–5-seriate, imbricate, gradate; receptacle glabrous, concave to flat,
II.5. Subtribe Gerberinae
Benth. & Hook. f. (1873).
Rosettiform herbaceous annuals or perennials,
rarely shrubs. Leaves alternate, rosulate. Inflorescences scapiform, peduncles usually elongating
114
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
as achenes ripen; capitula chasmogamous or
rarely cleistogamous; involucres hemispherical,
campanulate or turbinate; phyllaries 2–7-seriate;
receptacles epaleaceous, glabrous or rarely with
fimbriate scales; florets usually homomorphic and
all fertile; style shaft rarely papillose in upper part,
arms short, apices rounded. Achenes fusiform or
cylindrical, apices truncate or beaked, setuliferous
or rarely papillate; carpopodium annular or
absent; pappus setae 2–3-seriate, rarely uniseriate
or multiseriate, capillary, barbellate.
Scapigerous perennial, rarely annual, herbs. Leaves
rosulate, simple, often relatively few, lamina elliptic
to obovate, unlobed or lyrate-pinnatifid, entire,
finely toothed or lobed. Capitula solitary, terminal
on scapes, conspicuously or inconspicuously
radiate, heterogamous, nodding at first becoming
erect in flower and fruit, sometimes nodding in
66. Trichocline Cass.
Trichocline Cass., Bull. Sci. Soc. Philom. 1817: 13 (1817);
Zardini, Darwiniana 19: 618–733 (1975), rev.
Perennial scapigerous herbs or shrubs, rarely
caespitose. Leaves rosulate, lamina linear, linearlanceolate, oblanceolate, elliptic, oblong or
occasionally orbicular, lyrate or pinnatipartite
to pinnatisect, entire, lobed, dentate or rarely
undulate. Capitula solitary on scapes, radiate,
erect; involucre hemispherical; phyllaries 3–7seriate, outer series usually foliaceous, inner series
imbricate, gradate, rarely all phyllaries gradate;
receptacle flat or sometimes concave, epaleaceous,
smooth, with ridge around insertion point of
achenes or alveolate with fimbriate scales between
achenes. Ray florets female, uniseriate, yellow to
orangish yellow or rarely reddish, outer lip conspicuous, spreading, apices very short 3-toothed,
inner of two long linear spiralled lobes; staminodes
with sterile anthers; style base lacking basal node,
glabrous, shaft glabrous, arms short, undivided
or slightly divergent and spreading. Disc florets
numerous, hermaphrodite; corollas bilabiate,
outer lip short 3-toothed, inner of two short linear
lobes; basal anther appendages long-sagittate,
very long-papillose or pilose; style base lacking
basal node, glabrous, shaft glabrous, arms scarcely
divided to slightly recurved, rounded or obtuse,
very short-papillose. Achenes ovoid, densely
setuliferous; pappus setae multiseriate, persistent,
barbellate, sometimes with dilated apices, occasionally with crisped apices, white or off-white.
n = 18, 20. Twenty-one species, Argentina, Bolivia,
Brazil, Chile, Colombia, Paraguay, Peru, Uruguay.
67. Chaptalia Vent.
Fig. 24
Chaptalia Vent., Descr. Pl. Jard. Cels: tab. 61 (1802);
Burkart, Darwiniana 6: 505–594, pl. i–x (1944), reg. rev.;
Nesom, Brittonia 36: 396–401 (1984), part. rev.; Phytologia
78: 153–188 (1995), reg. rev.
Fig. 24. Compositae-Mutisieae. Chaptalia chapadensis.
A Rosette leaf. B Scapose inflorescence. C Ray floret. D Disc
floret. (Drawings by Margaret Tebbs)
Compositae
fruit or always erect, chasmogamous or sometimes
cleistogamous; involucre turbinate to campanulate
or hemispherical, elongating in fruit; phyllaries
imbricate, gradate, 3–6-seriate; receptacle flat to
slightly concave, foveolate/alveolate or smooth,
glabrous, epaleaceous. Florets many, usually
trimorphic, sometimes dimorphic without intermediate florets, marginal florets 1–2(–3)-seriate,
female; corollas usually white or creamy white,
rarely purplish, sometimes with purplish midstripe beneath limb, bilabiate, outer lip an enlarged
limb, apices short 3-toothed, inner present or
absent, if present, then of 2 relatively small linear
to narrowly lanceolate lobes; style shaft glabrous,
arms linear to elliptic, acute, papillose inside;
intermediate florets female, corollas often bilabiate
with distinct ray and sometimes with abortive
staminodes or sometimes corolla reduced to filiform tube surrounding style; central/disc florets,
when present, hermaphrodite, usually functionally
male, corollas white, cream or pinkish, sometimes
with a purplish stripe if bilabiate and appearing
radiate, bilabiate or actinomorphic with erect
or reflexed short lobes; basal anther appendages
caudate, tails usually entire; style base lacking
basal node, glabrous, shaft glabrous, arms short,
scarcely spreading. Fertile achenes fusiform or
rostrate with distinctive prolonged beak, usually
5-ribbed (4–12-ribbed), glabrous or setuliferous;
pappus setae uniseriate, numerous, connate into
small cup at apex of beak, barbellate, pale strawcoloured. n = 16, 24. Circa 60 species, southern
USA, West Indies, Central and South America,
with one species from Texas south to Argentina.
68. Leibnitzia Cass.
Leibnitzia Cass., Dict. Sci. Nat. 25: 420 (1825); Hansen,
Nordic J. Bot. 8: 61–76 (1988), reg. rev.; Nesom, Brittonia
35: 126–139 (1983), reg. rev.
Perennial scapose herbs. Leaves rosulate, appearing
concurrently with first capitula, lamina discolorous, entire to lyrate-lobed. Capitula solitary
on long scapes, usually chasmogamous, radiate,
heterogamous, sometimes cleistogamous, erect;
involucre turbinate; phyllaries imbricate, usually
3–4-seriate, gradate; receptacles flat to slightly
convex, foveolate. Florets all with fertile ovaries;
corollas white, cream, pink or purple-tinged,
bilabiate, marginal florets with an expanded limb,
female, those of cleistogamous heads with reduced
limb, disc florets tubular, hermaphrodite, those
of chasmogamous heads strongly bilabiate, those
115
of cleistogamous heads ± actinomorphic with
very short lobes; basal anther appendages caudate;
style base lacking basal node, glabrous, shaft
glabrous, arms short, scarcely divergent, rounded,
dorsally pilose. Achenes fusiform, short- or longrostrate in some species, moderately setuliferous,
setulae adpressed and ascending; carpopodium
a minute annular ring; pappus setae 2–3-seriate,
numerous, finely barbellate, capillary, persistent,
straw-coloured or purple. n = 23. Six species,
China, Guatemala, India, Japan, Nepal, Siberia,
Taiwan, and Mexico and the south-western USA.
69. Uechtritzia Freyn
Uechtritzia Freyn, Oesterr. Bot. Z. 42: 240 (1892); Hansen,
Nordic J. Bot. 8: 61–76 (1988), rev.
Scapose perennial herbs. Leaves rosulate, lamina
lyrate, pinnatifid or entire and ± sinuate. Capitula
solitary on terminal scape, radiate, usually homogamous; involucre hemispherical; phyllaries imbricate, gradate, 5–7-seriate; receptacle alveolate, margins fimbriate-laciniate. Ray florets uniseriate, female with staminodes; corolla pale pink or purple,
bilabiate, outer lip limb-like, short 3-toothed, inner of two long, sometimes coiled lobes, glabrous;
basal appendages of staminodes caudate, scarcely
pilose; style base lacking basal node, glabrous, shaft
glabrous, arms short, connate, short-papillose outside. Disc florets numerous, hermaphrodite, fertile;
corollas pink or purple, glabrous, bilabiate with two
subequal lips, outer short 3-toothed, inner of two
long tightly coiled lobes (or outer 2-toothed, inner
with 3 long coiled lobes); basal anther appendages
long-caudate, short-pilose; style base lacking basal
node, glabrous, shaft glabrous, arms short, scarcely
separated, short-papillose outside. Achenes often
obscurely 6–11-ribbed; pappus setae 2–3-seriate,
barbellate, apices more coarsely and densely barbellate, white to off-white. Three species, Armenia,
Turkey, Afghanistan, China, India and Kashmir.
70. Amblysperma Benth.
Amblysperma Benth., Enum. pl.: 67 (1837).
Perennial rosulate herbs. Leaves probably seasonal, lamina oblanceolate, sinuate to lobulate or
sometimes very broadly serrate with mucronate
lobes, sometimes entire. Capitula solitary on
terminal scapes, radiate; involucre turbinate to
campanulate; phyllaries 3–5-seriate, gradate,
imbricate; receptacle epaleaceous, glabrous, flat to
slightly convex, scarcely ridged between achenes.
116
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Florets numerous, all fertile, heteromorphic. Ray
florets functionally female, uniseriate; corollas
glabrous with prominent well-developed outer lip
forming ray, apex short 3-toothed, white above
and pink to brownish purple beneath, inner
of two long linear lobes, usually tightly coiled,
often somewhat dissimilar; basal appendages
of staminodes decurrent on the filament; style
base lacking basal node, glabrous, shaft glabrous,
arms adnate. Disc florets with poorly developed
outer lip of three longish teeth or sub-bilabiate
to actinomorphic with five subequal lobes; basal
anther appendages long-caudate, usually smooth;
style base lacking basal node, glabrous, shaft
glabrous, arms relatively short, scarcely divided,
short-papillate outside towards apices. Achenes
cylindrical, densely covered in long papillae with
rounded/blunt apices; pappus setae few-seriate,
persistent, barbellate, off-white. Two species
endemic to western Australia.
71. Perdicium L.
Perdicium L., Pl. Rar. Afric.: 22 (1760).
Perennial scapigerous herbs. Crown usually
white-lannose. Leaves rosulate, few, lamina narrowly obovate, coarsely lobed, sometimes almost
runcinate, lobes sparsely serrate. Capitula solitary,
terminal on scapes, disciform, heterogamous;
involucre campanulate, few-seriate; phyllaries
gradate, imbricate; receptacle epaleaceous, convex. Florets bilabiate, corollas white, glabrous;
marginal florets uniseriate, female; corollas with
short 3-toothed outer lip, inner of two long lobes;
staminodes absent; style shaft glabrous; style arms
moderately long, short-papillose outside, obtuse;
central/disc florets probably bilabiate at first, but
appearing long 5-lobed in some material; basal
anther appendages long-caudate, entire; style base
lacking basal node, but often with a nectary disc,
glabrous, shaft glabrous, cylindrical, arms short,
scarcely divided, short-papillose outside. Achenes
terete, densely papillate or glabrous, papillae with
obtuse apices; pappus setae multiseriate, barbellate, usually united into a distinct cupule-like
callus at base and detached as a unit. Two species,
South Africa.
72. Gerbera L.
Gerbera L., Opera Varia: 214/247 (1758), nom. cons.;
Hansen, Opera Bot. 78: 1–36 (1985), sect. rev.; Nordic J. Bot.
5: 451–453 (1985), sect. rev.; Nordic J. Bot. 8: 61–76 (1985),
sect. rev.; Humbert, Fl. Madag. III: 854–868 (1963), reg. rev.
Piloselloides (Less.) C. Jeffrey (1967).
Scapigerous perennial herbs from woody rootstocks. Rootstock/crowns lanate or villous. Leaves
rosulate, lamina elliptic, ovate or subcircular,
herbaceous or coriaceous, entire, serrulate, dentate, pinnatifid or pinnatisect. Capitula solitary on
terminal scapes, erect, heterogamous, bilabiateradiate; involucre broadly obconic, cylindrical to
broadly campanulate; phyllaries 2-seriate to multiseriate, numerous, imbricate, gradate; receptacle
flat to convex, epaleaceous, shallowly alveolate.
Corollas white, yellow, pink or red (paler-coloured
corollas often tinged pink or violet beneath), all
2-lipped, outer lip strap-shaped or shortly elliptic
and 2–3-toothed, inner of 2 small linear lobes.
Marginal florets female, radiate, submarginal
florets female, equally bilabiate, central florets
hermaphrodite, equally bilabiate; basal anther
appendages caudate, tails entire or ‘ciliate’; style
base lacking basal node, glabrous, shaft glabrous
except for portion just beneath style arms which
has scattered to moderate short papillae, style
arms of hermaphrodite florets short and broadly
lanceolate, rounded or sub-acute with short pollen
sweeping-hairs outside. Achenes fusiform to
narrowly flask-shaped, of outer florets sometimes
filiform to narrowly cylindrical and infertile, ±
attenuate above or distinctly rostrate, sometimes
very long-beaked, 4–10-ribbed, sparsely setuliferous, setulae, when without divided apices, acute
or obtuse; pappus multiseriate, setae minutely
barbellate, white, cream, straw-coloured, pinkish
or reddish. n = 23, 25. Circa 30 species, China,
Kenya, Malawi, Madagascar, Mozambique, South
Africa, Tanzania, Uganda, Yemen, Zimbabwe. One
species, G. hieracioides (Kunth) Zardini, is native
in South America (Ecuador and Peru) but its
inclusion in the genus is disputed (Jeffrey 1967;
Zardini 1974; Hansen 1985a, b, 1990, 1991b).
The genus has been divided into six sections by
Hansen (1985a, b, 1988, 1990; cf. Jeffrey 1967).
II.6. Subtribe Gochnatiinae
Benth. & Hook. f. (1873).
Gynodioeceous, hermaphrodite, gynomonoeceous
or polygamous subshrubs, shrubs or trees. Leaves
alternate or rosulate. Inflorescences scapiform,
paniculate or corymbose; capitula homogamous
(hermaphrodite or female) or heterogamous
(hermaphrodite and female, radiate or discoid;
Compositae
involucre campanulate or turbinate; phyllaries
3–10-seriate; receptacle glabrous, epaleaceous; florets few to many, female, male or hermaphrodite,
corollas bilabiate (1/3, 2/3, 1/4) or actinomorphic
and deeply 5-lobed; style arms short, glabrous.
Achenes cylindrical to turbinate; ribbed, densely
setuliferous; carpopodium cylindrical or annular;
pappus setae 1–3-seriate, barbellate, whitish,
yellowish or reddish.
73. Gochnatia Kunth
Fig. 25
Gochnatia Kunth in Humb., Bonpl. & Kunth, Nov. Gen.
Sp. (folio ed.) 4: 15 (1818); Cabrera, Revista Mus. La Plata,
Secc. Bot. 12: 1–160 (1971), rev.
Monoecious shrubs or trees. Leaves alternate,
simple, lamina narrowly lanceolate, elliptic,
ovate or obovate, entire, dentate or denticulate
(sometimes only in upper part of leaf), rarely
spiny. Capitula solitary, terminal or few to many
in dense terminal clusters or (sect. Hedraiophyllum) numerous in panicles, cymes, corymbs
or ‘glomerules’, small to relatively large, often
subtended by numerous reduced leaf-like bracts;
involucres cylindrical, turbinate or campanulate;
phyllaries 4–10-seriate (sometimes many-seriate
117
and decrescent in size down pedicel or towards
base), imbricate, gradate; receptacle epaleaceous,
sometimes alveolate or foveolate, glabrous or with
short hairs and sometimes glands. Florets few to
many (4–c. 150), hermaphrodite; corollas actinomorphic, profoundly 5-lobed, white to cream
or yellow, ± orangish; basal anther appendages
long-caudate, sometimes very long, finely pilose
or glabrous/entire; style base lacking basal node,
glabrous, shaft glabrous, arms short and scarcely
separating to somewhat bifid, rounded to truncate.
Achenes cylindrical to turbinate, often ribbed;
carpopodium a distinct annulus or short to
distinct cylinder; pappus setae 2(–3)-seriate, setae
persistent, barbellate, straw-coloured, yellowish
to reddish. n = 23. Circa 60 species, mostly from
Cuba and the Caribbean Islands, including Puerto
Rico, Haiti, Dominican Republic, Bahamas, few
species in South America (Brazil, Bolivia, Peru,
Argentina, Paraguay), and USA and Central
America (Mexico).
Six sections have been recognized by Cabrera
(1971), and eight by Freire et al. (2002).
74. Pentaphorus D. Don
Pentaphorus D. Don, Trans. Linn. Soc. London 16: 296
(1830).
Monoecious shrubs. Leaves alternate, sessile,
entire, coriaceous, glandular-punctate. Capitula
in dense terminal clusters, sessile or subsessile,
discoid; involucre cylindrical to turbinate; phyllaries 4–5-seriate, gradate, imbricate; receptacle
glabrous, epaleaceous. Florets few to several,
(3–)5–20, hermaphrodite; corollas deeply 5-lobed,
lobes erect to partially coiled; basal anther appendages caudate, entire or antrorsely laciniate;
style base lacking basal node, glabrous, shaft
glabrous, arms rounded, scarcely separate, glabrous. Achenes turbinate; carpopodium a pale or
dark cylinder; pappus setae numerous, biseriate,
subequal, ascending to spreading, flattened with
barbellate margins, pale straw-coloured. Two
species, Argentina, Chile.
Cabrera (1971) and Freire et al. (2002) treated
Pentaphorus as a section of Gochnatia. I prefer to
recognize it as a separate genus.
75. Richterago Kuntze
Fig. 25. Compositae-Mutisieae. Gochnatia floribunda.
A Flowering branch. B Floret. (Drawings by Margaret
Tebbs)
Fig. 26
Richterago Kuntze, Rev. Gen. Pl. 1: 360 (1891); Roque &
Pirani, Taxon 50: 1155–1160 (2001), rev.
Seris Less. (1830), nom. illegit.
Actinoseris Cabrera (1970).
118
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
or turbinate; phyllaries multiseriate, gradate, imbricate; receptacle flat to slightly convex, alveolate,
glabrous, epaleaceous. Ray florets when present
uniseriate, white, pink, or purplish; corolla bilabiate, outer lip 3- or sometimes 4-toothed, inner of
two long, tightly coiled lobes (or one, if outer lip
4-toothed); staminode bases caudate and pilose;
style base glabrous with basal node, shaft glabrous,
arms glabrous, moderately long, scarcely divergent. Disc florets numerous, multiseriate, fertile,
white, creamish or pink; corollas actinomorphic,
deeply 5-lobed; basal anther appendages caudate,
laciniate; style base with basal node, glabrous,
shaft glabrous, arms relatively short, sometimes
scarcely divergent. Achenes cylindrical; carpopodium annular, sometimes excentric; pappus
setae uniseriate, barbellate, white to off-white or
straw-coloured. Nine species, Brazil.
II.7. Hecastocleis Group
Tribe Hecastocleideae Panero & V.A. Funk
(2002).
Shrubs. Leaves dimorphic, primary alternate,
margins spiny, secondary fasciculate at bases of
brachyblasts, apices spiny. Inflorescences solitary,
terminal and surrounded by broad foliaceous
spinescent bracts forming a secondary involucre enclosing several capitula; capitula discoid,
single-flowered; involucre narrowly cylindrical;
phyllaries 3–4-seriate, spiny; receptacle glabrous,
epaleaceous; florets hermaphrodite; corollas
actinomorphic, deeply 5-lobed; style arms short.
Achenes glabrous; carpopodium inconspicuous;
pappus a scale-like corona.
Only one genus:
76. Hecastocleis A. Gray
Hecastocleis A. Gray, Proc. Amer. Acad. Arts Sci. 17: 221
(1881).
Fig. 26.
Compositae-Mutisieae. Richterago discoidea.
A Rosette leaf. B Inflorescence. C Floret. (Drawings by
Margaret Tebbs)
Characters of the group. n = 8. One species, H.
shockleyi A. Gray, USA (Nevada and California).
II.8. Nouelia Group
Subshrubs or shrubs, often single-stemmed.
Leaves alternate, rosulate, lamina linear, obovate
or spathulate, entire or denticulate. Capitula solitary on scapes or in few-headed panicles, radiate or
discoid; involucre campanulate to hemispherical
Large shrubs or treelets. Leaves alternate. Inflorescence terminal solitary capitula or capitula in
dense corymbs or glomerules; capitula homogamous, radiate or discoid; involucres turbinate or
campanulate; phyllaries 4–7-seriate; receptacles
epaleaceous; florets all hermaphrodite and fertile,
Compositae
ray florets bilabiate, disc florets and those of discoid
capitula actinomorphic and deeply 5-lobed; style
arms short to moderately long. Achenes cylindrical
to turbinate, setuliferous; carpopodium annular;
pappus setae 2–3-seriate, capillary, barbellate.
77. Nouelia Franch.
Nouelia Franch., J. Botanique 2, 5: 66 (1888).
Shrub to small tree. Leaves alternate, lamina elliptic, entire or minutely serrate. Capitula solitary, terminal, radiate, homogamous; involucre turbinate
to campanulate; phyllaries multiseriate (6–7), coriaceous, imbricate, gradate; receptacle flat to convex, glabrous, alveolate. Florets all hermaphrodite,
fertile, marginal florets uniseriate, bilabiate, outer
lip 3-toothed to long 3-lobed at apex, inner of two
long lobes, tightly rolled, disc florets numerous,
actinomorphic, deeply 5-lobed, lobes tightly rolled;
basal anther appendages pilose; style base glabrous,
gradually narrowing upwards into glabrous shaft,
arms short, glabrous. Achene with annular carpopodium; pappus setae 2–3-seriate, many, barbellate, fawn. n = 27 . One species, N. insignis Franch.,
China (Sichuan–Yunnan region).
78. Leucomeris D. Don
Leucomeris D. Don, Prodr. Fl. Nepal.: 169 (1825).
Large shrubs or small trees. Leaves alternate, lamina broadly lanceolate to elliptic, entire. Capitula
in a dense terminal cyme or panicle or in a dense
many-headed terminal glomerule; involucres narrowly turbinate; phyllaries 4–5-seriate; receptacle
small, epaleaceous, scarcely alveolate. Florets few
to several, (4–)6–10, hermaphrodite; corollas actinomorphic, deeply 5-lobed, white, lobes long, usually recurved, sometimes strongly coiled; basal anther appendages caudate, sometimes contorted, entire to very irregularly and sparsely short-laciniate;
style base lacking basal node but slightly swollen
towards base, shaft glabrous, arms moderate, linear, obtuse. Achenes cylindrical to turbinate, often ribbed; carpopodium a distinct pale cylinder;
pappus setae 2–3-seriate, somewhat flattened at
base, barbellate, straw-coloured yellowish to reddish. n = 27. Two species, Burma, China, India,
Nepal, Pakistan, Thailand, Vietnam.
II.9. Catamixis Group
Small shrubs. Leaves alternate, coriaceous.
Inflorescences corymbose; capitula homoga-
119
mous, ligulate; involucre turbinate; phyllaries
4–5-seriate, scarcely imbricate; receptacle epaleaceous; corollas yellow; apical anther appendages
narrow-triangular; style arms short, acute. Achenes densely long-setuliferous, apices acute;
carpopodium annular; pappus setae uniseriate,
long-barbellate, white.
Only one genus:
79. Catamixis T. Thomson
Catamixis T. Thomson, J. Linn. Soc., Bot. 9: 342 (1865).
Characters of the group. One species, C. baccharoides T. Thomson, India and the Himalayas.
II.10. Subtribe Tarchonanthinae
Cass. ex Dumort. (1829).
Dioecious trees or shrubs. Leaves alternate. Inflorescences terminal capitula or axillary panicles or
racemes, or glomerulous; capitula unisexual, discoid or disciform; involucres campanulate, hemispherical or globular; phyllaries 1–5-seriate; receptacle glabrous or with long silky hairs; corollas 5-lobed, white, lobes villous; apical anther appendages attenuate; style arms short, flat, acute.
Achenes of female florets fusiform, ribbed, densely
setuliferous; carpopodium annular; pappus absent
or setae uniseriate and barbellate.
80. Brachylaena R. Br.
Brachylaena R. Br., Trans. Linn. Soc. London 12: 115 (1817);
Beentje, Kew Bull. 55: 1–41 (2000), rev.
Semi-deciduous, deciduous or evergreen dioecious
trees or shrubs. Leaves alternate, lamina elliptic
to obovate, entire, denticulate, repand or coarsely
dentate to sinuate towards apex. Capitula numerous, terminal in axillary panicles or racemes, or
in clusters or glomerules, unisexual, disciform,
few- to several-flowered; phyllaries imbricate,
few-seriate (usually 3–5); receptacle epaleaceous,
small, scarcely ridged. Female capitula few- to
many-flowered, (1–)4–80, involucre campanulate;
corollas filiform, 5-lobed, lobes loosely coiled to
ascending; staminodes occasionally present; style
base glabrous, arms exserted, short to medium,
flat, spreading to coiled, acute, papillate outside;
achenes fusiform, 4–8-ribbed; carpopodium
a narrow annulus; pappus setae 2–3-seriate, setae
subequal or with several outer much shorter,
barbellate, pale straw-coloured. Male capitula
120
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
smaller than female, involucre turbinate or campanulate; florets few to many, (1–)4–40; corollas
funnel-shaped, 5-lobed, white, lobes relatively
short, usually coiled; basal anther appendages
moderately short-tailed, entire to slightly laciniate,
style base lacking basal node but with prominent
nectary, glabrous, shaft glabrous, arms short-bifid,
short-papillose, acute; achenes rudimentary;
pappus setae uniseriate, barbellate, off-white to
straw-coloured. Eleven species, Angola, Botswana,
Kenya, Madagascar, Mozambique, South Africa,
Swaziland, Tanzania, Uganda, Zimbabwe.
81. Tarchonanthus L.
Tarchonanthus L., Sp. Pl.: 842 (1753); Pope, Fl. Zamb. 6, 1:
9–11 (1992), reg. rev.; Beentje, Kew Bull. 54: 81–95 (1999),
rev.; Herman, Bothalia 32: 21–28 (2002), reg. rev.
Dioecious trees or shrubs, often aromatic, evergreen. Leaves alternate, lamina elliptic to narrowly
obovate, entire to coarsely irregularly serrate or
rarely apically 3-lobed towards apex. Capitula numerous in terminal or axillary panicles, unisexual,
discoid; involucre campanulate to hemispherical
or globular; phyllaries 1–3-seriate, imbricate, gradate; receptacle ± convex, epaleaceous, sometimes
with long silky hairs. Male capitula many-flowered;
corollas cream, tubular or funnel-shaped, longer
than female, lobes 5, relatively short, apices
recurved; basal anther appendages short-caudate,
entire, connate with neighbouring appendage;
style base lacking basal node but with prominent
nectary, glabrous, shaft glabrous, relatively short,
style simple or shortly bifid. Female capitula 1-,
2- or 3-flowered; corolla lobes 4–5, equalling
tube and usually recurved, anthers absent; style
base without node, glabrous, shaft glabrous,
arms exserted, short, flat, glabrous. Achenes
obovoid or ellipsoid, densely long-setuliferous;
pappus absent. Two species, Angola, Botswana,
Ethiopia, Kenya, Lesotho, Mozambique, Namibia,
Saudi Arabia, Somalia, South Africa, Swaziland,
Tanzania, Uganda, Yemen, Zambia.
II.11. Dicoma Group
Tribe Dicomeae Panero & V.A. Funk (2002).
Annual or perennial herbs, subshrubs, shrubs or
small trees. Leaves alternate. Inflorescences of
solitary or many terminal capitula, or terminal
and scapose, corymbose or racemose; capitula
homogamous and discoid, heterogamous and
radiate, or rarely heterogamous and disciform;
involucre campanulate to almost hemispherical
or urceolate; phyllaries 4–10-seriate, sometimes
spinescent; receptacles very rarely paleaceous;
ray florets, when present, uniseriate, usually
female, bilabiate (2/3); disc florets hermaphrodite,
numerous, actinomorphic; style shaft glabrous,
style arms short, scarcely divided, acute, obtuse
or rounded, sometimes with a subapical fringe of
hairs or papillae Achenes turbinate or fusiform to
cylindrical, glabrous or densely long-setuliferous;
carpopodium annular or apparently absent;
pappus setae 1–many-seriate, capillary or sometimes flattened and squamelliform, barbellate or
subplumose to plumose, rarely laciniate.
82. Oldenburgia Less.
Oldenburgia Less., Linnaea 5: 252 (1830); Bond, S. African
J. Bot. 53: 493–500 (1987), rev.
Shrubs or small trees. Leaves alternate, rosulate
at branch apices, simple, lamina thick and coriaceous, narrowly oblanceolate to oblanceolate
or narrowly obovate, entire. Capitula solitary or
in few-headed cymes, medium to large, homogamous, radiate, sessile or pedicellate; involucre
campanulate to almost hemispherical or urceolate;
phyllaries imbricate, ‘subgradate’ with shorter
outer but subequal middle and inner series, or
entirely gradate, 4–10-seriate; receptacle flat to
concave, epaleaceous, alveolate, alveole margins
somewhat fimbriate. Ray florets uniseriate, female;
corollas white or rarely pink, bilabiate, outer lip
an elongated limb, short 3-toothed, inner of two
long linear coiled lobes; basal anther appendages
long-caudate; style base slightly inflated, glabrous,
shaft glabrous, arms short, scarcely divided,
truncate. Disc florets hermaphrodite, numerous;
corollas white, actinomorphic, deeply 5-lobed,
lobes strongly coiled; basal anther appendages
extremely long-caudate, entire, rough or longlaciniate; style base slightly inflated but without
basal node, glabrous, shaft glabrous, apices short,
scarcely bifid, truncate or acute. Achenes fusiform
to cylindrical, ribbed; carpopodium a narrow
annulus; pappus setae uniseriate, long-barbellate
to subplumose or plumose, straw-coloured to
off-white. n = 18. Four species, South Africa.
83. Pasaccardoa Kuntze
Pasaccardoa Kuntze, Rev. Gen. 1: 354 (1891); Lisowski, Flor.
Geobot. Ann. 36, pars 1, suppl. 1: 535–541 (1991), reg. rev.;
Pope, Fl. Zamb. 6, 1: 26–29 (1992), reg. rev.
Compositae
121
Annual to perennial herbs or weak shrubs or
subshrubs. Leaves alternate, simple, lamina narrowly oblanceolate to obovate, finely denticulate
or subentire. Capitula solitary, terminal or in
axillary and terminal obscurely scorpioid cymes,
medium to large, radiate; involucre campanulate
to turbinate; phyllaries multiseriate, gradate, imbricate; receptacle convex, epaleaceous, markedly
alveolate, alveole margins fimbriate. Ray florets
uniseriate, neuter; corollas deep red, occasionally
creamy white, limb short 3-toothed, inner lip
absent. Disc florets hermaphrodite, fertile; corollas
actinomorphic, glabrous or glandular-punctate,
deeply 5-lobed, red or creamish; lobes long, usually
recurved; basal anther appendages long-caudate,
retrosely (upwardly) pilose/laciniate; style base
scarcely swollen, although sometimes with a basal
nectary, glabrous, shaft glabrous to just below
branching of arms and then pilose with a band of
short hairs/papillae, arms relatively short, acute,
glabrous. Achenes ribbed, base with a dense tuft
of setulae; pappus setae subequal or very unequal
with outer shorter, united at base, broadly winged
and scale-like and narrowly lanceolate, midrib
usually very distinct, finely laciniate, acute or
acuminate. Four species, tropical Africa (Angola,
Tanzania, Zaire, Zambia).
gradate; receptacle epaleaceous, moderately to
deeply alveolate, honeycombed, margins of alveoli
toothed, at least at angles. Ray florets, when present,
few, neuter; corolla bilabiate, outer lip short 3toothed and not larger than inner 2-toothed lip or
pseudoligulate with one long sinus and a 5-toothed
lip (only in Madagascan taxa), otherwise all florets
hermaphrodite, few to many; corollas white,
cream, yellow, violet to reddish, deeply divided
into 5 narrow equal lobes, only apices coiled; basal
anther appendages caudate, retrorsely long-pilose;
nectary usually conspicuous; style base lacking
basal node in some species (and these with obvious
nectary), glabrous, shaft glabrous, arms short,
scarcely separating, obtuse, short-papillose. Achenes turbinate, 5–10-ribbed, glabrous or glabrous
and with dense basal tuft of setulae, otherwise
densely setuliferous usually between ribs; pappus
usually multiseriate, outer setae coarse and barbellate or plumose, median often with narrow hyaline
margins and barbellate to almost scabrid along
rachis, inner (when present) scale-like with broad
hyaline margins, rachis scabrid, or pappus uniseriate, of spreading to ascending scale-like setae; setae
white, straw-coloured or rarely purplish. n = 11.
Between 32 and 65 species, South Africa and tropical Africa, Madagascar, north-eastern Africa, Asia.
84. Dicoma Cass.
85. Erythrocephalum Benth. & Hook. f.
Dicoma Cass., Bull. Soc. Philom.: 12 (1817); Humbert,
Fl. Madag. III: 844–851 (1963), reg. rev.; Lawalrée &
Mvukiyumwami, Bull. Jard. Bot. Nat. Belg. 52: 151–163
(1982), reg. rev.; Lisowski, Fragm. Flor. Geobot. Ann. 36,
pars 1, suppl. 1: 541–555 (1991), reg. rev.; Pope, Fl. Zamb.
6, 1: 29–44 (1992), reg. rev.; Ortiz & Tadesse, Ann. Missouri
Bot. Gard. 85: 440–459 (1998), reg. rev.; Jeffrey & Beentje,
Fl. Trop. E. Afr. Compositae, part 1: 13–19 (2000), reg. rev.
Macledium Cass. (1825).
Hochstetteria DC. (1838).
Cloiselia S. Moore (1906).
Erythrocephalum Benth. & Hook. f., Gen. Pl. 2: 488 (1873);
Lisowski, Fragm. Flor. Geobot. Ann. 36, pars 1, suppl. 1:
528–535 (1991), reg. rev.; Pope, Fl. Zamb. 6, 1: 20–26 (1992),
reg. rev.; Jeffrey & Beentje, Fl. Trop. E. Afr. Compositae,
part 1: 26–35 (2000), reg. rev.
Achyrothalamus O. Hoffm. (1893).
Perennial, sometimes acaulescent, or sometimes
annual herbs, subshrubs or shrubs. Leaves alternate, rarely rosulate (in one Madagascan species),
densely set and imbricate, lamina linear, linearlanceolate, oblanceolate or spathulate, entire to
serrulate. Capitula solitary, terminal or in terminal
corymbs or racemes, sometimes precocious in
acaulescent species, erect, homogamous and discoid or sometimes heterogamous and disciform,
rarely radiate; involucre campanulate or turbinate,
often with numerous gradate bracts running down
pedicel; phyllaries 4- to many-seriate, imbricate,
Annual herbs or subshrubs. Leaves alternate, narrowly linear-lanceolate to obovate or broadly obovate, dentate. Capitula one to many, terminal, often
on several short side branches, homogamous and
discoid or heterogamous and radiate, ray florets
hermaphrodite, functionally male; involucre hemispherical to campanulate; phyllaries imbricate,
outer series often with foliaceous, often coarsely
dentate apical appendages; receptacle paleaceous,
convex; paleae linear-lanceolate. Ray florets usually
present, uniseriate, few to many (5–16); corollas
usually red, sometimes white or pink, bilabiate,
outer lip an expanded 3-toothed limb, teeth relatively broad and long or very short, inner of two
long, usually erect linear lobes; basal anther appendages long-caudate and retrorsely long-pilose,
anther collar somewhat pronounced; style base and
122
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
shaft as in disc florets, arms short, truncate, appearing coronate with long papillae around apices.
Disc florets usually many (20–80), hermaphrodite;
corollas actinomorphic, red, deeply 5-lobed, lobes
erect; basal anther appendages long-caudate, longpilose, retrorsely so except at base, base somewhat
truncate, anther collar somewhat pronounced;
style base with prominent basal nectary, glabrous,
shaft glabrous, arms short to moderate, truncate,
usually appearing coronate with larger papillae
around edge, often with papillae along margins for
short distance. Achene usually somewhat inflated,
5-angled, straw-coloured; pappus setae uniseriate,
often few (3–5), flattened, caducous, barbellate,
straw-coloured. Circa 13 species, eastern, central
and southern tropical Africa.
86. Pleiotaxis Steetz
Pleiotaxis Steetz in Peters, Reise Mossamb., Bot. 2: 499
(1864); Jeffrey, Kew Bull. 21: 180–200 (1967), rev.; Lisowski,
Fragm. Flor. Geobot. Ann. 36, pars 1, suppl. 1: 501–527
(1991), reg. rev.; Pope, Fl. Zamb. 6, 1: 11–20 (1992), reg.
rev.; Jeffrey & Beentje, Fl. Trop. E. Afr. Compositae, part 1:
21–26 (2000), reg. rev.
Perennial herbs or more usually subshrubs. Leaves
alternate, upper margin of base usually coarsely
toothed, lamina linear, elliptic, ovate-lanceolate
or obovate, toothed, sometimes finely so, rarely
almost entire. Capitula solitary, terminal or few to
many in racemes or racemose panicles, discoid,
homogamous; involucre campanulate, turbinate
or cylindrical; phyllaries 5–7-seriate, gradate, imbricate, broad; receptacle epaleaceous, alveolate,
sometimes with scarcely or distinctly bordered
pits, flat to slightly convex. Florets numerous,
actinomorphic, hermaphrodite; corollas red or
yellowish-white, or white in ‘albino forms’, lobes
long, linear; style base glabrous, lacking basal node,
shaft glabrous, arms short to moderately long, usually short-papillate outside above branching point,
obtuse; basal anther appendages caudate, pilose;
Achenes 4- to many-ribbed; carpopodium annular; pappus setae multiseriate, usually flattened,
barbellate, sometimes coarsely so or subplumose,
rarely with dilated and flattened apices, off-white
to fawn. n = 10. Circa 26 species, tropical Africa.
87. Gladiopappus Humbert
Gladiopappus Humbert, Bull. Soc. Bot. France 95: 181
(1948).
Small branched shrub. Leaves alternate, simple,
lamina broadly obovate, entire. Capitula solitary,
terminal, sessile, surrounded by dense cluster
of leaves, radiate, homogamous; involucre campanulate; phyllaries c. 4-seriate; receptacle flat to
slightly convex, fimbriate between achenes (acc.
to Humbert – but not seen in Kew isotype). Ray
florets uniseriate, several (c. 9–10); corollas white,
2-lipped, outer lip a conspicuous limb with 3
short teeth, inner of 2 linear, coiled lobes; basal
anther appendages long-caudate and conspicuously densely retrorsely pilose. Disc florets many
(25–30), actinomorphic, 5-lobed; corollas with
white lobes and reddish-black tubes, lobes linear,
strongly coiled; basal anther appendages longcaudate, retrorsely pilose but less so than in ray
florets; style base with distinct glabrous basal node,
shaft glabrous, arms short. Achenes obovoid or
turbinate, terete; pappus setae biseriate, persistent,
flattened and squamelliform, white, outer series
shorter than inner, of variable length and width,
laciniate, acute, inner series subequal, laciniate and
more coarsely so towards apices, acute, sometimes
irregularly divided into 2 lobes. One species, G.
vernonioides Humbert, endemic to Madagascar.
II.12. Pertya Group
Tribe Pertyeae Panero & V.A. Funk (2002).
Herbs or scandent shrubs, rarely dioecious shrubs.
Leaves alternate. Inflorescences paniculate, spiciform or racemose, or of solitary capitula on
brachyblasts; capitula 1–16-flowered, discoid, very
rarely cleistogamous; involucres cylindrical or
campanulate; phyllaries 5–7-seriate; receptacles
epaleaceous; florets hermaphrodite and deeply 5lobed or zygomorphic with one deeper sinus; style
arms short, acute or obtuse to rounded. Achenes
fusiform, ribbed or terete, glabrous or setuliferous;
carpopodium annular; pappus setae 2–3-seriate,
rarely absent, barbellate or subplumose.
88. Pertya Sch. Bip.
Pertya Sch. Bip., Bonplandia 10: 109 (1862).
Perennial rhizomatous herbs, shrubs or scandent
shrubs. Leaves alternate, sometimes fasciculate
on brachyblasts, lamina subulate to lanceolate or
oblong to ovate, entire, dentate or divided. Capitula racemose, paniculate or solitary, generally
few-flowered, discoid, homogamous; involucre
campanulate; phyllaries multiseriate, imbricate,
membranous or coriaceous; receptacle flat, glabrous or villous, epaleaceous. Florets (1–)5(–16),
hermaphrodite, fertile; corollas actinomorphic,
Compositae
deeply 5-lobed, usually equally lobed, usually
white, sometimes slightly pinkish towards base;
lobes loosely coiled above; basal anther appendages
caudate, tails conspicuously pilose or finely laciniate; style base with distinct node, glabrous, shaft
glabrous, arms short, dorsally with short hairs
or papillae. Achenes obovoid-oblong, 10-ribbed;
carpopodium annular; pappus setae 2–3-seriate,
finely barbellate, off-white, fawn or pinkish.
n = 12, 13, 14. Fifteen species, Afghanistan, China,
Thailand, Japan, Taiwan.
89. Macroclinidium Maxim.
Macroclinidium Maxim., Bull. Acad. Imp. Sci. SaintPétersbourg 15: 375 (1871).
Perennial herbs. Leaves more or less rosulate
or alternate at base of inflorescence, lamina
large, obovate or trilobed, grossly dentate or
serrate. Capitula numerous in panicles, compound
corymbs or spikes, single- to several-flowered
(1 or 6–11), discoid, homogamous; involucres
5–7-seriate, cylindrical to campanulate; phyllaries imbricate; receptacle convex, glabrous or
setose, alveolate. Florets hermaphrodite, fertile;
corollas actinomorphic, deeply 5-lobed, white,
lobes long, usually coiled towards apices; basal
anther appendages caudate, tails conspicuously
pilose or finely laciniate; anther collar indistinct;
style base with conspicuous node, glabrous, shaft
glabrous, arms short, with short hairs or papillae
beneath, acute. Achenes terete or sometimes
striate; carpopodium annular; pappus 2–3-seriate,
setae barbellate, white, apices usually somewhat
flattened and dilated. Three species, Japan.
90. Ainsliaea DC.
Ainsliaea DC., Prodr. 7: 13 (1838).
Diaspananthus Miq. (1866).
Perennial herbs. Leaves usually alternate, lamina
linear to very narrowly lanceolate, elliptic, ovate or
orbicular, entire, crenate, dentate or lobed. Capitula many in spikes, racemes or elongated panicles,
solitary or in dense clusters, often nodding,
discoid, homogamous; involucres cylindrical;
phyllaries few-seriate to multiseriate, imbricate,
gradate; receptacle small, epaleaceous, glabrous.
Florets few (1–3), hermaphrodite, usually all fertile, rarely cleistogamous or sterile; corollas deep
rose-purple, sometimes white, actinomorphic
and deeply 5-lobed with lobes erect to reflexed,
or sometimes zygomorphic with deeper sinus
between two lobes and almost ligulate; basal
123
anther appendages caudate with connate shortpilose/laciniate or entire tails; style base scarcely
swollen or with distinct basal node, glabrous,
shaft glabrous, arms short, truncate, dorsally
with short hairs or papillae. Achenes fusiform to
oblanceolate, sometimes flattened; carpopodium
annular, usually whitish; pappus uniseriate, setae
plumose, brownish or purplish, rarely absent.
n = 6, 8, 12, 13, 14, 15. Circa 50 species, China,
India, Japan, Nepal, Taiwan, southeast Asia.
91. Myripnois Bunge
Myripnois Bunge, Enum. Pl. China bor. collegit.: 38 (1833).
Dioecious shrub. Leaves scattered and alternate
or few (1–5) verticillate on brachyblasts, lamina
simple, entire. Capitula solitary and sessile or very
short-pedicellate, on brachyblasts, usually subtended by bud and leaf-scales in 2–3 series. Male
and female capitula few-flowered, homogamous,
male smaller than female; involucre narrowly
campanulate to cylindrical, subtended by one
phyllary-like subinvolucral bract; phyllaries 5–7,
biseriate, both series of ± equal size, not imbricate;
receptacle small, naked, epaleaceous. Female
florets lacking anther cylinder; corollas purple,
5-lobed with much deeper sinus between fourth
and fifth lobe, sometimes subligulate; style base
not inflated, style glabrous in lower half and with
short hairs in upper part, arms short, divergent,
with short hairs outside. Male florets with abortive
achene; corollas purple, irregularly 3-lipped, two
of two short lobes and a third of the remaining
long lobe; style base with prominent nectary,
otherwise lacking basal node, glabrous, shaft
glabrous in lower part and short-pilose in upper
part beneath arms, arms slightly divergent, pilose
outside; basal anther appendages caudate, finely
laciniate or short-pilose. Achenes of female florets
long-setuliferous; carpopodium annular; pappus
setae barbellate to subplumose, white to off-white.
One species, M. dioica Bunge, Mongolia and China.
III. Tribe Cardueae Cass. (1819).5
A. Susanna and N. Garcia-Jacas
Perennial, biennial, or monocarpic herbs or
shrubs, less often annual herbs, very rarely small
5
The authors of the tribal account prefer to adopt the name Cardueae Cass. (1819) for the tribe, but the name Cynareae Lam. &
DC. (1806) is validly published under Articles 32.1 and 19.6 of the
124
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
trees, often spiny. Leaves alternate, frequently
rosetted, rarely in terminal whorls. Resin-ducts
always present in roots, less frequent in aerial
parts; laticiferous cells often present but only
in aerial parts. Capitula scapose-solitary or
diversely corymbose, often aggregate, usually
many-flowered, rarely glomerate in secondary
capituliform compound inflorescences and then
one-flowered. Involucral bracts in many rows,
spiny or unarmed, foliaceous or membranous,
often prolonged into a membranous, variously
fimbriate, lacerate or pectinate, spiny or unarmed
appendage. Receptacle variously chaffy or more
often setose, rarely naked (Alfredia pro parte,
Dolomiaea, Onopordum, Myopordon, Russowia
and Tugarinovia). Florets usually tubular, very
rarely peripheral florets ligulate (Atractylis and
Carlina); all fertile or the peripherals sterile
through abortion and radiant; sterile peripheral
florets often absent (especially in subtribe Carduinae). Corollas usually almost actinomorphic,
very rarely zygomorphic, divided into a tube and
a campanulate limb, straight or s-shaped. Anthers
sagittate, apically extending into a rigid, lignified,
lanceolate appendage, basally caudate, often
with long divisions; anther filaments glabrous or
papillose; in many derived groups (especially in
subtribe Centaureinae), the stamens are strongly
thigmotropic, making up an elaborate mechanism
of pollen presentation. Pollen tricolporate, oblate,
spherical or more or less prolate; ektexine formed
by two layers of columellae, sometimes caveate (in
subtribe Centaureinae), spiny (Fig. 27A), verrucate, scabrate (Fig. 27B) or almost smooth. Style
with a papillose-pilose thickening (functionally
a pollen brush) below the branches; stigmatic areas
confined to the inner surfaces of the style arms;
nectary present at the base of the style. Achenes
very variable, with parenchymatous pericarp
(in Carlininae, Cardopatiinae and Echinopsinae,
rarely in Carduinae) or radially lignified (in Carduinae and all Centaureinae), hirsute in subtribes
Carlininae, Cardopatiinae and Echinopsinae,
glabrous in most of Carduinae and Centaureinae.
Insertion areole basal, basal-lateral or lateral.
Pappus of scales or bristles, directly attached
to the pericarp wall (subtribes Cardopatiinae,
Carlininae, and Echinopsinae; Berardia, Staehelina
and the Xeranthemum group of Carduinae) or
connate by means of a parenchymatous ring to the
International Code of Botanical Nomenclature, and under Article
11.1 must be adopted as correct (C. Jeffrey, ed.).
Fig. 27. Compositae-Cardueae. Pollen grains. A Serratulatype (Cheirolophus sempervirens). B Centaurea scabiosatype (Centaurea prolongi; Garcia-Jacas 1992)
apical plate; pappus in two structurally different
rows (double pappus) in subtribe Centaureinae.
Pinnules shorter than width of palea (scabrate), as
long as width of palea (pinnulate) or much longer
and capillary (plumose). Apical caruncle present
in many genera of subtribe Carduinae, basal
elaiosome in subtribe Centaureinae, associated
with myrmecochory.
The tribe contains 73 genera and over 2,360
species.
Chromosome numbers show complex dysploid
series within natural groups. As a general rule,
chromosome numbers in subtribe Carlininae
range from x = 12 in the basal Atractylodes to
x = 10 in Carlina. In subtribe Echinopsinae, basic
chromosome numbers are x = 7, 14, 15 and 16. Subtribe Carduinae shows also complicated dysploid
patterns, not fully correlated with phylogeny. The
Xeranthemum group shows a dysploid series with
x = 6, 7, 10 and 11. Many groups (e.g. Onopordum,
Cynara and Cirsium) have x = 17, while x = 13
seems widespread in Saussurea and even x = 11
is present in Galactites. In subtribe Centaureinae,
the relationship between dysploidy and phylogeny
was thoroughly studied by Garcia-Jacas et al.
(1996). Basal groups have chromosome numbers
ranging from x= 13 to x = 15, whereas the complex
of genera with derived features (pollen usullay
caveate, crystals in the phyllaries, and hilum
basal) show base chromosome numbers from
x = 7 to x = 12. Secondary metabolites include
predominantly lipophilic compounds (especially
sesquiterpene lactones); hydrophilic compounds
are scarcely represented. Only some genera have
been investigated in depth (Centaurea, Cirsium).
Subtribes Carduinae and mainly Centaureinae
show higher structural differentiation than the
remaining subtribes (Wagner 1977). Geographical
Compositae
distribution is mainly Mediterranean in its widest
sense (including the Irano-Turanian region), with
a major centre of diversification in central Asia.
The mountains of central Asia (Tian-Shan, Pamir
and the Himalayas) constitute the eastern boundary for most of the genera. The tribe becomes less
frequent in central Africa, and disappears in equatorial and southern Africa. Three genera are native
to North America (Cirsium, Plectocephalus and
Saussurea), only two to temperate South America
(Centaurodendron and Plectocephalus), and two
species are (with reservations) native to Australia.
The tribe contains many subcosmopolitan and
noxious weeds (Acroptilon, species of Carthamus,
Carduus, Centaurea, Cirsium and Onopordum).
The traditional classification of Cardueae
into four subtribes (Carduinae, Carlininae,
Centaureinae and Echinopsinae) is extremely
controversial (reviewed by Dittrich 1977; Susanna
et al. 1995; Petit 1997); many doubts are centred
in subtribes Carlininae and Echinopsinae, often
treated as separate tribes, Carlineae and Echinopeae. However, molecular evidence of diverse
origin (cpDNA restriction sites, Jansen 1991;
rbcL, Kim et al. 1992; nuclear-ribosomal DNA,
Susanna et al. 1995; combined ribosomal-nuclear
and chloroplast DNA, Garcia-Jacas et al. 2002;
Susanna et al. 2006) consistently groups Cardueae in a robust monophyletic clade. A limited
cladistic-morphological analysis by Bremer (1994)
also supported the monophyly of the group.
However, none of these analyses has solved
the relative position of the subtribes; they have
suggested only that Carlininae, the restored subtribe Cardopatiinae, and Echinopsinae probably
form a grade basal to subtribes Carduinae to
Centaureinae; Centaureinae are a highly derived monophyletic group, and Carduinae are
a paraphyletic assemblage. These results suggest
that the compound inflorescence of Echinops is
without systematic relevance. The trend towards
grouping uniflowered capitula into compound
inflorescences is common in all subtribes of
Cardueae: in Carlininae (Atractylis polycephala
has only three flowers per head), in Cardopatiinae
(Cardopatium), in Carduinae (Cousinia congesta,
Cirsium congestum) and even in Centaureinae
(Centaurea aggregata). Subtribe Echinopsinae
should not include Xerantheminae, a separate tribe
in Cassini’s earliest classification, usually included
in Carlininae, moved to Carduinae by Petit (1997),
and moved again to Echinopsinae by Garcia-Jacas
et al. (2002). The Xeranthemum group is here
125
placed in Carduinae, together with Staehelina. In
the classical delimitation of the tribe (e.g. Bremer
1994), Staehelina and the Xeranthemum group
were placed in Carlininae.
Most recent studies have also confirmed the paraphyly of Carduinae when the monophyletic Centaureinae alone, or Centaureinae plus their sister
group (Cousinia and Saussurea groups), are moved
to a different subtribe. In view of the strong support
for the monophyly of the tribe, we have followed
the traditional classification in one tribe Cardueae,
even if broadly redefined and with the restoration
of subtribe Cardopatiinae. However, taking into account the very probable paraphyly of Carduinae,
this classification is a conservative approach and
not fully satisfactory. The alternative solution, i.e.
to combine subtribes Carduinae and Centaureinae,
would be impractical because the resulting Carduinae would encompass nearly 2,300 species representing more than 90% of the tribe.
We have organized the genera of Carduinae
in five informal groups. This classification follows
partly the suggestions of Dittrich (1977), Häffner
and Hellwig (1999) and Häffner (2000), strongly
modified on the basis of Garcia-Jacas et al. (2002)
and Susanna et al. (2006). However, the position
of some genera from central Asia is only tentative. All of them are poorly represented in Western
World herbaria (cf. we have not seen any material),
and the diagnoses and descriptions are not conclusive. In Centaureinae, we have organized the genera
into several informal groups which recent molecular studies have confirmed as natural. Regarding
generic limits, we have acted more as “lumpers”
than as “splitters” in Cousinia, Jurinea and Saussurea. Many small genera described from central
Asia fall within the broad range of variability of
one of these three large genera, and the lack of a recent, comprehensive revision of any of the three
makes the suggested segregates only hypothetical.
Finally, there is the problem of the treatment of
Centaurea. In the present account, the small group
previously classified as Centaurea sect. Centaurea
is named Rhaponticoides, and we have used Centaurea for the remainder of the genus (the Acrocentron, Cyanus and Jacea groups, as defined in
Garcia-Jacas et al. 2001).
Key to the Subtribes
1. Capitula one-flowered, clustered in second-order
spherical or hemispherical compound inflorescences
3. Echinopsinae (p. 128)
126
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
– Capitula not one-flowered, sometimes clustered in
flat-topped corymbs but never forming second-order
heads
2
2. Achenes soft, with parenchymatous pericarp,
sericeous, rarely glabrous. Receptacle with broad
scales, very rarely naked (Tugarinovia)
3
– Achenes hard, lignified, usually glabrous. Receptacle
with bristles, rarely naked
5
3. Capitula usually subtended by pectinate-pinnatisect
leaf-like bracts. Corolla lobes 1–3 mm long. Pappus of
long, plumose bristles, often connate at the bases into
broader, robust scales
1. Carlininae (p. 126)
– Capitula usually subtended by entire or dentate bracts
or on leafless pedicels. Corolla lobes usually longer
than 3 mm. Pappus of free bristles or scales, very rarely
connate at the bases
4
4. Pappus of two rows of short lanceolate scales. Corollas
filiform, blue
2. Cardopatiinae (p. 127)
– Pappus of one or two rows of pinnulate or plumose long
scales or bristles. Corollas not filiform, never blue
4. Carduinae (p. 129)
5. Insertion areole of the achenes straight, basal or basalabaxial, without elaiosome. Achenes often with apical
caruncle. Pappus rarely differentiated into two rows,
usually deciduous
4. Carduinae (p. 129)
– Insertion areole of the achenes concave, lateralabaxial, often with an elaiosome. Caruncle absent.
Pappus differentiated into two rows (sometimes
aborted), more often persistent
5. Centaureinae (p. 138)
III.1. Subtribe Carlininae Dumort. (1827).
Perennial herbs or shrubs, plants less often annual.
Leaves often spiny, deeply pinnatisect, rarely
entire. Capitula frequently subtended by pectinate
leaf-like bracts, homogamous or heterogamous
with radiate florets. Inner involucral bracts often
showy, radiant and coloured. Receptacle densely
covered with large scales, absent only in Tugarinovia. Anther filaments glabrous, appendages long
and sericeous. Corolla and style often very short.
Achenes with parenchymatous pericarp, densely
sericeous. Pappus of plumose bristles, often
connate into stout scales, persistent or deciduous.
3. Annual herbs with innermost involucral bracts spinytipped
94. Thevenotia
– Perennial or annual herbs with unarmed innermost
involucral bracts
92. Carlina
4. Leaves entire or 3–5-lobate, finely serrulate. Involucral
bracts green, foliose. Achenes with a basal glabrous
thickening
95. Atractylodes
– Leaves usually pinnatisect, spiny-toothed or rarely entire and unarmed. Involucral bracts scarious. Achenes
without basal glabrous thickening
93. Atractylis
Genera of Carlininae
92. Carlina L.
Fig. 28
Carlina L., Sp. Pl. 2: 828 (1753); Petit, Bull. Mus. Natl Hist.
Nat., B, Adansonia 9: 407–440 (1987), part. rev.; Meusel &
Kästner, Österr. Akad. Wissensch., Math.-Naturwissensch.
Kl., Denkschr. 127, 128 (1990–1994), rev.
Chamaeleon Cass. (1827).
Annual or perennial herbs or shrublets. Leaves
dentate to pinnatisect, spiny, rarely entire and
unarmed. Capitula homogamous or heterogamous. Outer involucral bracts similar to upper
leaves, dentate to pinnatisect, spinulose; innermost
narrowly linear, longer than florets, radiant and
brightly coloured, rarely not radiant. Receptacular
scales connate into a honeycombed structure
covering the receptacle and enclosing the achenes.
Florets from whitish to pink or purple, or yellow.
Corolla lobes papillose on the margins. Achenes
oblong-obconical, sericeous or villous. Pappus of
plumose, basally stout and connate scales. x = 10,
rarely x = 9. Twenty-eight species, central Europe,
Mediterranean region (2 species in the Canary
Islands, 1 species in the Caucasus).
Key to the Genera
1. Plants dioecious. Receptacle naked, foveoloate. Male
heads much smaller than female, both scapose on leafless pedicels laterally arising from a basal rosette of
leaves
96. Tugarinovia
– Plants monoecious. Receptacle with scales. Capitula
not scapose on leafless pedicels laterally arising from
a basal rosette of leaves
2
2. Receptacular scales connate, enclosing the achenes.
Innermost involucral bracts showy, rarely absent 3
– Receptacular scales free, not enclosing the achenes.
Innermost involucral bracts inconspicuous
4
Fig. 28. Compositae-Cardueae. Carlina acaulis. A Habit.
B Inner involucral bract. C Achene. D Plumose pappus hair.
(Font Quer 1962)
Compositae
93. Atractylis L.
Atractylis L., Sp. Pl. 2: 829 (1753); Meusel & Kästner, Österr.
Akad. Wissensch., Math.-Naturwissensch. Kl., Denkschr.
127, 128 (1990–1994), part. rev.; Petit, Bull. Soc. Bot. France
134: 165–184 (1987), part. rev.
Annual or perennial herbs or shrublets. Leaves
dentate to pinnatisect, spinulose, rarely unarmed.
Capitula heterogamous, sessile in involucels of
leaf-like bracts. Outer involucral bracts winged;
innermost linear-lanceolate, often appendiculate
with brown, scarious, wide appendages. Receptacle with large, scarious, laciniate-fimbriate
scales. Florets whitish to violet, pink, purple,
or yellow. Peripheral ligulate florets sometimes
present, with staminodes and stylodia. Achenes
oblong-obconical, sericeous. Pappus of plumose
free scales. x = 10. Thirty species, mainly Mediterranean (Europe and northern Africa); also Eurasia
and Canary Islands.
94. Thevenotia DC.
Thevenotia DC., Arch. Bot. (Paris) 2: 331 (1833).
Annual herbs. Leaves white-tomentose, coriaceous, somewhat fleshy, with prominent veins,
spiny-toothed, densely tomentose or lanate. Capitula homogamous, subtended by leaf-like bracts.
Receptacle with fimbriate scales, basally connate.
Outer involucral bracts dentate-spinulose, woolly;
middle bracts entire, oblong; innermost ending
in a dark purple spine. Florets purple. Achenes
oblong-obconical, sericeous. Pappus of plumose,
basally stout and connate bristles. Two species,
Irano-Turanian and central Asia semi-deserts.
127
x = 12. Five species, eastern Asia (Siberia, China,
Korea and Japan).
96. Tugarinovia Iljin
Tugarinovia Iljin, Izv. Glavn. Bot. Sada S.S.S.R. 27: 356
(1928); Dittrich et al., Bot. Jahrb. Syst. 108: 167–186 (1987),
rev.
Dioecious perennial herb with stout woody stock.
Leaves rosulate, deeply dentate, spiny. Capitula
on decumbent peduncles arising laterally from
the rosette, subscapose, homogamous, male
or female; male capitula smaller than female.
Receptacle naked, alveolate. Achenes obconical,
very densely sericeous. Pappus of scabrid bristles
basally connate into larger scales, double, inner
longer than outer. One species, T. mongolica Iljin,
montane steppes of Mongolia.
III.2. Subtribe Cardopatiinae Less. (1832).
Spiny perennial or annual herbs. Leaves spinydentate or pinnatisect. Capitula either few-flowered
and clustered in corymbs, or many-flowered. Involucral bracts with spiny pectinate-fimbriate
appendages. Anther filaments glabrous. Florets
deep blue, filiform. Style very shortly bilobed.
Achenes with parenchymatic pericarp, densely
sericeous; pappus double, of two rings of short
scales.
Key to the Genera
1. Robust perennials. Heads few-flowered, clustered in
flat-topped corymbs
97. Cardopatium
– Delicate annuals. Heads many-flowered, not clustered
in flat-topped corymbs
98. Cousiniopsis
95. Atractylodes DC.
97. Cardopatium Juss.
Atractylodes DC., Prodromus 7: 48 (1838); Zarembo &
Boyko, Comp. Newslett. 33: 61–72 (1999), part. rev.
Cardopatium Juss., Ann. Mus. Natl Hist. Nat. 6: 324 (1805).
Broteroa DC. (1836).
Perennial herbs. Leaves simple or pinnately 3to 5-lobed, marginally finely serrulate-spinulose.
Capitula often heterogamous. Outer involucral
bracts green and foliose, widely winged; medium
bracts with a narrow translucent margin; innermost linear-lanceolate. Receptacle with linear,
entire scales. Florets white, yellow, pink or purple;
female florets sometimes present. Anther filaments
thick, basally flattened. Apical appendages very
long, slender. Achenes linear or oblong, sericeous,
with a basal, glabrous, thickened lip. Pappus of
easily deciduous, basally connate plumose bristles.
Robust perennial herbs. Leaves pinnatisect, very
spiny. Capitula numerous in a flattened corymb
subtended by leaf-like spiny bracts, very small and
few-flowered, homogamous. Involucral bracts with
pectinate, very spiny appendages, gradually less divided inwards, outer leaf-like, innermost scarious
and lanceolate. Receptacle densely setose with very
thin fimbriae. Style shortly bilobed with a subapical
ring of longer hairs. Achenes obconical-oblong, villous. Pappus of apically fimbriate scales with a few
lateral fimbriae. x = 18. One or two species, eastern Mediterranean region, from the Caucasus to
Fig. 29
128
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
(C. Winkler) Nevski, semi-desert steppes of central
Asia.
III.3. Subtribe Echinopsinae (Cass.) Dumort.
(1829).
Perennial herbs, less often annuals, spiny or unarmed. Capitula one-flowered, aggregated in secondary inflorescences. Bracts spiny. Anther filaments glabrous, basal appendages short, laciniate.
Achenes with parenchymatous pericarp, densely
sericeous. Pappus of broad, short scales directly
attached to the apical plate.
Key to the Genera
1. Secondary capitula spherical. Leaves pinnatifid or pinnatisect, rarely entire, spiny
99. Echinops
– Secondary capitula hemispherical. Leaves entire, unarmed
100. Acantholepis
99. Echinops L.
Fig. 30
Echinops L., Sp. Pl. 2: 814 (1753); Garnatje et al., Folia
Geobot. 40: 407–419 (2005), mol. phylog.
Fig. 29. Compositae-Cardueae. Cardopatium corymbosum.
A Habit and leaf. B Involucral bract. C Floret. D Achene.
(Fiori and Paoletti 1895–1899)
Italy and northern Africa. Robust colonizing ruderal plants with an easily sprouting rootstock.
98. Cousiniopsis Nevski
Cousiniopsis Nevski, Trudy Bot. Inst. Akad. Nauk S.S.S.R.
ser. 1, 4: 288 (1937).
Small annual herb, branched from the base. Leaves
pinnatisect, spinulose. Capitula homogamous,
solitary, sessile, one or two basal, two more
subterminal, subtended by an involucel of leaf-like
bracts. Outer involucral bracts with a pectinatespinulose appendage, innermost with a brown,
scarious, cuspidate appendage. Receptacle densely
setose with laciniate scales. Anther appendages
laciniate, with divisions as long as the filament.
Corolla lobes rolled outwards. Achenes obovoid,
sericeous-villous. Pappus of membranous scales in
two series; outer linear-lanceolate, laciniate; inner
subulate and connate. One species, C. atractylodes
Stout annual or perennial herbs. Leaves entire or
dentate to pinnatisect, spiny. Capitula surrounded
by a tuft of bristles, aggregated in globose secondary capitula subtended by small bracts. Involucral bracts in many rows. Outer bracts strongly
keeled, winged, apically slightly fimbriate. Median
more broadly winged, spinose. Innermost usually
green or green-brown, polished, linear-lanceolate,
often partly or totally connate with only the apical appendages free, not spinose. Florets violetblue, red, greenish or white. Corolla lobes often
apically scarious, densely denticulate. Achenes oblong. x = 14, 15, 16. About 120 species, Eurasia,
northern and central Africa; introduced elsewhere.
100. Acantholepis Less.
Acantholepis Less., Linnaea 6: 88 (1831).
Unarmed annual herb. Leaves sessile, linearoblong, entire or marginally denticulate-spinose,
white-woolly. Capitula single-flowered, each capitulum surrounded by a tuft of white plumose hairs,
aggregated in semi-globose terminal secondary
capitula subtended by leaf-like bracts. Involucral
bracts in many rows, very densely woolly, especially apically, free. Outer bracts bristly. Florets
pink. Achenes linear-oblong, many-angled. x = 7.
One species, A. orientalis Less., central Asia.
Probably not different from Echinops.
Compositae
129
plate very often inclined adaxially. Pappus inserted
on a parenchymatous ring in the apical plate, simple or in many undifferentiated rows, deciduous.
Key to the Genera
1.
–
2.
–
3.
–
4.
–
5.
–
6.
–
7.
–
8.
–
9.
–
10.
–
11.
Fig. 30. Compositae-Cardueae. Echinops ritro. A Habit.
B Compound head. C Individual capitulum. D Floret. (Font
Quer 1962)
–
12.
–
III.4. Subtribe Carduinae (Cass.) Dumort.
(1827).
Perennial, biennial or annual spiny herbs or subshrubs, rarely unarmed. Capitula homogamous,
very rarely peripheral florets sterile and radiant.
Bracts usually spiny, innermost exappendiculate or
with rudimentary appendages. Achenes with radially sclerified pericarp (absent in Staehelina and
the Saussurea group), often with apical caruncle.
Insertion areole straight or lateral-abaxial. Apical
13.
–
14.
–
15.
–
16.
Leaves spiny or spinulose
2
Leaves unarmed
22
Receptacle naked, honeycombed (foveolate)
3
Receptacle with bristles
5
Robust biennial herbs with spiny-winged stems, rarely
acaulescent. Achenes tetrangular, often fringed or pitted
117. Onopordum
Perennial herbs with stems not spiny-winged; achenes
compressed, ridged
4
Leaves spiny-pinnatifid. Bracts with bright-coloured
appendages
118. Alfredia
Leaves entire, hastate, spiny-toothed. Bracts linear,
without appendages
121. Synurus
Capitula sessile on a broad tubular stem, clustered,
covered with woolly hairs
127. Schmalhausenia
Capitula not sessile on a tubular stem, not covered
with woolly indument
6
Receptacle with strongly twisted bristles. Pappus bristles free, individually deciduous
7
Receptacle with straight bristles. Pappus bristles
basally connate in a ring, deciduous as a single piece 8
Corollas glandular
128. Hypacanthium
Corollas eglandular
126. Cousinia
Pappus plumose
9
Pappus scabrid or barbellate
17
Receptacle fleshy. Involucral bracts orbicular. Achenes
tetragonous
108. Cynara
Receptacle not fleshy. Involucral bracts lanceolate.
Achenes rounded or compressed
10
Peripheral florets sterile, radiant. Pericarp hygrophanous
112. Galactites
Peripheral florets not radiant. Pericarp not hygrophanous
11
Anther filaments glabrous. Achenes ridged, with apical
rim
12
Anther filaments papillose. Achenes smooth, usually
without apical rim
13
Leaves linear, subglabrous. Pappus bristles s-shaped
119. Ancathia
Leaves broadly elliptic, lanate below. Pappus bristles
usually straight
124. Lamyropappus
Achenes broadly ovoid, not compressed, without nectary
113. Ptilostemon
Achenes oblong, compressed, often with apical nectary
14
Leaves green above, white-lanuginose beneath. Nectary cylindrical
116. Lamyropsis
Leaves glabrous or uniformly pilose. Nectary globose
15
Leaves white-veined. Corolla more deeply split on the
abaxial side
115. Notobasis
Leaves not white-veined. Corolla regularly split
16
Annual, much-branched plant. Capitula almost concealed by densely araneose bracts. Involucral bracts
with pectinate appendages
110. Picnomon
130
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
– Perennial or robust biennials, little-branched. Capitula not concealed by bracts. Involucral bracts ending
in a single spine
109. Cirsium
17. Outer involucral bracts with stout spines to 50 mm.
Leaves white-variegated
111. Silybum
– Outer involucral bracts with spines less than 30 mm.
Leaves not white-variegated
18
18. Anther filaments glabrous. Achenes without caruncle
19
– Anther filaments papillose. Achenes carunculate 21
19. Leaves linear or narrowly lanceolate, spinulosedentate
122. Syreitschikovia
– Leaves broadly lanceolate or oblong in outline,
spinose-pinnatisect
20
20. Achenes smooth. Capitula with a leaf-like involucel
120. Olgaea
– Achenes ridged. Capitula without involucel
118. Alfredia
21. Capitula on long leafless peduncles. Pappus bristles
s-shaped
114. Tyrimnus
– Peduncles foliose. Pappus bristles straight
107. Carduus
22. Achenes soft, parenchymatous
23
– Achenes hard, lignified
32
23. Pappus of up to 15 pinnulate scales. Capitula heterogamous
24
– Pappus of barbellate or plumose bristles. Capitula homogamous
27
24. Dwarf shrubs
106. Amphoricarpos
– Annuals
25
25. Achenes dimorphic, outer dorsiventrally compressed,
keeled or winged, innermost linear-oblong, subcylindrical
104. Chardinia
– Achenes not dimorphic
26
26. Outer florets with liguloid corollas. Innermost bracts
unarmed
103. Xeranthemum
– Outer florets not liguloid. Innermost bracts spinytipped
105. Siebera
27. Leaves lanate on both faces, with prominent veins below. Pappus bristles retrorsely twisted 101. Berardia
– Leaves usually green above, without prominent veins
below. Pappus bristles straight
28
28. Pappus very long, showy, exceeding involucral bracts
29
32
– Pappus not exceeding involucral bracts6
29. Annual plants. Leaves somewhat fleshy, sparsely
glandular-hirsute above
132. Polytaxis
– Perennial plants, very rarely annuals. Leaves not fleshy,
not glandular-hirsute above
30
30. Receptacle naked, foveolate
131. Dolomiaea
– Receptacle setose
31
31. Nectary prominent, detachable as a single piece with
the pappus. Leaves not decurrent. Achenes usually plicate, ribbed or foveolate
130. Jurinea
– Nectary not prominent. Leaves usually decurrent. Achenes ridged or costate
129. Saussurea
32. Dwarf shrubs. Leaves green above, white beneath. Pappus very long, exceeding involucral bracts, usually persistent
102. Staehelina
– Perennial herbs or shrubs. Leaves not bicoloured. Pappus short, never exceeding involucral bracts, deciduous
33
6
Except in 102. Staehelina
33. Achenes smooth, broadly ovoid, subglobose. Apical
plate slanted. Pappus detachable as a single piece
113. Ptilostemon
– Achenes often ridged, keeled or winged, oblanceolate,
never globose. Apical plate straight. Pappus bristles
individually deciduous
34
34. Leaves often hastate, very large, to 70 cm. Involucral
bracts usually hooked
125. Arctium
– Leaves small, never hastate. Involucral bracts without
hooks
126. Cousinia
Genera of Carduinae
Unplaced Genera
101. Berardia Vill.
Berardia Vill., Prosp. Hist. Pl. Dauphiné 27 (1779); Dittrich,
Ann. Naturhist. Mus. Wien 99, B: 329–342 (1996), rev.
Acaulescent, unarmed perennial herb. Leaves
rounded, entire or denticulate, densely woolly,
with veins prominent beneath, white above.
Capitula solitary, sessile, homogamous. Involucral bracts subulate, scarious, woolly, ending in
a slender flat point. Receptacle areolate. Florets
yellowish or pinkish. Staminal connective very
long, apiculate. Achenes oblong, glabrous, slightly
sulcate. Pericarp not sclerified. Pappus of scabrid
cylindric bristles retrorsely twisted, directly
attached to the apical plate. x = 18. One species,
B. subacaulis Vill., high Alps (France, Italy and
Switzerland).
102. Staehelina L.
Staehelina L., Sp. Pl. 2: 840 (1753).
Hirtellina Cass. (1827).
Unarmed dwarf shrubs or subshrubs. Leaves entire
or dentate-pinnatifid, linear to obovate, dark green
above, white-woolly beneath. Capitula corymbose or rarely solitary, homogamous. Involucral
bracts ovate to lanceolate, mucronate, sometimes
minutely hirsute. Receptacle with wide, basally
connate scales. Florets whitish or pink-purple.
Corolla lobes very long. Anther filaments glabrous;
basal appendages very long, sericeous. Achenes
linear-oblong, glabrous or sericeous, with minute
apical coronula. Pappus of bristles basally connate
into broader paleae, more or less divided apically
into pinnulate fimbriae (into capillary hairs in
Staehelina dubia L. and S. baetica DC.), always
overtopping involucre, sometimes deciduous in
a ring. x = 15, 17. Eight species, Mediterranean
region.
Compositae
131
III.4. a. Xeranthemum Group
Unarmed annual herbs, rarely dwarf shrubs.
Leaves always entire, velvety underneath. Capitula
heterogamous. Receptacle with large scarious
scales. Anther filaments glabrous, anther appendages short, laciniate. Corolla lobes very short.
Achenes often dimorphic, with pappus of long
tapering or subulate scales, rarely reduced to
a corona in Chardinia.
103. Xeranthemum L.
Fig. 31
Xeranthemum L., Sp. Pl. 2: 857 (1753).
Unarmed annual herbs. Leaves entire, ellipticlinear, usually greyish-hairy. Capitula solitary,
pedunculate. Involucral bracts scarious, ovatelanceolate to obovate; inner bracts elongated,
erect or spreading and radiate, white, pink-violet
or purple. Receptacle with linear-lanceolate,
scarious, densely glandular-punctate scales. Outer
florets sterile, with a very long pink stylodium
and corollas deeply cleft, bilabiate; central florets
perfect. Achenes narrowly obconical to obovoid,
sericeous. Pappus of wide, scarious, subulate,
apically pinnulate scales. x = 6, 7, 10. Five species,
Mediterranean region, from northern Africa to
western Asia.
104. Chardinia Desf.
Chardinia Desf., Mém. Mus. Hist. Nat. 3: 455 (1817).
Unarmed cleistogamous herb. Capitula solitary,
pedunculate. Involucral bracts ovate-lanceolate
to obovate, scarious. Receptacle with lanceolate,
subulate scales. Outer florets female, central florets
perfect. Corollas shorter than the scales of the
pappus. Achenes dimorphic; outer dorsiventrally
flattened with an adaxial keel, obovate, with broad,
lacerate-fimbriate wings apically elongated into
two horns, glabrous; central achenes obconical,
apically sulcate and minutely pilose, basally
sericeous. Pappus of the external achenes formed
by prolongations of the wings; in central achenes
a denticulate corona. x = 11. One species, C.
orientalis (L.) Kuntze, Turkey, Middle East, Iran,
Afghanistan and Pakistan.
105. Siebera J. Gay
Siebera J. Gay, Mém. Soc. Hist. Nat. Paris 3: 344 (1827).
Annual herbs. Capitula solitary or aggregated,
subtended by a small verticel of leaves. Involucral
bracts scarious, inner with a long lanceolate,
Fig. 31.
Compositae-Cardueae. Xeranthemum cylindraceum. A Habit. B Achene. (Feinbrun-Dothan 1977)
132
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
subulate appendage usually pink to purplish;
median bracts with a slender spine; outer bracts
with a minimal spine, finely tomentose. Outer
florets sterile, bilabiate; central florets perfect.
Achenes narrowly oblong-obconical, sericeous.
Pappus of scarious scales ending in a barbellate
or plumose bristle. x = 10. Two species, Turkey,
Middle East, Iran, Afghanistan and Pakistan.
106. Amphoricarpos Vis.
Amphoricarpos Vis., Fl. Dalmat. 2: 27 (1847); Schwarz,
Phyton (Horn) 14: 125–133 (1970), part. rev.
Unarmed dwarf shrubs. Leaves green above,
white-tomentose beneath, entire or denticulate.
Capitula solitary, pedunculate or scapose. Involucral bracts scarious, ovate-lanceolate to obovate.
Receptacle with linear-lanceolate, long-subulate,
apically twisted scales. Florets violet to pink or
white, outer florets female, central florets perfect
or functionally male. Achenes dimorphic; outer
achenes compressed, winged, sericeous; inner
achenes narrowly oblong-obconical, sericeous.
Pappus of subulate, bristle-like scales. x = 12. Four
species, Balkan Peninsula, Turkey and Caucasus.
9, 10, 11. About 90 species, Eurasia (mainly in
the Mediterranean region), also in northern and
central Africa (however, central African species
should belong to Cirsium; see Häffner and Hellwig
1999), introduced elsewhere.
108. Cynara L.
Cynara L., Sp. Pl. 2: 827 (1753); Wiklund, Bot. J. Linn. Soc.
109: 75–123 (1992), rev.
Arcyna Wiklund (2003).
Spiny stout perennial herbs. Leaves pinnatisect,
very spiny (unarmed in cultivated C. cardunculus L.). Capitula large, globose, solitary or clustered in lax corymbs. Involucral bracts oval, entire, coriaceous, usually spine-tipped. Receptacle
fleshy, densely setose. Florets purplish or violet.
Anther filaments papillose. Achenes glabrous, very
faintly angular; apical rim and nectary absent. Pappus of very long, plumose bristles, basally connate
in a ring, deciduous or persistent. x = 17. Eight
species, Mediterranean region. One species, C. cardunculus, widely cultivated from Roman times for
its edible fleshy receptacles.
109. Cirsium Mill.
III.4. b. Carduus Group
Spiny perennial, biennial or annual herbs or subshrubs, rarely unarmed. Leaves often decurrent;
stem frequently winged. Capitula homogamous,
rarely outer florets sterile. Florets usually purple,
pink or white, less often yellow. Anthers very
shortly caudate, with papillose filaments. Achenes
smooth, narrowly obovoid, oblong or orbiculate,
often with apical nectary. Pappus detachable as
a single piece.
107. Carduus L.
Carduus L., Sp. Pl. 2: 820 (1753); Kazmi, Mitt. Bot.
Staatsamml. München 5: 139–198, 279–550 (1963–1964),
rev.
Annual, biennial or perennial herbs with spinywinged stems. Leaves dentate-pinnatisect,
rarely entire, spiny (very rarely unarmed in
C. personatus Gaertn.). Involucral bracts longspinose-acuminate, rarely muticous. Florets red,
purple or pink. Achenes obovoid-oblong, smooth,
glabrous, umbonate, with an apical, globose,
obscurely 5-lobulate, small caruncle, rarely absent
(C. nutans L.). Pappus bristles from almost smooth
(C. collinus Waldst. & Kit.) to serrulate. x = 8,
Fig. 32
Cirsium Mill., Gard. Dict., ed. 4, 1 (1754).
Breea Less. (1832).
Cephalanophlos Fourr. (1869).
Perennial or biennial herbs, rarely to 4 m high
(Cirsium subcoriaceum (Less.) Sch. Bip.). Leaves
dentate-pinnatisect, spiny, sometimes entire, often
semi-amplexicaul. Stems less often than in Carduus
spiny-winged. Outer involucral bracts few, spiny;
inner bracts without spines, leaf-like, often appendiculate and coloured. Florets red, purple or pink,
rarely yellow. Achenes obovoid-oblong, smooth,
glabrous, with an apical rim and a small obconical caruncle. Pappus of plumose, usually deciduous
bristles. x = 17 (Eurasia) and x = 15 (North America). About 250 species, Eurasia, North America,
northern and eastern Africa, frequent in wet waste
grounds; some noxious cosmopolitan weeds (Cirsium vulgare (Savi) Ten., C. palustre (L.) Scop.).
110. Picnomon Adans.
Picnomon Adans., Fam. Pl. 2: 116 (1763), nom. cons. prop.
Acasma Vaill. (1754), nom. rej. prop.
Annual herb. Leaves dentate-pinnatifid, spinulose. Stem winged. Capitula solitary or few
clustered together, sessile, overtopped by the
Compositae
133
with long hairs, mimicking a double pappus
(the same as in Tyrimnus Cass. and Cirsium
ciliatum Moench). x = 17. Two or three species,
Mediterranean region, introduced elsewhere.
112. Galactites Moench
Galactites Moench, Methodus: 558 (1794), nom. cons.
Annual spiny herbs. Leaves pinnatifid-pinnatisect,
spiny. Capitula solitary or corymbose, heterogamous, radiate. Outer florets sterile and radiant,
flattened, patent, much exceeding the involucre,
purple or pink; central florets perfect, whitish.
Anther filaments concrescent with very short tails.
Achenes obovoid, slightly compressed, smooth,
glabrous, hygrophanous, with an apical rim and
a large hemispherical stipitate nectary. Pappus of
plumose, deciduous bristles. x = 11. Two species,
western Mediterranean region.
113. Ptilostemon Cass.
Ptilostemon Cass., Bull. Sci. Soc. Philom. Paris 1816: 200
(1816); Greuter, Boissiera 22: 9–215 (1973), rev.
Lamyra Cass. (1822).
Fig. 32.
Compositae-Cardueae. Cirsium eriophorum.
A Habit. B Involucral bract. C Floret. D Achene. (Folch
1986)
uppermost leaves. Receptacle densely setose.
Involucral bracts marginally spiny, densely covered with woolly-arachnoid hair. Florets purple.
Achenes obovoid-oblong, smooth, glabrous, with
a small apical rim. Caruncle small, 5-lobed.
Pappus plumose, deciduous, basally connate in
a ring. x = 16. One species, P. acarna (L.) Cass.,
Mediterranean region; introduced elsewhere.
Perennial or annual spiny herbs, rarely unarmed
shrublets. Leaves dentate-pinnatisect with divaricate spines, rarely entire and unarmed. Capitula
homogamous, sometimes functionally heterogamous. Leaves always white-variegated above and
snow-white beneath. Involucral bracts almost unarmed to spiny, innermost often coloured. Florets
red, purple, pink, perfect or the outer functionally
male. Anther filaments papillose. Achenes not or
only slightly compressed, smooth, glabrous. Pappus of plumose, deciduous bristles, basally connate
in a ring, in outer florets sometimes of barbellate
to scabrid bristles. x = 16, rarely x = 12. Fourteen species, dry stony places in the Mediterranean
region.
111. Silybum Vaill.
Silybum Vaill., Königl. Akad. Wissensch. Paris Phys. Abh.
5: 173 (1754), nom. cons.
Biennial herbs. Leaves coarsely dentate-pinnatifid,
semi-amplexicaul, white-veined or variegated,
spiny. Capitula solitary. Involucral bracts with
spinose margin and spine-tipped appendages.
Florets purple. Anther filaments adnate. Achenes
obovoid, compressed, smooth, glabrous, with
a white apical rim and a large apical nectary.
Pappus of scabrid, deciduous bristles, basally connate in a parenchymatous ring forming a cupule
114. Tyrimnus Cass.
Tyrimnus Cass., Bull. Sci. Soc. Philom. Paris 1818: 168
(1818).
Annual spiny herb with spiny-winged stems.
Leaves coarsely dentate-pinnatifid. Capitula solitary on long leafless peduncles, heterogamous.
Florets purple or pink; outer sterile, sometimes
absent. Anther filaments concrescent. Achenes
obovoid, sometimes flattened, with four ribs, glabrous, myxogenic in the concavities between ribs.
Pappus of smooth or minutely scabrid deciduous
134
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
bristles, inserted downwards and bent upwards,
basally connate in a very large parenchymatous
ring with long hairs on the inner rim. x = 17.
One species, T. leucographus Cass., Mediterranean
region.
115. Notobasis Cass.
Notobasis Cass., Dict. Sci. Nat. 25: 225 (1822), nom. cons.
prop.
Polyacantha Vaill. (1754), nom. rej. prop.
Annual herb. Leaves semi-amplexicaul, dentate to
pinnatisect, spiny. Stem not winged. Capitula heterogamous, usually in racemose clusters, sessile.
Involucral bracts sharply spine-tipped. Florets purple. Central florets perfect, peripheral florets sterile. Corolla more deeply cleft on the abaxial side.
Achenes obliquely and broadly obovoid, strongly
compressed, smooth, glabrous, with hygrophanous
surface. Apical rim and nectary absent. Pappus simple, of pluriseriate, plumose bristles, basally connate in a parenchymatous cupuliform ring. As in
Tyrimnus Cass. and Silybum Adans., inner margin
of the basal ring hirsute. x = 17. One species, N.
syriaca (L.) Cass., Mediterranean region.
116. Lamyropsis (Kharadze) Dittrich
Lamyropsis (Kharadze) Dittrich, Candollea 26: 98 (1971).
Perennial herbs. Leaves dentate-pinnatisect,
spinulose, green above, white beneath. Capitula
solitary or clustered. Involucral bracts spinulose,
with a short membranous appendage. Florets red,
purple or pink. Anther filaments papillose. Basal
appendages very long, filiform, with two narrow
divisions. Achenes oblong, sulcate, slightly compressed, with a smooth apical rim. Nectary 5-lobed.
Pappus plumose, deciduous, basally connate in
a ring. x = 13. Six species, eastern Mediterranean
region, Balkan Peninsula to Caucasus.
III.4. c. Onopordum Group
Robust biennial or perennial herbs, usually spinytoothed, rarely unarmed. Capitula homogamous.
Receptacular bracts very often absent; receptacle
foveolate, margins of foveoles with short scales,
rarely setose. Pappus bristles always deciduous and
basally connate in a ring.
117. Onopordum L.
Onopordum L., Sp. Pl. 2: 827 (1753).
Stout, erect, very spiny biennial herbs with
winged stems, rarely acaulescent. Leaves dentatepinnatisect or pinnatilobed, rarely undivided,
spiny. Capitula solitary or rarely corymbose.
Receptacle foveolate; foveoles with short marginal
scales. Involucral bracts very deeply serrulate,
spiny. Florets reddish, purple or pink. Anther
filaments minutely papillose. Achenes obovoidoblong, somewhat tetrangular, glabrous, often
transversally fringed, sometimes with a short
apical rim. Pappus of plumose, barbellate or
scabrid bristles. x = 17. About 60 species, western
and central Asia, Europe, northern Africa and the
Canary Islands; introduced elsewhere.
118. Alfredia Cass.
Alfredia Cass., Bull. Sci. Soc. Philom. Paris 1817: 33 (1817).
Xanthopappus C. Winkl. (1893).
Perennial spiny herbs. Leaves long-petiolate,
entire-hastate, lobed or spiny-dentate, green
above, silver-white beneath. Capitula solitary or
corymbose, often nodding at anthesis. Involucral
bracts with a large membranous appendage.
Receptacle foveolate in A. cernua Cass., setose in
A. acantholepis Kar. & Kir. and A. nivea Kar. &
Kir.; unknown in remaining species. Florets pink
or cream-white. Corolla lobes apically thickened
and incurved. Achenes obovoid, glabrous, ridged,
with a denticulate apical rim. Pappus of scabridbarbellate bristles. x = 13. Four species, central
and eastern Asia.
119. Ancathia DC.
Ancathia DC., Arch. Bot. (Paris) 2: 331 (1833).
Perennial herb. Leaves linear or linear-lanceolate,
entire, spinulose. Capitula solitary, terminal.
Involucral bracts linear to subulate, innermost brown-purple. Receptacle setose. Florets
purple-orange. Anther filaments glabrous. Anther
appendages convolute with very long tails. Achenes
deeply sulcate-ridged and minutely verrucose,
foveolate between ridges, with a long denticulate
apical rim, umbonate. Pappus of plumose s-shaped
bristles. One species, A. igniaria (Spreng.) DC.,
mountains of central Asia, China and Mongolia.
120. Olgaea Iljin
Olgaea Iljin, Bot. Mater. Gerb. Glavn. Bot. Sada S.S.S.R. 3,
36: 142 (1922).
Perennial herbs. Leaves coriaceous, dentatepinnatisect, spiny; cauline leaves semi-amplexicaul.
Compositae
Capitula solitary or clustered, with a leaf-like involucel. Involucral bracts spiny, innermost with
short, scarious, coloured appendage. Receptacle
densely setose. Florets red, blue, white or purple.
Corolla lobes apically somewhat thickened, anther
filaments glabrous, appendages long and laciniate.
Achenes linear-oblong, glabrous, smooth, with
a long dentate apical rim. Pappus of barbellate
bristles apically widened-incrassate. x = 13.
Sixteen species, Afghanistan, central and eastern
Asia (China and Mongolia). Very closely related to
Alfredia Cass.
121. Synurus Iljin
Synurus Iljin, Not. Syst. Herb. Hort. Bot. U.S.S.R. 6: 35
(1923).
Perennial herbs. Leaves very big (30–40 cm),
long-petiolate, entire-hastate, spiny-dentate, green
above, silver-white beneath. Capitula solitary,
often nodding at anthesis. Involucral bracts linear,
spiny-tipped, densely araneose. Receptacle foveolate. Florets pink. Corolla lobes apically thickened
and incurved. Achenes obovoid, glabrous, ridged,
with a denticulate apical rim. Pappus of scabridbarbellate bristles. x = 13. Four species, eastern
Asia, from Mongolia to Japan. Very closely related
to Alfredia.
135
setose. Florets purple. Achenes glabrous, slightly
compressed. Pappus of serrulate bristles. One
species, T. lomonossowii (Trautv.) Kitag. & Kitam.
Mongolia.
Probably a synonyn of Olgaea Iljin, as suggested by Bremer (1994). However, corollas of
Olgaea are zygomorphic, whereas Takeikadzuchia
was described as having actinomorphic corollas.
[Note added in proof: morphological as well as
molecular data clearly show that Takeikadzuchia
is part of Olgaea s.l.]
124. Lamyropappus Knorring & Tamamsch.
Lamyropappus Knorring & Tamamsch., Bot. Mater. Gerb.
Bot. Inst. Komarova Akad. Nauk S.S.S.R. 16: 466 (1954).
Perennial herbs. Leaves elliptic, entire, spinosedentate, green above, white-woolly beneath;
cauline leaves sessile. Capitula long-pedunculate,
very large (3–5 cm), solitary. Involucral bracts
numerous, small, spiny. Receptacle setose. Florets
pink. Corollas very long with linear corolla lobes.
Anther filaments glabrous. Achenes obovoidoblong, glabrous, with longitudinal ridges and
marked apical rim. Pappus of long plumose bristles. One species, L. schakaptaricus (B. Fedtsch.)
Knorring & Tamamsch., central Asia.
122. Syreitschikovia Pavlov
Syreitschikovia Pavlov, Repert. Spec. Nov. Regni Veg. 31:
192 (1933).
Perennial herbs or dwarf shrubs. Basal leaves
rosetted, petiolate; cauline leaves sessile, coriaceous, green above, grey-tomentose beneath,
spinulose-dentate. Capitula solitary, terminal.
Involucral bracts linear, acute, spinulose, often
reddish. Receptacle setose. Florets pink. Anther
filaments glabrous. Achenes obpyramidal, compressed, glabrous, with a small denticulate apical
rim. Pappus double; outer of filiform bristles;
inner longer and paleaceous with bristles apically
incrassate, barbellate-scabrid. x = 12. Two species,
central Asia.
123. Takeikadzuchia Kitag. & Kitam.
Takeikadzuchia Kitag. & Kitam., Acta Phytotax. Geobot. 3:
102 (1934).
Perennial herbs. Leaves spinulose, decurrent.
Capitula globose, homogamous. Involucral bracts
linear, spinulose-denticulate. Receptacle densely
III.4. d. Arctium Group
Biennial or perennial herbs, rarely annuals, spiny
or less often unarmed. Receptacle with twisted
scales. Anther filaments slightly papillose or glabrous. Cypselae streaky (with wavy fringes), very
often winged, without nectary. Pappus of free deciduous bristles.
The limits between Arctium and Cousinia
have long been unclear. Most recent work has
demonstrated that Arctium and Cousinia can be
segregated by molecular, chromosome and pollen
characters. This does not accord with morphology: Schmalhausenia and Hypacanthium, part
of Arctium on the basis of the above characters,
are morphologically much closer to Cousinia. As
there is an “Arctioid” group of Cousinia, there is
a “Cousinioid” group of Arctium. Here, the traditional generic division into four genera is followed,
transferring subgenus Cynaroides of Cousinia to
Arctium. However, it is highly probable that all
four genera will have to be broadly redefined.
136
125. Arctium L.
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Fig. 33
Arctium L., Sp. Pl. 2: 816 (1753); Duistermaat, Gorteria,
suppl. 3 (1996), rev.; Susanna et al., Collect. Bot. 26: 101–118
(2003), mol. phylog., gener. delim.
Anura (Juz.) Tschern. (1962).
Unarmed biennial herbs. Leaves very large, broadly
ovate, serrulate, rarely entire, sparsely tomentose.
Capitula often heterogamous, corymbose or in
racemose clusters, dispersed as a unit. Involucral
bracts spreading to reflexed, usually apically
hooked. Florets purple; outermost often with very
long, brightly coloured, “radiant” anther tubes.
Achenes dimorphic; outer achenes linear, strongly
recurved, unwinged; central ones obovoid-oblong,
winged or sulcate, glabrous. Pappus of scabrid
bristles. x = 18. Circa 27 species (including the
“arctioid” species of Cousinia); some species probably hybridogenic, temperate Eurasia, introduced
elsewhere.
126. Cousinia Cass.
Cousinia Cass., Dict. Sci. Nat. 47: 503 (1827); Tscherneva,
The Cousinia of the SSSR, Leningrad (1988), system.;
Susanna et al., Collect. Bot. 26: 101–118 (2003), mol.
phylog., gener. delim.
Lipskyella Juz. (1936).
Tiarocarpus Rech. f. (1972).
Perennial, biennial or rarely annual herbs or
shrublets. Leaves dentate-spinose to spinosepinnatisect, often decurrent or semi-amplexicaul.
Capitula solitary, corymbose, or variously clustered, homogamous. Involucral bracts scarious,
very spiny. Inner bracts often with a large, scarious,
coloured appendage of same colour as florets.
Florets red, purple, pink, yellow. Achenes obovoidoblong, often winged, compressed, glabrous, rarely
ridged or even wrinkled (C. dolicholepis Schrenk),
with a very reduced apical rim (sometimes absent).
Apical plate extremely small. Pappus of scabrid
bristles, rarely absent. x = 9, 10, 11, 12, 13. About
600–700 species, central and western Asia, mainly
in steppes and semi-deserts.
127. Schmalhausenia C. Winkl.
Schmalhausenia C. Winkl., Trudy Imp. S.-Peterburgsk. Bot.
Sada 12: 281 (1892).
Spiny perennial herb. Basal leaves pinnatisect,
cauline pinnatifid, very spiny, pubescent beneath,
subglabrous above. Capitula small, homogamous,
few-flowered, clustered in subspherical terminal
corymbs or sessile on a very broad (2–3 cm)
fistulose stem. Involucral bracts densely covered
with white woolly hairs concealing the capitula,
ending in long pungent spines. Florets purple.
Corolla lobes apically thickened and recurved.
Achenes 3- or 4-angled, with a denticulate apical
rim. x = 18. One species, S. nidulans (Regel)
Petrak, central Asia.
128. Hypacanthium Juz.
Hypacanthium Juz., Trudy Bot. Inst. Akad. Nauk S.S.S.R.
ser. 1, 3: 324 (1936).
Fig. 33. Compositae-Cardueae. Arctium lappa. A Habit. B
Involucral bract. C Achene. (Font Quer 1962)
Spiny perennial herb. Leaves spinose-bipinnatisect,
green above, grey-tomentose beneath. Capitula
homogamous in lax corymbs, umbilicate. Involucral bracts scarious, spiny; innermost with
a scarious, coloured appendage. Florets purple;
corolla glandular-punctate. Anther appendages
lacerate. Achenes obovoid, compressed, ribbed,
without apical rim; apical plate much reduced.
Compositae
Three species, central Asia. Very closely related to
Schmalhausenia.
III.4. e. Saussurea Group
Unarmed perennial herbs or subshrubs; only two
annual herbs. Leaves entire or pinnatisect, often
silver-white below and glabrous above, sometimes
hirsute-scabrid. Capitula cylindrical or globose,
often paniculate, homogamous. Anther filaments
glabrous. Achenes not lignified, soft. Pappus of
very long (overtopping involucral bracts), showy,
plumose bristles, basally connate in a ring; sometimes with a shorter, pinnulate deciduous pappus
connate to a globose nectary. A difficult group because of unclear generic boundaries between Saussurea and Jurinea, and the high number of small
segregates, here included in the two larger genera.
137
Unarmed perennial herbs or shrublets. Leaves entire or dentate to pinnatifid, often white-woolly beneath. Capitula scapose. Florets pink, purple, rarely
whitish. Achenes tetrangular, obconical, usually
squamulose, verrucate, tuberculate or pitted. Apical plate with a large nectary (reduced or lacking
in J. humilis DC.), rounded, laterally ribbed. Apical
rim patent, usually crenate, rarely toothed or absent
(J. depressa (Steven) C.A. Mey.). Pappus of scabrid
to barbellate or plumose, biseriate bristles, inner
basally enlarged and broader, deciduous or rarely
persistent. x = 17, 18. About 200 species, western
and central Asia, Europe, northern Africa. Diplazoptilon Ling is probably part of Jurinea sensu lato.
129. Saussurea DC.
Saussurea DC., Ann. Mus. Natl Hist. Nat. 7: 156 (1810);
Lipschits, Rod Saussurea DC., Leningrad (1979), rev.
Hemistepta Bunge (1833).
Aucklandia Falc. (1845).
Cavea W.W. Sm. et Small (1917).
Unarmed perennial herbs, rarely annual. Leaves
entire to pinnatisect, decurrent. Capitula solitary,
corymbose or paniculate, homogamous. Involucral bracts entire, not spinose. Receptacle with
large scales, these apically divided into narrow,
twisted segments. Florets purple or pink. Achenes
obconical or obpyramidal, costate, ridged, rarely
pitted or plicate; usually with an apical rim. Pappus
biseriate, outer of short, scabrid, free, deciduous
bristles, sometimes absent; inner much longer,
deciduous or persistent. x = 13. About 300 species,
temperate Eurasia, North America, doubtfully
native in Australia.
130. Jurinea Cass.
Fig. 34
Jurinea Cass., Bull. Sci. Soc. Philom. Paris 1821: 140 (1821).
Outreya Jaub. & Spach (1843).
Aegopordon Boiss. (1846).
Jurinella Jaub. & Spach (1847).
Modestia Kharadze & Tamamsch. (1956).
Pilostemon Iljin (1961).
Perplexia Iljin (1962).
Hyalochaete Dittrich & Rech. f. (1979).
Anacantha Soják (1982).
Lipschitziella Kamelin (1993).
Himalaiella Raab-Straube (2003).
Fig. 34. Compositae-Cardueae. Jurinea pjatajeviae. A Habit.
B Floret. C Achene with pappus. D Achene without pappus,
showing nectary. (Iljin 1960)
138
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
131. Dolomiaea DC.
Dolomiaea DC., Guill. Arch. Bot. 2: 330 (1833).
Bolocephalus Hand.-Mazz. (1938).
Vladimiria Iljin (1939).
Frolovia Lipsch. (1954).
Unarmed perennial herbs, often acaulescent.
Leaves in a basal rosette, ovate to lanceolate,
often white-woolly beneath. Capitula subscapose.
Florets pink or purple, rarely whitish. Achenes
tetrangular, obconical, sulcate and ribbed. Apical
plate with a large nectary, rounded, laterally
ribbed. Apical rim small, toothed. Pappus of
scabrid, biseriate bristles, inner basally enlarged
and longer, detachable as a single piece. Twelve
species, central Asia. Very closely related to Jurinea.
132. Polytaxis Bunge
Polytaxis Bunge, Del. Sem. Hort. Dorpat.: 8 (1843).
Unarmed annual herbs. Leaves entire, elliptic,
fleshy, glandular-villous. Capitula homogamous,
solitary or laxly corymbose. Receptacle setose.
Involucral bracts few, laxly imbricate. Florets pink.
Achenes oblong, sulcate, crenate-papillose in the
furrows, sparsely pubescent, with a tuft of hairs
at the base of the ridges, with apical rim. Pappus
biseriate, persistent; outer bristles smooth, inner
plumose, twice as long. Two species, Afghanistan,
central Asia. Possibly not generically different
from Saussurea.
III.5. Subtribe Centaureinae (Cass.) Dumort.
(1827).
Perennial, biennial or annual unarmed herbs,
shrubs or very rarely treelets, rarely spiny. Capitula
often heterogamous with sterile radiant florets,
rarely homogamous. Involucral bracts often with
a diversely scarious, fimbriate, pectinate, spiny
or unarmed appendage; innermost bracts always
with a scarious appendage. Achenes with sclerified
pericarp. Insertion areole concave, lateral-adaxial,
very rarely (Crupina) straight, often with an elaiosome. Apical plate straight. Pappus inserted on
a parenchymatous ring in the apical plate, double,
formed by two rows of differently pinnulate
bristles, rarely single by abortion, deciduous or
persistent.
Key to the Genera
1. Subshrubs with green virgate branches and small linear leaves 2–3 mm wide, or leafless
2
– Treelets, shrubs, perennial herbs or annuals without
green virgate branches, leafy
4
2. Heads heterogamous. Peripheral florets usually large,
showy, radiate, with staminodes. Achenes with elaiosome
135. Psephellus
– Heads homogamous. Achenes without elaiosome 3
3. Plants leafless. Achenes densely sericeous
149. Karvandarina
– Plants basally leafy. Achenes glabrous 158. Nikitinia
4. Small pachycaul tree with leaves to 25 cm long clustered in terminal whorls
140. Centaurodendron
– Shrubs, shrublets, perennial or annual herbs, not
treelets
5
5. Innermost involucral bracts ending in a pinkish,
showy, shortly spiny appendage. Achenes with long,
oblique insertion areole
141. Schischkinia
– Innermost involucral bracts without pinkish spiny appendages. Achenes without long, oblique insertion
areole
6
6. Robust unarmed or spinescent annual plant, usually
cultivated. Capitula subtended by wide, leaf-like, entire or dentate bracts. Florets orange-saffron or bright
yellow
159. Carthamus
– Wild plants with capitula not subtended by leaf-like
bracts. Florets not saffron
7
7. Receptacle naked, foveolate
8
– Receptacle setose
9
8. Dwarf tragacanthoid shrub or dwarf rosetted perennial. Bracts of the involucre spiny-tipped
153. Myopordon
– Annual plant. Bracts of the involucre with a cartilaginous margin
148. Russowia
9. Plants usually spiny
10
– Plants unarmed
13
10. Very spiny annual plants. Achenes usually dimorphic
159. Carthamus
– Spiny perennials. Achenes all similar
11
11. Florets blue, rarely yellow. Perennial herbs, rarely
shrublets
161. Carduncellus
– Florets always yellow. Shrubs or subshrubs
12
12. Leaves broadly sheathing, ending in a trifurcate spine.
Capitula to 1.5 cm. Pappus of bristles basally connate
in a ring, deciduous
162. Femeniasia
– Leaves not reduced to a spine. Capitula 2.5–3 cm wide.
Pappus persistent
160. Phonus
13. Involucral bracts with very wide membranous-scarious, usually lacerate appendage. Achenes
ridged or tuberculate, peripheral often dorsiventrally
compressed; pappus deciduous (Rhaponticum group)
14
– Involucral bracts with narrow entire hyaline margin,
mucronate, spinose, pectinate or filiform. Achenes always laterally compressed, never tuberculate; pappus
usually persistent
17
14. Large shrub to 4 m high
155. Ochrocephala
– Annual or perennial plants, not shrubby
15
15. Leaves with margins and veins minutely denticulate,
very scabrid. Insertion areole two-lipped; lower lip
incrassate, white; upper lip with two white recurved
costae
151. Callicephalus
Compositae
– Leaves not densely scabrid. Insertion areole inconspicuous
16
16. Small, inconspicuous annuals to 30 cm high. Involucral bracts spinulose
152. Oligochaeta
– Perennial herbs with muticous bracts. Achenes subequal or dimorphic
150. Rhaponticum
17. Achenes with six translucent costae, outer ones
arcuate-falcate, innermost straight 147. Plagiobasis
– Achenes without translucent costae, all subequal 18
18. Achenes diversely ridged, ribbed or foveolate, rarely
smooth, often sericeous. Insertion areole with
a whitish incrassate margin (Volutaria group)
19
– Achenes smooth or sulcate, never foveolate, rarely
sericeous. Insertion areole without incrassate margin
23
19. Involucral bracts with very long filiform appendages
146. Tricholepis
– Involucral bracts shortly mucronate or muticous 20
20. Robust plants to 150 cm tall
21
– Plants to 80 cm
22
21. Robust annual. Stem deeply striate. Capitula homogamous, small (to 15 mm), clustered in paniculate inflorescences
143. Goniocaulon
– Robust perennial to 150 cm tall, rarely smaller biennial
or annual. Stem not striate. Capitula heterogamous, to
2 cm, terminal, corymbose or solitary 144. Mantisalca
22. Involucral bracts narrowly triangular, with a small appendage, acuminate or shortly spiny 142. Volutaria
– Involucral bracts broadly oval, muticous, with only
a hyaline margin
145. Amberboa
23. Capitula homogamous
24
– Capitula heterogamous
26
24. Outer involucral bracts similar to upper leaves. Achenes markedly 4-angled
161. Carduncellus
– Outer involucral bracts not similar to upper leaves.
Achenes not 4-angled
25
25. Achenes sulcate; insertion areole almost basal
157. Klasea
– Achenes almost smooth; insertion areole markedly lateral
156. Serratula
26. Achenes not or only slightly compressed, dark brown
with darker band at basis. Insertion areole basal or
oblique. Inner row of pappus of 5–10 very short scales
134. Crupina
– Achenes compressed, without darker band at basis.
Insertion areole lateral. Inner row of pappus of many
bristles or scales
27
27. Very scabrid annual herbs. Peripheral sterile florets
with large staminodes. Achenes with cream-white,
shiny base; insertion areole deeply lateral with a rugulose elaiosome
136. Stizolophus
– Annual or perennial herbs or shrubs not very scabrid.
Peripheral radiant florets with small staminodes, very
often lacking. Base of the achenes not cream-white;
elaiosome, if present, smooth
28
28. Shrubs or shrublets, rarely perennial herbs. Achenes
without elaiosome. Pappus deciduous
29
– Subshrubs, perennial or annual herbs. Achenes usually
with elaiosome. Pappus persistent
30
29. Achenes smooth, slightly falcate, to 4 mm. Insertion
areole with five small teeth
137. Cheirolophus
– Achenes sulcate, straight, to 8 mm. Insertion areole
not toothed
154. Centaurothamnus
139
30. Appendages of bracts long-plumose. Sterile florets
flattened. Achenes with three concentric apical ridges
133. Zoegea
– Appendages of bracts not long-plumose. Sterile flowers not flattened. Achenes without apical concentric
ridges
31
31. Plants with more or less fleshy, densely glandular
leaves. Achenes sericeous, outer sterile and oblong
163. Crocodylium
– Leaves not fleshy and glandular. Achenes glabrescent
or sparsely pilose, outer ones rarely sterile and, if so,
then linear-arcuate
32
32. Sterile marginal florets without staminodes
164. Centaurea
– Sterile marginal florets with staminodes
33
33. Sterile marginal flowers indistinct. Appendages of
bracts usually reduced to cartilaginous margin.
Achenes broadly ovate-elliptical, with basal whitish
band
139. Rhaponticoides
– Sterile marginal florets big and showy. Appendages of
bracts well-developed, pectinate or fimbriate. Achenes
oblong, without whitish basal band
34
34. Sterile marginal florets with narrow divisions and
minute sterile achenes. Achenes arcuate, asymmetrical, ridged. Pappus always present, deciduous
138. Plectocephalus
– Sterile marginal florets with wide divisions, without
sterile achenes. Achenes symmetrical, smooth. Pappus
persistent, often much reduced or absent
135. Psephellus
Genera of Centaureinae
Basal Genera
133. Zoegea L.
Zoegea L., Mant. pl. 1: 15 (1767).
Unarmed, scabrid annual herbs. Capitula solitary,
heterogamous. Involucral bracts purple-stained;
outer and median with long palmate-fimbriate
appendage; innermost bracts with scarious,
lanceolate appendage, yellow above. Outer florets
sterile, exceeding the involucre, large, flattened,
orange-yellow, pink-purple or white. Corolla
tube saccate. Achenes obovoid, strongly compressed, transversely and concentrically sulcate
and sparsely sericeous above. Insertion areole
with elaiosome. Pappus double; outer of long
scabrid bristles in many rows; inner of short scales,
apically lacerate. x = 15. Three species, Turkey,
Middle East, Irano-Turanian region and central
Asia.
134. Crupina (Pers.) DC.
Crupina (Pers.) DC., Ann. Mus. Natl Hist. Nat. 7: 157 (1810);
Couderc-Le Vaillant, Thèse Doctorale, Université de ParisSud (1984), rev.
140
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Unarmed annual herbs. Leaves pinnatisect,
dentate, with glochidiate hairs and sparse deeply
branched hairs. Capitula heterogamous. Involucral
bracts unarmed, lanceolate, mucronate. Florets
pink-purple, outer sterile. Corolla pilose. Achenes
cylindrical or only basally compressed, sparsely
sericeous, dark brown or blackish with a basal
darker band, and a deep furrow above which
forms an apical rim. Insertion areole exactly
basal or sublateral. Pappus double, persistent;
outer of dark bristles, inner of 5–10 small scales.
x = 14, 15, 29. Three species, Mediterranean
and Irano-Turanian region reaching eastwards to
central Asia, northern Africa, widely naturalized
in the western United States (California, Oregon,
Idaho and Washington).
135. Psephellus Cass.
Psephellus Cass., Dict. Sci. Nat. 43: 488 (1826).
Aetheopappus Cass. (1827).
Phaeopappus Boiss. (1845) pro parte.
Hymenocephalus Jaub. & Spach (1847).
Perennial herbs or dwarf desert shrubs. Leaves
usually pinnatisect, sometimes entire-dentate,
often silver-white beneath and green above.
Capitula long-pedunculate, heterogamous. Involucral bracts with triangular lacerate-fimbriate
appendages, silvery or blackish, often scarious and
very large, totally covering the bracts, sometimes
plicate. Outer florets sterile, with staminodes,
radiant, with a broad limb divided into 5–10
subequal lobes. Achenes oblong, compressed,
with elaiosome; hilum lateral. Pappus double,
sometimes very short or lacking. x = 15. Circa 100
species, western Siberia, Iran, Caucasus, Ukraine,
Crimea, Turkey. Some species widely cultivated as
ornamentals.
136. Stizolophus Cass.
Stizolophus Cass., Dict. Sci. Nat. 44: 36 (1826).
Annual herbs. Leaves entire or lyrate-pinnatifid,
scabrid; lobes ending in a short mucro, otherwise
unarmed. Capitula solitary or laxly corymbose,
heterogamous, globose, markedly contracted
above. Involucral bracts very small and many,
with ciliate appendages ending in a slender spine.
Receptacle setose. Florets yellow; outer sterile
with large staminodes, central florets perfect;
style very long (3–4 mm). Achenes oblong, faintly
ridged, glabrous. Insertion areole very deeply
carved, with a rugulose elaiosome. Hilum caudate.
Pappus deciduous, double; outer bristles with
long pinnules, inner bristles shorter with few
pinnules. x = 15. Two species, Turkey, Caucasus
and Irano-Turanian region.
137. Cheirolophus Cass.
Cheirolophus Cass., Dict. Sci. Nat. 51: 55 (1827); Susanna
et al., Pl. Syst. Evol. 214: 147–160 (1999), mol. phylog.
Paleocyanus Dostál (1971).
Shrubs or subshrubs (Ch. uliginosus (Brot.) Dostál
perennial herb). Leaves entire or pinnatisect, often
densely glandular. Capitula indistinctly heterogamous, many-flowered, usually globose. Involucral bracts leaf-like with a small, shortly fimbriate appendage, often tinged red. Flowers usually
pink, more rarely whitish or pale yellow. Outer florets sterile, with staminodes, scarcely radiant; central florets perfect. Achenes linear-oblong; detachment scar with five small teeth, without elaiosome;
hilum basal. Pappus a single row of easily deciduous bristles. x = 15. Twenty-five species, western Mediterranean region (extending eastwards to
Malta), northern Africa and Macaronesia.
138. Plectocephalus D. Don
Plectocephalus D. Don in Sweet, Brit. Fl. Gard. 4: 51 (1830);
Hind, Curtis’s Bot. Mag. 13: 3–7 (1996), rev.
Phalacrachena Iljin (1937).
Unarmed annual or perennial herbs, densely
glandular. Leaves with minutely denticulate,
cartilaginous margins and veins. Capitula terminal, solitary or laxly corymbose, heterogamous.
Involucral bracts membranous, veined, with very
large decurrent pectinate-fimbriate appendages,
never spiny. Marginal florets long-radiant, with
very narrow corolla lobes, sterile, with staminodes.
Corolla sometimes sparsely villous and glandular.
Achenes obovoid, arcuate, markedly asymmetrical, glabrous, faintly many-ribbed, with a basal
narrowing near the areole; insertion areole lateral.
Hilum basal. Pappus obscurely double, deciduous.
Five species, relict disjoint area: two species in
North America, two in South America, two in
Russia and Ukraine and one in eastern Africa. One
species, P. americanus D. Don, cultivated as an
ornamental (“basket flower”).
139. Rhaponticoides Vaill.
Rhaponticoides Vaill., Königl. Akad. Wissensch. Paris Phys.
Abh. 5: 165 (1754); Agababian, Bot. Inst. Acad. Sci. Armenia, Erevan (1992), rev. ut Centaurea.
Compositae
Centaurea L. (1753) pro minima parte.
Bielzia Schur (1866).
Perennial herbs. Leaves deeply divided, rarely entire, with denticulate lobes, sparsely araneose or
glabrous, often somewhat cartilaginous. Capitula
terminal, usually solitary, umbilicate-globose, heterogamous. Involucral bracts leaf-like, not appendiculate, with a membranous entire margin, rarely
lacerate. Florets pink or yellow. Anther filaments
papillose. Achenes broadly oblong, blackish, glabrous, with a cartilaginous and whitish insertion
areole and a small elaiosome. Hilum caudate. Pappus double; inner row of shorter and broader bristles. x = 13?, 15. Seventeen species, Irano-Turanian
and Mediterranean region, eastern Europe.
140. Centaurodendron Johow
Centaurodendron Johow, Estud. Fl. Juan Fernández 63
(1896).
Yunquea Skottsb. (1929).
Small pachycaul tree with soft wood. Leaves
terminal on the stems, very large (to 30 cm), obovate, with broad (alate) semi-amplexicaul petiole;
margins serrate. Capitula comparatively small (to
2 cm), clustered in dense corymbs, heterogamous.
Outer bracts with pectinate appendages, innermost with a broad round cucullate appendage.
Receptacle setose. Florets purple, outer sterile and
radiant. Achenes ovoid, obscurely tetragonous,
glabrous. Pappus double, easily deciduous. Two
species, Juan Fernández Islands.
141
III.5. a. Volutaria Group
Annual or perennial herbs. Capitula heterogamous,
very rarely homogamous. Sterile radiant florets
usually large and showy, with staminodes. Achenes diversely ribbed, ridged or foveolate, rarely
smooth; insertion areole very deep, concave, with
incrassate, cartilaginous margin and a small elaiosome. Hilum basal. Pappus of scales, rarely of bristles.
142. Volutaria Cass.
Volutaria Cass., Bull. Sci. Soc. Philom. Paris 1816: 200
(1816).
Cyanopsis Cass. (1817).
Volutarella Cass. (1826).
Unarmed annual herbs, rarely subperennial (V.
muricata (L.) Maire). Leaves entire to dentatepinnatisect, scabrid. Capitula solitary or laxly
corymbose, heterogamous (rarely homogamous).
Involucral bracts green, with marked veins, with
a mucronate or spinulose linear appendage, often
blackish. Outer florets pink, purple or violet,
sterile, tubular; central florets of the same colour
(in blue-rayed species, rarely yellow). Achenes
obovoid-oblong, ribbed, transversely rugulose
between the ribs. Insertion areole lateral, with
incrassate, cartilaginous margin, with elaiosome.
Pappus of linear-oblanceolate persistent scales.
x = 14, 16. Eighteen species, dry stony places in the
Mediterranean and Irano-Turanian region, from
Arabia and Iran to Morocco.
143. Goniocaulon Cass.
Goniocaulon Cass., Bull. Sci. Soc. Philom. Paris 1817: 33
(1817).
141. Schischkinia Iljin
Schischkinia Iljin, Repert. Spec. Nov. Regni Veg. 38: 73
(1935).
Annual herb. Leaves linear with palmate spines. Capitula clustered, sessile, heterogamous. Innermost
involucral bracts acute, coloured and showy. Florets
yellowish; outer sterile with oblong sterile achenes,
central with tube 1/10 as long as limb. Achenes oblong, much compressed, glabrous; insertion areole
very long, oblique, with rudimentary elaiosome.
Pappus double; outer as long as the achene, inner
very short, of a solitary, long, broad scale. Pappus
of the external sterile achenes reduced to a single
long palea. One species, S. albispina (Bunge) Iljin,
semi-desert steppes of central Asia.
Robust annual herb to 1.5 m high with deeply striate stem. Leaves entire, serrate. Capitula to 15 mm,
homogamous, clustered in dense panicles. Involucral bracts with wide scarious margin, apiculate.
Receptacle setose. Florets pink. Achenes oblong,
longitudinally ridged, glabrous, with a small apical
rim. Insertion areole lateral, with incrassate, cartilaginous, white margin and a small elaiosome.
Pappus obscurely double with very shortly pinnulate, wide scales. x = 16. One species, G. glabrum
Cass., India, Pakistan, eastern Africa.
144. Mantisalca Cass.
Mantisalca Cass., Bull. Sci. Soc. Philom. Paris 1818: 141
(1818).
142
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Microlonchus Cass. (1826).
147. Plagiobasis Schrenk
Unarmed annual, biennial or perennial herb.
Leaves dentate-pinnatifid, strongly scabrid.
Capitula solitary, pedunculate, heterogamous. Receptacle setose. Involucral bracts coriaceous, with
a narrow hyaline margin, apically ending in a triangular, scarious, black, spinescent appendage.
Outer florets sterile, with staminodes, purple,
longer than the perfect central florets of the same
colour. Achenes obovoid, ribbed and transversely
rugulose between the ribs, glabrous. Pappus of
scales. x = 11. One species, M. salmantica (L.)
Briq. & Cavill. Extremely variable, especially in
habit; some species of doubtful value described on
the basis of these differences. Dry places, ditches
and roadsides in the Mediterranean region, from
Turkey to Morocco.
Plagiobasis Schrenk, Bull. Cl. Phys.-Math. Acad. Imp. Sci.
Saint-Pétersbourg 3: 109 (1845).
145. Amberboa Vaill.
Amberboa Vaill., Königl. Akad. Wissensch. Paris Phys. Abh.
5: 182 (1754), nom. cons.; Iljin, Izv. Bot. Sada Akad. Nauk
S.S.S.R. 30: 101–116 (1932), part. rev.
Unarmed annual or rarely biennial herbs. Leaves
entire to dentate-pinnatisect. Capitula solitary,
pedunculate, heterogamous. Outer and middle involucral bracts without appendages or with small
brown, scarious, triangular appendage. Outer
florets sterile, radiant, with a very expanded 5–
8-lobed tubular limb. Florets violet-pink. Corolla
lobes apically incrassate, recurved, darker. Achenes
obovoid-oblong, densely sericeous, brown, ribbed,
transversely rugulose, with a cartilaginous apical
rim. Pappus double, persistent, scaly. x = 16. Six
species, eastern Europe, Caucasus, south-western
Asia. One species, A. moschata (L.) DC., widely
cultivated as an ornamental.
146. Tricholepis DC.
Tricholepis DC., Arch. Bot. (Paris) 2: 515 (1833).
Unarmed annual or perennial herbs. Leaves linear,
entire or sometimes dentate-pinnatifid. Capitula
solitary, pedunculate, terminal, homogamous.
Involucral bracts numerous, with a very long
subulate-acicular and hair-pointed or spine-tipped
appendage. Florets pink, purple, lilac or yellow. Achenes obovoid-oblong, striate or faintly
many-ribbed, glabrous. Pappus of two similar
rows of wide, pinnulate scales. x = 16. Circa
20 species, central Asia, Afghanistan, Myanmar,
Nepal, Pakistan and India.
Unarmed perennial herb. Leaves elliptic-oblong,
entire, serrate; cauline leaves sessile. Capitula
terminal, globose, homogamous. Involucral bracts
orbicular with a wide hyaline margin. Receptacle
setose. Florets pink. Achenes dimorphic; peripheral falcate, central linear-oblong, faintly costate,
rounded apically. Pappus deciduous, of wide,
long pinnulate scales. One species, P. centauroides
Schrenk, central Asia.
148. Russowia C. Winkl.
Russowia C. Winkl., Trudy Imp. S.-Peterburgsk. Bot. Sada
11: 282 (1890).
Unarmed annual herb. Leaves interruptedly
pinnatisect with linear-lanceolate lobes. Capitula
laxly corymbose. Involucral bracts muticous, with
scarious-hyaline margin and three dark, translucent, cartilaginous veins. Receptacle alveolate.
Florets pink. Achenes linear-oblong, smooth,
sparsely sericeous. Insertion areole lateral, very
deep, with spirally incrassate margins, with
elaiosome. Pappus obscurely double, of long, narrow, pinnulate bristles. One species, R. sogdiana
(Bunge) B. Fedtsch., central Asia.
149. Karvandarina Rech. f.
Karvandarina Rech. f., Österr. Akad. Wissensch., Math.Naturwissensch. Kl., Anz. 87: 198 (1950).
Unarmed virgate subshrub. Leaves small, very
often totally lacking, entire or dentate, basally
hairy, otherwise glabrous. Capitula laxly corymbose, homogamous. Involucral bracts scarious,
strongly veined, with a narrow hyaline, decurrent
margin and a triangular, red-brown, scarious, not
spiny appendage. Florets pink with straight corolla
tube. Achenes obovoid-oblong, subcompressed,
sericeous-villous, apically truncate, with very
narrow apical rim; insertion areole basal-central.
Pappus double, outer of scabrid bristles, inner of
few, basally wider bristles. One species, K. aphylla
Rech. f., Aellen & Esfand., Iran, Pakistan.
III.5. b. Rhaponticum Group
Unarmed perennial herbs, rarely annuals. Capitula
homogamous. Involucral bracts with very large,
unarmed scarious appendages, usually silvery-
Compositae
white. Achenes often dimorphic, diversely ribbed
or papillose, with small insertion areole with
a rudimentary elaiosome; hilum basal. Pappus
usually deciduous in a ring.
150. Rhaponticum Vaill.
Rhaponticum Vaill., Königl. Akad. Wissensch. Paris Phys.
Abh. 5: 177 (1754); Dittrich, Candollea 39: 45–49 (1984),
nomencl.
Leuzea DC. (1815).
Stemmacantha Cass. (1817).
Acroptilon Cass. (1827).
Unarmed perennial herbs. Leaves entire or lobedpinnatisect, usually snow-white beneath and green
above. Capitula large, solitary. Involucral bracts
membranous, with an entire or lacerate appendage;
inner bracts with a subulate appendage. Florets redpurple, whitish or yellow, with narrow corolla lobes.
Achenes obovoid-oblong, more or less ribbed with
the ribs ending in 8–10 teeth forming a crenate apical rim, glabrous. Pappus obscurely double; outer
of scabrid to shortly plumose, deciduous bristles;
inner bristles wider and longer than outer. x = 12,
13. Twenty-six species, mountains of Europe, Asia,
very doubtfully native in Australia. One species, Rh.
repens (L.) Hidalgo, is an extremely noxious weed
(“Russian knapweed”).
143
long-decurrent hyaline margin ending in a small,
woolly, spinescent appendage. Florets pink-purple
or whitish. Style arms very long, rolled outwards. Achenes dimorphic; outer dorsiventrally
compressed, often epappose; inner obconical,
laterally compressed, smooth or faintly ribbed
and rugulose, glabrous, with apical rim. Pappus
double, outer of scabrid bristles; inner pappus of
1–5 longer, wider bristles. x = 14. Four species,
Irano-Turanian region, Caucasus, Afghanistan,
India.
153. Myopordon Boiss.
Myopordon Boiss., Diagn. Pl. Orient. ser. 1, 6: 107 (1846);
Wagenitz, Ber. Deutsch. Bot. Gesell. 71: 271–277 (1958), rev.
Compact, unarmed or spiny subshrubs or perennial herbs, sometimes acaulescent. Leaves entire
or pinnatisect. Capitula solitary. Involucral bracts
with a wide scarious brown appendage ending in
a short thin spine. Receptacle foveolate with short
scales. Florets reddish, purple or yellowish. Anther filaments papillose; basal appendages shortly
laciniate. Achenes oblong, longitudinally striate,
transversely rugose (keeled, plicate or pitted), subcompressed, with apical rim. Pappus of barbellatesmooth bristles. Five species, western Asia.
154. Centaurothamnus Wagenitz & Dittrich
151. Callicephalus C.A. Mey.
Callicephalus C.A. Mey., Verz. Pfl. Casp. Meer 66 (1831).
Unarmed annual herb. Leaves linear-pinnatisect,
scabrid, with denticulate veins. Capitula solitary,
terminal, heterogamous. Involucral bracts with
a large appendage ending in a short brown mucro.
Florets pink-purple, small; outer sterile, very
short. Achenes obpyramidal, angled, rugulose,
with a denticulate apical rim. Insertion areole
two-lipped; lower lip incrassate, white; upper lip
with two white recurved costae. Pappus double,
outer of scabrid bristles, inner of (4–8) wider, very
long, subulate scales. x = 14. One species, C. nitens
(M. Bieb.) C.A. Mey., central and western Asia.
152. Oligochaeta (DC.) K. Koch
Oligochaeta (DC.) K. Koch, Linnaea 17: 42 (1843); Wagenitz,
Veröff. Geobot. Inst. ETH Stiftung Rübel Zürich 37: 315–329
(1962), rev.
Unarmed annual herbs, minutely glandular. Leaves
dentate or dentate-pinnatifid. Capitula solitary,
homogamous. Involucral bracts with a triangular,
Centaurothamnus Wagenitz & Dittrich, Candollea 37: 111
(1982).
Unarmed shrub. Leaves entire, lanceolate, whitelanose with marked veins beneath, tomentose or
glabrescent above when old. Capitula solitary,
terminal, heterogamous. Involucral bracts scarious, six-veined, with small, dark-brown, woolly
appendages. Receptacle setose. Florets rose, outer
sterile and radiant. Achenes linear-oblong, ridged,
glabrous. Pappus double, persistent. x = 14. One
species, C. maximus (Forssk.) Wagenitz & Dittrich,
Yemen.
155. Ochrocephala Dittrich
Ochrocephala Dittrich, Bot. Jahrb. Syst. 103: 467–480 (1983).
Tall shrub to 2 m high. Leaves elliptic or ovate, to
18 cm long and 8 cm wide, sparsely floccose. Capitula very large, 45 mm long and to 40 mm wide,
heterogamous, corymbose. Involucral bracts with
large scarious, whitish, entire or lacerate unarmed
appendages, innermost acuminate. Peripheral florets sterile, central florets perfect, with very short
144
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
corolla tube. Achenes oblong, costate. Pappus double, inner row with wider bristles. One species, O.
imatongensis (Philipson) Dittrich, eastern and central Africa, India.
III.5. c. Serratula Group
Unarmed perennial herbs or shrublets. Capitula
homogamous, very rarely heterogamous. Appendages of bracts rudimentary. Achenes usually
ridged or ribbed. Insertion areole almost basal,
small, without elaoisome. Hilum basal. Pappus
obscurely double, easily deciduous.
156. Serratula L.
basis of the stem. Capitula narrowly subcylindrical, terminal, homogamous. Involucral bracts
membranous with cartilaginous margin and a very
short black mucro. Receptacle subcylindrical,
convex, densely setose. Florets pink. Achenes
narrowly linear-oblong, 4–5-ribbed, glabrous,
with a small apical rim. Pappus double, deciduous;
outer of pinnulate bristles, inner of shorter apically
lacerate scales. One species, N. leptoclada (Bornm.
& Sint.) Iljin, Iran, central Asia. Note added in
proof: according to Martins and Hellwig, Taxon
54:633–638 (2005), Nikitinia should be placed in
Klasea.
Serratula L., Sp. Pl. 2: 816 (1753).
III.5. d. Carthamus Group
Unarmed perennial herbs. Leaves entire to
dentate-lobed or frequently pinnatifid-pinnatisect.
Capitula solitary or corymbose, homogamous.
Involucral bracts leaf-like, often reddish or pinkish, veined, with an appendage usually reduced to
a slender spine; inner bracts scarious, coloured.
Receptacle setose. Florets pink or purple. Achenes
linear-oblong, glabrous, faintly ribbed. Pappus
of pinnulate bristles in two rows; inner bristles
basally wider and as long as outer ones. x = 11.
Two to four species, Eurasia, rare in the south.
Carthamus group; López González, Anales Jard. Bot.
Madrid 47: 11–34 (1990), rev.; Vilatersana et al., Pl. Syst.
Evol. 221: 89–105 (2000), mol. phylog.
157. Klasea Cass.
Klasea Cass., Dict. Sci. Nat. 35: 173 (1825).
Schumeria Iljin (1960).
Unarmed perennial herbs. Leaves entire or serrate,
sometimes pinnatisect. Capitula homogamous.
Involucral bracts leaf-like, veined, with an appendage usually reduced to an apical slender
spine, very often reflexed, rarely muticous; rarely
appendages well developed, broadly scarious.
Florets pink, whitish or yellowish, with narrow
corolla lobes. Achenes linear-oblong, sulcate, often
apically rugulose between the ridges, glabrous.
Insertion areole almost basal. Pappus of barbellate or barbellate-plumose persistent bristles,
double; inner row raised above outer. x = 15.
Sixty-five species, Europe, Mediterranean region,
south-western Asia and northern Africa.
158. Nikitinia Iljin
Nikitinia Iljin, Bot. Mater Gerb. Bot. Inst. Komarova Akad.
Nauk S.S.S.R. 20: 356 (1960).
Unarmed, virgate, much-branched subshrub with
green branches. Leaves linear, congested at the
Annual or perennial herbs, rarely shrublets,
usually spiny. Leaves pinnatifid, rarely entire.
Capitula homogamous. Achenes compressed,
very hard, often angulose, glabrous, sometimes
dimorphic. Insertion areole very small, without
elaiosome. Hilum lateral. Pappus double, persistent, sometimes basally connate in a ring and
deciduous.
159. Carthamus L.
Carthamus L., Sp. Pl. 2: 830 (1753); Hanelt, Feddes Repert.
Spec. Nov. Regni Veg. 67: 41–180 (1963), rev.
Kentrophyllum Neck. ex DC. (1810).
Annual, spiny, rarely unarmed (C. tinctorius
L.) herbs. Leaves pinnatifid or pinnatisect, usually spiny. Capitula terminal, solitary or laxly
corymbose. Involucral bracts foliose, very spiny.
Innermost bracts usually without cucullate appendages. Florets yellow or pink, rarely orange.
Achenes obpyramidal, markedly dimorphic: outer
usually epappose, small, oblong-obconical, rugose;
inner pappose, large, oblong-obconical, almost
smooth. Pappus double, persistent, formed by two
rings of wide scales; inner ring shorter. x = 10,
11, 12, 32. Twenty species, Iran, Caucasus, eastern
Mediterranean region to central Asia.
Species of sect. Atractylis are colonizing weeds
very widely distributed in the Mediterranean
region and introduced to Australia, California,
and Chile. One species, C. tinctorius L. (safflower),
widely cultivated for oil, dye (substitute of saffron)
and as an ornamental.
Compositae
160. Phonus Hill
145
Phonus Hill, Veg. Syst. 4: 5 (1762).
Evol. 234: 15–26 (2002), mol. phylog., gener. delim.
Aegialophila Boiss. & Heldr. (1849).
Very spiny shrubs to 200 cm. Branches covered
by remains of leaves. Leaves spinose-dentate
or pinnatifid. Capitula terminal, solitary, large,
to 3 cm wide, many-flowered. Involucral bracts
foliose, green, spiny, innermost without cucullate
appendages. Florets yellow. Achenes obconical,
angulose, glabrous. Pericarp undifferentiated.
Pappus a single row of persistent bristles. x = 12.
Two species, southern Spain, northern Africa.
Annual or perennial herbs. Leaves interrupted
runcinate-pinnatisect, often fleshy, densely glandular. Capitula heterogamous. Involucral bracts
oval, with a very large entire or fimbriate-dentate
scarious appendage, sometimes spiny. Outer
florets sterile, large, radiant, with acheniodes;
inner perfect. All florets with many stalked glands,
especially on the tube. Achenes oblong-obconical,
densely sericeous; insertion areole very small,
161. Carduncellus Adans.
Carduncellus Adans., Fam. Pl. 2: 116 (1763).
Perennial or rarely annual herbs, often acaulescent. Leaves pinnatifid or pinnatisect, spiny, rarely
unarmed, glabrous or sparsely pubescent. Capitula solitary. Involucral bracts foliose, green, with
spiny fimbriate appendages, innermost with cucullate appendages. Florets usually pale or light
blue, very seldom yellowish. Achenes obconical,
angulose, apically with transversal ridges. Pappus
double, sometimes single by abortion of the outer
ring, rarely absent, deciduous or rarely persistent.
x = 12. Twenty-seven species, Iberian Peninsula
and northern Africa, one species widely distributed
in the Mediterranean region eastwards to Greece.
162. Femeniasia Susanna
Femeniasia Susanna, Collect. Bot. (Barcelona) 17: 83
(1987).
Spiny subshrub forming large pulvinules. Stems
and leaves fleshy when young. Leaves sheathing,
apically transformed into trifid spines. Capitula
small, to 1.5 cm wide, terminal. Involucral bracts
green, leaf-like, with a short spiny appendage;
innermost bracts with a rudimentary appendage.
Florets yellow. Achenes broadly obconical or
obscurely obpyramidal, very small, blackish,
glabrous. Pappus a single row of pinnulate bristles,
basally connate, deciduous as a single piece.
x = 12. One species, F. balearica (J.J. Rodr.)
Susanna, windy and rocky seashores of Menorca
(Balearic Islands).
III.5. e. Crocodylium Group
163. Crocodylium Vaill.
Fig. 35
Crocodylium Vaill., Königl. Akad. Wissensch. Paris Phys.
Abh. 5: 163 (1754) [“Crocodilium”]; Font et al., Pl. Syst.
Fig. 35. Compositae-Cardueae. Crocodylium syriacum. A
Habit. B Floret. C Achene. Drawing by E. Macpherson
146
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
sublateral. Pappus double, outer bristles with
pinnulae longer than width of bristle, inner row
shorter with apically lacerate bristles. x = 11.
Three species, coastal sands, Greece, Aegean
Islands, Turkey, Egypt and Middle East.
III.5. f. Centaurea Group
164. Centaurea L.
Fig. 36
Centaurea L., Sp. Pl. 2: 909 (1753) pro maxima parte, nom.
cons.; Susanna et al. (1995), mol. phylog.; Wagenitz &
Hellwig, in Proceedings of the International Compositae
Conference, Kew, 1994, Royal Botanic Gardens, Kew:
491–510 (1996), evol.; Garcia-Jacas et al., Pl. Syst. Evol.
185–199 (2000), mol. phylog.
Cnicus L. (1753), nom. cons.
Cyanus Mill. (1754).
Jacea Mill. (1754).
Colymbada Hill (1762).
Acosta Adans. (1763).
Calcitrapa Adans. (1763).
Seridia Juss. (1779).
Melanoloma Cass. (1823).
Chartolepis Cass. (1826).
Tomanthea DC. (1837).
Ptosimopappus Boiss. (1845).
Phaeopappus Boiss. (1846).
Hyalaea Jaub. & Spach (1847).
Cheirolepis Boiss. (1849).
Stephanochilus Coss. & Durieu ex Benth. (1873).
Chrysopappus Takht. (1938).
Grossheimia Sosn. & Takht. (1945).
Wagenitzia Dostál (1973).
Annual, biennial or perennial herbs or shrubs,
usually unarmed. Capitula heterogamous, rarely
homogamous. Involucral bracts scarious, rarely
leaf-like, with a variable apical appendage, spiny or
unarmed, membranous, appendage rarely absent.
Florets blue, pink, purple, orange or yellow, rarely
white. Sterile florets radiant and showy, sometimes
very long, rarely reduced. Achenes oblong, compressed, rarely obconical (sect. Stephanochilus),
brown-blackish, smooth, very rarely ridged
(sect. Cnicus and Stephanochilus), subglabrous or
sparsely pilose. Hilum lateral. Insertion areole with
elaiosome. Pappus double; outer bristles pinnulate
or rarely plumose, inner much shorter, epinnulate,
lacerate above; rarely pappus of pinnulate stiff
setae (sect. Cnicus ). x = 7, 8, 9, 10, 11, 12. Circa
250 species, Eurasia, especially Irano-Turanian
and Mediterranean region.
Fig. 36.
Compositae-Cardueae. Centaurea benedicta.
A Habit. B Achene. (Font Quer 1962)
Some species cultivated as ornamentals (C.
montana L., C. ragusina L.). Centaurea cyanus L.
is a cosmopolitan weed, associated with cereal
cultivation; other species naturalized as noxious
weeds in Australia and North America (C. diffusa
Lam., C. diluta Aiton, C. maculosa Lam., C.
melitensis L. and C. solstitialis L.).
Carduoid Genera
of Uncertain Placement
C. Jeffrey
165. Cavea W.W. Sm. & Small
Cavea W.W. Sm. & Small, Trans. Bot. Soc. Edinburgh 27: 119
(1917); Ling & Chen, Acta Phytotax. Sin. 10: 92–102, cum ic.
(1965), morph., distrib.
Perennial, sometimes dioecious herb. Leaves alternate, ovate-lanceolate, pubescent. Capitula solitary,
Compositae
terminal, large, heterogamous and disciform or homogamous and unisexual. Phyllaries few-seriate,
imbricate, herbaceous. Receptacle plane, epaleate.
Florets all regular; corollas hairy with robust, acute,
multicellular hairs, those of pistillate (outer in heterogamous capitula) 3–4-lobed; styles of pistillate
florets without articulation, arms spathulate, with
marginal, apically confluent stigmatic areas; corollas of functionally staminate (inner in heterogamous capitula) larger, limb campanulate, deeply
5-lobed; anthers shortly calcarate, ecaudate; anther collar poorly developed; endothecial cell wall
thickenings inconspicuous or absent; style undivided. Achenes oblong, terete or subquadrangular,
densely hirtellous with elongate, multicellular twin
hairs. Pappus of numerous, uniseriate, purple, barbellate bristles, of vestigial achenes of staminate
florets shorter, with apically slightly expanded bristles. One species, C. tanguensis (J.R. Drumm.) W.W.
Sm. & Small, north-eastern India, south-western
China.
In spite of its strikingly carduoid facies (the
species was originally placed in Saussurea), Cavea
may prove to belong elsewhere in Compositae, most
likely in or near Inuleae; palynological, carpological and molecular data are lacking.
166. Dipterocome Fisch. & Mey.
Dipterocome Fisch. & Mey., Ind. Sem. Hort. Petrop. 1: 26
(1835); Praglowski & Grafstrom, Bot. Notiser 133: 177–188
(1980), palynol.; Reese, Bot. Jahrb. Syst. 110: 325–419
(1989), carpol.
Annual herb, branches prostrate. Leaves alternate,
linear, entire, glabrous. Capitula small, sessile,
axillary or congested at stem base, few-flowered,
heterogamous. Involucre ovoid; phyllaries few,
oblong, with hyaline margins, outer smaller.
Receptacle epaleate. Marginal florets pistillate,
sub-2-seriate; corollas bilabiato-radiate, adaxial
lip minute. Inner florets functionally staminate;
corolla slender, shortly 5-lobed; filaments connate;
anthers ecalcarate, ecaudate; pollen spinulose,
ecaveate, with infratectal bacula well developed;
style undivided. Achenes terete, outcurving,
dorsally spiny, apically bicornute, sometimes
dimorphic, with outer less curved and apically
echinate. Pappus of a few, flattened, scabrid bristles.
One species, D. pusilla Fisch. & Mey., Palestine to
Afghanistan.
The carduoid pollen and achene features are
probably plesiomorphic and this strange genus
may, like Gymnarrhena, lie at or below the base of
147
Cichorioideae as a relic of a distinct line. Molecular
data are lacking.
IV. Tribe Gymnarrheneae
Panero & V.A. Funk (2002).
C. Jeffrey
Perennial rosulate, acaulescent, amphicarpic
herbs. Leaves forming a dense rosette, sessile,
lamina narrowly lanceolate to narrowly ovate,
denticulate, acute to attenuate. Capitula of two
kinds, subterranean and subaerial. Subterranean
capitula homogamous, pistillate, cleistogamous;
florets enclosed in the involucral bracts; corolla
vestigial. Subaerial capitula (apparently syncalathia) congested in the centre of the leaf
rosette, heterogamous, disciform. Involucral
bracts imbricate in several series, chartaceous,
whitish, acute; receptacle convex, marginally
bristly; functionally staminate florets in small
groups, loosely connected on very short pedicels,
interspersed among the pistillate florets; corollas
small, 3–4-lobed, whitish; stamens 3–4, anthers
calcarate, ecaudate, without apical appendage;
pollen spiny, ecaveate, with massive foot layer,
thick columellae and fine net above the columellae;
style undivided, truncate, with obtuse hairs apically; ovary vestigial. Pistillate florets solitary, each
enclosed in a prominent, stiff, white and green
bract; corolla filiform, basally suberized with age;
style arms long, with rounded apices. Achenes of
subterranean capitula laterally flattened, blackish,
sparsely hairy, pappus absent or of short, basally
flattened, somewhat scale-like bristles. Achenes
of pistillate florets of subaerial capitula ovoid,
ciliate, villous with long twin hairs; pappus of
long-lanceolate, ciliate, acutely acuminate scales.
Achenes of functionally staminate florets vestigial,
pappus of a few irregularly lacerate scales or
absent. Monotypic.
167. Gymnarrhena Desf.
Gymnarrhena Desf., Mem. Mus. Hist. Nat. Paris 4: 1, t. 1
(1818).
Characters of the tribe. n = 10. One species, G.
micrantha Desf., North Africa, Middle East.
Gymnarrhena has usually been included, although sometimes with reservation, in Inuleae.
Skvarla et al. (1977) pointed out that the pollen
grain structure did not support such a placement
148
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
and noted some similarity in wall stratification to
certain Cynareae. Bremer (1994) included it in Cichorioideae, without tribal assignment. Panero and
Funk (2002), in establishing the tribe Gymnarrheneae, referred it to a new, monotypic subfamily
Gymnarrhenoideae because in their cladograms,
based on comparative DNA sequence data of several chloroplast DNA genes and markers, totalling
some 13.380 bp, Gymnarrhena was consistently located by itself below the other cichorioid tribes and
above the pertyoid and carduoid clades.
V. Tribe Moquinieae H. Rob. (1994).
Subtribe Pseudostifftiinae H. Rob.,
R.M. King & F. Bohlmann (1980).
H. Robinson
Monoecious or gynodioecous, moderately
branched shrubs or trees; stems not fistulose.
Leaves alternate, shortly petiolate, coriaceous,
obovate, cuneate, entire, venation pinnate, with
glandular dots. Inflorescence terminal, pyramidally thyrsoid; peduncles 1–3 mm long. Heads
homogamous; involucre narrowly campanulate,
bracts in 4–5 series, gradate, inner bracts deciduous; receptacle epaleaceous. Florets 1–5 per head;
corollas regular, lavender to purple, narrowly
funnelform, glanduliferous, undivided limb short,
lobes 5, linear, smooth; anthers calcarate, shortly
tailed; endothecial cells with broad vertical thickened band usually narrowed at each end to one
point; apical appendages 3–4 times as long as wide,
with cell walls not thickened, anthers aborted in
functionally female florets. Pollen spherical, tricolporate, echinate, non-caveate, with strong solid
bacula randomly distributed in areas not directly
under spines, not grouped or single under spines
(Fig. 37). Style base broadly abruptly noduliferous;
upper part of style becoming broadened, upper
part of shaft and outer surface of arms scabrid,
inner surfaces of branches totally stigmatic.
Achenes densely setuliferous, 10–17-costate,
idioblasts obvious or obscure, raphids obscure,
phytomelanin lacking; carpopodia annuliform
to stopper-shaped, with subquadrate cells in
3–17 series, the walls thickened; pappus of many
capillary bristles, in c. 2 series, outer irregularly
somewhat shorter. The presence of triterpenoides
and guaianolides has been reported (Bohlmann
et al. 1982; Bohlmann and Jakupovic 1990).
Fig. 37.
Compositae-Moquinieae. SEM micrographs
of pollen grains. Pseudostifftia kingii. A Non-lophate
tricolporate grain, colpar view, scale bar = 10 µm. B Broken grain showing solid bacula positioned irregularly in
relation to spines, scale bar = 5 µm
Two genera and two species in Brazil.
The tribe consists of two genera originally
placed in other tribes – Moquinia in Mutisieae
(Cabrera 1977) and Pseudostifftia in Vernonieae
(Robinson 1979). The two were placed together first
by Gamerro (1990) in Vernonieae, and a separate
tribe was established by Robinson (1994).
Moquinieae differ from Mutiseae s.str. by the
smaller and thinner anther appendage and the
spinose pollen with simple tectum in the former.
Relationship is not considered close. Moquineae
are apparently close to Vernonieae, but differ by
their thickened scabrid styles similar to those of
Arctotideae, rather than these being thin with long
sweeping hairs as found in Vernonieae. The pollen
also differs by the random positions of the bacula,
not directly under the spines as in Vernonieae and
Liabeae.
Key to the Genera
1. Inflorescence with strongly racemiform or spiciform
branches; heads with 4 or 5 florets; pappus yellow;
stems and leaf undersides whitish or pale yellowishtomentose; plants sometimes gynodioecious
168. Moquinia
– Inflorescence with corymbiform branches; heads with
1 floret; pappus white; stems and leaf undersides usually yellowish-lepidote; plants monoecious
169. Pseudostifftia
168. Moquinia DC.
Fig. 38
Moquinia DC., Prodr. 7: 22 (1837), nom. cons., non Spreng.
(1828).
Spadonia Less., Syn. Comp. 99 (1832), non Fr. (1829).
Gynodioecious; stem hairs white, arachnoid. Leaf
lower surface whitish or pale yellowish-tomentose.
Inflorescences with densely racemiform branches.
Involucral bracts c. 25, c. 4–5-seriate. Florets 4 or 5.
Compositae
Achene idioblasts elongate, sometimes in series;
carpopodium stopper-shaped, cells in 13–17 series;
pappus bristles c. 60, yellowish, with some tips
broadened. One species., M. racemosa (Spreng.)
DC., Bahia and Minas Gerais, Brazil.
169. Pseudostifftia H. Rob.
Pseudostifftia H. Rob., Phytologia 44: 444 (1979).
Monoecious; stem and leaf hairs appressed, symmetrically T-shaped, cap-cells broadly fusiform,
stalks slender. Inflorescences with corymbiform
branches. Involucral bracts c.18, c. 5-seriate. Floret 1. Achene setulae with some intermixed uniseriate hairs, idioblasts obscure; carpopodium an-
Fig. 38. Compositae-Moquinieae. Moquinia racemosa.
A Habit showing blunt-tipped, abaxially tomentose,
alternate leaves. B Head; note homogamous condition with
regular corollas. C Corolla showing tips of anthers and
style; note deeply divided lobes. D Corolla in long section,
one lobe removed, showing anthers and style. E Style,
showing basal node, swollen upper shaft, showing scabrid
surface consisting of short sweeping hairs on upper shaft
and backs of lobes, and undivided stigmatic surface inside
of branches. F Anther showing long calcarate and shortly
tailed bases, and apical appendage. G Achene with setulae
and capillary pappus. (Drawing by A. Tangerini)
149
nuliform, cells in 5 or fewer series; pappus bristles
c. 100, white, tips not broadened. One species, P.
kingii H. Rob., Bahia, Brazil.
VI. Tribe Vernonieae Cass. (1819).
H. Robinson
Annual or perennial herbs, subshrubs, shrubs,
scandent shrubs or trees; with simple, T-shaped
or stellate hairs. Leaves usually alternate, rarely
opposite or ternate; blades usually undivided.
Inflorescences cymes, corymbiform with cymose
branches, or sometimes spicate; heads homogamous, sessile or pedunculate, with or without
foliose bracts at base of involucre; involucral bracts
in 3–9 series, usually gradate, rarely subequal or
decussate; receptacle usually glabrous, sometimes
with pales, spines or partitions. Florets 1–400
in a head, perfect; corollas mostly funnelform
with tubes usually broadened above well below
filament insertion, less often narrow up to filament
insertion, limbs usually actinomorphic with lobes
longer than wide, rarely zygomorphic with some
lobes longer, lobes usually erect; anther thecae usually calcarate, often tailed, apical appendage flat,
thin to somewhat stiffened, with or without glands.
Pollen (Fig. 39) spherical; commonly tricolporate
and echinate with perforated tectum continuous
between colpi, a form obviously derived from
lophate forms in at least some subtribes. Many
forms are lophate with well-developed muri and
variously lacking perforated tectum in lacunae or
sometimes over whole grain. Some are lophate with
walls crossing colpi or are completely triporate.
Polar lacunae are sometimes irregular with lines
of colpi completely obscured. Lophate grains are
usually echinate but sometimes psilate. Bacula are
solid and usually stout, directly under the spines,
but are sometimes weak and easily detached from
foot-layer. Details of lophate forms are particularly
useful in classification. Nectary usually glabrous.
Style base with or without sclerified or expanded
node, glabrous, upper shaft and outer surfaces
of branches with sweeping hairs, hairs with or
without septae, branches spreading tangentially,
stigmatic papillae covering whole inner surface
of branches. Achenes usually prismatic, rarely
angled, 3–20-costate, rarely obcompressed in
outer row, sometimes inner achenes differing in
setulae or pappus, walls usually with resiniferous
idioblasts on surface or raphids internally, rarely
150
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Fig. 39. Compositae-Vernonieae. SEM micrographs of
pollen grains. A Vernonanthura fuertesii. Non-lophate,
colpar view. B Lepidaploa salzmannii. Echinolophate,
tricolporate with polar lacuna, polar view. C Phyllocephalum scabridum. Lophate, triporate, emicropunctate.
D Blanchetia heterotricha. Broken grain showing solid
bacula under spine. Scale bars: A, C = 10 µm, B = 13.6 µm,
D = 2 µm. (A, C, D Robinson and Marticorena 1986;
B Robinson 1999a)
with phytomelanin; carpopodium stopper-shaped
to turbinate, rarely obsolete; pappus usually of
long capillary bristles, usually with outer series of
shorter bristles or squamellae, rarely coroniform,
squamellose without bristles, with flattened or
twisted segments, or lacking. Flavones, flavonols,
sesquiterpene lactones, nerolidol derivatives,
5-alkylcoumarins, and potentially commercially
useful epoxy oils in the seeds have been reported.
Sesquiterpenes include glaucolides, hirsutinolides,
furoheliangolides and elemanolides (Bohlmann
and Jakupovic 1990).
The tribe contains 118 genera and more than
1,000 species, mostly in tropical parts of the world.
Vernonieae are part of subfamily Cichorioideae in the more restricted sense, with closest
relationship to Arctotideae, Cichorieae, Liabeae
and Moquinieae, all with usually calcarate anther
bases and stigmatic papillae across the whole
inner surface of the style branches. The tribe
differs from Arctotideae and Moquinieae by the
narrow styles and long sweeping hairs, from
Cichorieae by the usual lack of milky sap and
the usually actinomorphic corollas, from Liabeae
and Arctotideae-Eremothamneae (included in
Arctotideae in this volume) by the usually bluish
or reddish flowers and the lack of ray florets,
and from Liabeae further by the usually alternate
leaves, the solid bacula in the pollen and the
often lophate pollen. Moquinieae differ further
by the pollen bacula not being directly under
single spines. Distinction of Vernonieae pollen
from that of Liabeae and a limited SEM survey of
Vernonieae pollen types can be seen in Robinson
and Marticorena (1986) and Robinson (1999b).
Vernonieae are notable for the frequent extreme
cymose forms of their inflorescences, involving
seriate or scorpioid cymes where each head is
produced on a lateral branch below the preceding
head. As such, the heads often look sessile and
lateral in a straight or arching series. The tribe
is more interesting for the truly spicate form
of inflorescence in Pithecoseris and a species of
Chresta. The latter is a departure from the basically
cymose form found in all other Asteraceae.
Vernonieae were one of the original tribes named
by Cassini (1819), and they have been consistently
recognized in subsequent treatments (Bentham
1873a; Hoffmann 1894; Cronquist 1955; Jones
1977; Bremer 1994). Until Robinson and Brettell
(1973c) and Robinson (1977), Vernonieae were
usually treated as a close relative of Eupatorieae
because of the homogamous rayless heads with
bluish to reddish florets, but the latter tribe is
a member of subfamily Asteroideae, as detailed
in Robinson (1977). The tribe Vernonieae has
had a few problems in delimitation, but it is now
known to include Stokesia with its liguliform
corollas (Jones 1977), Distephanus with its yellow
corollas and trinervate leaves (Robinson and Kahn
1986), and the Hawaiian, mostly apomictic Hesperomannia (Kim et al. 1998). Now excluded are
two South African genera – Hoplophyllum, related
to Eremothamnus and a member of Arctotideae
(Karis 1992; Robinson 1992), and Corymbium
(Bohlmann and Jakupovic 1990) which is treated
as a separate tribe in this volume. Internally,
Vernonieae have suffered from an excessively
paraphyletic core-genus concept, with Vernonia
defined only by what it is not. Recent efforts to correct the generic concepts are included in studies by
Robinson (1996, 1999b) mostly for the Americas,
and Robinson (1999a) for the palaeotropics.
At present, the majority of the American
subtribes, including subtribes Vernoniinae,
Centratherinae, Chrestinae, Leiboldiinae, Lychnophorinae, Piptocarphinae, Sipolisiinae and
Stokesiinae, are considered as comparatively
closely related. The single species in Pacourininae
is individually distinctive but probably also closely
Compositae
related. The pantropical Elephantopinae seem
closest to the tropical American Rolandrinae. In
the western hemisphere, only subtribe Trichospirinae is apparently of remote relationship. In
the palaeotropics, subtribe Gymnantheminae has
many morphological parallels with the American
Piptocarphinae, but has chromosome numbers and
some chemistry in common with the other eastern
hemisphere subtribes, Erlangeinae and Centrapalinae. The basically palaeotropical Erlangeinae
are particularly distinct in the triporate pollen
and the presence of 5-alkyl coumarins. The hemispheric geographical separation in the tribe is not
complete, with the distinctive Manyonia of Vernoniinae found in Africa, and the equally distinctive
Acilepidopsis, Mesanthophora and Telmatophila
apparently of Erlangeinae found in South America.
Many taxomonic treatments exist for members
of Vernonieae in various parts of the world, but
most of these do not follow recent generic limits.
Many of these treatments are listed in Robinson
(1999a, b). An index to American species of Vernonieae, giving present dispositions, is included in
Robinson (1999b).
Key to the Subtribes
and Unplaced Genera
1. Achenes flattened with pair of widely divergent cornute projections at top
15. Trichospirinae (p. 174)
– Achenes without a pair of large cornute projections at
top
2
2. Corollas liguliform, strongly zygomorphic with deepest sinus towards centre of head
3
– Corollas not or scarcely zygomorphic, without single
deepest sinus centred towards centre of head
4
3. Heads separate, on long peduncles
10. Stokesiinae (p. 165)
– Heads in compound clusters
8. Elephantopinae (p. 163)
4. Heads compound or sessile in compact glomerules or
spikes, not in obvious cymes
5
– Heads separate or on obvious cymose branches
12
5. Florets 4 in each head
6
– Florets not consistently 4 in each head
8
6. Leaves with veins sublongitudinal; heads in axils of
broad, imbricated upper leaves
4. Chrestinae (Soaresia)
– Leaves with veins not sublongitudinal; heads
subtended by bracts forming secondary involucre 7
7. Pappus biseriate, of bristles and outer broad squamae;
achene raphids subquadrate
8. Elephantopinae (Caatinganthus)
– Pappus of a single series of rudimentary awns; achene
raphids elongate
12. Erlangeinae (Telmatophila)
8. Branches of inflorescence often with decurrent wings
171. Gorceixia
– Branches of inflorescence not winged
9
151
9. African; pappus cupuliform
12. Erlangeinae (Muschleria)
– South American; pappus with bristles, straps, or scales
10
10. Florets 1 in each head of cluster; pappus a short crown
or short scales; pollen lophate 9. Rolandrinae (p. 164)
– Florets usually more than 1 in each head; pappus of
elongate segments; pollen lophate or non-lophate 11
11. Pollen usually lophate; raphids often elongate; inflorescences sometimes narrowly spicate with strongly
acropetal maturation
4. Chrestinae (p. 160)
– Pollen non-lophate; raphids always subquadrate; inflorescence not spicate or rarely spicate with large heads
6. Lychnophorinae (p. 161)
12. Tubes of corollas long and covered with obvious stipitate glands
13
– Corollas without long tubes covered with stipitate
glands
14
13. Leaves pinnately divided into linear segments; heads
not surrounded at base by many spreading foliose
bracts
12. Erlangeinae (Rastrophyllum)
– Leaves not pinnately divided into linear segments;
heads surrounded at base by many spreading foliose
bracts
5. Centratherinae (p. 160)
14. Inflorescence never seriately nor scorpioid cymose,
heads not borne secundly nor sessile in series of leaf
axils; anthers never with glands; mostly palaeotropical
or adventive in America
15
– Inflorescence variable, sometimes seriately cymose or
scorpioid or heads borne sessile in series of leaf axils;
anthers sometimes with glands; almost all American
17
15. Trees or shrubs; inner involucral bracts sometimes
deciduous; pollen never triporate
14. Gymnantheminae (p. 173)
– Herbs, rarely weak trees or shrubs; involucral bracts
persistent; pollen sometimes triporate
16
16. Coarse, usually perennial herbs or subshrubs; anther
appendages with somewhat thickened cell walls; hairs
of stems simple or unevenly T-shaped; achenes c. 10costate; 5-alkylcoumarins not present
13. Centrapalinae (p. 171)
– Annual or perennial herbs or weak shrubs or trees;
anther appendages thin and usually transparent; hairs
of stems simple to symmetrically or asymmetrically Tshaped; pollen triporate to tricolporate; achenes 4–6or 8–12-costate; 5-alkylcoumarins sometimes present
12. Erlangeinae (p. 165)
17. Large emergent aquatic herbs; pollen triporate without organized patterns of polar lacunae
11. Pacourininae (p. 165)
– Not large aquatic herbs; pollen tricolporate or with
polar lacunae in regular pattern
18
18. Receptacles with well-developed pales or spines 19
– Receptacles without obvious pales or spines
20
19. Hairs simple; achenes without phytomelanin in the
walls; Mexican, Central American
1. Leiboldiinae (p. 152)
– Hairs stellate or stellate at base, or T-shaped; achenes
usually with phytomelanin in the walls; Brazil
7. Sipolisiinae (p. 162)
20. Pappus bristles not sclerified at base, easily deciduous;
heads with more than 100 florets
1. Leiboldiinae (p. 152)
152
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
– Pappus bristles or elements sclerified at base, sometimes fragile; heads usually with 1–75 florets
21
21. Inner involucral bracts deciduous; hairs of stems stellate or lepidote; shrubs, scandent shrubs or trees 22
– Inner involucral bracts persistent; hairs of stems simple or T-shaped; often herbaceous
23
22. Leafy branches often ericoid or with leaf bases imbricated; leaves alternate; heads with 1–75 florets
6. Lychnophorinae (p. 161)
– Leafy branches not ericoid, leaf bases not imbricated;
leaves alternate or opposite; heads with usually fewer
than 20 florets
3. Piptocarphinae (p. 158)
23. Apical anther appendages with thin-walled cells, appendages or connectives sometimes with glands or
small hairs; pollen lophate or non-lophate
2. Vernoniinae (p. 153)
– Apical anther appendages with cell walls thickened
at least near margin, without glands or hairs; pollen
non-lophate
24
24. Involucral bracts spiniform; pappus of laciniate
scales, bearing glands; raphids of achene wall
elongate
170. Acanthodesmos (p. 152)
– Involucral bracts not spiniform; pappus usually of
bristles or scales, without glands; raphids of achene
wall subquadrate
6. Lychnophorinae (p. 161)
Unplaced Genera
170. Acanthodesmos C.D. Adams & M.C. du
Quesnay
Acanthodesmos C.D. Adams & M.C. du Quesnay, Phytologia
21: 405 (1971).
VI.1. Subtribe Leiboldiinae H. Rob. (1999).
Herbs or shrubs. Receptacles sometimes paleaceous. Anther appendages sclerified, glabrous.
Pappus bristles usually not sclerified at base,
easily deciduous. Pollen tricolporate, non-lophate.
Characteristic sesquiterpenes: glaucolides.
Key to the Genera
1. Pappus with a single row of long white bristles; style
with a distinct basal node; achene with scattered idioblasts
2
– Pappus with more than one row of short coloured
deciduous bristles; style base without node; achenes
closely covered by idioblasts
3
2. Receptacle with pales; achenes without apical annulus
inside pappus bristles
172. Bolanosa
– Receptacle without pales; achenes with raised apical
annulus inside unsclerified bases of pappus bristles
173. Leiboldia
3. Achenes without raised apical annulus inside rows of
pappus bristles; involucral bracts with often broadened and ornate tips
174. Lepidonia
– Achenes with raised apical annulus inside rows of pappus bristles; involucral bracts with narrow tips
175. Stramentopappus
Genera of Leiboldiinae
172. Bolanosa A. Gray
Bolanosa A. Gray, Smithsonian Contr. Knowl. 3: 82 (1852).
Subshrubs, branchlets distichous, hairs simple.
Vestigial spine-like bracts borne opposite lower
leaves, leaves white-tomentose below. Heads sessile
opposite upper leaves; involucral bracts 15–16, tips
mostly setiform; receptacles paleaceous; florets
12–13; anther appendages sclerified; style base
conical; sweeping hairs blunt, mostly septate. Achene 10-costate, raphids elongate; pappus laciniate,
glanduliferous. Pollen non-lophate. One species, A.
distichus C.D. Adams & M.C. du Quesnay, Jamaica.
Perennial rhizomatous herbs; white tomentum of
simple hairs. Inflorescences corymbiform. Involucral bracts c. 40, in c. 3 series, outer tomentose, foliiform, inner scarious, reddish-tipped; receptacular
pales sheathing. Florets c. 45; anther bases rounded;
style with node; sweeping hairs pointed. Achenes
6–7-costate, setuliferous, many idioblasts, raphids
elongate; pappus of broad bristles. One species, B.
coulteri A. Gray, Mexico.
171. Gorceixia Baker
173. Leiboldia Schltdl ex Gleason
Gorceixia Baker, J. Bot. 20: 225 (1882).
Leiboldia Schltdl ex Gleason, Bull. New York Bot. Gard. 4:
161 (1906).
Leiboldia Schltdl (1847), nom. nud.
Vernonia sect. Leiboldia (Schltdl) Benth. (1873).
Trees; stems partially winged; hairs stellate. Inflorescence corymbiform, of secondary heads containing many sessile heads; secondary involucre
of canescent bracts; involucre cylindrical, of 5 or
6 lanceolate subequal bracts. Florets 5; corollas
glabrous; anther tails small; style without node;
sweeping hairs sharp. Achenes tetragonous, glabrous, raphids elongate; pappus a collar. Pollen
non-lophate. One species, G. decurrens Baker, eastern Brazil.
Shrubs; hairs simple, contorted. Heads mixed with
short bracts, pedunculate; involucral bracts c. 100
in c. 6 series. Florets 100–120; anthers broadly
tailed; style with node; sweeping hairs obtuse,
often septate. Achenes 4–5-angled, idioblasts
sparse, raphids short-oblong; pappus whitish, with
callose ring, single capillary series not sclerified
Compositae
at base, easily deciduous. n = 19. One species, L.
serrata (D. Don) Gleason, Mexico.
174. Lepidonia S.F. Blake
Lepidonia S.F. Blake, J. Wash. Acad. Sci. 26: 454 (1936);
Turner, Brittonia 33: 401–412 (1981), rev.; Robinson & Funk,
Bot. Jahrb. Syst. 108: 212–228 (1987), emend.
Shrubs; hairs simple, multiseptate. Heads single or
few in axils of leaves or terminal. Involucral bracts
c. 100, tips with appendages; receptacle paleate
or glabrous. Florets c. 100; anther bases rounded;
style without node; sweeping hairs sharp, septate.
Achenes 4–5-ribbed, idioblasts numerous, raphids
subquadrate; pappus yellowish, short, multiseriate, unsclerified at base, deciduous. n = 19 . Seven
species, Mexico, Central America.
175. Stramentopappus H. Rob. & V.A. Funk
Stramentopappus H. Rob. & V.A. Funk, Bot. Jahrb. Syst. 108:
227 (1987).
Shrubs; hairs simple, multiseptate. Heads few, pedunculate, overtopped by foliose branches; involucral bracts 100–130, in c. 8 series. Florets c. 110;
anthers without tails; style without node; sweeping hairs sharp, often septate. Achenes 5-angled,
idioblasts dense, raphids subquadrate, apex with
callose ring; pappus multiseriate, of short, yellowish, deciduous bristles, unsclerified at base. n = 19.
One species, S. pooleae (B.L. Turner) H. Rob. & V.A.
Funk, Mexico (Oaxaca).
VI.2. Subtribe Vernoniinae Cass. ex Dumort.
(1829).
Herbs, weak shrubs, or vines. Inflorescence often seriate- or scorpioid-cymose; inner involucral
bracts usually persistent. Anther appendages thinwalled, often with glands or hairs. Pollen mostly
tricolporate. Characteristic sesquiterpenes: glaucolides.
Key to the Genera
1. Peripheral achenes broadly obcompressed, winged on
lateral margins
2
– Achenes not broadly obcompressed nor laterally
winged
3
2. Heads sessile in dense scorpioid or seriate cymes; involucral bracts with projecting keel or wing outside;
pollen not lophate; corollas zygomorphic
182. Dipterocypsela
153
– Heads pedunculate; involucral bracts acuminate, without a winged keel outside; pollen lophate; corollas actinomorphic
186. Heterocypsela
3. Pappus lacking or reduced to a cartilaginous sleeve 4
– Pappus with bristles, straps, or awns
5
4. Shrubs; corollas with 5 lobes; pappus lacking; pollen
tricolporate, with three intercolpar-aligned polar lacunae
185. Harleya
– Annuals; corollas with 3 or 4 lobes; pappus a cartilaginous sleeve; pollen triporate, with single polar lacuna
195. Sparganophoros
5. Pappus of long broadened straps or awns
6
– Pappus with capillary bristles
7
6. Pappus of subulate awns; anther bases long-calcarate;
not an aquatic herb
194. Stilpnopappus
– Pappus of canaliculate straps; anther without long
calcarate bases, with basal spurs shorter than collar;
aquatic herb
199. Xiphochaeta
7. Receptacle deeply pitted, pits enclosing complete bodies of achenes
176. Albertinia
– Receptacles plain to weakly alveolate, not deeply pitted
8
8. Inflorescences densely scorpioid-cymose, with apices
usually curved
9
– Inflorescences not truly scorpioid with scorpioid tips,
only seriate-cymose to thyrsoid or corymbiform 10
9. Petioles narrowly inserted on stem; older heads deciduous leaving only subtending bracteoles; pappus
bristles not clavate distally; anther thecae without sclerified tails; achenes with bulging idioblasts on surface
180. Cyrtocymura
– Petioles usually broadly winged to base, broadly inserted; older heads not deciduous, involucral bracts
persistent; pappus bristles broadened near tips; anther
thecae with sclerified tails; achenes without differentiated idioblasts on surface
184. Eirmocephala
10. Plants African; inflorescence seriate-cymose; pollen
non-lophate; achenes strongly 5-costate, with very
densely disposed subquadrate raphids
189. Manyonia
– Plants American; inflorescence variable, simple to
complex cymes, thyrsoid, seriate-cymose or with
solitary heads; pollen lophate or non-lophate; achenes
weakly 5- or 8–10-costate, with subquadrate to
elongate raphids mostly in one layer
11
11. Pollen echinate, never lophate; inflorescence a simple
cyme or corymbiform, never strongly seriate-cymose
12
– Pollen usually lophate; inflorescence often seriatecymose or heads in series of leaf axils, sometimes
individually pedunculate
17
12. Inflorescence a single spreading cyme; base of plant
decumbent, not xylopodial
198. Vernonia
– Inflorescence thyrsoid or more complex, with cymose
or corymbiform branches; plant bases erect or xylopodial, not decumbent
13
13. Scandent, inflorescence pyramidally thyrsoid, with
primary branching at c. 90° angles; secondary
branches often racemiform
196. Trepadonia
– Erect to subscandent plants; inflorescences thyrsoid
with cymose branches, secondary branches not noticeably racemiform
14
14. Corolla lobes much shorter than throat 179. Cololobus
– Corolla lobes as long as or longer than throat
15
154
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
15. Corollas without hairs inside limb; plants not or rarely
subscandent; raphids of achene subquadrate
197. Vernonanthura
– Corollas often with hairs inside limb; plants sometimes subscandent; raphids of achenes elongate
16
16. Corollas with whole inside of limb covered with short
one-celled hairs
203. Dasyanthina
– Corollas often with long, slender, multicellular hairs
inside of throat, no hairs on inside of lobes
192. Quechualia
17. Corollas slightly zygomorphic with longest lobe centred towards centre of head; basal nodes of inflorescence often with 2 or 3 branches; anther thecae with
tails
190. Mattfeldanthus
– Corollas actinomorphic; basal nodes bearing heads
with at most one branch; anther thecae without tails
18
18. Heads with 4–10 florets
19
– Heads with 10–70 florets
20
19. Hairs of stems T-shaped; pollen with three intercolparaligned polar lacunae
191. Pseudopiptocarpha
– Hairs of stems simple, often arachnoid; pollen with
single lacuna at pole
193. Stenocephalum
20. Pollen with colpi reaching poles and with 3 distinct
equatorial lacunae across intercolpi; heads often large,
mostly 1 cm or more long; style base usually without
basal node
188. Lessingianthus
– Pollen with or without polar lacunae, intercolpi with
only two distinct equatorial lacunae; heads of various
sizes, usually less than 1 cm long; style base with or
without basal node
21
21. Stems, leaves, peduncles, and involucres loosely
sericeous with yellowish hairs
22
– Plants with pubescence variable, never loosely
sericeous with yellowish hairs, sometimes stems,
leaves, peduncles or involucres sericeous with white
hairs
23
22. Anther appendages usually with numerous glands;
pollen with muri weakly attached to foot-layer, sometimes easily detached; heads sessile except at ends of
seriate-cymose branches
178. Chrysolaena
– Anthers without glands; pollen with muri strongly attached to foot-layer by stout baccula; heads often solitary or individually short-pedunculate in clusters
188. Lessingianthus subg. Oligocephalus
23. Heads usually sessile except at ends of cymose
branches
187. Lepidaploa
– Heads mostly long-pedunculate
24
24. Heads with subinvolucre of foliose bracts; leaves
4–6 cm wide; pollen with 3 intercolpar-aligned lacunae meeting at pole; raphids of achene subquadrate
177. Aynia
– Heads without subinvolucre of foliose bracts, with only
multiseriate involucre of erect-patent subulate bracts;
leaves less than 4 cm wide; pollen with colpi reaching
the pole; raphids of achene elongate 183. Echinocoryne
Genera of Vernoniinae
176. Albertinia Spreng.
Albertinia Spreng., Neue Entdeck. Pflanzenk. 2: 133 (1821)
[1820].
Symblomeria Nutt. (1840).
Branching shrubs; hairs T-shaped with multicellular stalks. Inflorescence loosely corymbiform,
heads pedunculate; involucral bracts 55–60 in c. 3
series; deep receptacular pits enclosing achenes.
Florets 45–50; anther bases rounded; style with
node; sweeping hairs broad-acicular. Achene c.
10-costate, raphids subquadrate; pappus capillary, tawny. Pollen non-lophate. One species, A.
brasiliensis Spreng., eastern Brazil.
177. Aynia H. Rob.
Aynia H. Rob., Proc. Biol. Soc. Wash. 101: 959 (1988).
Perennial herbs; hairs simple. Peduncles usually
long. Heads subtended by large foliose bracts,
involucral bracts c. 100 in 4–5 series, 10–25 mm
long. Florets c. 50; anthers tails short, truncate;
style with node; sweeping hairs pointed. Achene
10-ribbed, raphids subquadrate; pappus capillary,
with squamellae. Pollen echinolophate, with
intercolpus-aligned polar lacunae. One species, A.
pseudascaricida H. Rob., Peru.
178. Chrysolaena H. Rob.
Chrysolaena H. Rob., Proc. Biol. Soc. Wash. 101: 956 (1988);
Robinson, Proc. Biol. Soc. Wash. 105: 657–663 (1992), key.
Vernonia subsect. Flexuosae Cabrera (1944).
Vernonia series Verbascifoliae S.B. Jones (1979).
Perennial, usually xylopodial herbs, sericeous
or lanate with simple yellowish hairs. Inflorescence seriate-cymose, heads sessile; involucral
bracts/florets 1–2/1. Florets 10–65; anther bases
obtuse; apical appendage usually with glands; style
without prominent node; sweeping hairs broadacicular. Achenes 5-costate, densely sericeous,
with glands, raphids elongate; pappus capillary,
with squamellae. Pollen echinolophate, single
polar lacunae. n = 10, 17. Nine species, Brazil to
Argentina and Peru.
179. Cololobus H. Rob.
Cololobus H. Rob., Proc. Biol. Soc. Wash. 107: 557 (1994).
Subshrubs; hairs simple or asymmetrically Tshaped. Inflorescence thyrsoid; involucral bracts,
c. 6-seriate, c. 35, in outer 4 series pubescent, inner
mostly glabrous. Florets 20–30; corollas glabrous,
lobes very short; anther base obtuse; style with
node; sweeping hairs blunt, often septate. Achenes
8–10-costate, with setulae and glands, raphids
subquadrate; pappus capillary, with squamae.
Pollen non-lophate. Three species, eastern Brazil.
Compositae
180. Cyrtocymura H. Rob.
Fig. 40
Cyrtocymura H. Rob., Proc. Biol. Soc. Wash. 100: 849
(1987).
Perennial herbs; hairs simple. Inflorescences
scorpioid-cymose with crowded sessile heads,
deciduous with age; involucral bracts 20–30, in
c. 3 series. Florets 14–30; corolla lobes sericeous;
anther bases rounded; style with node; sweeping hairs broadly acicular. Achenes 10-costate,
setuliferous, idioblasts bulging, raphids elongate;
pappus capillary, outer squamellae persistent.
Pollen non-lophate. n = 17 . Six species, Mexico,
West Indies to Brazil and Argentina.
155
181. Dasyanthina H. Rob.
Dasyanthina H. Rob., Proc. Biol. Soc. Wash. 106: 778 (1993).
Perennial herbs 2–4 m high; stem hairs T-shaped.
Inflorescences rounded-thyrsoid; peduncles slender. Involucral bracts c. 60 in 5–6 series. Florets c.
25; corolla inside with unicellular hairs; anther tails
unsclerified, connective with glands; stylar node
annular; sweeping hairs pointed. Achenes 8–10ribbed, setuliferous, raphids elongate; pappus capillary, squamellae persistent. Pollen non-lophate.
Two species, eastern Brazil.
182. Dipterocypsela S.F. Blake
Dipterocypsela S.F. Blake, J. Wash. Acad. Sci. 35: 36 (1945).
Fleshy herbs; hairs symmetrically T-shaped. Seriate cymes with crowded sessile heads. Involucral
bracts c. 12 in c. 2 series, subequal, winged. Florets
c. 12; corollas zygomorphic, inner lobes longer; anther bases rounded; apical appendage with gland;
style with node; sweeping hairs fusiform, septate.
Achenes obcompressed, winged, glabrous, raphids
subquadrate; pappus bristles short, deciduous.
Pollen sublophate. One species, D. succulenta S.F.
Blake, Colombia.
183. Echinocoryne H. Rob.
Echinocoryne H. Rob., Proc. Biol. Soc. Wash. 100: 586
(1987).
Perennial herbs; sericeous with straight hairs.
Heads pedunculate; involucral bracts c. 110–500 in
6–9 series, linear, straight, pungent. Florets 15–60;
anther bases rounded; style with node; sweeping hairs acicular. Achenes 5-costate, sericeous,
raphids elongate; pappus capillary, with squamellae. Pollen echinolophate, no polar lacunae. Six
species, Brazil.
184. Eirmocephala H. Rob.
Fig. 40. Compositae-Vernonieae. Cyrtocymura scorpioides.
A Habit showing alternate leaves and scorpioid cymes; note
bracts at base of one branch after heads have fallen. B Head,
note homogamous condition with regular corollas. C Spinulose receptacle surface and spreading persistent involucral
bracts. D Corolla with tips of anthers and style. E Corolla
in long section, note deeply divided corolla lobes, calcarate
anther bases, nectary and node at base of style, sweeping
hairs on upper style shaft and backs of style branches, and
undivided stigmatic surface inside of style branches. F Achene with ribs and setulae, and pappus with capillary inner
series and outer series of shorter bristles or squamellae.
(Robinson 1999a)
Eirmocephala H. Rob., Proc. Biol. Soc. Wash. 100: 853
(1987).
Perennial herbs; hairs simple. Inflorescence
seriate- or densely scorpioid-cymose. Heads sessile, persistent; involucral bracts 24–65, in c. 4 series. Florets 7–35; anthers tailed; apical appendage
often glanduliferous; style with node; sweeping
hairs acicular. Achenes 10-veined, setuliferous,
without idioblasts, raphids elongate; pappus
capillary, with squamellae. Pollen non-lophate or
156
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
subtriporate with polar lacuna. n = 16, 17. Three
species, Central America to central Andes.
188. Lessingianthus H. Rob.
185. Harleya S.F. Blake
Perennial herbs, sometimes xylopodial; hairs
simple. Inflorescences simple or seriate-cymose.
Heads sessile or pedunculate; involucral bracts
45–100 in 4–6 series. Florets 15–50; anther bases
rounded; style usually without node; sweeping
hairs sharp. Achenes 5-costate, without glands,
raphids subquadrate or elongate; pappus capillary,
with squamellae. Pollen echinolophate, without
polar lacunae, with strong bacula. n = 17. More
than 102 species, South America, mostly Brazil and
Argentina, one or two species north to Colombia
and Venezuela.
Harleya S.F. Blake, J. Wash. Acad. Sci. 22: 379 (1932).
Shrub, bases decumbent; arachnoid hairs contorted. Leaves discolorous. Inflorescences deflected
at nodes, with sessile, axillary clusters of narrow
heads; involucral bracts 30–35, in 5 series. Florets
6–8; corollas hairless; anthers without tails; style
without node; sweeping hairs acicular. Achenes
5-angled, pustulate, raphids subquadrate; pappus
lacking. Pollen echinolophate with intercolpusaligned polar lacunae. One species, H. oxylepis
(Benth.) S.F. Blake, Central America.
Lessingianthus H. Rob., Proc. Biol. Soc. Wash. 101: 939
(1988).
189. Manyonia H. Rob.
186. Heterocypsela H. Rob.
Heterocypsela H. Rob., Phytologia 44: 442 (1979).
Perennial herbs; hairs symmetrically T-shaped. Peduncles 5–30 mm long; involucral bracts c. 70 in c.
6 series, caudate-acuminate. Florets c. 60–70; anther bases blunt; apical appendages with glands;
style with node; sweeping hairs sharp. Outer achenes obcompressed, margins winged, raphids subquadrate, pappus bristles deciduous; inner achenes prismatic, setuliferous, pappus more persistent. Pollen subtriporate. One species, H. andersonii H. Rob., eastern Brazil.
187. Lepidaploa (Cass.) Cass.
Lepidaploa (Cass.) Cass. in Cuvier, Dict. Sci. Nat. 36: 20
(1825); Robinson, Proc. Biol. Soc. Wash. 103: 464–498
(1990), emend., list; Robinson et al., Phytologia 46: 421–436
(1980), lectotyp.
Vernonia subg. Lepidaploa Cass. (1817).
Flustula Rafin. (1838) [1836].
Herbs or shrubs, rarely annual; sericeous or
tomentose, hairs simple or T-shaped. Leaves
usually alternate. Inflorescences seriately cymose.
Heads usually sessile; involucral bracts 20–70, in
3–6 series. Florets (8–)10–35; anther bases obtuse;
apical appendages rarely with glands; style with
node; sweeping hairs broadly acicular. Achenes
8–10-ribbed, raphids elongate; pappus capillary,
with squamellae. Pollen echinolophate, muri
weakly attached. n = 17. More than 130 species,
tropical America.
Manyonia H. Rob., Proc. Biol. Soc. Wash. 112: 224 (1999).
Perennial herbs; stems hispidulous, hairs simple. Inflorescence seriate-cymose, peduncles
2–3 mm long. Involucral bracts c. 100, outer 3–4
series spreading, subulate. Florets c. 35; anthers
without tails; style with node; sweeping hairs
acicular. Achenes 5-costate, setulae and idioblasts
in furrows, raphids subquadrate; pappus setae
fragile, uniseriate, squamellae persistent. Pollen
non-lophate. One species, M. peculiaris (Verdc.),
H. Rob., Tanzania.
190. Mattfeldanthus H. Rob. & R.M. King
Mattfeldanthus H. Rob. & R.M. King, Willdenowia 9: 10
(1979).
Shrubs; hairs simple. Inflorescence base pseudotrichotomous, branches seriate-cymose, bracts foliiform. Heads sessile; involucral bracts c. 100, c.
7-seriate. Florets 14–16; outer corollas zygomorphic, innermost lobe longer; anther tails lobed;
style with node; sweeping hairs acicular. Achenes
c. 10-costate, sericeous, raphids elongate; pappus
capillary, with squamellae. Pollen echinolophate,
without polar lacunae. Two species, eastern Brazil.
191. Pseudopiptocarpha H. Rob.
Pseudopiptocarpha H. Rob., Proc. Biol. Soc. Wash. 107: 561
(1994).
Shrubs or subshrubs; hairs appressed, symmetrically T-shaped. Heads axillary, clustered, sessile;
involucral bracts 25–30 in c. 5 series. Florets 8–10;
anther tails short; style with node; sweeping hairs
Compositae
sharp. Achenes weakly 10-costate, setulae, glands,
and idioblasts scattered, raphids subquadrate; pappus bristles capillary, with squamellae. Pollen echinolophate, with intercolpus-aligned polar lacunae.
Two species, Colombia.
192. Quechualia H. Rob.
Quechualia H. Rob., Proc. Biol. Soc. Wash. 106: 780 (1993).
Erect or scrambling shrubs; hairs asymmetrically
T-shaped, often proliferated. Inflorescences thyrsoid. Heads pedunculate; involucral bracts 60–90
in 5–6 series. Florets 30–55; corolla limb often
with multicellular hairs inside; anther connectives
with glands, tails denticulate; style with node;
sweeping hairs subacicular. Achenes 8–10-ribbed,
setuliferous, raphids elongate; pappus capillary,
with squamellae. Pollen non-lophate. n = 17. Four
species, Peru to Argentina.
193. Stenocephalum Sch. Bip.
Stenocephalum Sch. Bip., Jahresber. Pollichia 20/21: 385
(1863); Robinson, Proc. Biol. Soc. Wash. 100: 578–583
(1987), rev.
Perennial herbs; hairs simple, often arachnoid.
Leaves pale tomentose below. Heads axillary or
in panicles; involucral bracts 15–22 in 3–4 series.
Florets 4–7(–10); anther bases rounded; style
with node; sweeping hairs short, sharp. Achenes
10-ribbed, setuliferous, without glands, idioblasts,
or raphids; pappus capillary, squamellae present.
Pollen echinolophate with polar lacuna. n = 17.
Five species, Central America and tropical South
America.
194. Stilpnopappus Mart. ex DC.
Stilpnopappus Mart. ex DC., Prodr. 5: 75 (1836).
Strophopappus DC. (1836).
Perennial herbs or shrubs; hairs simple. Inflorescences axillary or seriate-cymose. Heads sessile,
1 to few per node; involucral bracts 20–50 in
3–4 series. Florets 6–50; anther bases obtuse;
style with node; sweeping hairs acicular. Achenes
8-costate, projecting rim at top, setuliferous,
idioblasts scattered, raphids subquadrate; pappus
of lanceolate awns. Pollen non-lophate or lophate
with intercolpar-aligned polar lacunae. Twenty
species, Brazil, Venezuela.
157
195. Sparganophoros Vaill.
Sparganophoros Vaill., Königl. Akad. Wissensch. Paris Phys.
Abh. 5: 368 (1754).
Struchium P. Browne (1756).
Decumbent annuals; hairs sparse, simple. Heads
axillary, clustered; involucral bracts 20–25 in c. 2
subequal series. Florets 50–70; corolla lobes 3 or
4; anther bases rounded, connective with glands;
style with node; sweeping hairs sharp. Achenes
3–5-angled, without setulae, idioblasts numerous,
raphids elongate; pappus a cartilaginous sleeve.
Pollen subtriporate, with polar lacuna. n = 16. One
species, S. sparganophora (L.) O. Kuntze, pantropical, widely adventive.
196. Trepadonia H. Rob.
Trepadonia H. Rob., Proc. Biol. Soc. Wash. 107: 564 (1994).
Vines; hairs mostly symmetrically T-shaped.
Leaves alternate or opposite. Inflorescence branching often at 90°, branchlets often subracemose.
Involucral bracts c. 25 in c. 5 series. Florets
8–10; corollas glabrous; anther bases obtuse;
style with node; sweeping hairs acicular. Achenes
10-costate, setulae scattered, raphids subquadrate;
pappus capillary, squamellae persistent. Pollen
non-lophate. Two species, Peru.
197. Vernonanthura H. Rob.
Vernonanthura H. Rob., Phytologia 73: 66 (1992).
Subshrubs to small trees, sometimes xylopodial;
hairs simple or T-shaped. Inflorescences thyrsoid.
Heads sessile to long-pedunculate; involucral
bracts 6–30(–60) in 4–10 series. Florets 4–30;
corollas without hairs; anther bases obtuse or
tailed; appendages often with glands or hairs; style
with node; sweeping hairs short-acute. Achenes
8–10-costate, setuliferous or with idioblasts;
pappus capillary, with squamellae. Pollen nonlophate. n = 17. Sixty-five or more species, tropical
America.
198. Vernonia Schreb.
Vernonia Schreb., Gen. 2: 541 (1791), nom. cons.; Jones &
Faust, N. Amer. Fl. 2, 10: 180–195 (1978), reg. rev.
Behen Hill (1762).
Perennial herbs, bases decumbent; hairs simple or symmetrically T-shaped. Inflorescences
cymose with branches longer than central axis.
Heads usually pedunculate; involucral bracts c.
158
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
50 in 5–6 series. Florets 8–120; anthers without
tails; appendages often with glands; style with
node; sweeping hairs acute, sometimes septate.
Achenes 5–10-costate, with setulae, glands and/or
idioblasts, raphids subquadrate; pappus capillary,
with squamellae. Pollen non-lophate. n = 17.
Twenty-two species, south-eastern United States,
Bahamas, to central Mexico, 2 distinctive species
in South America.
199. Xiphochaeta Poepp.
Xiphochaeta Poepp., Nov. Gen. Sp. 3: 44, pl. 250, 8–11 (1842).
Aquatic short-lived herbs; hairs simple, appressed.
Heads sessile, 1(–3) in axils; involucral bracts 70–
80 in 3–4 series. Florets c. 30; anther bases not calcarate; appendages with glands; style with node;
sweeping hairs acicular. Achenes 5-costate, setulae
and idioblasts scattered, raphids elongate; pappus
segments c. 10, canaliculate, non-costate. Pollen
echinolophate. One species, X. aquatica Poepp.,
Amazon and Orinoco basins.
VI.3. Subtribe Piptocarphinae H. Rob.,
F. Bohlmann & R.M. King (1980).
Woody shrubs, trees or vines; hairs stellate or
lepidote. Inner involucral bracts deciduous; anther
appendage slightly indurate, glabrous. Achene
raphids short or subquadrate. Pollen tricolporate, non-lophate. Characteristic sesquiterpenes:
usually glaucolides.
Key to the Genera
1. Pappus a collar, without bristles or scales
205. Huberopappus
– Pappus of capillary bristles or scales
2
2. Corollas divided to base of limb, throat lacking, lobes
beginning at top of basal tube
3
– Corollas not divided to base of limb, distinct throat
present
4
3. Heads with single florets; leaves alternate
202. Cuatrecasanthus
– Heads with 9–12 florets; leaves opposite
207. Joseanthus
4. Plants lepidote; pappus with laciniate outer collar
204. Ekmania
– Plants with mostly stellate hairs or nearly glabrous,
sometimes lepidote; pappus without fused outer collar
5
5. Inner pappus of deciduous broadened segments or
flattened bristles; style base usually without node 6
– Pappus with many rather persistent capillary bristles;
style base with node
8
6. Branchlets of inflorescence without foliose bracts;
pubescence not heterotrichous
209. Piptocoma
– Branchlets of inflorescence with distinct foliose bracts
larger than heads; pubescence heterotrichous or
evanescent
7
7. Dark uniseriate hairs prominent among stellate hairs;
axis of inflorescence not deflected at nodes; receptacle with thin partitions; pappus with c. 20 deciduous
flattened bristles
200. Blanchetia
– Dark uniseriate hairs small or evanescent; axis of inflorescence deflected at nodes; receptacle without thin
partitions; pappus with only c. 8–10 broad, linear inner segments
206. Irwinia
8. Corolla with many hairs inside limb
203. Dasyandantha
– Corolla glabrous inside
9
9. Tails of anthers, when present, not sclerified; only inner involucral bracts deciduous; trees or shrubs
201. Critoniopsis
– Tails of anthers sclerified, blunt or sharp; most involucral bracts deciduous; erect or scandent shrubs or trees
208. Piptocarpha
Genera of Piptocarphinae
200. Blanchetia DC.
Blanchetia DC., Prodr. 5: 75 (1836).
Shrubs, with intermixed long dark multicellular hairs and pale stellate hairs. Inflorescences
corymbiform with foliose bracts; involucral bracts
25–30 in c. 4 series; receptacle with thin partitions.
Florets 8–10; filaments inserted near sinuses;
anthers not tailed; style without node; sweeping
hairs pointed. Achenes 10-costate, hairless; pappus
deciduous, c. 20 flattened bristles. One species, B.
heterotricha DC., north-eastern Brazil.
201. Critoniopsis Sch. Bip.
Critoniopsis Sch. Bip., Jahresber. Pollichia 20/21: 430 (1863);
Cuatrecasas, Bot. Jahrb. Syst. 77: 52–84 (1956), reg. rev.;
Jones, Brittonia 25: 86–115 (1973), reg. rev.; Robinson,
Proc. Biol. Soc. Wash. 106: 606–627 (1993), emend., list.
Turpinia Lex. ex LaLlave & Lex. (1824), nom. illegit.
Monosis DC. sect. Eremosis DC. (1836).
Tephrothamnus Sch. Bip. (1863).
Eremosis (DC.) Gleason (1906).
Shrubs or trees; hairs often stellate. Leaves
alternate or opposite, petiole often lobed or
winged. Inflorescences terminal; involucral bracts
18–25(–35) in 4–6 series. Florets 1–11(–15 or 20);
corolla throat present, lobes often recurved, with
small glands; anthers often with unsclerified tails;
style usually with node; sweeping hairs obtuse.
Achenes 5–10-costate; pappus capillary. n = 17.
Seventy-six species, Mexico, Central America,
Andes to Brazil.
Compositae
DNA studies (Keeley and Chan, pers. comm.)
show that Tephrothamnus and Eremosis are worthy
of separate generic rank.
159
some with single awn. One species, H. maigualidae
Pruski, Venezuela (Bolivar-Amazonas border).
206. Irwinia G.M. Barroso
202. Cuatrecasanthus H. Rob.
Irwinia G.M. Barroso, Rodriguesia 32: 11 (1980).
Cuatrecasanthus H. Rob., Revista Acad. Colomb. Ci. Exact.
17: 209 (1989).
Subscandent shrub; mixed evanescent, small, multicellular and pale stellate hairs. Inflorescence axis
deflected at nodes, branches with foliose bracts.
Involucral bracts 18–20 in c. 4 series. Florets 5;
filaments at sinuses; anther bases rounded; style
without node; sweeping hairs obtuse. Achenes 6–7costate, glabrous; pappus 8–10 deciduous, twisted
straps, outer bristles short. One species, I. coronata
G.M. Barroso, north-eastern Brazil.
Weak shrubs; hairs with apical cell enlarged at
base. Inflorescence terminal; heads sessile in
glomerules; involucral bracts c. 15 in 5–6 series.
Florets 1; corolla throat lacking; anthers with
fimbriate tails; style with node; sweeping hair tips
rounded. Achenes 10-costate, glanduliferous and
spiculiferous, raphids lacking; pappus capillary,
outer setae short. Three species, Ecuador, Peru.
203. Dasyandantha H. Rob.
Dasyandantha H. Rob., Proc. Biol. Soc. Wash. 106: 778
(1993).
Small trees; stems sublanate, hairs simple. Leaves
to 30 cm long. Inflorescence thyrsoid-paniculate.
Heads sessile in glomerules. Involucral bracts c.
30, in c. 4 series. Florets c. 12; corolla throat enlarged, pilosulous inside; anthers with unsclerified
tails; style with node; sweeping hairs blunt-tipped.
Achenes 8-ribbed, setuliferous; pappus capillary,
fragile. One species, D. cuatrecasasiana (Aristeg.)
H. Rob., Venezuelan Andes.
204. Ekmania Gleason
Ekmania Gleason, Bull. Torrey Bot. Club 46: 250 (1919).
Lepidote shrubs. Leaves discolorous below. Inflorescence corymbose; heads with small foliose
subinvolucral bract; involucral bracts c. 30 in
4–5 series. Florets c. 12; anther bases rounded;
style without node; sweeping hairs acute, often
septate. Achenes 10-costate, glanduliferous; outer
persistent pappus a laciniate collar, few deciduous
flattened bristles inside. One species, E. lepidota
Griseb., Cuba.
205. Huberopappus Pruski
207. Joseanthus H. Rob.
Joseanthus H. Rob., Revista Acad. Colomb. Ci. Exact. 17: 210
(1989).
Shrubs or trees; hairs sometimes T-formed. Leaves
opposite. Inflorescences densely corymbose. Heads
shortly pedunculate; involucral bracts 20–30 in c.
4–5 series. Florets 9–12; corolla densely pilosulous, without throat; anthers with fimbriate tails;
style with node; sweeping hairs obtuse. Achenes
3–8-costate, with glands and small setulae; pappus
capillary, with squamellae. Five species, Colombia,
Ecuador.
208. Piptocarpha R. Br.
Piptocarpha R. Br., Observ. Comp. 121 (1817) [1818].
Carphobolus Schott in Spreng. (1827).
Vernonia Schreb. sect. Vanillosma Less. (1831).
Monanthemum Griseb. (1861), nom. illegit.
Scandent scrambling shrubs or trees; hairs stellate
or lepidote. Leaves alternate or opposite. Heads
in axillary or terminal clusters; involucral bracts
18–30 in 3–4 series. Florets 3–20; corolla lobes
often recurved; anther tails sclerified; style with
node; sweeping hairs blunt, often septate. Achenes 3–5-costate, glabrous, with idioblasts, raphids
short-oblong; pappus capillary. n = 17. Forty-three
species, tropical America.
Huberopappus Pruski, Novon 2: 19 (1992).
209. Piptocoma Cass.
Shrubs; hairs stellate. Inflorescences of few shortpedunculate heads and small foliiform bracts.
Involucral bracts 14–17 in 3–4 series. Florets
19–22; anther bases rounded; style without node;
sweeping hairs obtuse, often septate. Achenes 3–5angled, c. 10-striate, glabrous; pappus collarform,
Piptocoma Cass., Bull. Soc. Philom. Paris 1817: 10 (1817);
Pruski, Novon 6: 96–102 (1996), emend.
Dialesta Kunth (1818).
Odontoloma Kunth (1818).
Pollalesta Kunth (1818).
Adenocyclus Less. (1829).
160
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Trees and shrubs; hairs stellate to lepidote. Inflorescences corymbiform. Involucral bracts 5–25 in
4–5 series, partly deciduous. Florets 1–12; corolla
throat very short; anther bases rounded; style
without node; sweeping hairs mostly blunt, often
septate. Achenes weakly 7–10-costate, idioblasts
present; pappus with deciduous linear segments,
persistent outer scales. Eighteen species, Greater
Antilles, northern South America.
VI.4. Subtribe Chrestinae H. Rob. (1999).
Inflorescence compound, sometimes spicate, maturing acropetally. Anther appendage thin-walled,
usually glabrous; style without node. Pollen tricolporate, lophate. Characteristic sesquiterpenes:
glaucolides.
Key to the Genera
1. Leaves deeply lobed; inflorescence a spike of heads
without obvious bracts; peduncle swollen and fistulose
211. Pithecoseris
– Leaves entire or shallowly dentate; inflorescence not
or only slightly spicate; without swollen peduncle 2
2. Clusters of heads borne in axils of large imbricate upper leaves or bracts; leaves broad with many sublongitudinal veins
212. Soaresia
– Heads in glomerules or umbels or small spikes, not
in axils of series of large leaves or bracts; leaves with
pinnate venation or narrow with few longitudinal veins
210. Chresta
Genera of Chrestinae
210. Chresta Vell. ex DC.
Chresta Vell. ex DC., Prodr. 5: 85 (1836).
Vernonia Schreb. sect. Pycnocephalum Less. (1831).
Pycnocephalum (Less.) DC. (1836).
Stachyanthus DC. (1836), nom. rej.
Eremanthus Less. sect. Pycnocephalum (Less.) Baker (1873).
Glaziovianthus G.M. Barroso (1947).
Argyrovernonia MacLeish (1984), nom. nov. for Stachyanthus DC.
Perennial herbs and subshrubs; hairs symmetrically or asymmetrically T-shaped or forming
dense felt. Inflorescences long-peduculate, clusters
globose, subumbellate or spicate; involucral bracts
10–20 in 5–6 series. Florets 2–12; corolla tube
long; anther bases rounded; appendage sometimes
with glands; sweeping hairs fusiform. Achenes
5-angled, raphids short or elongate; pappus
capillary, sometimes caducous. Pollen lophate or
non-lophate. Eleven species, Brazil, Bolivia.
211. Pithecoseris Mart. ex DC.
Pithecoseris Mart. ex DC., Prodr. 5: 84 (1836).
Coarse biennial herbs; hairs symmetrically Tshaped. Leaves deeply pinnatifid. Inflorescence
a dense spike, peduncle inflated, fistulose. Involucral bracts c. 8 in c. 3 subequal series. Florets 3–7;
anther bases rounded; sweeping hairs fusiform.
Achenes 5-costate, setuliferous or glabrous,
raphids elongate; capillary pappus caducous,
usually with squamellae. Pollen echinolophate,
with polar lacuna. One species, P. pacourinoides
Mart. ex DC., eastern Brazil.
212. Soaresia Sch. Bip.
Soaresia Sch. Bip., Jahresber. Pollichia 20/21: 376 (1863),
nom. cons., non F. Allem. (1851).
Bipontia S.F. Blake (1937), nom. nov. for Soaresia Sch. Bip.
Subshrubs, with xylopodia; stems felted with
fusiform hairs. Leaves imbricate, silvery velvety,
veins nearly longitudinal. Rows of sessile heads in
upper axils. Involucral bracts c. 12 in c. 3 series.
Florets 4; corollas sericeous; anthers tailed; sweeping hairs acicular. Achene 5-ribbed, sericeous,
raphids elongate; pappus awns c. 15, subulate,
basally winged. Pollen echinolophate, with polar
lacuna. One species, S. velutina Sch. Bip., Brazil
(Goias, Minas Gerais).
VI.5. Subtribe Centratherinae H. Rob.,
F. Bohlmann & R.M. King (1980).
Short-lived herbs. Heads single, terminal, with
foliose spreading subinvolucral bracts. Corolla
tube slender, densely stipitate-glanduliferous.
Style without node. Achene raphids subquadrate.
Pollen tricolporate, non-lophate. Characteristic
terpenoids: furoheliangolides.
Key to the Genera
1. Perennial herbs or shrubs branching mostly near base,
with or without a pappus; achenes glabrous or stiffly
setuliferous
213. Centratherum
– Weak annual herbs; without a pappus; achenes with
small, divergently tipped setulae 214. Oiospermum
213. Centratherum Cass.
Centratherum Cass., Dict. Sci. Nat. 7: 384 (1817); Kirkman,
Rhodora 83: 1–24 (1981), rev.; Robinson, Phytologia 46:
143–145 (1980), emend.
Ampherephis Kunth (1818).
Crantzia Vell. (1827).
Compositae
161
Perennial herbs or shrubs branching near base;
hairs T-shaped and simple multicellular. Involucral bracts 30–40, in c. 4 series, with distal margins scarious. Florets 40–100+; anthers not tailed;
sweeping hairs acicular. Achenes 8–10-ribbed; pappus of short deciduous awns or lacking. n = 16, 32.
Three species, tropical America, Philippines, Australia, adventive.
Genera of Lychnophorinae
214. Oiospermum Less.
215. Anteremanthus H. Rob.
Oiospermum Less., Linnaea 4: 339 (1829).
Anteremanthus H. Rob., Proc. Biol. Soc. Wash. 105: 646
(1992).
Annual herbs; hairs T-shaped. Leaf blades hairless,
glandular-dotted. Involucral bracts c. 45 in c. 4 series. Florets c. 25; anther bases rounded; sweeping
hairs short-pointed, few septa. Achenes 10-ribbed,
pilosulous, setulae tips flagelliform or hooked; pappus lacking. One species, O. involucrata (Spreng.)
Less., north-eastern Brazil.
VI.6. Subtribe Lychnophorinae
Benth. & Hook. f. (1873).
Rosettiform herbs to small trees. Heads often compound. Anther appendage indurate, glabrous; style
without node. Achene raphids subquadrate. Pollen
tricolporate, non-lophate. Characteristic sesquiterpenes: furoheliangolides.
Key to the Genera
1. Acaulescent; peduncles directly from the basal rootstock
222. Prestelia
– Leafy stems present
2
2. Pappus with broad, strap-shaped or subulate
segments, easily deciduous
3
– Pappus with capillary bristles, persistent
5
3. Leaves with clasping bases; involucral bracts with pungent acuminate tips; heads with c. 80 florets; achenes
10-ribbed
223. Proteopsis
– Leaves not clasping at base; involucral bracts not
acuminate at tips; heads with 1–23 florets; achenes 4or 5-angled, sometimes faintly 8–10-ribbed
4
4. Inflorescence with rounded or axillary glomeruli of
heads; heads with 1–12 florets
218. Lychnophora
– Inflorescence with spiciform cluster of heads; heads
with 9–23 florets
219. Lychnophoriopsis
5. Herbs with leaves mostly in basal rosette 220. Minasia
– Shrubs or trees with cauline leaves
6
6. Ericoid shrubs, often with small densely spirally inserted leaves; heads solitary in axils or on branch tips
221. Piptolepis
– Broad-leaved shrubs or trees; heads in corymbiform
cymes, axillary clusters, or glomeruli
7
7. Heads terminal, not in clusters, large, with c. 60 florets
215. Anteremanthus
– Heads in leaf axils or in dense clusters, with 1–10(–24–
26) florets
8
8. Upper leaf surfaces strongly bullate; clusters of heads
axillary; corollas with long and narrow tube and throat
216. Chronopappus
– Upper leaf surfaces weakly roughened or plane; heads
in terminal clusters or glomeruli; corollas short
217. Eremanthus
Shrubs; hairs mostly T-formed. Inflorescence
thyrsoid, with gradate foliaceous bracts. Involucral
bracts c. 60 in 5–6 series. Florets c. 60; corolla tube
short, lobes with contorted hairs; anther bases
rounded; sweeping hairs pointed, often septate.
Achene 8–10-ribbed, densely long-setulose; pappus capillary, with squamellae. One species, A.
hatschbachii H. Rob., Brazil (Minas Gerais).
216. Chronopappus DC.
Chronopappus DC., Prodr. 5: 84 (1836).
Shrubs; stem felted with thick-walled fusiform and
stellate hairs. Leaves bullate, abaxial tomentum
of contorted stellate-based hairs. Heads sessile
in upper axils, c. 8 pubescent subinvolucral
bracts; involucral bracts c. 8–10, rather deciduous.
Florets 8–10; anther bases rounded; sweeping
hairs pointed. Achenes 10-angled, glabrous; inner
pappus of flattened bristles. One species, C. bifrons
(DC. ex Pers.) DC., Brazil (Minas Gerais).
217. Eremanthus Less.
Eremanthus Less., Linnaea 4: 317 (1829); MacLeish, Ann.
Missouri Bot. Gard. 74: 265–290 (1987), rev.
Vanillosmopsis Sch. Bip. (1861).
Sphaerophora Sch. Bip. (1863), nom. illegit.
Paralychnophora MacLeish (1984), nom. nov. for
Sphaerophora Sch. Bip.
Shrubs and trees; stems felted or woolly or
stellate-lepidote. Heads usually in 1 to many
dense, globular clusters; involucral bracts 10–40 in
4–7 usually gradate series, innermost deciduous.
Florets 1–9, 24–26; corolla tube short; anther bases
rounded; sweeping hairs pointed, often septate.
Achenes 3–5-angled, or 10(–20)-costate; pappus
bristles 3–5-seriate, broadened to filiform. n = 15,
18. Circa 25 species, south-eastern to north-eastern
Brazil, eastern Bolivia.
162
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
218. Lychnophora Mart.
Lychnophora Mart., Denkschr. Königl.-Baier. Bot. Gesell.
Regensburg 2: 148 (1822).
Haplostephium Mart. ex DC. (1836).
Lychnocephalus Mart. ex DC. (1836).
Usually candelabriform or rosettiform shrubs or
small trees; stem felted with fusiform hairs. Leaves
often crowded, with stellate or asymmetrical Tshaped hairs below. Heads sessile in globular or
hemispherical clusters. Involucral bracts c. 12–25
in 4–6 series, innermost deciduous. Florets 1–12;
corollas glabrous; anther bases rounded; sweeping
hairs sharp, mostly septate. Achenes 4–5-angled
and 8–10-veined, glabrous; pappus of flattened,
usually twisted bristles or straps, often with squamae. n = 17. Circa 30 species, south-eastern to
north-eastern Brazil.
219. Lychnophoriopsis Sch. Bip.
Lychnophoriopsis Sch. Bip., Jahresber. Pollichia 20/21: 375
(1863); Robinson, Proc. Biol. Soc. Wash. 105: 640–652
(1992), key.
Episcothamnus H. Rob. (1981).
Weakly candelabriform small trees, some hairs
long-stalked T-shaped; stems felted with fusiform
hairs. Leaves densely spiralled, tomentose below. Axillary heads sessile in elongate clusters.
Involucral bracts c. 30–60 in c. 6 series. Florets
9–23; anther tails small; sweeping hairs sharp,
often septate. Achenes 8–10-ribbed, 4–5-angled,
glabrous; pappus segments strap-shaped or
bristle-tipped, with squamellae. n = 17. Four
species, south-eastern Brazil.
220. Minasia H. Rob.
8–18(–25) in 3–4 series. Florets 8–18; anther tails
short, lobed; sweeping hairs short-acute. Achenes
strongly 10–12-costate, furrows with setulae and
glandular dots; pappus bristles flattened, bases
often broad, sometimes with shorter setae. Five or
six species, south-eastern Brazil.
222. Prestelia Sch. Bip.
Prestelia Sch. Bip., Festschr. Naturf. Gesell. Emden 73
(1864).
Perennial acaulous subshrubs, rootstock broad;
axils lanulose; hairs stellate-based. Leaves linear.
Inflorescences 1 to many pedunculate compound
heads, secondary involucre of c. 5 subequal bracts.
Individual heads 5–10 in cluster, sessile; involucral
bracts c. 12 in c. 2 series. Florets 5–6; anthers not
tailed; sweeping hairs acute. Achenes c. 8-costate,
scattered setulae, glands and idioblasts; pappus
bristles in 2–3 series, no squamellae. One species,
P. eriopus Sch. Bip., south-eastern Brazil.
223. Proteopsis Mart. & Zucc. ex Sch.Bip.
Proteopsis Mart. & Zucc. ex Sch.Bip., Jahresber. Pollichia
20/21: 378 (1863).
Coarse perennial herbs; silvery with appressed
elliptic-stellate hairs. Leaves smaller above, bases
clasping. Inflorescences a dense cluster of large
heads. Involucral bracts c. 60 in c. 6 series,
strongly pungent-acuminate. Florets c. 80; anthers
with broad tails; sweeping hairs acute. Achenes
10-ribbed, glabrous; pappus of few deciduous
awns. One species, P. argentea Mart. & Zucc. ex
Sch. Bip., south-eastern Brazil.
Minasia H. Rob., Proc. Biol. Soc. Wash. 105: 648 (1992).
Perennial rosettiform silvery herbs; hairs T-shaped.
Inflorescences scapose, branched, with crowded or
pedunculate heads; involucral bracts 50–60 in c.
5 series, with blunt tomentose tips. Florets 20–30;
anther with minute tails; sweeping hairs pointed,
septa few. Achene c. 8-veined, at least some setulae; pappus capillary, with squamellae. Five species,
Brazil (Minas Gerais).
VI.7. Subtribe Sipolisiinae H. Rob. (1999).
Coarse herbs; hairs mostly stellate or stellate at
base. Receptacles paleate or spinose. Anther appendages partly indurate, glabrous; style without
node. Achene usually with phytomelanin, raphids
usually lacking. Pollen tricolporate, non-lophate.
Characteristic terpenoids: furoheliangolides, few
eudesmane derivatives.
221. Piptolepis Sch. Bip.
Piptolepis Sch. Bip., Jahresber. Pollichia 20/21: 380 (1863),
nom. cons.
Ericoid shrubs; hairs appearing granular, stellate,
sometimes also thick-walled, straight. Heads
single, sessile or pedunculate; involucral bracts
Key to the Genera
1. Receptacles with deciduous pales; corollas with short
basal tube
2
– Receptacles with long spines, without pales; corollas
with elongate tube
3
Compositae
2. Stems and leaves with floccose tomentum; corolla
lobes with hairs; pappus of deciduous capillary
bristles
224. Bishopalea
– Stems and leaves with appressed tomentellum; corolla
lobes with spicules, no hairs; pappus of twisted flattened bristles
225. Heterocoma
3. Pubescence lepidote; achenes without phytomelanin
in walls, with subquadrate raphids
226. Hololepis
– Pubescence stellate or with hairs stellately armed near
base; achenes with phytomelanin in walls, without
raphids
4
4. Large caulescent plants; heads in clusters 227. Sipolisia
– Mostly acaulescent plants; heads solitary on long peduncles
228. Xerxes
163
227. Sipolisia Glaz. ex Oliv.
Sipolisia Glaz. ex Oliv., Hooker’s Icon. Pl. 23, t. 2281 (1894).
Perennial herbs or shrubs; stems lanate, leaves
tomentose. Inflorescence branches bearing leaves
only around terminal cluster of 1–6 large sessile
heads. Involucral bracts 55–100, lanceolate in c. 6
series; receptacles spinose. Florets 25–50; anther
tails broadly lobed; sweeping hairs sharp. Achenes
10-grooved, glabrous, with phytomelanin; pappus
bristles deciduous, flattened. One species, S.
lanuginosa Glaz. ex Oliv., south-eastern Brazil.
Genera of Sipolisiinae
224. Bishopalea H. Rob.
Bishopalea H. Rob., Phytologia 48: 211 (1981).
Erect woolly stems, 1.5–5 m. Heads few, peduncles
to 10 mm long, few foliose subinvolucral bracts;
involucral bracts 30–35 in c. 4 series; linear pales
persistent. Florets c. 20; corolla tubes short, apical
hairs on lobes; anther bases pointed; sweeping hairs
pointed, septate. Achenes 10-costate, glabrous, with
phytomelanin; pappus capillary, deciduous. One
species, B. erecta H. Rob., Brazil (Bahia).
225. Heterocoma DC.
228. Xerxes J.R. Grant
Xerxes J.R. Grant, Nordic J. Bot. 14: 287 (1994), nom. nov.
Alcantara Glaz. ex G.M. Barroso (1969), non Alcantarea
(Morren ex Mez) Harms
Acaulescent perennial herbs; stems lanate, leaves
tomentose. Leaf bases clasping. Inflorescences of
long, axillary peduncles bearing single heads, with
foliose subinvolucral bracts, involucral bracts c. 100
in c. 4 series; receptacle spinose. Florets c. 75; anthers tailed; sweeping hairs acute, septate. Achenes
10-costate, glabrous, with phytomelanin; pappus
of deciduous bristles. Two species, south-eastern
Brazil.
Heterocoma DC., Ann. Mus. Natl Hist. Nat. 16: 190, t. 7
(1810).
Subshrub; hairs appressed; leaves crowded, bases
vaginate. Large head in each upper axil; few foliose
subinvolucral bracts; involucral bracts c. 100, in
c. 4 series; pales linear, persistent. Florets c. 50;
corolla tube short; anthers not tailed; sweeping
hairs pointed, often septate. Achenes 10-costate,
with phytomelanin; pappus elements twisted,
strap-shaped, deciduous. One species, H. albida
(DC.) DC., south-eastern Brazil.
VI.8. Subtribe Elephantopinae Less. (1830).
226. Hololepis DC.
Key to the Genera
Hololepis DC., Ann. Mus. Natl Hist. Nat. 16: 155, 189 (1810).
Shrubs or trees; hairs symmetrically T-shaped.
Petioles long. Heads axillary, solitary, longpedunculate; subinvolucral bracts c. 8, foliiform,
trinervate; involucral bracts c. 30 in c. 4 series;
receptacular spines linear. Florets 25–35; anther
tails lobed; sweeping hairs short-pointed to obtuse.
Achenes 4-angled, glabrous, raphids subquadrate;
pappus of flattened persistent bristles, 2–3-seriate.
Three species, south-eastern Brazil.
Herbs. Involucre decussate with 4, 6 or 8 bracts; florets 4, corollas often zygomorphic, base of anthers
sometimes not calcarate; apical appendage thin,
glabrous. Achene raphids elongate. Pollen tricolporate to subtriporate, usually lophate. Characteristic
terpenoids: dilactones.
1. Corollas actinomorphic; plants without basal rosettes
229. Caatinganthus
– Corollas zygomorphic, with deepest sinus centred towards centre of head; plants with leaves partly in basal
rosette
2
2. Pappus of five contorted bristles; n = 13
232. Pseudelephantopus
– Pappus with straight awns or bristles; n = 11
3
3. Pappus with usually 5 strong bristles, rarely many bristles; pollen lophate
230. Elephantopus
– Pappus of many capillary bristles; pollen not lophate,
with many crowded echinate crests 231. Orthopappus
164
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Genera of Elephantopinae
229. Caatinganthus H. Rob.
Caatinganthus H. Rob., Smithsonian Contr. Bot. 89: 15
(1999).
Short-lived herbs, tomentum arachnoid. Leaves
linear. Branches spiciform, 3–20 adaxial heads,
biseriate. Outer involucral bracts 4–6, foliiform
distally, inner 4, scariose. Anther bases rounded;
style with node; sweeping hairs acicular. Achene
10-ribbed, long-setuliferous, raphids subquadrate;
pappus setae c. 10 and scales 10, large. Pollen
lophate, single polar lacuna. Two species, northeastern Brazil.
230. Elephantopus L.
Elephantopus L., Sp. Pl. 814 (1753).
Elephantosis Less. (1829).
Perennials; pilose, hairs simple, stiff. Leaves mostly
basal. Heads in bracteate glomerules; involucral
bracts 8. Florets 2–4; corollas zygomorphic, with
deep inner sinus; anthers not calcarate, not tailed;
style without node; sweeping hairs acicular. Achenes 10-costate, setulose, raphids elongate; pappus bristles 5–15(40–81), straight, basally winged.
Pollen triporate, echinolophate. n = 11. Twelve or
more species, pantropical.
231. Orthopappus Gleason
Orthopappus Gleason, Bull. New York Bot. Gard. 4: 237
(1906).
Perennials, hairs simple, stiff, yellow. Leaves mostly
basal. Inflorescence a spiciform cluster of narrow
heads; involucral bracts 8. Corollas zygomorphic,
inner sinus deepest; anthers not long-calcarate;
style with node; sweeping hairs acicular. Achenes
5-costate, setuliferous, idioblasts along costae,
raphids elongate; pappus bristles 2–3-seriate,
inner basally winged. Pollen tricolporate, with
crowded echinate ridges. n = 11. One species, O.
angustifolius (Sw.) Gleason, tropical America.
232. Pseudelephantopus Rohr
Pseudelephantopus Rohr, Skr. Naturhist.-Selsk. Kjb. 2: 213
(1792), nom. et orth. cons.
Distreptus Cass. (1817).
Matamoria LaLlave & Lex. (1824).
Spirochaeta Turcz. (1851).
Perennial herbs; hairs stiff, simple. Leaves mostly
basal. Inflorescences spiciform with clustered ses-
sile heads. Involucral bracts 8. Corollas zygomorphic with deep inner sinus; anther shortly calcarate; style without node; sweeping hairs acicular. Achenes 10-costate, setuliferous, with glands,
idioblasts along costae, raphids short-oblong; pappus bristles 5, contorted, base broadened. Pollen
triporate, lophate. n = 13. Two species, tropical
America.
VI.9. Subtribe Rolandrinae Cass. ex Dumort.
(1829).
Heads sessile in globose glomerules. Involucral
bracts 2–6; florets 1 per head. Achene raphids
minute or lacking. Pollen triporate, echinolophate.
Characteristic sesquiterpenes: glaucolides.
Key to the Genera
1. Shrubs; corolla lobes glabrous; pappus of short scales;
inflorescence with sessile axillary globose clusters of
heads, without group of subtending bracts
233. Rolandra
– Herbs; corolla lobes with some hairs; pappus of few
short bristles; inflorescence a pedunculate globose
cluster of heads subtended by foliose bracts
234. Spiracantha
233. Rolandra Rottb.
Rolandra Rottb., Soc. Med. Havn. Collect. 2: 256 (1775).
Shrubs; hairs simple. Leaves white-tomentose
below. Inflorescences axillary, sessile. Involucral
bracts 2, awned. Corolla 4-lobed, glabrous; anther
bases rounded, with sterile margin; style without
node; sweeping hairs acicular. Achenes 5-veined,
without setulae, with glands, many poorly differentiated idioblasts; pappus a persistent ring
of jagged scales. One species, R. argentea Rottb.,
tropical America.
234. Spiracantha Kunth
Spiracantha Kunth in Humb., Bonpl. & Kunth, Nov. Gen.
Sp., folio ed. 4: 22 (1818).
Weak herbs; hairs simple. Leaves white-tomentose
below. Inflorescence of pedunculate, 3–4-bracted
glomerules; involucral bracts c. 6, inner awned.
Corolla with filaments near sinuses, lobes 4 or 5;
anther bases obtuse; style without node; sweeping hairs acicular. Achenes fusiform, weakly 5–6veined, few glands and setulae above; pappus of
short bristles. n = 8. One species, S. cornifolia
Kunth, central and northern South America.
Compositae
VI.10. Subtribe Stokesiinae H. Rob. (1999).
Heads pedunculate in lax cymes; corollas mostly
ligulate; anther appendages glabrous; style
branches glanduliferous, hairs acicular. Pollen
tricolporate with crosswalls in colpi; rhomboidal lacunae. Characteristic sesquiterpenes:
elemanolides.
Only one genus:
235. Stokesia L’Hér.
Stokesia L’Hér., Sert. Angl. 27 (1789).
Cartesia Cass. (1816).
Perennial herbs; sparsely pilose, hairs simple.
Leaves mostly basal. Involucral bracts 40–50 in
3–4 series, outer with long spinose-margined
foliiform appendages. Florets 60–70; anther bases
rounded; style without node. Achenes with 3–4
sides, glands scattered, raphids few, narrowly
rhomboidal; pappus of 4 or 5 awn-like deciduous
scales. n = 7. One species, Stokesia laevis (L.)
Greene, south-eastern United States.
VI.11. Subtribe Pacourininae H. Rob. (1999).
Aquatic herbs. Heads axillary, large. Anther appendages sclerified, glabrous; style without node,
sweeping hairs acicular. Achenes with corky surface. Pollen triporate, lophate, psilate, emicropunctate.
Only one genus:
236. Pacourina Aubl.
Pacourina Aubl., Hist. Pl. Guiane 2: 800, t. 316 (1775).
Hairs simple, small. Leaf bases clasping, margins
spinose-toothed. Heads single and sessile in axils;
involucral bracts c. 50 in c. 3 series, broad, green,
margins whitish. Florets c. 50; corolla lobes sclerified distally; anther tails toothed. Achene c. 10costate, idioblasts in furrows, raphids none; pappus bristles short, deciduous, squamellae persistent. One species, P. edulis Aubl., Central and South
America south to Argentina.
VI.12. Subtribe Erlangeinae H. Rob. (1999).
Involucral bracts persistent. Anther appendages
thin (sclerified in Acilepidopsis), glabrous; sweeping hairs usually acicular. Achene raphids usually
elongate. Pollen tricolporate and non-lophate
or often triporate, lophate with irregular polar
165
lacunae, sometimes emicropunctate. Chemistry:
5-alkylcoumarins, glaucolides, guaianolides.
Key to the Genera
1. Receptacles with few to many pales
2
– Receptacles without pales
5
2. Pappus of separate or partially united segments; pollen
lophate, triporate
3
– Pappus lacking, a crenulated corona or a cupule; pollen
non-lophate, tricolporate
4
3. Pales only at periphery of receptacle; stem hairs
spreading; leaves opposite or whorled, alternate
above; corolla lobes with many hairs; florets c. 15 in
a head
239. Ageratinastrum
– Receptacle with pales throughout; stem hairs
sericeous; leaves spirally inserted; corolla lobes with
few hairs; florets c. 80 in a head 247. Dewildemania
4. Achenes 10-ribbed, densely glanduliferous; petioles
narrow, 1–2 cm long; corolla lobes sericeous with stiff
hairs; florets 10–15 in a head; corollas whitish
248. Diaphractanthus
– Achenes 5-ribbed, glabrous; petioles short; corolla
lobes hairless; florets c. 35 in a head; corollas purple
261. Omphalopappus
5. Stem hairs symmetrically T-shaped
6
– Hairs of stems simple or asymmetrical or lacking 13
6. Pappus lacking, coroniform, or of few short prongs or
deciduous setae, not with many long bristles
7
– Pappus of many capillary bristles, at least as long as
basal tube of the corolla
10
7. Leaves filiform; anther thecae long and narrow, linear,
to 3 mm long
264. Paurolepis
– Leaves with broadened blades; anther thecae not linear
8
8. Leaves sessile; corolla lobes usually with few to many
T-shaped hairs; achenes usually glabrous
251. Gutenbergia
– Leaves petiolate; corolla lobes without T-shaped hairs;
achene with tips of setulae anchor-shaped or hooked
9
9. Pappus none; corolla lobe tips with strong spreading
spines
255. Iodocephalis
– Pappus of 4 or 5 short prongs; corolla lobe tips without
spines
259. Msuata
10. Weak trees or shrubs; achenes 10-ribbed; T-shaped
hairs often with long, multiseptate stalks 240. Ambassa
– Herbs; achenes 4- or 5-ribbed or terete; T-shaped hairs
usually with short stalks
11
11. Annuals or short-lived perennials; pollen lophate, triporate
245. Cyanthillium
– Perennials; pollen non-lophate, tricolporate
12
12. Corolla lobes with many symmetrical T-shaped hairs;
anthers with short or no tails
253. Hilliardiella
– Corolla lobes without symmetrical T-shaped hairs; anthers with basal tails
263. Orbivestus
13. Achenes with 4–6 ribs or costae
14
– Achenes with 8–12 angles or ribs, rarely smooth and
slightly obcompressed
21
14. Pappus of copious long capillary bristles
15
– Pappus with only a ring of free or united scales,
a corona, or absent, with few or no inner caducous
bristles, without numerous long bristles
16
166
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
15. Annual herbs; hairs asymmetrically T-shaped; anther
266. Polydora
bases without tails; n = 9
– Perennial herbs; hairs not T-shaped; anther bases with
tails
269. Vernoniastrum
16. Leaves deeply pinnatifid; achenes densely minutely tuberculate
267. Rastrophyllum
– Leaves not deeply pinnatifid; achenes not densely
minutely tuberculate
17
17. Outer pappus of partially to totally free scales; inner
pappus of short brownish bristles 243. Brachythrix
– Pappus absent or with outer pappus a ring or corona
and inner pappus white
18
18. Involucral bracts greenish with sclerified pale or reddish margins; leaves alternate or opposite or whorled;
achenes with long and broad ribs; pollen tricolporate,
lophate or non-lophate
242. Bothriocline
– Involucral bracts with margin thinner or less differentiated; leaves alternate; achenes with short or narrow
ribs; pollen sometimes triporate
19
19. Corolla glabrous; pappus a high collar or lacking
250. Ethulia
– Corolla lobes with hairs; pappus often with caducous
bristles
20
20. Heads with 20–100 florets; leaves often broadly sessile
(Africa)
249. Erlangea
– Heads with 4 florets; leaves narrow (Brazil)
268. Telmatophila
21. Pappus short, usually not as long as basal tube of
corolla
22
– Pappus of many capillary bristles nearly as long as
corolla
26
22. Inflorescence with one or more dense glomeruli of
sessile heads; pappus coroniform
260. Muschleria
– Inflorescence not densely glomerulate; pappus of
scales or short bristles or lacking
23
23. Pappus of short scales; pollen tricolporate, nonlophate; achenes with few setulae
246. Decastylocarpus
– Pappus of short bristles or lacking; pollen triporate,
lophate; achenes glabrous
24
24. Lower involucral bracts spreading and foliose or with
spreading and foliose tips; achenes somewhat obcompressed, strongly rounded at top to a narrow corolla
and pappus insertion; raphids subquadrate
265. Phyllocephalum
– Lower involucral bracts not spreading and foliose nor
with reflexed foliose tips, with only narrow reflexed
herbaceous tips; achenes prismatic, not narrowed at
top below pappus and corolla; raphids oblong to elongate
25
25. Corolla lobes reflexed when mature; anther thecae
partially exserted, pale; style base with distinct node;
Africa
256. Kinghamia
– Corolla lobes not reflexed; anther thecae strongly
exserted, blackish; style base without node; Thailand
257. Koyamasia
26. Pappus bristles subplumose
262. Oocephala
– Pappus bristles only scabrid or barbellate
27
27. Heads with 4–13 florets; anther thecae with broad or
short tails
28
– Heads with 25–100 florets; anther bases without tails
30
28. Leaves mostly in basal rosette, large, tomentose below;
involucral bracts with spine-tips 254. Hystrichophora
– Leaves mostly cauline; involucral bracts without spine
at tip
29
29. Plants with decumbent bases; heads in obvious cymes;
stems, leaves, involucral bracts, corollas, and achenes
with many yellowish or reddish glandular dots; anther
bases with sharp sclerified tails, with sclerified apical
appendage; South America
237. Acilepidopsis
– Plants erect from xylopodia; heads in clusters at
nodes; plants without numerous reddish or yellowish
glands; anther tails and appendages not sclerified;
Africa
244. Cabobanthus
30. Plants glabrous; heads pedunculate from middle of internodes of inflorescence branches; leaves with sessile
auriculate bases; South America 258. Mesanthophora
– Plants pubescent; heads sessile or peduncles from axils
of bracts or leaves; leaves usually petiolate at base; Asia
or Africa
31
31. Inflorescences scapiform; stems and bracts with some
red stipitate glands with multicellular tips
241. Bechium
– Inflorescences with many heads sessile or in panicles;
without red stipitate glands with multicellular tips 32
32. Outer pappus of short bristles, not broad scales; hairs
of stems not angled, with tips erect; setulae of achenes
split to base into unequal halves; Asia 238. Acilepis
– Outer pappus of short broad scales or squamellae;
hairs of stems L-shaped, transverse cap cells mounted
at one end; setulae of achene s not split to base into
unequal halves
266. Polydora
Genera of Erlangeinae
237. Acilepidopsis H. Rob.
Acilepidopsis H. Rob., Phytologia 67: 289 (1989).
Perennial herbs decumbent from thickened rhizome; with numerous glandular dots, hairs forming felt on stems. Inflorescence branches seriately
cymose. Heads sessile; involucral bracts c. 30. Florets 8–13, corolla lobes reflexed, sericeo-pilosulous;
anther bases and appendages sclerified; style without node. Achene 10-costate; raphids lacking; pappus capillary. Pollen triporate, lophate, emicropunctate. One species, A. echitifolia (Mart. ex DC.)
H. Rob., Argentina, Bolivia, Brazil, Paraguay.
238. Acilepis D. Don.
Acilepis D. Don., Prodr. Fl. Nepal 169 (1825).
Lysistemma Steetz (1864).
Xipholepis Steetz (1864).
Erect perennial herbs, stem hairs multiseptate at
base. Heads pedunculate or sessile and single at
nodes; involucral bracts 50–200 in 6–12 series. Florets 25–80; corolla lobes gland-dotted; anther base
blunt; style with node. Achene 8–10-ribbed, setulae
halves very unequal; pappus capillary, pollen tripo-
Compositae
rate, emicropunctate. Ten or more species, mostly
India.
239. Ageratinastrum Mattf.
Ageratinastrum Mattf., Notizbl. Bot. Gart. Berlin-Dahlem
11: 412 (1932), nom. nov.
Ageratina O. Hoffm. (1900), non Ageratina Spach (1841).
Shrubs or subshrubs; indumentum heterotrichous, hairs rarely symmetrically T-shaped.
Leaves mostly opposite or whorled. Inflorescences
corymbose. Heads sessile; involucral bracts c. 30
in 3–4 series; pales few. Florets c. 15; corolla with
sinuous hairs; anther bases rounded; style with
node. Achene 4–5-ribbed; pappus scales united or
free, sometimes with 5–7 setae. Pollen triporate,
lophate, emicropunctate. Five species, tropical
Africa.
167
gins paler or redder. Florets 3–100; anther base
rounded; style with node. Achenes with 4 massive ribs, idioblasts covering furrows, raphids subquadrate; pappus with caducous setae or collarform. Pollen tricolporate, echinolophate, or nonlophate. n = 20 . Thirty or more species, tropical
Africa.
243. Brachythrix Wild & Pope
Brachythrix Wild & Pope, Kirkia 11: 25 (1978).
Perennial herbs; hairs simple, multiseptate, sometimes unevenly T-shaped. Heads solitary or in cymose clusters; involucral bracts c. 65 in c. 5 series. Florets c. 40; anther bases with sterile lobe;
style with node. Achenes 4–5-angled, raphids subquadrate; pappus bristles short, deciduous, outside with scales or continuous corona. Pollen nonlophate. Six species, Africa.
240. Ambassa Steetz
Ambassa Steetz in Peters, Naturwissensch. Reise Mossambique Bot. 363 (1864).
Weak shrubs or small trees; hairs long-stalked,
symmetrically T-shaped. Peduncles 2–10 mm long.
Involucral bracts c. 40 in c. 4 series. Florets 20–25;
anther bases rounded; style with node. Achene 10costate, setuliferous, idioblasts numerous; pappus
of fragile bristles, narrowed below, persistent squamulae. Pollen triporate, lophate, subpsilate. Two
species or more, eastern Africa.
244. Cabobanthus H. Rob.
Cabobanthus H. Rob., Proc. Biol. Soc. Wash. 112: 228
(1999).
Perennial herbs, xylopodial; hairs simple, multiseptate. Heads clustered at upper axils; involucral
bracts c. 35 in c. 5 series. Florets c. 10; anthers shorttailed; style with node. Achenes 8–10-costate, with
many equal-celled setulae; pappus capillary, with
shorter outer bristles. Pollen triporate, lophate,
emicropunctate. Two or more species, tropical
Africa, Tanzania to Zambia.
241. Bechium DC.
Bechium DC., Prodr. 5: 70 (1836).
Erect or horizontally proliferating herbs; hairs
sericeous, red stipitate glands with multicellular
tips on stems and bracts. Inflorescences scapiform;
heads 1 to many; involucral bracts c. 30 in c.
3 series. Florets 25–50; anther bases rounded,
style with node; sweeping hairs pointed, septate.
Achenes 10-costate, setulae uneven, many idioblasts; pappus capillary, with squamae. Pollen
non-lophate. Two or more species, Madagascar.
242. Bothriocline Oliv. ex Benth.
Bothriocline Oliv. ex Benth., Hooker’s Icon. Pl. 12: 1133
(1873).
Volkensia O. Hoffm. (1894).
Annual or perennial; hairs simple or asymmetrically T-shaped. Leaves alternate, opposite or ternate. Involucral bracts 25–50 in 3–5 series, mar-
245. Cyanthillium Blume
Cyanthillium Blume, Bidjr. Fl. Ned. Ind. 889 (1826).
Isonema Cass. (1817), nom. illegit., non R. Br. (1810).
Cyanopsis Blume ex DC. (1836), nom. illegit. et superfl.
Vernonia Schreb. sect. Tephrodes DC. (1836).
Seneciodes L. ex T. Post & O. Kuntze (1903).
Triplotaxis Hutch. (1914).
Short-lived herbs; hairs asymmetrically or symmetrically T-shaped. Leaves narrowly petiolate,
membranaceous. Heads pedunculate; involucral
bracts, c. 30 in 3(–5) series, green. Florets 15–94;
corolla with straight hairs; anthers without tails;
style with node. Achenes 5-ribbed or terete,
with idioblasts; pappus capillary, slender-tipped,
squamellae persistent, rarely with callose ring.
Pollen triporate, echinolophate. n = 9, 11, 18.
More than seven species, Indian Ocean, Indonesia,
south-eastern and eastern Asia, tropical Africa,
widely adventive.
168
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
246. Decastylocarpus Humb.
Decastylocarpus Humb., Mém. Soc. Linn. Normandie 25:
30, 280 (1923).
Herbs; stem hairs multiseptate or with erect cap
cells. Leaves without glands. Heads few, subsessile.
Involucral bracts 35 in c. 4–5 series, green. Florets 25–30; corollas with glands, lobes sericeous;
anthers without tails; style without node. Achenes
10-costate, no idioblasts, many glands, few setulae,
raphids subquadrate; pappus coroniform, dentate.
Pollen non-lophate. One species, D. perrieri Humbert, Madagscar.
247. Dewildemania O. Hoffm.
Dewildemania O. Hoffm. in De Wild., Ann. Mus. Congo
sér. 1, Bot. sér. 4, 1: 10 (1903).
Perennial herbs; hairs simple, multiseptate. Heads 1
to many; peduncles bracteiferous; involucral bracts
60–80 in 2–7 series; receptacle paleaceous. Florets c.
80; corollas with stipitate glands; anther untailed;
style with node. Achenes 4–5-sided, setuliferous,
with idioblasts, raphids subquadrate; 5 or more
long pappus scales and alternating broad scales.
Pollen triporate, lophate, emicropunctate. Seven
species, tropical Africa.
248. Diaphractanthus Humb.
Diaphractanthus Humb., Mém. Soc. Linn. Normandie 25:
31, 280 (1923).
Annual herbs; stems sericeous. Petioles narrow,
blade with glands. Heads crowded, pedunculate;
involucral bracts c. 30, in c. 4 series, green, margins scarious; receptacle paleaceous. Florets 10–15;
corollas with glands, lobes sericeous; anthers untailed; style with node. Achenes 10-costate, glands
dense, no idioblasts, raphids not visible; pappus
4–5-dentate, coroniform. Pollen non-lophate. One
species, D. homolepis Humb., Madagascar.
249. Erlangea Sch. Bip.
Erlangea Sch. Bip., Flora 36: 34 (1853).
Stephanolepis S. Moore (1900).
Haarera Hutch. & E.A. Bruce (1932).
Short-lived herbs; hairs simple, multicellular.
Leaves sessile. Heads pedunculate; involucral
bracts 20–200 in 3–7 series. Florets c. 25–100+;
corolla tips with hairs and often stalked glands;
anther slightly tailed; style with node. Achenes
3–6-angled, mostly glabrous; pappus lacking or
with caducous setae. Pollen triporate, lophate,
emicropunctate, or tricolporate, non-lophate.
n = 10. Nine or more species, tropical Africa.
250. Ethulia L. f.
Ethulia L. f., Dec. Pl. Horti Upsal. 1 (1762); Gilbert & Jeffrey,
Kew Bull. 43: 165–193 (1988), rev.
Hoehnelia Schweinf. (1892).
Short-lived herbs; stem hairs with erect cap cells.
Leaves short-petiolate. Inflorescence cymose,
heads shortly pedunculate; involucral bracts 15–40
in 2–3 usually subequal series. Florets 3–100;
corollas with glands, apices glabrous; anther
bases plain; style without node; sweeping hairs
short-acute. Achenes 2–6-ribbed, glanduliferous,
raphids short-oblong; pappus lacking or coroniform. Pollen non-lophate. Nineteen species,
Africa, southern Asia, Indonesia, one species
widely adventive.
251. Gutenbergia Sch. Bip. ex Walp.
Gutenbergia Sch. Bip. ex Walp., Repert. 2: 703 (1843).
Erlangea Sch. Bip. sect. Platylepis S. Moore (1902).
Short-lived herbs; stem hairs symmetrically
T-shaped, arms filiform. Leaves opposite, ternate
or alternate. Heads pedunculate; involucral bracts
20–40, in 3–5 series. Florets 4–50; corollas usually
with T-shaped hairs; anther bases rounded; style
scarcely nodular. Achenes 4–5, 8–12-costate,
usually glabrous, raphids subquadrate; pappus
lacking or short deciduous setae. Pollen triporate,
echinolophate. n = 10. At least 13 species, Africa.
252. Herderia Cass.
Herderia Cass., Ann. Sci. Nat. 17: 421 (1829).
Procumbent annuals; finely sericeous. Leaves
small. Heads solitary, terminal; involucral bracts
3–4-seriate, subequal, outer 15–25 more filiform,
25–30 inside. Florets c. 30; corollas 1.5 mm long,
minutely stipitate-glanduliferous; anther base
blunt; style with node. Achenes 4-ribbed, glabrous, idioblasts reddish; pappus of 5–9 scales
and 3–5 short bristles. Pollen triporate, lophate.
One species, H. truncata Cass., Senegal, Gambia,
Ghana, Nigeria, tropical western Africa.
253. Hilliardiella H. Rob.
Hilliardiella H. Rob., Proc. Biol. Soc. Wash. 112: 229 (1999).
Perennial herbs; stems with symmetrically Tshaped hairs. Heads pedunculate; involucral bracts
Compositae
169
c. 25–40 in 3–4 series, subpersistent. Florets 12–20;
corollas with T-shaped hairs; anthers scarcely
tailed; style with node. Achenes 4–5-costate,
densely setuliferous and idioblastiferous; pappus
setae slender, subpersistent, with shorter outer
series. Pollen non-lophate. n = 9, 10. Eight or more
species, mostly southern and eastern Africa.
Achenes strongly 10-ribbed, glabrous, idioblasts
in grooves; pappus with short deciduous setae or
none. Pollen triporate, echinolophate. Five species,
tropical Africa.
254. Hystrichophora Mattf.
Perennial herbs; stem hairs simple, multiseptate.
Petioles narrow. Heads solitary, long-pedunculate;
involucral bracts c. 90 in 4–5 series, mostly
herbaceous and reflexed. Florets c. 90; anther
thecae strongly exerted, dark, bases rounded;
style without node. Achene 10-costate, glabrous,
with idioblasts, raphids minute, oblong; pappus
setae short, deciduous. Pollen triporate, lophate,
emicropunctate. 2n = 54. One species, K. calcarea
(Kitamura) H. Rob., Thailand.
Hystrichophora Mattf., Notizbl. Bot. Gart. Berlin-Dahlem
13: 288 (1936).
Herbs with basal rosette. Leaves to 40 ×25 cm,
margins spinulose-toothed, tomentellous below. Inflorescences axillary, appearing terminal,
branches often paired, one subsessile, with
glomerules of few heads, adaxial branch pedunculate, with globose glomerules of 30–50 narrow
heads; involucral bracts 6–7-seriate, spine-tipped.
Florets 5–6; corollas glabrous; anther base with
sterile margin; style with node (?). Achenes
10-costate, setuliferous, with idioblasts, raphids
elongate or lacking; inner pappus squamae c. 12,
caducous, 4–5 mm long, squamules persistent.
Pollen triporate, lophate. One species, H. macrophyllum Mattf., Tanzania (Lindi), Mueraplateau
(Rondo).
255. Iodocephalis Thorel ex Gagnep.
Iodocephalis Thorel ex Gagnep., Notul. Syst. (Paris) 4: 16,
24 (1920).
Annuals; hairs symmetrically T-shaped. Petioles
to 5 mm, blades glandular-dotted. Peduncles
5–10 mm long; involucral bracts c. 20, 3–7-seriate,
acuminate, arachnoid-pubescent. Florets 25–70;
corolla tips strongly spinose; anther bases obtuse
to acuminate; style without node. Achenes 5–7–
10-costate, with glands, ribs setuliferous, setulae
anchor-shaped or hooked, raphids subquadrate;
pappus none. Pollen triporate, lophate. Two
species, Laos, Thailand and Vietnam, mostly along
banks of Mekong River.
256. Kinghamia C. Jeffrey
Kinghamia C. Jeffrey, Kew Bull. 43: 274 (1988); Robinson,
Proc. Biol. Soc. Wash. 112: 220–247 (1999), rev.
Perennial herbs; hairs simple, multiseptate. Heads
pedunculate; involucral bracts 100–200 in 5–6 series, outer subulate, inner broad, apiculate. Florets
50–80; corolla lobe tips recurved; anthers pale,
partially exerted, base obtuse; style with node.
257. Koyamasia H. Rob.
Koyamasia H. Rob., Proc. Biol. Soc. Wash. 112: 234 (1999).
258. Mesanthophora H. Rob.
Mesanthophora H. Rob., Novon 2: 172 (1992).
Glabrous perennial herbs. Leaves sessile, shortly
decurrent, auriculate, glandular-dotted below.
Racemiform leafy cymes with peduncles from
middle of internodes; involucral bracts c. 100 in
c. 5 series, linear-lanceolate, erect-patent. Florets
90–100; anther bases rounded; style with node.
Achenes 8–10-costate, costae scabrid; pappus bristles deciduous, capillary, squamellae persistent.
Pollen triporate, lophate, emicropunctate. Two
species, Bolivia, Paraguay.
259. Msuata O. Hoffm.
Msuata O. Hoffm. in Engler & Prantl, Nat. Pflanzenfam.
IV/5: 388 (1894).
Subshrubs; hairs symmetrically T-shaped. Petioles
narrow. Heads short-pedunculate; involucral
bracts 16–20, 2-seriate. Florets c. 26; corollas glanduliferous; anther bases rounded; style without
node; sweeping hairs round-tipped. Achenes 4- or
5-ribbed, sides with glands and idioblasts, setulae
beside ribs, with anchor-like points, raphids subquadrate; pappus crown with 4 or 5 short prongs.
Pollen non-lophate. One species, M. buettneri O.
Hoffm., tropical Africa, Congo.
260. Muschleria S. Moore
Muschleria S. Moore, J. Bot. 52: 89 (1914).
Rhizomatous shrubs; hairs simple, multiseptate.
Leaves linear, tomentose below. Inflorescence ter-
170
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
minal, 1 to few glomeruli of sessile heads. Involucral
bracts c. 20 in 4–5 series. Florets 4–6; corolla glanduliferous, lobes with multiseriate spines; anther
bases sagittate, not tailed; style with node. Achenes
10-costate, glands between costae, idioblasts numerous; pappus coroniform. Pollen non-lophate.
n = 9. One species, M. angolensis S. Moore, Angola, southern tropical Africa.
261. Omphalopappus O. Hoffm.
Omphalopappus O. Hoffm. in Engler & Prantl, Nat. Pflanzenfam. IV/5: 234 (1890–1891).
Gossweilera S. Moore (1908).
Perennial herbs; hairs simple, multiseptate. Inflorescences virgately corymbiform. Heads pedunculate, hemispherical; involucral bracts c. 50 in c. 3 series, many subequal; receptacle paleaceous. Florets
c. 35; anther thecae reddish, bases plain; style thickened without node. Achenes 5-ribbed, glabrous,
without idioblasts, raphids subquadrate; pappus
a denticulate corona or cupule. Pollen non-lophate.
Two or three species, Angola.
262. Oocephala (S.B. Jones) H. Rob.
Oocephala (S.B. Jones) H. Rob., Proc. Biol. Soc. Wash. 112:
281 (1999).
Vernonia Schreb. subsect. Oocephalae S.B. Jones (1981).
Shrubs; hairs simple, weakly L-shaped. Inflorescences corymbiform. Heads shortly pedunculate;
involucral bracts 35–40 in c. 7 series. Florets c. 15;
anther base rounded; style base with ring. Achenes 8-ribbed, setuliferous on ribs; idioblasts in
furrows; pappus biseriate, outer shorter, inner setiform, subplumose. Pollen triporate, lophate. Two
species, tropical Africa.
263. Orbivestus H. Rob.
Orbivestus H. Rob., Proc. Biol. Soc. Wash. 112: 230 (1999).
Vernonia Schreb. subg. Orbisvestus S.B. Jones (1981).
Shrubs or subshrubs; hairs T-shaped. Leaf blade
sparsely pilose. Inflorescence laxly corymbiform.
Heads pedunculate; involucral bracts 25–30 in c. 4
series. Florets 8–16; corollas essentially glabrous;
anther bases tailed; style with node. Achenes 4–5costate, setuliferous, with idioblasts; pappus setae
deciduous, with persistent squamae. Pollen nonlophate. n = 9, 20. Four or more species, tropical
and southern Africa.
264. Paurolepis S. Moore
Paurolepis S. Moore, J. Bot. 55: 102 (1917).
Perennial herbs; hairs symmetrically T-shaped.
Leaves filiform. Inflorescence diffuse, peduncles
elongate. Involucral bracts c. 25, c. 4-seriate. Florets
c. 10; corollas sericeous, hairs T-shaped; anther
thecae linear, base rounded; style with node.
Achenes 4-sided, setulae on ribs, glands in furrows, raphids subquadrate; pappus scales 4, erose,
subdeciduous. Pollen triporate, echinolophate.
Three species, Africa.
265. Phyllocephalum Blume
Phyllocephalum Blume, Bijdr. Fl. Ned. Ind. 888 (1826); Kirkman, Rhodora 83: 1–24 (1981), rev.; Robinson, Proc. Biol.
Soc. Wash. 112: 220–247 (1999), emend.
Decaneurum DC. ex Wight (1833), non Decaneurum DC.
(1833).
Lamprachaenium Benth. in Benth. & Hook. f. (1873).
Short-lived herbs; hairs flagelliform, multicellular.
Leaf blades white-tomentose below. Heads 1 to few,
pedunculate; with spreading, foliaceous subinvolucre or tips of outer involucral bracts; involucral
bracts c. 40, subequal, 2–3-seriate. Florets 75–150;
anther bases sharp, diverging; style with node. Achenes obcompressed, oblong, narrowed to corolla,
glabrous, 10-costate or smooth and shiny, raphids
subquadrate; pappus bristles short, coloured, caducous. Pollen triporate, lophate, emicropunctate.
n = 9. Circa ten species, India, Java.
266. Polydora Fenzl
Polydora Fenzl, Flora 27: 312 (1844).
Crystallopollen Steetz (1863).
Annuals; hairs L-shaped. Inflorescence laxly
thyrsoid-paniculate. Heads pedunculate; involucral bracts c. 80, c. 7-seriate, margins scarious,
tips black. Florets c. 30; corolla lobes hairless;
anthers without tails; style with node. Achenes
5–8–10-ribbed, setuliferous, idioblasts scattered;
pappus bristles greenish, yellowish or tawny, rarely
white, with squamellae. Pollen triporate, lophate.
n = 9. Eight species, tropical Africa.
267. Rastrophyllum Wild & Pope
Rastrophyllum Wild & Pope, Kirkia 10: 325 (1977).
Annuals; hairs eccentrically T-shaped. Leaves
deeply pinnatifid. Heads in corymbiform clusters,
peduncles to 5 mm. Involucral bracts 30–40 in
Compositae
2–5 series. Florets 25–35(?); corolla tubes covered
with stipitate glands. Achenes 4-sulcate, surface
minutely tuberculate; pappus lacking. Two species,
tropical Africa.
268. Telmatophila Mart. ex Baker
Telmatophila Mart. ex Baker in Mart., Fl. Brasil. 6, 2: 170
(1873).
Perennials; sericeous with simple hairs. Inflorescences axillary, secondary heads surrounded by
foliaceous bracts. Heads cylindrical; subequal involucral bracts and florets 4. Corollas with long
hairs distally; style base without node. Achenes
oblong-ovoid, c. 8-costate, with idioblasts, long setulae deeply divided, raphids elongate; pappus of
c. 8 short, laciniate scales. Pollen triporate. One
species, T. scolymastrum Mart. ex Baker, northeastern Brazil.
269. Vernoniastrum H. Rob.
Vernoniastrum H. Rob., Proc. Biol. Soc. Wash. 112: 233
(1999).
Vernonia Schreb. sect. Lepidella Oliv. & Hiern (1877), non
Lepidella Tiegh. (1911), nec Lepidella E.J. Gilbert (1925).
Perennial herbs; hairs simple-multiseptate or
slightly uneven at base. Heads single or grouped.
Involucral bracts c. 50, c. 3-seriate. Florets c. 50;
corolla pilose distally; anther tails short; style with
node. Achenes 4–5-angled, with setulae, idioblasts
often in transverse bands, raphids elongate;
pappus of capillary bristles and squamellae. Pollen
triporate, lophate. n = 10. Eight species, tropical
Africa.
VI.13. Subtribe Centrapalinae H. Rob.
(1999).
Coarse herbs. Leaves alternate. Involucral bracts
persistent. Anther appendages indurated, glabrous;
sweeping hairs pointed. Pollen tricolporate, usually
lophate. Characteristic sesquiterpenes: elemanolides.
Key to the Genera
1. Pappus coroniform, of short deciduous bristles or of
only small scales, not of long bristles
2
– Pappus of 5-numerous long capillary or flattened bristles
4
2. Receptacle with pales; Australia 278. Pleurocarpaea
– Receptacle without pales; Asia
3
171
3. Hairs of stems simple, multiseptate; pappus none or
an annulus or corona
270. Adenoon
– Hairs of stems mixed, simple plus T-shaped; pappus
a lobate corona or of short deciduous segments
273. Camchaya
4. Pappus of flattened bristles; corollas purple to white;
style without basal node, with high sheathing nectary
5
– Pappus of capillary bristles; corollas violet or blue;
style base with node, nectary short
6
5. Corolla lobes shorter than throat; pappus bristles nu272. Baccharoides
merous; pollen lophate; n = 10
– Corolla lobes about as long as throat; pappus bristles
c. 5; pollen not lophate
277. Neurolakis
6. Leaves in basal rosette; West Indies 275. Lachnorhiza
– Leaves not restricted to basal rosette; Africa
7
7. Heads surrounded by greatly enlarged foliiform bracts
below involucre; leaves often with longitudinal venation
271. Aedesia
– Heads not surrounded by greatly enlarged foliiform
subinvolucral bracts; venation never longitudinal 8
8. Achenes with weak ribs or costae, without lines of
idioblasts along costae; pollen lophate without spurs
projecting into colpi; corollas lavender or purple to
274. Centrapalus
blue; n = 9
– Achenes with strong ribs or costae, usually with lines
of idioblasts crowded along sides of costae; pollen
lophate with spur walls projecting into colpi; corollas blue; n = 10
276. Linzia
Genera of Centrapalinae
270. Adenoon Dalzell
Adenoon Dalzell, in Hooker’s J. Bot. Kew Gard. Misc. 2: 344
(1850).
Perennials; hairs simple, multiseptate. Inflorescence corymbose. Heads long-pedunculate;
involucral bracts c. 50 in 4–5 series; receptacle
fimbriate. Florets c. 50; corolla tube pilose; anthers
shortly tailed; style without node, nectary long;
sweeping hairs acicular. Achene 10-costate, glabrous, raphids elongate; pappus none. Pollen with
spur walls, lophate, without polar lacunae. One
species, A. indicum Dalzell, India.
271. Aedesia O. Hoffm.
Aedesia O. Hoffm., in Engler & Prantl, Nat. Pflanzenfam.
Nachtr. 1: 321 (1897).
Perennials, glabrous. Solitary terminal heads surrounded by large leaves; involucral bracts c. 50 in
4–5 series, apical margin toothed; receptacle with
few pales. Florets c. 50(?); corolla tube long, throat
none; anther bases rounded; style without node;
sweeping hairs acicular. Achene 10-costate, setulose, raphids subquadrate; pappus capillary, multi-
172
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
seriate. Pollen echinolophate. Three species, tropical Africa.
272. Baccharoides Moench
Baccharoides Moench, Methodus 528 (1794); Isawumi et al.,
Grana 35: 205–230 (1996), tax.
Ascaricida Cass. (1817), nom. illegit. superfl.
Candidea Tenore (1839).
Stengelia Sch.Bip. ex Steetz (1864).
Perennials, hairs short-stalked, cap-cells erect.
Involucral bracts 25–100, 4–8-seriate, apices
often expanded. Florets 25–100; corolla base
slender, abruptly expanded into cylindrical limb,
throat as long as erect lobes; anther tails small;
style without node, nectary tubular; sweeping
hairs acicular. Achenes 8–20-ribbed, ribs usually
setulose, raphids elongate; pappus pluriseriate,
setae flattened, sometimes with squamellae. Pollen
echinolophate, with polar lacuna. n = 10. Circa 25
species, tropical Africa, southern Asia.
273. Camchaya Gagnep.
Camchaya Gagnep., Notul. Syst. (Paris) 4: 14 (1920).
Koyama, Acta Phytotax. Geobot. 35: 49–58 (1984), rev.
Thorelia Gagnep. (1920), nom. cons., non Thorelia H.F.
Hance (1877), nom. rej.
Thoreliella C.Y. Wu (1975), nom. nov. for Thorelia Gagnep.
Annuals; hairs T-shaped and simple. Heads few,
pedunculate; involucral bracts 20–130, 3–5-seriate,
herbaceous and spinose only at tips. Florets
12–130; anther bases rounded; style without node,
nectary long; sweeping hairs acicular. Achenes
10-costate, with glands, many idioblasts, raphids
short-rhomboid; pappus with corona or short
deciduous segments; pollen triporate with crosswalls, echinolophate. Five species, Thailand, China
(Yunnan), Laos.
274. Centrapalus Cass.
Centrapalus Cass., Bull. Soc. Philom. Paris 1817: 10 (1817).
Vernonella Sonder (1850).
Annuals or perennials, often scapigerous; stem
hairs simple, multiseptate. Involucral bracts 125–
150, c. 5-seriate, linear, green, often denticulate.
Florets c. 100; corollas blue to purplish, lobes
sometimes fringed; anther not tailed; style with
node; sweeping hairs acute. Achenes weakly
10-costate, setuliferous, raphids oblong; pappus
capillary. Pollen lophate or non-lophate. n = 9.
Nine species or more, Africa.
275. Lachnorhiza A. Rich.
Lachnorhiza A. Rich. in Sagra, Hist. Fis. Cuba, Bot. 11: 34
(1850).
Rosettiform perennials; many cells of longest hairs
basally spurred. Peduncles long, with 1 or 2 heads.
Involucral bracts c. 25, c. 3-seriate. Florets 10–
15; corollas glanduliferous; anther bases obtuse;
style without node; sweeping hairs acicular. Achenes 10-costate, with glands, raphids oblong; pappus bristles persistent, few spicules. Pollen echinolophate, emicropunctate. One species, L. piloselloides A. Rich., Cuba.
276. Linzia Sch. Bip. ex Walpers
Linzia Sch. Bip. ex Walpers, Rep. 2: 948 (1843).
Vernonia Schreb. sect. Azureae S.B. Jones (1981).
Perennials; hairs simple, multicellular. Involucral
bracts 50–150 in 5–6 series, pectinate-denticulate
along upper margins. Florets c. 20–50; corollas
bluish, lobes apically strigulose; anther base
rounded; style base with annulus; sweeping hairs
sharp. Achenes strongly 10-costate, setuliferous,
costae often bordered with idioblasts, raphids
subquadrate; pappus bristles persistent, outer
short. Pollen lophate, emicropunctate. n = 10.
Seven or more species, Africa.
277. Neurolakis Mattf.
Neurolakis Mattf., Bot. Jahrb. Syst. 59 Beibl. 133: 12 (1924).
Unbranched perennials; hairs stalked-L-shaped
plus simple multiseptate. Heads 1 to few, terminal,
large, pedunculate; involucral bracts c. 60, c.
4-seriate, subequal. Florets c. 50; corolla tube
filiform, abruptly campanulate above; anther tails
broad; style without node; nectary sheathing;
sweeping hairs acicular. Achenes 10-ribbed,
setuliferous, raphids elongate; pappus bristles c.
5, short, flattened, caducous. Pollen non-lophate.
One species, N. modesta Mattf., Cameroon, Chad.
278. Pleurocarpaea Benth.
Pleurocarpaea Benth., Fl. Austral. 3: 460 (1867).
Perennials; hairs simple, submoniliform, rather Lshaped. Heads few, pedunculate; involucral bracts
c. 12, 2–3-seriate; receptacle paleaceous. Florets
8–13; anther bases rounded; style without node;
sweeping hairs acute. Achenes 10-grooved, furrows
with hairs and glands, raphids lacking; pappus bristles few, short, deciduous, outer fimbriae persis-
Compositae
tent. Pollen echinolophate, with polar lacuna. One
species, P. denticulata Benth., northern Australia.
VI.14. Subtribe Gymnantheminae H. Rob.
(1999).
Shrubs, vines or trees. Leaves alternate. Inner
involucral bracts sometimes deciduous. Corolla
lobes sometimes reflexed. Anther appendage indurate, glabrous; sweeping hairs often blunt. Pollen
tricolporate, usually non-lophate. Characteristic
sesquiterpenes: elemanolides and guaianolides.
Key to the Genera
1. Apomictic, with styles not or scarcely divided at tip,
stigmatic tissue lacking; outer surfaces of style only
with ends of surface cells protruding (prorulose) or
with short spicules, without sweeping hairs; pollen
with short spinules (Hawaii)
283. Hesperomannia
– Sexually reproducing plants with style branches
well-developed; stigmatic tissue on whole inner
surface of style branches; long sweeping hairs present;
pollen with large spines or lophate with murae in
well-developed reticulum
2
2. Pappus of many capillary or flattened bristles which
are shorter than or only about as long as the body of
the achene
3
– Pappus bristles or awns longer than bodies of achenes
4
3. Receptacles with pales
280. Centauropsis
– Receptacles without pales
286. Oliganthes
4. Heads with large, rounded, green, outer involucral
bracts; pollen lophate with three intercolpar-aligned
polar lacunae
285. Myanmaria
– Heads without large, rounded, green, outer involucral
bracts; pollen often not lophate, without intercolparaligned polar lacunae
5
5. Subshrubs; pappus bristles broad and flat, multiseriate; achenes sericeous with long uniseriate hairs
284. Lampropappus
– Shrubs, trees or vines; pappus bristles capillary, sometimes slightly flattened, mostly in 1–2 series; achenes
usually with short or biseriate setulae
6
6. Inflorescence pinnately branched, frondiform
279. Brenandendron
– Inflorescence not pinnate, not frondiform
7
7. Leaves often trinervate; corollas usually yellow; style
base with large abrupt node
281. Distephanus
– Leaves pinnately veined; corollas not yellow; style base
without node or with small gradually enlarged node
282. Gymnanthemum
Genera of Gymnantheminae
279. Brenandendron H. Rob.
Brenandendron H. Rob., Proc. Biol. Soc. Wash. 112: 244
(1999).
173
Trees; stems velutinous, hairs fusiform or
contorted. Leaves large, sometimes lobed. Inflorescences frondiform, branchlets racemiform
or spiciform. Involucral bracts to 110 in 7–9
series, innermost deciduous. Florets 30–50; anther
bases long-tailed; style with node; sweeping hairs
blunt-tipped. Achenes 8–10-costate, glabrous, with
idioblasts, raphids short to moderately elongate;
pappus capillary, subpersistent. n = 9. Three
species, tropical western Africa, Congo, Central
African Republic.
280. Centauropsis Bojer ex DC.
Centauropsis Bojer ex DC., Prodr. 5: 93 (1836).
Shrubs; stems lepidote or glabrous. Heads 1 to
few, subsessile to long-pedunculate; involucral
bracts 25–50, in 4–6 series, inner deciduous,
sometimes appendaged; receptacle paleaceous.
Florets 25–50+; corolla glabrous; anthers with
broad tails; style without node; sweeping hairs
subacute. Achenes 10- or 20-ribbed, glabrous,
raphids elongate; pappus capillary, shorter than
achene body. Eight species, Madagascar.
281. Distephanus Cass.
Distephanus Cass., Bull. Soc. Philom. Paris 1817: 151 (1817);
Robinson & Kahn, Proc. Biol. Soc. Wash. 99: 493–301
(1986), emend., list.
Gongrothamnus Steetz (1862).
Newtonia O. Hoffm. (1892), non Newtonia Baill. (1888).
Antunesia O. Hoffm. (1873), nom. nov. for Newtonia O.
Hoffm.
Shrubs, vines, or trees; hairs arachnoid, contorted,
or asymmetrically T-shaped. Leaves often trinervate. Inflorescences terminal; involucral bracts
21–24(–75), 4–6(–7)-seriate. Florets 10–16(–75);
corollas usually yellow, mostly glabrous; anther
tails often sclerified; style node abruptly enlarged;
sweeping hairs obtuse. Achenes (5–)10(–12)ribbed, raphids elongate; pappus capillary, with
squamellae, persistent. Pollen non-lophate or
lophate. Forty or more species, Africa, Indian
Ocean, India, south-western China.
282. Gymnanthemum Cass.
Gymnanthemum Cass., Bull. Soc. Philom. Paris 1817: 10
(1817).
Decaneurum DC. (1833), nom. illegit. superfl.
Monosis DC. in Wight (1834).
Vernonia Schreb. sect. Strobocalyx Blume ex DC. (1836).
Plectreca Raf. (1838) (“1836”).
174
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Strobocalyx (Blume ex DC.) Spach (1841).
Punduana Steetz (1864).
Shrubs or trees; stems often felted, hairs rarely
asymmetrically T-shaped. Inflorescences densely
corymbiform; involucral bracts 25–35, 4–5-seriate,
inner bracts often deciduous. Florets 1–50; anther
base broadly tailed; style usually with node;
sweeping hairs pointed or blunt. Achenes 5–10costate, raphids short, elongate or lacking; pappus
bristles persistent, capillary, with squamellae.
Pollen non-lophate or lophate. n = 10, 20, 2n = 20,
30. More than 43 species, Africa, southern Asia,
Indonesia.
New studies show that Monosis DC. in Wight
(1834), with c. seven species, is a separate genus
with obovate or oblanceolate leaves and lophate
pollen. Strobocalyx (Bl. ex DC.) Spach should also
be resurrected for various Asian and Malaysian
species.
283. Hesperomannia A. Gray
Hesperomannia A. Gray, Proc. Amer. Acad. Arts 6: 554
(1865).
Mostly apomictic shrubs or trees; hairs matted plus
some moniliform. Heads 2–10; involucral bracts c.
50 in 6–7 series. Florets c. 25; corollas yellow; anther tails long, blunt; apical appendages lanceolate;
style without node, tip mostly undivided, prorulose. Achenes glabrous, without raphids; pappus
capillary, longer than achene. Pollen minutely spiculiferous. Three species, Hawaii.
284. Lampropappus (O. Hoffm.) H. Rob.
Lampropappus (O. Hoffm.) H. Rob., Proc. Biol. Soc. Wash.
112: 245 (1999).
Vernonia Schreb. sect. Lampropappus O. Hoffm. (1896).
Vernonia Schreb. subsect. Turbinellae S.B. Jones (1981).
Subshrubs; stems tomentose, hairs with contorted
cells. Leaves tomentose below. Inflorescences
corymbiform. Involucral bracts c. 30, c. 4–5seriate, broad. Florets c. 20; corollas actinomorphic or inner lobe longest; anther bases rounded;
style with node; sweeping hairs obtuse. Achenes 5-costate, villous, hairs uniseriate, raphids
mostly subquadrate; pappus setae 3-seriate,
broad, flat. Three species, Angola, Congo, Malawi,
Zambia.
285. Myanmaria H. Rob.
Myanmaria H. Rob., Proc. Biol. Soc. Wash. 112: 244 (1999).
Shrubs; hairs simple, multiseptate. Inflorescences
corymbiform. Heads pedunculate; outer c. 20 involucral bracts broad, foliiform, 2–3-seriate; inner
bracts linear. Florets 35–50; anthers broadly tailed;
style without node; sweeping hairs obtuse. Achenes 10-costate, with idioblasts, costae setuliferous,
raphids subquadrate; pappus bristles 2–3-seriate,
persistent. Pollen echinolophate, with intercolparaligned polar lacunae. One species, M. calycina
(DC.) H. Rob., Myanmar.
286. Oliganthes Cass.
Oliganthes Cass., Bull. Sci. Philom. Paris 1817: 10 (1817).
Vernonia Schreb. sect. Trianthaea DC. (1836).
Trianthaea (DC.) Spach (1841).
Shrubs or trees; stem hairs simple, lacking, or
T-shaped with contorted cap-cells. Inflorescences
corymbiform. Involucral bracts 20–40, 4–6-seriate,
sometimes appendaged. Florets 3–150; corolla
throat half as long as lobes; anther base plain or
short-appendaged; style without node; sweeping
hairs acute, few septa. Achenes 10–15-costate,
glabrous, raphids subquadrate; pappus bristles
flattened, deciduous, fimbriate corona often
present. Nine species, Madagascar.
VI.15. Subtribe Trichospirinae Less. (1831).
Creeping herbs. Vegetative leaves alternate, in
inflorescence subopposite. Achenes strongly
compressed, cuneate, bicornute distally, small
awns around corolla base, spiculiferous on surface.
Pollen tricolporate, non-lophate.
Only one genus:
287. Trichospira Kunth
Trichospira Kunth in Humb., Bonpl. & Kunth, Nov. Gen. Sp.
Pl., ed. folio 4: 21 (1818).
Perennials; hairs simple, arachnoid. Leaves sessile,
tomentose below. Heads axillary, sessile. Involucral
bracts c. 12, subequal, receptacle with few pales.
Florets c. 10; corollas 4-lobed; anthers with spurs
short; style without node; sweeping hairs acute,
often septate. Achenes with 2 marginal ribs and
2 or 3 weaker ones on surfaces, raphids lacking.
One species, T. verticillata (L.) S.F. Blake, tropical
America.
Compositae
VII. Tribe Liabeae (Cass. ex Dumort.)
Rydb. (1927).
V.A. Funk, H. Robinson and M.O. Dillon
Mostly perennial herbs or shrubs, sometimes
scandent, some annuals or small trees; latex
usually present; hairs simple; usually white tomentose on stems and on abaxial surfaces of
leaves, sometimes only on tips of involucral bracts,
elsewhere sometimes pilose or strigose with stiff
hairs. Leaves opposite, sometimes in a rosette,
usually petiolate or with a petioliform base, rarely
sessile; leaf bases often with pseudostipules or
nodal disc, sometimes fused into sheath; leaf
blades linear to broadly triangular, ovate, oblanceolate, or elliptical; venation trinervate, pinnate,
or palmate. Inflorescences simple or subcymose,
sometimes forming a thrysoid panicle (rarely
sessile). Heads usually on short to long peduncles;
involucre usually subimbricate with bracts in many
gradate series; receptacle alveolate, often with
projecting crests or points, rarely paleaceous. Ray
florets usually present, usually yellow, occasionally
reddish to purple or white, 3–c. 320, fertile; limb
present, usually linear to elliptic-oblong; style
branches elongate, sometimes spiralled. Disc
florets usually yellow, rarely red, purple or white,
3–c. 150, perfect, tube or lower throat pilosulous
or glandular, lobes elongate, usually linear; anther
collars mostly with short-oblong cells; thecae
usually pale, blackened in subtribe Munnoziinae;
bases calcarate, sometimes tailed and fringed or
digitate; apical appendage thin, flat, usually longer
than wide, not constricted at base. Pollen spherical, 25–50 µm in diameter in fluid, tricolporate,
echinate, spines regularly to somewhat irregularly
arranged, never lophate, the bacula tubular or in
circle under spines, sometimes reduced and grains
caveate (Fig. 41). Style base glabrous, usually
enlarged; upper style shaft and backs of branches
with sweeping hairs; branches with stigmatic
papillae over entire inside surface, with little or no
appendage, branches usually shorter than hairy
upper style shaft. Cypselae usually prismatic or
subterete, with 5–10 ribs, less often 4- or 2-ribbed;
variously with biseriate setulae, uniseriate hairs,
stipitate glands or glandular dots; pappus usually
with numerous long inner capillary bristles and
short outer series of squamellae, sometimes with
scales or plumose bristles or absent. Chromosome
numbers known from 12 genera, base numbers
x = 7, 9, 10, 12, 14, 16 and 18 (Robinson et al. 1985).
175
A tribal base number of x = 9 was most probable
from the data cited by Turner et al. (1967), and was
proposed by Robinson et al. (1985).
Tribe Liabeae contains approximately 190
species arranged in 16 genera, and is distributed in
the montane Neotropics at 50–4,750 m elevation.
The greatest generic and species concentration
is in western South America; 13 of the 16 genera
occur in northern Peru. Most species of Liabeae
are found in open areas in mid-elevation forests.
A few species in several genera are found in
disturbed habitats associated with rivers, roadcuts
or treefalls. More rarely, species occupy seasonally
dry scrub or desert habitats. A few members are
found in essentially alpine habitats well above
forested zones, including subparamo, paramo,
jalca and puna (> 3,000 m).
Obvious features shared by many but not
all members of Liabeae include leaves which are
opposite, often strongly trinervate with tomentose
undersurfaces, yellow ray and disc florets, and
the frequent occurrence of latex. The tribe shares
the deeply lobed disc corollas, long-spurred or
calcarate anther bases, continuous stigmatic
surfaces on the inside of the style branches, and
spherical spinose pollen characteristic of members
of subfamily Cichorioideae along with Cichorieae,
Vernonieae and Arctotideae (Robinson and Funk
1987; Jansen et al. 1991; Bremer 1994). Although
Fig. 41. Compositae-Liabeae. SEM micrographs of pollen
grains. A, B Whole grains. A Microliabum humile. Form with
unevenly disposed spines. B Chrysactinium acaule. Form
with evenly disposed spines. C, D Broken grains. C Dillandia perfoliata. Bacula grouped under spines. D Munnozia
tenera. Base of spine fused into cylinder. Scale bars: A,
B = 10 µm, C, D = 1 µm
176
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
monophyly of the subfamily is in doubt, Liabeae,
Vernonieae, Cichorieae and Arctotideae appear to
form a monophyletic group within the subfamily,
and the sister group of Liabeae can be found in
one of these tribes (Robinson and Funk 1987;
Kim et al. 2003). These four tribes have long
sweeping hairs which cover the outer surface of
the style branches and the upper style shaft. Latex
is found in at least some genera of Cichorieae,
Arctotideae and Liabeae, but it is rarely noted
for Vernonieae. Cichorieae and Vernonieae have
homogamous heads without rays, while presence
of rays seems plesiomorphic in Liabeae and
Arctotideae. Lophate pollen and bluish florets
are common in Cichorieae and Vernonieae, both
are lacking in Liabeae and Arctotideae lacks the
bluish florets. The history of the classification of
Liabeae reflects the difficulty in tribal placement
encountered by early workers. Cassini (1823, 1825,
1830), Lessing (1832), de Candolle (1836), Weddell
(1855–1857), and Bentham (1873a) all variously
treated groups of taxa which are now placed in
this tribe as members of Vernonieae, Heliantheae,
Helenieae, Senecioneae and Mutisieae. Bentham’s
classification, which placed most of the taxa in
one genus, Liabum, in tribe Senecioneae, was
essentially adopted by Hoffmann (1894). Rydberg
(1927) was the first to propose tribal status for
the genera from the North American Flora area.
However, despite his work which recognized
genera from Mexico, Central America and the
West Indies, and isolated works by Blake (1935),
Cabrera (1954), and Sandwith (1956), who recognized the affinities of other genera to Liabeae,
Bentham’s classification was retained more or less
intact and accepted by many modern floristic
and taxonomic workers (Cronquist 1955; D’Arcy
1975; Carlquist 1976). Nash and Williams (1976)
accepted Bentham’s concept of a single genus, but
they placed the genus in Vernonieae. Robinson and
co-workers published a series of papers bringing
the genera together into one tribe (Robinson and
Brettell 1973a, 1974; Robinson 1983a with included additional references). Nordenstam (1977)
followed Robinson’s tribal circumscription in
a tribe separate from Senecioneae. Robinson
(1983a) provided the first modern view of the
whole tribe, including a detailed review of previous
classification efforts and relevant literature. Since
the review of Robinson (1983a), there have been
several significant changes in generic limits in the
tribe. Two new genera have been described from
northern Peru, Dillandia (Funk and Robinson
2001) and Bishopanthus (Robinson 1983b); Austroliabum has been reduced to synonymy under
Microliabum (Robinson 1990b); and Liabellum has
been placed into synonymy with Sinclairia (Turner
1989) and then resurrected (Robinson 1990a).
Key to the Genera
1. Abaxial surfaces of leaves green, without tomentum;
leaves and stems with stiff hairs with enlarged bases 2
– Abaxial surfaces of leaves covered with dense white or
grey tomentum; leaves and stems without stiff hairs
with enlarged bases
3
2. Coarse perennial herbs, subshrubs, shrubs, or small
trees; leaves with 5–9 palmately arranged veins; cypselae 4-sided with 4 ribs
301. Erato
– Small erect, decumbent, or creeping herbs; leaves 3nervate; cypselae compressed with 2 ribs
303. Philoglossa
3. Anther thecae dark brown or black
4
– Anther thecae pale yellow or very light brown
5
4. Pappus white; small herbaceous perennials; leaves in
a rosette or grouped close together on a short stem
300. Chrysactinium
– Pappus sordid or reddish; usually lax subshrubs,
sometimes annual or perennial herbs; leaves cauline
302. Munnozia
5. Heads sessile, subtended by a rosette of leaves or nearly
sessile with peduncles of less than 2 cm long
298. Paranephelius
– Heads on peduncles more than 2 cm long
6
6. Leaves pinnately veined
7
– Leaves 3-nervate
10
7. Herbs; heads few; disc florets 10–55; leaves sessile,
subperfoliate or perfoliate
8
– Trees, shrubs, or vines; heads many; disc florets 3–
34; leaves usually with distinct unwinged to broadly
winged petioles
9
8. Adaxial surfaces of leaves rugose to nearly smooth;
corollas reddish-orange to red; involucral bracts eximbricate, linear-lanceolate with attenuate tips
299. Pseudonoseris
– Adaxial surfaces of leaves bullate; corollas yellow to
yellow and purple; involucral bracts subimbricate,
lanceolate with acute tips
291. Dillandia
9. Trees or shrubs; petioles without wings, bases of petiole pairs fused into a sheath; raphids of cypsela wall
elongate
292. Ferreyranthus
– Vines or shrubs; petioles with or without wings, sometimes included in perfoliate leaf base but not fused into
a sheath; raphids of cypsela wall subquadrate
296. Oligactis
10. Pappus lacking
289. Cacosmia
– Pappus of bristles, awns and/or squamellae
11
11. Pappus of plumose bristles; disc corollas red
290. Chionopappus
– Pappus of capillary bristles, awns and/or squamellae;
all corollas yellow
12
12. Heads terminal and solitary on leafy stems; leaf bases
fused into a sheath
288. Bishopanthus
– Heads few to many but not solitary; leaf bases without
a sheath
13
Compositae
13. Inflorescences with all branches alternate; receptacle
scarcely alveolate, without hairs, squamellae or projections; Argentina and Bolivia
295. Microliabum
– Inflorescences with at least basal branches opposite;
receptacles with hairs, squamellae or projections; Bolivia northwards to Mexico
14
14. Plants without latex; style branches of disc florets
longer than hispidulous part of distal style shaft; anther bases digitate; rays 20–120; involucral bracts 50–
150 in 5 series; raphids of cypsela walls subquadrate
294. Liabum
– Plants with latex; style branches of disc florets shorter
than hispidulous part of distal style shaft; anther
bases minutely crenulate; rays usually absent or when
present 4–25; raphids of cypsela walls elongate
15
15. Herbs; rays always absent; leaf blades shallowly to
deeply lobed; petioles winged to base, sometimes perfoliate; disc florets 25–30; inner pappus of 40–50 capillary bristles, outer of 20–40 squamellae 293. Liabellum
– Shrubs or vines; rays usually present (absent in a few
species); leaf blades entire or serrate; petiole bases
simple, not winged and without pseudostipules; disc
florets 5–30; inner pappus of 30–40 bristles, outer of
10–15 squamellae
297. Sinclairia
VII.1. Subtribe Liabinae Cass. ex Dumort.
(1829).
Plants perennial; with or without latex; leaves opposite, stems or leaf undersides always tomentose;
anther thecae pale, usually tailed and fringed or
digitate at base; florets usually yellow, rarely red or
purple; style branches usually shorter than hairy
upper shaft; cypselae prismatic or subterete. Pollen
spines unevenly grouped.
288. Bishopanthus H. Rob.
Bishopanthus H. Rob., Phytologia 54: 64 (1983).
Shrub; with latex. Leaf bases fused into sheath;
blades 3-nervate, bullate. Heads solitary, pedunculate, campanulate; involucral bracts c. 25, in 2 subequal series; epaleaceous. Ray florets c. 20; limbs
linear. Disc florets c. 25; bases of anther thecae
short-tailed, short-fringed. Cypselae with setulae,
uniseriate hairs, and glands, raphids subquadrate;
pappus bristles c. 35, with short outer bristles. One
species, B. soliceps H. Rob., Peru.
289. Cacosmia Kunth
Cacosmia Kunth in H.B.K., Nov. Gen. et Sp., ed. folio 4: 227
(1818); Nordenstam, Bot. Notiser 130: 279–286 (1977), rev.;
Robinson, Fl. Ecuador 190, 2: 1–63 (1978), reg. rev.
Clairvillea DC. (1836).
Shrub; usually with latex; stems densely pubescent.
Leaf bases fused into sheath; blades 3–5-nervate,
177
bullate. Inflorescence densely corymbiform; heads
cylindrical; involucral bracts 20–25, 5–6-seriate,
outer bracts ranked, inner bracts not in ranks; receptacle epaleaceous. Ray florets 5, limbs broad.
Disc florets 5–6; bases of anther thecae without
tails, minutely digitate. Cypselae 3–5-angled, glabrous, raphids elongate; pappus lacking. n = 7 + 1
or 2. Three species in Ecuador, one into Peru.
290. Chionopappus Benth.
Chionopappus Benth. in Benth. & Hook. f., Gen. Pl. 2: 485
(1873).
Shrubs; latex not noted; stems arachnoidtomentose. Leaf bases fused into sheath, blades
3-nervate. Inflorescences simple dichasia; heads
campanulate; involucral bracts 50–55, c. 5-seriate;
paleae strap-shaped. Ray florets c. 40. Disc florets
75–125; corollas red, glabrous; bases of anther
thecae short-tailed. Cypsela 8–10-ribbed, setulae
minute, raphids elongate; pappus bristles 8–10,
plumose. n = c. 9. One species, C. benthamii S.F.
Blake, Peru.
291. Dillandia V.A. Funk & H. Rob.
Dillandia V.A. Funk & H. Rob., Syst. Bot. 26: 218 (2001).
Moderate-sized to small herbs less than 60 cm tall;
latex not noted; stems arachnoid-tomentose. Leaf
bases sessile, subpetiolate or perfoliate; surface bullate; blades pinnately veined. Inflorescences of 1–2
heads or, more frequently, a 3–7-headed subumbel; heads campanulate, involucral bracts 25–40,
5–6-seriate; receptacle without chaff. Ray florets
15–40, limbs oblong to narrowly oblong. Disc florets 10–30, corollas yellow to yellow and purple,
tubes pilose, anther thecae pale, bases rounded.
Cypselae (immature) 7–10-ribbed, densely setulose, with a few subquadrate raphids; pappus bristles 10–30, sometimes shorter outer bristles. Three
species, Peru.
292. Ferreyranthus H. Rob. & Brettell
Ferreyranthus H. Rob. & Brettell, Phytologia 28: 50
(1974); Dillon & Sagastegui, Arnaldoa 2: 7–23 (1994), rev.;
Robinson, Fl. Ecuador 190, 2: 1–63 (1978), reg. rev.
Shrubs or weak trees; without latex; stems
arachnoid-tomentose. Leaf bases fused into
sheath; blades pinnately veined. Inflorescences
densely corymbiform; heads broadly campanulate;
involucral bracts 45–55, c. 5-seriate; receptacle
squamuliferous. Ray florets 8–12, limbs short.
178
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Disc florets 12–25, corollas glandular-dotted;
bases of anther thecae short-tailed, strongly
fringed. Cypselae c. 10-ribbed, with setulae and
glands, raphids elongate; pappus bristles 10–15,
squamellae narrow. n = c. 18. Eight species, Peru,
one into Ecuador.
293. Liabellum Rydb.
Liabellum Rydb., N. Amer. Fl. 34, 4: 294 (1927).
Small xylopodial herbs; with latex; stems short,
arachnoid-tomentose. Leaf bases perfoliate, sessile,
blades scarcely to deeply lobed, 3-nervate. Heads
terminal, 1–3 on elongate peduncles, broadly campanulate; involucral bracts 20–40, c. 4-seriate; receptacle sometimes puberulous. Rays lacking. Disc
florets 25–30; bases of anther thecae scarcely crenulate. Cypsela c. 10-ribbed, setulae long, raphids
elongate; pappus bristles 40–50, squamiform setae
20–40. Five species, Mexico.
294. Liabum Adans.
Liabum Adans., Fam. 2: 131 (1763); Robinson, Fl. Ecuador
190, 2: 1–63 (1978), reg. rev.
Starkea Willd. (1803).
Andromachia Humb. & Bonpl. (1809).
Allendea La Llave & Lex. (1824).
Perennial herbs; without latex; stems arachnoidtomentose. Leaf bases connected across node, often forming nodal disc; blades 3-nervate. Inflorescence partly subumbellate; heads broadly campanulate; involucral bracts 50–150, c. 5-seriate; receptacle ridged or bristly. Ray florets 20–120. Disc florets
10–85; style branches long; anther thecae bases digitate. Cypselae c. 10-ribbed, raphids subquadrate;
pappus bristles 17–40, squamellae short, narrow.
n = c. 12 to 39, many counts are c. 18. Forty-five
species, Mexico, Central America, Greater Antilles,
and Andes of South America.
295. Microliabum Cabrera
Microliabum Cabrera, Bol. Soc. Argent. Bot. 5: 211 (1955),
nom. nov. for Liabellum Cabrera
Liabellum Cabrera (1954), non Liabellum Rydb. (1927).
Angelianthus H. Rob. & Brettell (1974), nom. nov. illegit.
superfl. for Liabellum Cabrera
Austroliabum H. Rob. & Brettell (1974).
Annual to perennial herbs or subshrubs; with latex;
stems white-tomentose and glandular-hairy. Leaf
bases broadened or with pseudostipules or nodal
discs; blades 3-nervate. Heads solitary or in cymes,
broadly campanulate; involucral bracts 30–75, 2–
4-seriate, subequal to gradate; receptacle epaleaceous. Rays 10–30; limbs narrowly elliptical to linear. Disc florets 15–175; bases of anther thecae with
few or no teeth. Cypselae 8–10-ribbed, setuliferous, raphids elongate; pappus bristles or lamellae
8–16, outer squamellae 8–16. Six species, southern
Bolivia, northern Argentina.
296. Oligactis (Kunth) Cass.
Oligactis (Kunth) Cass., Dict. Sci. Nat. 36: 16 (1825);
Robinson, Fl. Ecuador 190, 2: 1–63 (1978), reg. rev.
Andromachia sect. Oligactis Kunth (1818).
Shrubs and vines, without latex. Leaf bases sometimes confluent across nodes; blades pinnately
veined. Inflorescence densely corymbiform; heads
narrowly to broadly campanulate; involucral
bracts 16–55, 4–5-seriate; receptacle ridged and
squamuliferous. Ray florets 3–18, limbs short. Disc
florets 3–34; style branches rather long; anther
thecae bases digitate. Cypselae 5–8-ribbed, with
glands and contorted setulae, raphids subquadrate;
pappus bristles 20–35, tips often broad, squamellae
7–10. n = 18, 39. Fifteen species, mostly northern
Andes, one to Costa Rica.
297. Sinclairia Hook. & Arn.
Sinclairia Hook. & Arn., Bot. Beech. Voy. 433 (1841).
Megaliabum Rydb. (1927).
Sinclairiopsis Rydb. (1927).
Subshrubs, shrubs or vines; with latex; stems
arachnoid-tomentose. Leaves opposite or ternate,
sometimes seasonally deciduous, bases without
sheaths or pseudostipules; blades 3-nervate.
Inflorescences laxly to densely corymbiform
or pyramidal; heads narrowly to broadly campanulate; involucral bracts 18–45, 3–5-seriate;
receptacle glabrous, spinulose or puberulous.
Ray florets absent or 4–25. Disc florets 5–30;
anther thecae bases minutely crenulate. Cypselae
c. 8–10-ribbed, glabrous or setuliferous, raphids
elongate; pappus bristles 30–40, squamellae 15–20.
n = 15 to 18. Thirty species, Mexico, Central
America, one to western Colombia.
VII.2. Subtribe Paranepheliinae H. Rob.
(1983).
Plants rosettiform or short-stemmed, fibrousrooted, usually with latex; roots often swollen and
fleshy; leaves often pinnately lobed, undersurface
Compositae
179
with white tomentum; involucre gradate; ray floret
limbs linear; disc corollas funnelform, tubes 2–3
times as long as lobes; thecae pale, base not tailed
nor noticeably lobed; style branches longer than
scabrid upper style shaft; cypselae prismatic or
fusiform, raphids elongate. Pollen spines mostly
evenly dispersed.
298. Paranephelius Poepp.
Paranephelius Poepp., Nov. Gen. et Sp. 3: 42 (1843);
Robinson, Fl. Ecuador 190, 2: 1–63 (1978), reg. rev.
Liabum sect. Paranephelius (Poepp.) Benth. in Benth. and
Hook. f. (1873).
Acaulescent perennial herbs. Leaf bases not
sheathing; blades trinervate to pinnately veined.
Heads sessile, broadly campanulate; involucral
bracts 40–50, c. 4-seriate; receptacle ridged. Ray
florets 20–35; style branches spiralled. Disc florets
20–33. Cypsela c. 10-ribbed, glabrous or with
some arachnoid hairs; pappus bristles 45–80, outer
series indistinct. n = 9, 14, 15. Seven species, Peru,
Bolivia, one to northern Argentina.
299. Pseudonoseris H. Rob. & Brettell
Pseudonoseris H. Rob. & Brettell, Phytologia 28: 59 (1974).
Small perennial herbs. Leaf bases sessile, subauriculate; blades pinnately veined. Inflorescence
scapose or subscapose; heads pedunculate, broadly
campanulate; involucral bracts c. 40, c. 4-seriate;
receptacle subglabrous. Ray florets yellow or
yellow-orange, 15–20. Disc florets orange-yellow
or red, 25–55. Cypselae c. 10-ribbed, with sparse
setulae or tomentum; pappus bristles 25–30,
squamiform outer series. n = 12(?). Three species,
Peru.
VII.3. Subtribe Munnoziinae H. Rob. (1983).
Plants usually with latex; heads often longpedunculate; ray floret limbs linear; disc corolla
throat broadened at base; thecae black, base not
tailed nor noticeably lobed; style branches much
shorter than hairy upper style shaft, blunt; cypselae
prismatic to biconvex, raphids subquadrate. Pollen
spines evenly dispersed.
Fig. 42. Compositae-Liabeae. Chrysactinium wurdackii.
A Habit, note opposite leaves. B Leaf, note tomentose
undersurface. C Head, note heterogamous heads. D Outer
involucral bract. E Inner involucral bract. F Ray floret
corolla and style. G Hairs on tube of ray floret. H Apex
of ray corolla. I, J Disc corolla, note continuous stigmatic
surface on inside of style branches, calcarate anther bases,
and deeply divided corolla. K Style of disc floret. L Cypsela
and pappus. (Drawing by A. Tangerini)
Perennial rhizomatous herbs, rosettiform or shortstemmed; stems evanescent-tomentose. Leaf bases
cuneate or petioliform; blade 3-nervate, tomentose.
Heads solitary, long-pedunculate, broadly campanulate; involucral bracts 40–60, 4–5-seriate; receptacle squamulose. Ray florets 30–60. Disc florets
30–60; anther collar cells annulate. Cypselae 8–10ribbed; pappus bristles 30–60, white, no squamellae. n = 12 to 16. Six species, Ecuador, northern
Peru.
300. Chrysactinium (Kunth) Wedd. Fig. 42
Chrysactinium (Kunth) Wedd., Chlor. And. 1: 212 (1857);
Robinson, Fl. Ecuador 190, 2: 1–63 (1978), reg. rev.; Funk &
Zermoglia, Syst. Bot. 24: 323–338 (1999), rev.
Andromachia sect. Chrysactinium Kunth (1818).
301. Erato DC.
Erato DC., Prodr. 5: 318 (1836); Robinson, Fl. Ecuador 190,
2: 1–63 (1978), reg. rev.
Munnozia subg. Erato (DC.) H. Rob. & Brettell (1974).
180
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Coarse perennial herbs; stems and leaves pilose
or strigose. Leaves with oblong pseudostipules;
blades palmately 5–9-veined, surfaces green.
Inflorescence cymose to subumbellate; heads
broadly campanulate; involucral bracts 40–100, c.
4-seriate, tips sometimes tomentose; receptacle
foveolate and ridged. Ray florets 75–230. Disc
florets 20–150. Cypselae 4-ribbed, glabrous or
hispidulous; pappus of 25–50 bristles or short
awns, no outer series. n = 9. Five species, Costa
Rica to Peru.
302. Munnozia Ruiz & Pav.
Munnozia Ruiz & Pav., Fl. Peru Chil. Prodr. 108 (1794);
Robinson, Fl. Ecuador 190, 2: 1–63 (1978), reg. rev.
Alibum Less. (1832).
Prionolepis Poepp. (1845).
Kastnera Sch. Bip. (1853).
Liabum subg. Chrysastrum Willd. ex Sch. Bip. (1853).
Munnozia subg. Kastnera (Sch. Bip.) H. Rob. & Brettell
(1974).
Annual or perennial herbs or subshrubs. Petioles
sometimes winged, bases often pseudostipulate;
blades 3-nervate to pinnately veined, usually tomentose abaxially. Inflorescence terminal, more or
less corymbose; heads broadly campanulate; involucral bracts 17–70, 2–4-seriate, subequal to gradate; receptacle often squamulose. Florets yellow,
rarely whitish to lavender. Ray florets 6–70. Disc florets 9–85. Cypselae 6–10-ribbed, setuliferous; pappus bristles sordid to reddish, 5–55, with squamellae. n = 12 to 36, most frequently 10, 11 and 12.
Forty-six species, mostly from the Andes, few in
Costa Rica, Panama.
303. Philoglossa DC.
Philoglossa DC., Prodr. 5: 567 (1836); Robinson, Fl. Ecuador
190, 2: 1–63 (1978), reg. rev.
Small erect to creeping herbs; stems and leaves
pilose to strigose. Leaf bases with oblong pseudostipules; blades 3-nervate, surfaces green. Involucres broadly campanulate; bracts 20–30, 3–4seriate, subequal to gradate. Ray florets 21–70. Disc
florets 30–60, yellow, rarely deep purple or brown.
Cypselae compressed, 2-ribbed, mostly glabrous;
pappus deciduous squamellae or awns, or lacking.
n = 18. Five species, southern Colombia to Bolivia.
VIII. Tribe Cichorieae Lam. & DC.
(1806).
Tribe Lactuceae Cass. (1819).
H.W. Lack
Perennial to annual herbs, rarely subshrubs, shrubs
or rosette trees, very rarely scandent vines, characterized by ligulate capitula and abundant milky
latex. Leaves mostly alternate, entire, lobed to pinnatisect, very rarely spiny. Involucral bracts imbricate in several series or equal and in a single
row. Capitula always ligulate and homogamous,
solitary or aggregated in synflorescences, sometimes in secondary heads. Involucral bracts various, receptacle mostly epaleate and naked, rarely
scaly-bristly or paleate. All florets with a 5-lobed
ligule, perfect, of various colours although predominantly yellow. Anthers basally calcarate and
caudate, apical appendage elongate and obtuse, filaments smooth. Pollen echinolophate or echinate.
Style slender, mostly with long, slender branches,
hairs on the shaft and the branches. Achenes and
pappus of various forms.
About 86 genera in 12 subtribes, number of
species differing widely due to different views on
their circumscription, in particular in Hieracium,
Pilosella and Taraxacum. Excluding the latter three
genera, the tribe contains c. 1,500 species. A predominantly northern hemisphere tribe but found
on all continents, with comparatively few strictly
tropical species, the latter as a rule in open habitats. Most taxa occur in rather dry to somewhat
humid areas, and are almost absent from aquatic
habitats. The tribe comprises many widespread,
successful and noxious weeds, in particular species
of Chondrilla, Crepis, Hypochaeris, Sonchus, Taraxacum and Youngia. In addition, numerous species
have become naturalized in areas very far from
their native habitats, e.g. European Sonchus arvensis on Fiji, and Mediterranean Crepis pusilla in
southern Australia.
A group notoriously poor in diagnostic characters, with convergent evolution having led to very
similar forms in different genera. This applies in
particular to the indumentum, with glabrous forms
in several groups, the achene, with heteromorphic
fruits in a few subtribes, and the pappus, which has
been reduced or lost in several groups.
Knowledge of the tribe is extremely unbalanced. Very limited information is available
on several genera from central, eastern and
south-eastern Asia, whereas the European,
Compositae
Mediterranean and North American groups have
been well studied. Recently described, enigmatic
genera include Faberiopsis, Phitosia and Spiroseris.
The position of Thamnoseris, an endemic of the
Desaventuradas Islands off the coast of South
America, may be outside the Cichorieae.
Recent molecular studies have brought significant new insights into the systematic relationships
of the tribe (e.g. Kim et al. 1999a, b, on Sonchinae),
and indicate the need for a completely new view
of the group. Certain taxonomic conclusions from
these studies have been cautiously integrated into
the present treatment, mostly referring to Lactucinae, Sonchinae and the former Dendroseridinae.
The recent studies by Chinese workers, in particular on Lactucinae, have also been considered in
this account, although they are exclusively based
on classical morphology and almost no herbarium
material was available to the author for comparison. The circumscription of Lactucinae is still incompletely understood, and is based tentatively on
achene characters here.
Very many publications on the taxonomy of
Cichorieae are available (Babcock and Stebbins
1943; Stebbins 1953; Carlquist 1960; Jeffrey 1966;
Sell 1975; Tomb 1976, 1977; Blackmore 1981, 1982;
Voytenko 1989b; Bremer 1993, 1994; Whitton
et al. 1995; Blackmore and Persson 1996; Reveal
1997; Tegel 2002; Lee et al. 2003) but no synthetic
approach on a global level has been achieved so
far. The circumscription of subtribes and genera
differs widely in quality and precision, with several
genera in Crepidinae and Lactucinae being very
vaguely delimited.
Key to the Subtribes7
1. Stems winged, resin ducts and latex canals always
present
1. Scolyminae (p. 182)
– Stems never winged, only latex canals present
2
2. Florets blue, pappus of minute, irregularly shaped,
round to acute scales or absent 2. Cichoriinae (p. 182)
7
Previous workers (e.g. Stebbins 1953; Jeffrey 1966; Bremer
1993) have refrained from providing a key to the subtribes
of Cichorieae, basically for two reasons: (1) the philosophy to
place a taxon ‘nearest to those genera which it most nearly
resembles in respect to the largest number of characteristics’
(Stebbins 1953), which has traditionally been applied to Cichorieae, does not easily lead to a clear and workable key; (2)
the pappus – ‘the most useful single character for subdividing the tribe’ (Stebbins 1953) and ‘of cardinal value’ (Bremer
1994) – has undergone very many transformations, including
repeated losses in several subtribes.
181
– Florets mostly yellow, sometimes violet, pink or white;
if blue, then pappus neither of minute, irregularly
shaped, round to acute scales nor absent
3
3. Pappus of very few broad, lanceolate scales, receptacle
mostly setose
3. Catananchinae (p. 182)
– Pappus of various forms or absent, but never of only
very few broad, lanceolate scales, receptacle very rarely
setose
4
4. Pappus in all achenes absent
5
– Pappus at least in central achenes well developed 16
5. Florets white
361. Atrichoseris
– Florets of various colours but not white
6
6. Involucral bracts in a single row, basally fused
7
– Involucral bracts almost never in a single row, never
basally fused
8
7. Outer achenes with strong hooks or glochids
385. Koelpinia
– Outer achenes without strong hooks or glochids
349. Phalacroseris
8. Outer achenes stellately patent
381. Rhagadiolus
– Outer achenes not stellately patent
9
9. Inner bracts becoming hard and woody in fruit
10
– Inner bracts not becoming hard and woody in fruit 11
10. Inner bracts becoming connate in fruit
309. Acanthocephalus
– Inner bracts not becoming connate in fruit
315. Heteracia epapposa
11. Achenes deeply furrowed
12
– Achenes not deeply furrowed
13
12. Densely hirsute plant with very long hairs
368. Hispidella
– Plants without indumentum
347. Krigia cespitosa
13. Branched herbs
14
– Scapose plants
15
14. Achenes apically with two horns 321. Lapsanastrum
– Achenes apically without appendages 320. Lapsana
15. Perennial plant, rosette of deeply dentate leaves
372. Aposeris
– Annual plant, rosette of spathulate leaves
373. Arnoseris
16. Inner achenes apically with crown-like structure
374. Garhadiolus
– Inner achenes apically different
17
17. At least some pappus setae thick, sometimes with long
lateral pinnulae
24
– All pappus setae slender, denticulate or barbellate, but
never with long lateral pinnulae
18
18. Pappus comprising very thin, flexible rays
6. Sonchinae (p. 189)
– Pappus never comprising very thin flexible rays 19
19. Pollen distinctly orange, New World
7. Microseridinae p.p. (p. 191)
– Pollen never orange, predominantly Old World
20
20. Pappus barbellate or of five rigid awns and slender
setae
21
– Pappus not barbellate, always homogenous
22
21. Mostly annual herbs, receptacle paleate or scaly, heads
with numerous florets
9. Malacothricinae p.p. (p. 193)
– Mostly perennial herbs, receptacle naked, heads with
few florets
8. Stephanomeriinae p.p. (p. 192)
22. Achenes basally constricted into a small, smooth annulus, achenes often compressed
5. Lactucinae (p. 187)
182
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
– Achenes basally never constricted, rarely compressed
23
23. Plants often with small, soft, branched hairs, pappus
setae brittle
10. Hieraciinae p.p. (p. 194)
– Plants without small, soft, branched hairs, pappus setae non-brittle
4. Crepidinae p.p. (p. 183)
24. Pappus of bristle-tipped, awn-like scales
348. Microseris
– Pappus of setae, the latter barbellate or with long side
projections
25
25. Pappus setae truly feather-like or barbellate, New
World
26
– Pappus setae mostly with side projections pointing in
various directions, Old World
27
26. Mostly perennial herbs, receptacle naked, heads with
very few florets
8. Stephanomeriinae p.p. (p. 192)
– Mostly annual herbs, receptacle paleate or scaly, heads
with numerous florets
9. Malacothricinae p.p. (p. 193)
27. Pappus rays with stiff lateral side projections consisting of a single, giant tubular cell, plant often with
coarse indumentum of large hairs
11. Hypochaeridinae p.p. (p. 195)
– Pappus rays with soft lateral side projections consisting of a row of flattened cells (pappus woolly), plants
with soft indumentum of small hairs
12. Scorzonerinae p.p. (p. 198)
VIII.1. Subtribe Scolyminae Less. (1832).
Very spiny thistle-like herbs with winged stems,
resin ducts and latex canals. Exclusively Old World
subtribe, comprising a single genus.
304. Scolymus L.
Scolymus L., Sp. Pl. 813 (1753); Vazquez, Anales Jard. Bot.
Madrid 58: 83–100 (2000), rev.
Annual to perennial herbs, very spiny with winged
stems, receptacle paleate, paleae broadly winged
and deciduous, enclosing the dorsiventrally
compressed achenes, florets yellow, pappus of stiff
scabrous bristles or absent. x = 10, diploids. Three
species, Macaronesia, Mediterranean area.
VIII.2. Subtribe Cichoriinae Cass. ex Dumort.
(1829) s. str.
Florets blue, pappus of minute, irregularly shaped,
round to acute scales or absent. Exclusively Old
World subtribe, comprising a single genus.
305. Cichorium L.
Fig. 43
Cichorium L., Sp. Pl. 813 (1753); Wagenitz & Bedarff, Davis
& Hedge Festschr.: 11–21 (1989), rev.; Kiers et al., Syst. Bot.
24: 645–659 (1999); Kiers, Gorteria suppl. 5: 1–77 (2001),
mol. phylog.
Fig. 43.
Compositae-Cichorieae. Cichorium intybus.
A Habit. B Middle cauline leaf. C Upper part of flowering
branch. D Floret. E Floret, upper part of corolla cut off.
F Mature achene, part of pappus further enlarged. G Part
of receptacle. (Ross-Craig 1962)
Annual to perennial herbs, lower part of outer involucral bracts fleshy at flowering time and hard at
fruiting time, involucral bracts in two rows, florets
few, achenes obovoid to cylindrical. x = 9, diploids.
Six species, Europe, Mediterranean area, Near East.
Cichorium endivia L. (endive) and C. intybus L.
(chicory) cultivated for their edible leaves.
VIII.3. Subtribe Catananchinae K. Bremer
(1993).
Perennial to annual herbs, receptacle setose or
paleate, achenes without ribs or beak. Pappus of
few lanceolate to triangular scales, sometimes
prolonged into bristles. Exclusively Old World
tribe, comprising three genera.
Compositae
Key to the Genera
1. Pappus consisting of five triangular, stiff woody bracts,
involucral bracts not scarious at margins
308. Rothmaleria
– Pappus of lanceolate scales apically prolonged into
bristles, involucral scales scarious at margins
2
2. Margins of involucral bracts silvery or shiny, leaves
parallel-veined, receptacle bristly 306. Catananche
– Margins of involucral bracts neither silvery nor shiny,
leaves not parallel-veined, receptacle peripherally
paleate
307. Hymenonema
Genera of Catananchinae
beaked, rarely compressed, pappus rays scabridbarbellate, usually several cells in diameter at
base, pappus rarely absent. Old World subtribe,
with Crepis and Taraxacum also in America. The
generic limits between Crepidiastrum, Ixeridium,
Ixeris and Youngia are unclear, and hybrids
between them have been reported. Phitosia is
tentatively included in this subtribe, although it
may well belong to another group.
Key to the Genera8
1. Achenes all or some muricate with projections, tubercules or scales on upper part below beak; plants with
or without deflexed hairs on the stems
2
– Achenes not muricate on upper part below beak, at
most hairy or minutely scabrid on the ribs, beaked or
unbeaked; plants without deflexed hairs on the stems
7
2. Plants with taproot and many-flowered capitula solitary and terminal on leafless hollow scapes; corollatube with tuft of long hairs at the apex; x = 8
325. Taraxacum
– Plants not with the above combination of characters;
corolla-tube with short, one-celled hairs; x = 7, 5, 4 or
3
3
3. Pappus absent or coroniform or only the inner achenes
with a pappus of short bristles
4
– Pappus present, of long bristles
5
4. Internal involucral bracts becoming thickened, hardened and finally connate in fruit
309. Acanthocephalus
– Internal involucral bracts not becoming thickened,
hardened and connate in fruit
315. Heteracia
5. Achenes with a corona of 5–6 short, rounded scales
below the beak
326. Willemetia
– Achenes not as above
6
6. Achenes heteromorphic, outer more or less beakless
and clasped at the base by the inner involucral bracts,
with deciduous pappus, inner with long beak and persistent pappus
316. Heteroderis
– Achenes homomorphic or, if heteromorphic, then not
as above
310. Chondrilla
7. Pappus of long bristles absent from all achenes
8
– Pappus of long bristles present on at least the central
achenes
9
8. Achenes with 20 slender ribs; perennial herbs (Europe)
320. Lapsana
– Achenes with 10–13 ribs; annual or biennial scapose
herbs (E Asia)
321. Lapsanastrum
9. Involucral bracts lax, outer foliaceous, inner blackish;
achenes 4-ribbed (Pakistan)
324. Spiroseris
– Involucral bracts imbricate, or all of more or less equal
length, or in two series with the outer much shorter
than the inner; achenes not as above
10
10. Achenes heteromorphic, outer different in form, indumentum or pappus from inner
11
– Achenes homomorphic
12
306. Catananche L.
Catananche L., Sp. Pl. 812 (1753).
Annual or perennial herbs, involucral bracts scarious except the midvein, receptacle bristly, bristles persistent, florets blue or yellow, pappus of
few lanceolate, apically elongated scales. x = 9,
8, diploids and tetraploids. Six species, Mediterranean area.
307. Hymenonema Cass.
Hymenonema Cass., Bull. Sci. Soc. Philom. Paris 1817: 34
(1817).
Perennial herbs, involucral bracts with somewhat
scarious margins, receptacle paleate on the periphery, paleae persistent, florets yellow, achenes obconical, pilose, pappus of lanceolate, apically elongated scales. x = 9, diploids. Two species, endemic
to Greece.
308. Rothmaleria Font Quer
Rothmaleria Font Quer, Brotéria (Ci. Nat.) 9: 151 (1940);
Lack et al., Willdenowia 10: 37–49 (1980), rev.
Perennial scapose herb, involucral bracts with
somewhat scarious margins, receptacle paleate,
paleae persistent, florets yellow, achenes obconical,
pappus consisting of 5(6) triangular, stiff, woody
scales. x = 9, diploid. Monotypic, R. granatensis
(Boiss. ex DC.) Font Quer, endemic to the Granada
area (Spain).
VIII.4. Subtribe Crepidinae Cass. ex Dumort.
(1827).
Involucral bracts usually in two unequal rows,
capitula mostly with many flowers, florets mostly
yellow, achenes terete to fusiform or obovoid, often ribbed, transversely muricate-tuberculate and
183
8
By C. Jeffrey.
184
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
11. Outer achenes prismatic, hairy, with caducous pappus,
inner glabrous with persistent pappus (central Asia)
319. Lagoseriopsis
– Outer and inner achenes otherwise distinct
(widespread)
312. Crepis
12. Capitula numerous, aggregated into large synflorescences; dwarf alpine perennial herbs (Sinohimalaya)
323. Soroseris
– Capitula few to numerous, but not aggregated into
large synflorescences; if dwarf alpine perennial herbs,
then capitula few
13
13. Involucral bracts in several series, imbricate or subequal
14
– Involucral bracts in two series, outer much shorter
than inner
15
14. Achenes fusiform, involucral bracts imbricate (Sinohimalaya)
314. Dubyaea
– Achenes elliptic, smooth, involucral bracts subequal
(tropical Africa)
313. Dianthoseris
15. Achenes without distinct apical beak, at most gradually narrowed in upper part into a short rostrum or,
if with distinct beak, then columnar or fusiform, not
compressed
16
– Achenes with distinct apical beak, more or less distinct
in colour and/or texture from achene body, more or
less compressed
19
16. Achenes narrowed towards the apex
17
– Achenes not narrowed towards the apex, with 10 equal
ribs; x = 5
311. Crepidiastrum
17. Achenes columnar or fusiform, with more or less equal
ribs; plants with indumentum, rarely glabrous
18
– Achenes more or less compressed, with 12–15 slender unequal ribs, the lateral often narrowly wing-like;
plants glabrous
327. Youngia
18. Plants glabrous; pappus bristles broadened at base;
322. Phitosia
x = 9 (Taygetos Mts, Greece)
– Plants with at least some indumentum; pappus bristles
not broadened at base; x = 8, 7, 6, 5, 4 or 3 (widespread)
312. Crepis
19. Achenes with filiform beak
20
– Achenes with stout beak
21
20. Achenes with 10 narrowly wing-like, smooth ribs
318. Ixeris
– Achenes with 9–12 low, shortly more or less spinulose
ribs
317. Ixeridium
21. Plant creeping, glabrous, leaves deeply palmately divided into 3–5 lobes
318. Ixeris
– Plant erect, leaves otherwise; x = 5 311. Crepidiastrum
Genera of Crepidinae
309. Acanthocephalus Kar. & Kir.
Acanthocephalus Kar. & Kir., Bull. Soc. Imp. Naturalistes
Moscou 15: 127 (1842); Voytenko, Bot. Zhurn. (Moscow &
Leningrad) 74: 1241–1257 (1989), fruit morph.
Annual herbs, involucral bracts in two rows,
inner connate, becoming hard and woody in fruit,
florets yellow, achenes at least basally enclosed
by woody involucre, pappus absent or of short
scabrid-barbellate rays. x = 3, hexaploid (?). Two
species, Near East, central Asia, China.
310. Chondrilla L.
Chondrilla L., Sp. Pl. 796 (1753); Iljin, Bjull. Otd.
Kaučukonosov 3: 1–61 (1930), rev.
Biennial or perennial herbs, often with much
reduced foliage, heads few-flowered, involucral
bracts in two rows, florets yellow, achenes clearly
differentiated into main body and beak, main
body ribbed, distally tuberculate or with small
corona, beak often deciduous with the pappus of
long, scabrid-barbellate rays. x = 7, 5, diploids,
triploids and tetraploids. Circa 25 species, Eurasia,
Mediterranean area.
311. Crepidiastrum Nakai
Crepidiastrum Nakai, Bot. Mag. (Tokyo) 34: 147 (1920); Pak
& Kawano, Mem. Fac. Sci. Kyoto Univ., ser. Biol. 15: 29–61
(1992), rev.
Paraixeris Nakai (1920).
Perennial to annual herbs, occasionally subshrubs,
taprooted, 5–19 flowers per capitulum, involucral
bracts in two rows, flowers yellow to white, achenes fusiform, somewhat flattened, obtusely 10–
20-winged, pappus of scabrid-barbellate, white or
brownish, deciduous rays. x = 5, diploids. Thirteen
species, eastern Asia, Far East.
312. Crepis L.
Fig. 44
Crepis L., Sp. Pl. 805 (1753); Babcock, Univ. Calif. Publ.
Bot. 21: x, 1–197, 22: x, 199–1030 (1947), rev.; Merxmüller,
Mitt. Bot. Staatssamml. München 7: 271–275 (1968), rev.;
Duvigneaud, Lejeunia n.s. 71: 1–8 (1973), rev.
Zacintha Mill. (1754).
Perennial to annual herbs, involucral bracts in
two rows, those of outer row often much shorter
than inner, receptacle naked, very occasionally
paleate, florets yellow, rarely white or pink, achenes
fusiform, not compressed, ribbed, gradually tapering into beak, homomorphic or rarely dimorphic
with the inner achenes beaked or outer achenes
compressed. Pappus of scabrid-barbellate rays.
x = 8, 7, 6, 5, 4, 3, diploids, tetraploids, hexaploids,
octoploids. About 200 species, all continents except
Australia, in America south to northern Mexico.
313. Dianthoseris Sch. Bip. ex A. Rich.
Dianthoseris Sch. Bip. ex A. Rich., Tent. Fl. Abyss. 1: 468
(1848).
Nannoseris Hedberg (1957), nom. illegit.
Perennial dwarf acaulescent herb from afro-alpine
habitats, capitula sessile, usually broader than long,
Compositae
185
315. Heteracia Fisch. & C.A. Mey.
Heteracia Fisch. & C.A. Mey., Index Sem. Hort. Petrop. 1:
29 (1835); Voytenko, Bot. Zhurn. (Moscow & Leningrad)
74: 1241–1257 (1989), fruit morph.
Annual herbs, involucral bracts in two rows,
florets yellow, marginal achenes irregularly ribbed,
obconical, epappose and persistent, median
obpyramidic, beaked, epappose, inner achenes
obpyramidic, long-beaked, provided with pappus
of fine scabrid-barbellate rays. x = 4, diploids. Two
species, Crimea, Near East, central Asia, China.
316. Heteroderis (Bunge) Boiss.
Heteroderis (Bunge) Boiss., Fl. Orient. 3: 793 (1875);
Voytenko, Bot. Zhurn. (Moscow & Leningrad) 74: 1241–
1257 (1989), fruit morph.
Annual herb, involucral bracts in 2 rows, florets
yellow, achenes dimorphic, outer thickly 5–6ribbed, unbeaked, pappus absent, enclosed by
persistent star-like-arranged involucral bracts,
inner achenes caducous, clearly differentiated into
main body and long beak, body many-ribbed,
distally markedly tuberculate, with pappus of fine
scabrid-barbellate rays. x = 3, diploid. Possibly
monotypic, Near East, central Asia.
Fig. 44. Compositae-Cichorieae. Crepis biennis. A Habit.
B Lower leaf. C Middle cauline leaf. D Upper part of flowering stem. E Floret, part of pappus removed. F Achene.
G Transverse section of achene. (Ross-Craig 1962)
317. Ixeridium (A. Gray) Tzvelev
Ixeridium (A. Gray) Tzvelev in Kom., Fl. S.S.S.R. 29: 388
(1964); Pak & Kawano, Mem. Fac. Sci. Kyoto Univ., ser. Biol.
15: 29–61 (1992), rev.
among the rosette of leaves, solitary or few, involucral bracts in several rows, subequal, florets yellow
with very short ligule, pappus of scabrid-barbellate
rays. x = 2 , diploid and tetraploid (?). Monotypic,
East Africa; D. schimperi A. Rich.
Perennial herbs, taprooted, capitula very small,
with 5–12 flowers, involucral bracts in 2 rows, receptacle naked, florets yellow, white to purplish,
achenes fusiform, beaked, c. 10-ribbed, pappus of
yellow to dirty-white scabrid-barbellate rays. x = 8,
7, 5, diploids, triploids, tetraploids and hexaploids.
Thirteen species, Asia, Malesia to New Guinea.
314. Dubyaea DC.
318. Ixeris (Cass.) Cass.
Dubyaea DC., Prodr. 7:247 (1838); Stebbins, Mem. Torrey
Bot. Club 19, 3: 1–76 (1940), rev.; Shih, Acta Phytotax. Sin.
31: 432–450 (1993), rev.
Ixeris (Cass.) Cass. in Cuvier, Dict. Sci. Nat. 25: 623 (1822);
Stebbins, J. Bot. (London) 75: 43–51 (1937), rev.; Pak &
Kawano, Mem. Fac. Sci. Kyoto Univ., ser. Biol. 15: 29–61
(1992), rev.
Chorisis DC. (1838).
Perennial caulescent or acaulescent herbs, involucral bracts in several rows, receptacle naked, flowers yellow, pink, bluish or purple, achenes fusiform,
with 5–10 prominent ribs and 1–6 lesser ones, pappus rays scabrid-barbellate, yellow, white or rufous. x = 8 , diploids. Fourteen species, Himalayan
Mountains, China.
Perennial to annual herbs, stoloniferous or taprooted, involucral bracts in 2 rows, 14–51 flowers per capitulum, receptacle naked, florets yellow,
white or purplish, achenes less than 5 mm long,
fusiform, slightly compressed, c. 10-winged, pap-
186
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
pus of white, persistent scabrid-barbellate rays. x =
9, 8, 7, 6, diploids, triploids, tetraploids, hexaploids.
Ten species, Asia, Malesia to New Guinea.
319. Lagoseriopsis Kirp.
Lagoseriopsis Kirp. in Kom., Fl. S.S.S.R. 29: 726 (1964);
Voytenko, Bot. Zhurn. (Moscow & Leningrad) 74: 1241–
1257 (1989), fruit morph.
Annual herb, involucral bracts few, in 2 rows, capitula narrowly cylindrical with 4–6 flowers, receptacle naked, florets yellow, achenes heteromorphic,
outer prismatic, hairy, persistent, pappus readily
deciduous, inner fusiform apically without tubercles or a crown-like structure, pappus persistent, of
scabrid-barbellate rays. Monotypic, central Asia; L.
popovii (Krasch.) Kirp.
320. Lapsana L.
Lapsana L., Sp. Pl. 811 (1753); Pak & Bremer, Taxon 44:
13–21 (1995), rev.
Perennial herbs, heads small, involucral bracts in
two rows, receptacle naked, florets yellow, achenes
often slightly flattened, beaked, many-ribbed,
rounded at apex, pappus absent. x = 7, diploid.
Monotypic, Europe; L. communis L.
321. Lapsanastrum J.-H. Pak & K. Bremer
Lampsanastrum J.-H. Pak & K. Bremer, Taxon 44: 19 (1995);
Pak & Bremer, Taxon 44: 13–21 (1995), rev.
Biennial to annual scapose herbs, involucral bracts
in two series, florets yellow, achenes oblong, hairy,
10–13-ribbed, ribs unequal, sometimes prolonged
into apical hornlike appendages, pappus absent.
x = 8, diploid. Four species, eastern Asia.
322. Phitosia Kamari & Greuter
Phitosia Kamari & Greuter, Bot. Hron. 13: 14 (2000); Kamari
& Greuter, Bot. Hron. 13: 11–36 (2000), rev.
Perennial herb, very similar to Crepis, but with
a totally deviant chromosome number, lack of indumentum, and basally rather broad pappus rays.
x = 9, diploid. Monotypic, endemic to the Taygetos
Mountains in Greece; C. crocifolia (Boiss. & Heldr.)
Kamari & Greuter. Possibly not belonging to Crepidinae.
323. Soroseris Stebbins
Soroseris Stebbins, Mem. Torrey Bot. Club 19, 3: 27 (1940);
Stebbins, Mem. Torrey Bot. Club 19, 3: 1–76 (1940), rev.;
Shih, Acta Phytotax. Sin. 31:432–450 (1993),rev.
Stebbinsia Lipsch. (1956).
Dwarf alpine perennial herbs, stems hollow, capitula on short peduncles, numerous, aggregated into
large synflorescences, involucral bracts in 2–3 rows,
receptacle naked, 4–25 flowers in each capitulum,
florets yellow or white, achenes oblong, apically
contracted, pappus rays scabrid-barbellate, yellow,
rufous or white. x = 8, diploid. Nine species, Himalayan Mountains, China.
324. Spiroseris Rech. f.
Spiroseris Rech. f., Fl. Iran. 122: 338 (1977).
Incompletely known perennial herb, involucral
bracts laxly arranged, outer foliaceous, inner
blackish, florets yellow, achenes fusiform, 4ribbed, pappus a single row of scabrid-barbellate,
basally connate rays. Monotypic, known only
from three collections, endemic to Pakistan; S.
phyllocephala Rech. f., possibly related to Dubyaea
DC.
325. Taraxacum Weber
Taraxacum Weber in F.H. Wigg., Prim. Fl. Holsat. 56 (1780),
nom. cons.; Sterk et al., Biblioth. Koninkl. Ned. Natuurhist.
Veren. 42: i–ii, 1–348 (1987), rev.
Perennial scapose herbs with taproot, leaves in
basal rosette, scape leafless, hollow, involucral
bracts in two unequal series, the outer shorter,
often reflexed at tips, receptacle naked, capitula
with very many flowers, florets yellow, achenes
obovoid, fusiform, ribbed, distally scabrid or
warty-tuberculate, with distinct long, slender
beak, pappus of many scabrid rays. x = 8, diploids,
triploids, tetraploids, pentaploids, hexaploids,
septemploids, octoploids, decaploids. Cosmopolitan, but mainly in the northern hemisphere,
a single section in southern South America,
Australia and New Zealand. Numerous clones
described as species.
326. Willemetia Neck.
Willemetia Neck., Willemetia 1 (1777–1778); Kirschnerová
& Kirschner, Taxon 45: 627–630 (1996), rev.
Calycocorsus F.W. Schmidt (1795).
Perennial herbs with creeping rootstock, scapose
or stem branched, involucral bracts in two rows,
receptacle naked, flowers yellow, achenes beaked,
ribbed, with five squamules forming a distinct apical collar, abruptly narrowing to a fragile, thin ros-
Compositae
trum, pappus of scabrid bristles. x = 5, diploids.
Two species, Europe, Caucasus, east to Iran.
327. Youngia Cass.
Youngia Cass., Ann. Sci. Nat. (Paris) 23: 88 (1831); Babcock
& Stebbins, Publ. Carnegie Inst. Wash. 484:1–106 (1937),
rev.
Perennial, biennial to annual herbs, involucral
bracts in two rows, usually 8 inner bracts, becoming
carinate in fruit, receptacle naked, 5–30 flowers per
capitulum, florets yellow, achenes less than 5 mm,
unequally many-ribbed, pappus of many scabridbarbellate rays, white, yellow, fuscous to cinereous. x = 8, 6, 5, diploids. Circa 30 species, Asia.
VIII.5. Subtribe Lactucinae Cass. ex Dumort.
(1827).
Capitula mostly slender and with few flowers, florets often non-yellow, achenes often compressed
and ribbed, often with a beak, carpopodium
funnel-shaped, callose, pappus rays fine, at least
some basally with very few cells in diameter. Old
World tribe, Lactuca and Prenanthes also in North
America.
Key to the
Genera9
1. Pappus monomorphic, without outer ring of
extremely short setae
2
– Pappus dimorphic, with outer ring of extremely short
setae
11
2. Capitula sessile, aggregated into compact synflorescences
339. Syncalathium
– Capitula sessile or pedunculate, in more or less lax to
spiciform inflorescences
3
3. Achenes compressed, with broad, thin, wing-like margins
336. Pterocypsela
– Achenes columnar to compressed but without broad,
thin, wing-like margins
4
4. Achenes without distinct apical beak, at most gradually narrowed in upper part into a short rostrum
5
– Achenes with distinct apical beak more or less distinct
in colour and/or texture from achene body
331. Lactuca p.p.
5. Capitula with more than 7 florets or, if fewer, then
plants scandent
6
– Capitula with 5–7 florets
10
6. Achenes more or less cylindrical, at most a little narrowed towards apex, apex truncate
7
– Achenes more or less fusiform, narrowed towards apex
9
7. Achenes with 4–5 ribs or more or less terete; rhizomatous perennials
335. Prenanthes p.p.
9
By C. Jeffrey.
187
– Achenes with 10–12 ribs; rhizomatous perennials or
taprooted biennials
8
8. Rhizomatous perennials (E Asia)
331. Lactuca p.p.
– Taprooted biennials (E Asia, North America)
332. Nabalus
9. Achenes more or less strongly compressed, ribs on
margins often inflated
331. Lactuca p.p.
– Achenes not or little compressed, ribs narrow, equal
334. Paraprenanthes
10. Achenes not compressed, with 4–5 ribs
335. Prenanthes p.p.
– Achenes compressed, with 10–20 ribs 333. Notoseris
11. Achenes with unequal ribs, margins thickened, often
rugose
12
– Achenes with equal ribs, margins not thickened 13
12. Florets 10–40
329. Chaetoseris
– Florets 3–5
337. Stenoseris
13. Achenes compressed, without beak and with 12–18
ribs, or with filiform beak and 4–12 unequal ribs 14
– Achenes little or not compressed, with short to long,
stout beak and with 12–18 more or less equal ribs
328. Cephalorrhynchus
14. Achenes with filiform beak
15
– Achenes not beaked, with 12–18 ribs 330. Cicerbita
15. Achenes with 12 unequal ribs
331. Lactuca p.p.
– Achenes with 4 strongly unequal ribs, the two lateral
wing-like, the two facial weak 338. Steptorhamphus
Genera of Lactucinae
328. Cephalorrhynchus Boiss.
Cephalorrhynchus Boiss., Diagn. Pl. Orient. ser. 1, 1(4): 28
(1844).
Perennial to biennial herbs with tuberous rootstocks, involucral bracts in several rows, receptacle naked, florets yellow, white, violet or purple,
achenes fusiform, beaked, not compressed, pappus
capillary. x = 9, 8, diploids. South-eastern Europe
to China, 14 species.
329. Chaetoseris C. Shih
Chaetoseris C. Shih, Acta Phytotax. Sin. 29: 398 (1991); Shih,
Acta Phytotax. Sin. 29: 394–417 (1991), rev.
Perennial to annual herbs, involucral bracts in 3–5
rows, receptacle naked, 10–40 flowers in each capitulum, florets purple to violet, rarely yellow, achenes compressed and beaked, margins thickened
and winged, pappus rays capillary. Eastern Asia, 18
species.
330. Cicerbita Wallr.
Cicerbita Wallr., Sched. Crit. 433 (1822).
Perennial herbs, rhizomatous or taprooted, leaves
clasping the stem, involucral bracts in several
rows, receptacle naked, florets blue, lilac or violet,
188
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
sometimes yellow, achenes compressed, not
beaked, pappus rays capillary. x = 9, 8, diploids.
Eurasia, c. 5 species.
331. Lactuca L.
Fig. 45
Lactuca L., Sp. Pl. 795 (1753); Stebbins, Bull. Jard. Bot. État
14: 333–352 (1937), rev.; Tuisl, Ann. Naturhist. Mus. Wien
72: 587–638 (1968), reg. rev.; Feráková, The genus Lactuca
L. in Europe (Bratislava, 1975), reg. rev.; Dethier, Bull. Jard.
Bot. Belg. 52: 367–382 (1982), reg. rev.; Koopman et al.,
Amer. J. Bot. 85: 1517–1530 (1998), mol. phylog.
Scariola F.W. Schmidt (1795).
Mulgedium Cass. (1824).
Mycelis Cass. (1824).
Lagedium Soják (1961).
Lactucella Nazarova (1990).
Faberiopsis Shih (1996).
Perennial to annual herbs, rarely scandent or
subshrubby, heads often in panicles, involucral
bracts in several rows, receptacle naked, florets
yellow, bluish, violet or white, achenes compressed, abruptly constricted, beaked, pappus rays
capillary, white to yellowish. x = 9 , 8, 6, diploids,
triploids and tetraploids. Northern hemisphere, in
Asia to New Guinea, in North America to Mexico,
in tropical Africa to South Africa, c. 60 species.
Lactuca sativa L. (lettuce) is cultivated for its
edible leaves. The single specimen known so far of
Faberiopsis Shih & Y.L. Chen is characterized by
blue florets and is unique in Cichorieae by possessing deeply trisect ligules with a tridentate central
and two simple lateral lobes (Shih and Chen 1996).
Since this may be an aberrant form, Faberiopsis is
tentatively regarded here as a synonym of Lactuca.
332. Nabalus Cass.
Nabalus Cass. in Cuvier, Dict. Sci. Nat. 34: 94 (1825); Milstead, A revision of the North American species of Prenanthes, Thesis, Purdue University (1964), reg. rev.; Sennikov,
Novosti Sist. Vyssh. Rast. 32:177–181 (2001), rev.
Perennial herbs, involucral bracts 5–15, capitula
with 5–38 flowers, receptacle naked, florets blue,
lavender, white and yellow, pappus capillary, yellow to red-brown. x = 8, diploids. Eastern Asia,
North America, 18 species.
333. Notoseris C. Shih
Notoseris C. Shih, Acta Phytotax. Sin. 25: 196 (1987); Shih,
Acta Phytotax. Sin. 25: 189–203 (1987), rev.
Perennial herbs, involucral bracts in 3–5 rows, purplish, receptacle naked, 3–5 flowers in each capitulum, florets purple, achenes compressed, purple
to reddish-brown, truncate, pappus rays capillary.
Eastern Asia, 12 species.
334. Paraprenanthes Chang ex C. Shih
Paraprenanthes Chang ex C. Shih, Acta Phytotax. Sin. 26:
418 (1988); Shih, Acta Phytotax. Sin. 26: 382–393, 418–428
(1988), reg. rev.
Fig. 45. Compositae-Cichorieae. Lactuca saligna. A Habit
of young plant. B Lower part of older plant. C Floret, part
of pappus removed. D Achene and transverse section of
achene. E Upper part of fruiting stem. (Ross-Craig 1963)
Perennial to annual herbs, involucral bracts in 2–3
rows, receptacle naked, 7–15 flowers in each capitulum, florets pink or purple, achenes black, beakless,
pappus rays capillary. Fifteen species, eastern Asia
to Malesia.
Compositae
335. Prenanthes L. s. str.
Prenanthes L., Sp. Pl. 797 (1753); Stebbins, Bull. Jard. Bot.
État 14: 333–352 (1937), reg. rev.; Milstead, A revision of
the North American species of Prenanthes, Thesis, Purdue
University (1964), reg. rev.
Faberia Hemsl. (1888).
Perennial herbs, one species scandent, leaves clasping the stem, heads in panicles with up to 5 flowers, nodding, involucral bracts in 2–3 rows, receptacle naked, florets pink, purplish to blue, achenes compressed, not beaked, pappus rays capillary, not thickened at the base. x = 9, diploids and
tetraploids. Eight species, Eurasia, south to tropical
Africa.
336. Pterocypsela C. Shih
Pterocypsela C. Shih, Acta Phytotax. Sin. 26: 385 (1988);
Shih, Acta Phytotax. Sin. 26: 382–393, 418–428 (1988), reg.
rev.
Perennial to annual herbs, involucral bracts in 4–5
rows, receptacle naked, florets yellow, achenes compressed, with two broad wings, pappus rays capillary. Seven species, eastern Asia, China, Malesia to
New Guinea. Pterocypsela indica (L.) C. Shih (Indian lettuce) is cultivated for its edible leaves.
337. Stenoseris C. Shih
Stenoseris C. Shih, Acta Phytotax. Sin. 29: 411 (1991); Shih,
Acta Phytotax. Sin. 29: 394–417 (1991), rev.
Perennial herbs, involucral bracts 3, in a single row,
receptacle naked, 3(–5) flowers in each capitulum,
florets purple to violet, achenes compressed with
wings, pappus rays capillary. Six species, eastern
Asia.
338. Steptorhamphus Bunge
Steptorhamphus Bunge, Mém. Acad. Imp. Sci. SaintPétersbourg Divers Savans 7: 381 (1854).
Perennial herbs, mostly tuberous, involucral bracts
in several series, receptacle naked, flowers yellow
or violet, achenes compressed and winged with
long, slender beak, pappus rays capillary. x = 8,
diploids. Eight species, south-eastern Europe to
western Himalaya.
339. Syncalathium Lipsch.
Syncalathium Lipsch., Akad. N.S. Sukatschevu 75 Let.: 358
(1956); Ling Yong, Acta Phytotax. Sin. 10: 283–289 (1965),
rev.; Shih, Acta Phytotax. Sin. 31: 432–450 (1993), rev.
189
Perennial to biennial, acaulescent or caulescent
herbs, capitula sessile, numerous, aggregated
into an often compact synflorescence, involucral
bracts 3–5(6) in a single row, pink to pale green,
receptacle naked, 3–6 flowers in each capitulum,
florets pink or purple, achenes oblong, apically
contracted, compressed, pappus rays capillary.
Eight species, Himalayas, China.
VIII.6. Subtribe Sonchinae K. Bremer (1993).
Perennial to annual herbs, subshrubs or shrubs.
Receptacle naked, achenes ellipsoid-fusiform or
oblong-obovoid, mostly without beak, carpopodia
never annular-callose. Pappus of setae or dimorphic with setae and cottony hairs, always flexible.
Cosmopolitan tribe.
Key to the Genera
(in part based on Kilian 1997)
1. Rosette trees or rosette shrubs of the Juan Fernandez
Islands or succulent subshrubs of the Desventuradas
Islands
2
– Perennial to annual herbs, rosette subshrubs of Canary
Islands
3
2. Rosette trees or rosette shrubs of the Juan Fernandez
Islands
341. Dendroseris
– Succulent subshrubs of the Desaventuradas Islands
345. Thamnoseris
3. Pappus (sub)homomorphic, rays at most gradually
differing in thickness from centre towards margin 4
– Pappus dimorphic with few inner setae and numerous
outer downy or (rarely) cottony hairs
5
4. Peducles expanded upwards; marginal achenes always
truncate, glabrous and with thick, tuberculose projections; most achenes with 4 main ribs, only the very
marginal with 5 main ribs
343. Reichardia
– Peduncles never expanded upwards; marginal achenes
truncate, cuspidate or rostrate, smooth or transversely
wrinkled (but never sculptured as above), glabrous,
scabrid or papillose; all or most achenes with 5 main
ribs, otherwise marginal achenes pappillose
342. Launaea p.p.
5. All achenes with 4 main ribs and (sub)equal in ornamentation, apex shape, or size; peduncles and involucres often glandular and/or (glabrescent) white
tomentose; outer involucral bracts without distinct
scarious margin and white tip
6
– Most achenes (except innermost) with 5 main ribs,
otherwise marginally (densely) papillose; inner(most)
achenes mostly different from marginal in ornamentation, apex shape or size; peduncle and involucre
never glandular nor tomentose; outer involucral bracts
mostly with distinct (though sometimes thin) scarious
margin and/or white, mucronate tip
342. Launaea p.p.
190
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
6. Plants (sub)scapose and stoloniferous; all achenes
smooth and subcolumnar to slender subfusiform,
even marginal little compressed
340. Aetheorhiza
– Plants never (sub)scapose and stoloniferous; at least
marginal achenes wrinkled and/or distinctly compressed
344. Sonchus
Genera of Sonchinae
340. Aetheorhiza Cass.
Aetheorhiza Cass. in Cuvier, Dict. Sci. Nat. 48: 425 (1827);
Rechinger, Phyton (Horn) 16: 211–220 (1974), rev.
Perennial scapose, stoloniferous herb with tubers
along the rhizomes. Flowers yellow, achenes
fusiform, apically attenuate, slightly compressed,
4-ribbed, pappus of many setae. x = 9, diploid.
Monotypic, Europe, Mediterranean area; Ae. bulbosa (L.) Cass.
343. Reichardia Roth
Reichardia Roth, Bot. Abh. Beobacht. 35 (1787); Gallego
et al., Lagascalia 9:159–217 (1980), rev.
Perennial to annual herbs, leaves marginally somewhat spinulose, peduncles expanded upwards,
involucral bracts with scarious margins, capitula
with many flowers, florets yellow, marginal achenes truncate, glabrous with thick, tuberculose
projections. x = 9, 8, 7, diploids. Eight species,
Macaronesia, Mediterranean area, Near East,
south to tropical Africa.
344. Sonchus L.
Fig. 46
Sonchus L., Sp. Pl. 793 (1753); Boulos, Bot. Notiser 125:
287–305 (1972), 126: 155–196 (1973), 127: 7–37, 402–451
(1974), rev.; Perez de la Paz, Bot. Macaronés. IV Ci. 1: 51–65
341. Dendroseris D. Don
Dendroseris D. Don, Philos. Mag. Ann. Chem. 11: 388
(1832); Carlquist, Brittonia 19: 99–121 (1967), anat.;
Sanders et al., Opera Bot. 92: 195–215 (1987), evol.;
Crawford et al., Syst. Bot. 17: 676–682 (1992), mol. phylog.;
Kim et al., Syst. Bot. 21: 417–432 (1996), mol. phylog.;
Kim et al., Pl. Syst. Evol. 215: 85–99 (1999), mol. phylog.;
Esselman et al., Amer. J. Bot. 87: 591–596 (2000), mol.
phylog.
Rosette trees and shrubs with leaf rosettes at
apex of stems, involucral bracts in several rows,
receptacle naked or bristly, capitula variously
arranged, sometimes solitary, florets whitish or
orange-yellow, achenes compressed, rarely winged.
x = 9, tetraploids. Eleven species, endemic to Juan
Fernandez Islands (Chile).
342. Launaea Cass.
Launaea Cass. in Cuvier, Dict. Sci. Nat. 25: 321 (1822); Kilian, Englera 17: 1–478 (1997), rev.
Paramicrorhynchus Kirp. (1964).
Hexinia H.L. Yang (1992).
Perennial to annual herbs, small shrubs or subshrubs. Capitula mostly slender and with c. 7–40
flowers, involucral bracts mostly with membranous
or translucent margins, florets yellow, pappus of setaceous rays or setaceous and cottony rays. x = 9,
8, 7, 6, diploids, tetraploids, hexaploids. Fifty-four
species, Old World, one species in Australia.
Fig. 46. Compositae-Cichorieae. Sonchus arvensis. A Habit.
B Middle cauline leaf. C Upper part of flowering stem. D Involucral bracts. E Floret, part of pappus removed. F Achene.
G Part of achene surface. H Transverse section of achene.
(Ross-Craig 1963)
Compositae
(1976), rev.; Aldridge, Bot. Macaronés. IV Ci. 2: 25–58,
81–93 (1976), rev.; Lander, Telopea 1: 129–135 (1976),
rev.; Aldridge, Bot. J. Linn. Soc. 76: 249–285 (1978), anat.;
Aldridge in Bramwell, Plants and islands: 279–291 (1979),
evol.; Kim et al., Syst. Bot. 21: 417–432 (1996), mol. phylog.;
Kim et al., Proc. Natl. Acad. Sci. U.S.A. 93: 7743–7748
(1996), mol. phylog.; Kim et al., Pl. Syst. Evol. 215: 85–99
(1999), mol. phylog.; Kim et al., Pl. Syst. Evol. 215:101–118
(1999), mol. phylog.
Atalanthus D. Don (1829).
Sventenia Font Quer (1949).
Kirkianella Allan (1961).
Babcockia Boulos (1965).
Embergeria Boulos (1965).
Taeckholmia Boulos (1967).
Lactucosonchus (Sch. Bip.) Svent. (1969).
Actites Lander (1976).
Wildpretia U. and A. Reiffenberger (1996).
Perennial to annual herbs; on oceanic islands
often shrubs to subshrubs with leaves in rosettes
at apex of stems. Capitula with many flowers,
variously arranged, involucral bracts without
translucent margins, receptacle naked, florets yellow, at least marginal achenes wrinkled, pappus of
setaceous and downy rays. x = 9, 8, 7, 5, diploids,
triploids, tetraploids, hexaploids, octoploids,
2n = 90, 126 reported for Kirkianella novaezelandiae (Hook. f.) Allan. Circa 80 species, Old
World, with very few species in Australia and New
Zealand.
345. Thamnoseris Phil.
Thamnoseris Phil., Anales Univ. Chile 1875:191 (1875);
Skottsberg, Göteb. Kungl. Vetensk. Samhälles Handl. ser. B,
Mat. Naturvetensk. Skr. ser. 5, 5(6): 11–88 (1937), rev.
An incompletely known succulent subshrub with
leaf rosette at apex of stems, capitula aggregated into globular synflorescences, the latter
on branched terminal shoots, florets yellow (?).
Monotypic, Desventuradas Islands; Th. lacera
(Phil.) Johnst., probably outside Cichorieae.
VIII.7. Subtribe Microseridinae Stebbins
(1953).
Pollen distinctly orange, style branches short,
blunt, pappus various, rarely absent. An exclusively
New World subtribe, except for Microseris found
also in Australia and New Zealand.
191
Key to the Genera
1.
–
2.
–
3.
–
4.
–
5.
–
6.
–
Pappus absent
2
Pappus well developed
3
Scapose perennial herb
349. Phalacroseris
Branched annual herb
347. Krigia cespitosa
Basal portion of pappus rays always broad
348. Microseris
Basal portion of pappus rays never broad
4
Capitula on leafless scapes
346. Agoseris
Flowering stems branched
5
Pappus consisting of long inner bristles and small
outer scales
347. Krigia
Pappus different
6
Involucral bracts in a single row, native to South America
350. Picrosia
Involucral bracts in several rows, native to North
America
351. Pyrrhopappus
Genera of Microseridinae
346. Agoseris Rafin.
Agoseris Raf., Fl. Ludov. 58 (1817).
Perennial to annual herb, scapose, involucral bracts
in 2–4 series, receptacle naked, florets yellow to redorange, achenes fusiform, tapered to beak, pappus
of scabrid-barbellate bristles. x = 9, diploids and
tetraploids. Circa 10 species, western North America, Chile and Argentina.
347. Krigia Schreb.
Krigia Schreb., Gen. Pl. 532 (1791), nom. cons.; Kim &
Turner, Brittonia 44: 173–198 (1992), rev.
Perennial to annual herbs, involucral bracts in 1–2
rows, florets yellow or orange-yellow, pappus double, single or absent, when present of few scales
or of few inner bristles surrounded by few outer
scales. x = 9, 6, 5, 4, diploids, triploids, tetraploids,
hexaploids and dodecaploids. Seven species, North
America.
348. Microseris D. Don
Microseris D. Don, Philos. Mag. Ann. Chem. 11: 388 (1832);
Chambers, Contr. Dudley Herb. 4: 207–312 (1955), rev.;
Jansen et al., Amer. J. Bot. 78: 1015–1027 (1991), mol. phylog.; Vijverberg et al., Amer. J. Bot. 86: 1448–1463 (1999),
mol. phylog.
Uropappus Nutt. (1841).
Apargidium Torr. & A. Gray (1843).
Nothocalais (A. Gray) Greene (1886).
Stebbinsoseris K.L. Chambers (1991).
Perennial to annual herbs, leaves mostly basal, involucral bracts in 2 to several series, receptacle
192
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
naked, florets white to orange, achenes cylindric
to fusiform, not beaked, with c. 10 ribs, pappus
of 5 to many narrowly lanceolate, bristle-tipped,
awn-like scales. x = 9, diploids and tetraploids.
Fourteen species, North America, Chile, Australia,
New Zealand.
349. Phalacroseris A. Gray
Phalacroseris A. Gray, Proc. Amer. Acad. Arts 7: 364 (1868).
Perennial scapose herbs, involucral bracts mostly
in a single row, basally connate, ligules yellow, fruit
4-angled, smooth, pappus absent. x = 9, diploid.
Monotypic, California; Ph. bolanderi A. Gray.
– Achenes columnar, mostly 3–6 mm
359. Stephanomeria
3. Pappus with 5 rigid awns
352. Chaetadelpha
– Pappus without rigid awns
4
4. Shrub
5
– Subshrubs, taprooted perennials or annuals
6
5. Shrub with fleshy, brownish, tomentose stems (San
Clemente Island)
355. Munzothamnus
– Shrub with succulent leaves (Chichuachuan Desert,
Mexico)
354. Marshalljohnstonia
6. Subshrubs, branches becoming sharp thorns
356. Pleiacanthus
– Perennial or annual herbs, unarmed
7
7. Lower leaves opposite, achenes 8–10-sulcate
358. Shinnersoseris
– Lower leaves alternate, achenes variously sculptured,
but never 8–10-sulcate
353. Lygodesmia
350. Picrosia D. Don
Picrosia D. Don, Trans. Linn. Soc. London 16:183 (1832).
Genera of Stephanomeriinae
Perennial herbs, involucral bracts in a single series, receptacle naked, flowers white, pink or violet,
achenes fusiform, with long beak, pappus of many
scabrid bristles. x = 7, diploids. Two species, South
America.
352. Chaetadelpha A. Gray ex S. Watson
351. Pyrrhopappus DC.
Pyrrhopappus DC., Prodr. 7: 144 (1838), nom. cons.;
Northington, Spec. Publ. Mus. Texas Tech. Univ. 6: 1–38
(1974), rev.; Turner & Kim, Amer. J. Bot. 77: 845–850 (1990),
rev.
Perennial or biennial herbs, subscapose, involucral
bracts in two series, receptacle naked, flowers
yellow, achenes subfusiform, long tapering into
filiform beak, smooth towards the tip of the
body, pappus of thin brownish to reddish rays,
surrounded at base by a short villous ring. x = 6,
diploids and tetraploids. Three species, North
America.
VIII.8. Subtribe Stephanomeriinae Stebbins
(1953).
Capitula few-flowered, non-yellow flowers, achenes
narrowly terete to fusiform, pappus of scabrid or
plumose rays. Exclusively New World subtribe.
Key to the Genera
1. Pappus rays plumose
2
– Pappus rays barbellate or pappus of 5 rigid awns and
slender bristles
3
2. Achenes mostly beaked, at least 8 mm
357. Rafinesquia
Chaetadelpha A. Gray ex S. Watson, Amer. Naturalist 7: 301
(1873); Tomb, Madroño 21: 459–462 (1972), rev.
Perennial herb, involucral bracts in 2 series,
receptacle naked, 5 flowers per head, ligule pale
lavender-white, pappus dimorphic with many fine
rays and five rigid, thick awns. x = 9, diploid.
Monotypic, endemic to south-western USA;
Ch. wheeleri A. Gray.
353. Lygodesmia D. Don
Lygodesmia D. Don, Edinburgh New Philos. J. 1829: 311
(1829); Tomb, Brittonia 24: 223–228 (1972), rev.; Tomb, Syst.
Bot. Monogr. 1: 1–51 (1980), rev.
Prenanthella Rydb. (1906).
Perennial or annual herbs, rarely subshrubs, capitula few-flowered, involucral bracts in two unequal
rows, flowers pink, purplish or white, pappus of distichously barbellate rays. x = 9, 7, diploids. Nine
species, southern Canada to northern Mexico.
354. Marshalljohnstonia Henr.
Marshalljohnstonia Henr., Syst. Bot. 1: 171 (1976); Henrickson, Syst. Bot. 1: 169–180 (1976), rev.
Shrub, leaves fleshy, involucral bracts in several
rows, receptacle naked, 10–18 flowers per capitulum, florets pink to purple, cylindric achenes,
pappus of slightly flattened, distichously barbellate
bristles, pollen echinate. x = 9, diploid. Monotypic,
endemic to the Chichuachuan Desert in Mexico;
M. gypsophila Henr.
Compositae
355. Munzothamnus Raven
Munzothamnus Raven, Aliso 5: 345 (1963); Lee et al., Amer.
J. Bot. 89: 160–168 (2002), mol. phylog.
Shrub with fleshy, brownish tomentose stems,
leaves tufted at branch tips, inflorescences densely
stalked-glandular, florets light purple, pappus of
non-plumose bristles. x = 8, diploid. Monotypic,
endemic to San Clemente Island (California); M.
blairii (Munz & Johnst.) Raven.
356. Pleiacanthus (T. Nutall) Rydb.
Pleiacanthus (T. Nutall) Rydb., Fl. Rocky Mount. 1023
(1918); Lee et al., Amer. J. Bot. 89: 160–168 (2002).
Subshrub, woolly at base, branches becoming
thick thorns, leaves linear to scale-like, florets
3–5, pink, pappus of non-plumose bristles. x = 8,
diploid. Monotypic, south-western United States;
P. spinosus Rydb.
357. Rafinesquia Nutt.
Rafinesquia Nutt., Trans. Amer. Philos. Soc. n.s. 7:429
(1841), nom. cons.
Annual herbs, leaves basal and cauline, involucral
bracts in 3–4 series, outer series often with
membranous margins, receptacle naked, florets
white to cream, often rose-tinged, achenes smooth
or tubercled, weakly ribbed, tapered to a beak,
pappus of stiff distichously plumose rays. x = 8,
diploids. Two species, California to New Mexico.
358. Shinnersoseris Tomb
Shinnersoseris Tomb, Sida 5: 186 (1973); Tomb, Sida 5: 183–
189 (1973), rev.
Annual herb, lower leaves opposite, capitula fewflowered, involucral bracts in two unequal rows,
ligules violet-pink, c. 1 mm wide, pappus of fine
scabrid rays. x = 6, diploid. Monotypic, restricted
to central North America; S. rostrata (A. Gray)
Tomb.
359. Stephanomeria Nutt.
Stephanomeria Nutt., Trans. Amer. Philos. Soc. n.s. 7:427
(1841), nom. cons.; Gottlieb, Madroño 21: 463–481 (1972),
rev.
Subshrubs, perennial to annual herbs, involucral
bracts in 2 to several rows, receptacle naked,
capitula few-flowered, ligules lavender, pink or
whitish, pappus of stiff distichously plumose
rays. x = 8, diploids and tetraploids. Twenty-two
193
species, British Columbia to Baja California, east
to Texas.
VIII.9. Subtribe Malacothricinae K. Bremer
(1993).
Perennial to annual herbs, receptacle generally
bristly-scaly, rarely paleate, pappus of slender, barbellate or plumose rays, rarely absent. Exclusively
New World subtribe.
Key to the Genera
1.
–
2.
–
3.
–
4.
–
5.
Pappus absent
361. Atrichoseris
Pappus present
2
Pappus plumose
360. Anisocoma
Pappus various, but not plumose
3
Prostrate herbs
363. Glyptopleura
Plants different
4
Achenes beaked
5
Achenes columnar
364. Malacothrix
Perennial herbs, receptacle paleate, pappus brownish
365. Pinaropappus
– Annual herbs, receptacle naked, pappus whitish
362. Calycoseris
Genera of Malacothricinae
360. Anisocoma Torr. & A. Gray
Anisocoma Torr. & A. Gray, Boston J. Nat. Hist. 5: 111 (1845).
Annual scapose herb, involucral bracts in several
series, with broad papery-transparent margins, receptacle naked, ligules pale yellow, pappus of stiff
distichously arranged plumose rays. x = 9, diploid.
Monotypic, restricted to south-western USA and
northern Mexico; A. acaulis Torrey & A. Gray.
361. Atrichoseris A. Gray
Atrichoseris A. Gray, Syn. Fl. N. Am. 1, 2: 410 (1884).
Annual herb, leaves mostly basal, involucral
bracts in 2–4 series, receptacle naked, ligules
white, fragrant, achenes with 5 thick, white, corky
ribs, pappus absent. x = 9, diploid. Monotypic,
restricted to south-western USA; A. platyphylla
A. Gray.
362. Calycoseris A. Gray
Calycoseris A. Gray, Smithsonian Contr. Knowl. 5: 104
(1853).
Annual herb, leaves basal and cauline, involucral
bracts scarious-margined, in 2 series, receptacle
minutely bristly, ligules yellow to white, achenes
194
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
tapered to short beak, pappus of slender rays.
x = 9, diploids. Two species, south-western USA to
north-western Mexico.
363. Glyptopleura Eaton
Glyptopleura Eaton, Bot. King’s Exped. 207 (1871).
Annual herb, forming small tufts, stems semiprostrate, involucral bracts in 2 series, receptacle
naked, 7–16 flowers per capitulum, ligules cream
to pale yellow, achenes with 5 ribs alternating
with 5 rows of pits, abruptly short-beaked, pappus
of slender rays. x = 9, diploid. Monotypic, restricted to western North America; G. marginata
Eaton.
Key to the Genera
1. Pappus absent, peduncles swollen in fruit
368. Hispidella
– Pappus well developed, peduncles not swollen
2
2. Perennial, stoloniferous herbs
3
– Annual or perennial herbs, never stoloniferous
4
3. Achenes 4-angled, plants nearly glabrous
369. Hololeion
– Achenes c. 10-ribbed, plants always densely hairy
370. Pilosella
4. Receptacle ciliate with long hairs or scales
366. Andryala
– Receptacle naked or shortly fimbriate
6
5. Style branches long
367. Hieracium
– Style branches short
371. Tolpis
Genera of Hieraciinae
366. Andryala L.
364. Malacothrix DC.
Malacothrix DC., Prodr. 7: 192 (1838); Williams, Amer.
Midl. Naturalist 58: 494–512 (1957), rev.; Davis, Madroño
44: 223–244 (1997), reg. rev.
Perennial to annual herbs, involucral bracts in
3–6 series, receptacle naked or with fragile bristles,
ligules yellow or white, achenes fusiform, truncate,
pappus of slender rays. x = 9, 8, 7, diploids and
tetraploids. Twenty-one species, western North
America to southern South America.
365. Pinaropappus Less.
Pinaropappus Less., Syn. Gen. Comp. 143 (1832); McVaugh,
Contr. Univ. Michigan Herb. 9: 359–484 (1972), reg. rev.
Perennial herbs, scapose or with leaves in basal
part of stem, receptacle paleaceous, ligules white or
pink, achenes fusiform, tapering into short beak,
pappus of numerous brownish capillary rays. x = 9,
8, diploids and tetraploids. Eight species, southern
USA to Guatemala.
VIII.10. Subtribe Hieraciinae
Cass. ex Dumort. (1827).
Leaves of various form, never entire and parallelnerved, mostly with both branched and unbranched hairs. Achenes obovoid-obconical,
not compressed, without beak. Pappus bristles
scabrid-barbellate, never thick or plumose, rarely
absent. Old World subtribe, with Hieracium also
in America.
Andryala L., Sp. Pl. 808 (1753).
Pietrosia Nyárády ex Sennik. (2000).
Perennial to annual tomentose herbs with
branched and unbranched hairs, receptacle ciliate
or scaly, florets yellow or orange, style branches
long, achenes terete to obconical, glabrous,
about 10-ribbed, pappus scabrid to barbellate.
x = 9, diploids. Circa 25 species, Macaronesia,
Mediterranean region.
367. Hieracium L.
Fig. 47
Hieracium L., Sp. Pl. 799 (1753); Beaman, Syst. Monogr. 29:
1–77 (1990), reg. rev.
Stenotheca Monn. (1829).
Perennial herbs with branched stocks, but without
stolons, leaves and stems with branched and unbranched hairs, involucral bracts in several rows,
receptacle naked, florets yellow or rarely reddish,
style branches long, achenes cylindric, 10-ribbed,
ribs apically confluent into an obscure ring, pappus
of scabrid-barbellate fragile bristles in two rows.
x = 9, diploids, triploids, tetraploids, pentaploids.
Depending on species concept, c. 90 to more than
1,000 species. Eurasia, North and South America.
Numerous clones described as species.
368. Hispidella Barnadez ex Lam.
Hispidella Barnadez ex Lam., Encycl. 3: 134 (1789).
Annual herb with branched and unbranched hairs,
involucral bracts in a single row, receptacle hairy,
peduncles in fruit expanded upwards, florets yellow, inner florets much smaller, brownish, style
branches long, achenes 10-ribbed, pappus absent.
Compositae
195
370. Pilosella Vaill.
Pilosella Vaill., Königl. Akad. Wissensch. Paris Phys. Abh.
5: 703 (1754).
Perennial herbs with stolons, with branched and
unbranched hairs, capitula on more or less leafless
scapes, involucral bracts in several rows, receptacle naked, florets yellow, style branches long,
achenes cylindric, 10-ribbed, each rib projecting
above to form a finely scalloped apex, pappus of
fragile, scabrid-barbellate bristles in a single row.
x = 9, diploids, triploids, tetraploids, pentaploids,
hexaploids, heptaploids, octoploids, nonaploids,
decaploids. Depending on species concept, c.
20–80 species. Eurasia, northern Africa. Numerous
clones described as species.
371. Tolpis Adans.
Tolpis Adans., Fam. Pl. 2: 112 (1763); Mort et al., Taxon 52:
511–518 (2003), mol. phylog.
Fig. 47. Compositae-Cichorieae. Hieracium umbellatum.
A Habit. B Middle and upper part of plant. C Leaf shape
variation. D Involucral bracts. E Floret, part of pappus removed, pappus hair further enlarged. F Achene. (Ross-Craig
1963)
x = 9, diploid. Monotypic, restricted to Spain and
Portugal; H. hispanica Barnadez ex Lam.
369. Hololeion Kitam.
Hololeion Kitam., Acta Phytotax. Geobot. 10:301 (1941).
Perennial stoloniferous herbs, nearly glabrous,
involucral bracts in two rows, receptacle naked,
flowers yellow, style branches long, achenes
4-angled, pappus of scabrid-barbellate bristles.
x = 8, diploids. Three species, eastern Asia.
The stoloniferous growth form is regarded as
the main argument against the recent proposal by
Sennikov and Illarinova (2001) to include Hololeion
in Crepis, where this character is never found.
Perennial to annual herbs, rarely subshrubs,
leaves mostly basal, involucral bracts in 2–3 rows,
receptacle naked, florets yellow, often becoming
green when dry, style branches short, inner florets
often much smaller and brownish, achene ribbed,
truncate at apex, pappus of scabrid-barbellate
bristles or very short scales or both. x = 9,
diploids, tetraploids, hexaploids. Circa 15 species,
Macaronesia, Mediterranean region east to Iran,
and south through eastern Africa to southern
Africa.
VIII.11. Subtribe Hypochaeridinae Less.
(1832).
Leaves of various form, never entire and parallelveined, mostly hirsute or hispid, never with soft
branched hairs. Pappus rays thick, often plumose
with many to very few stiff pinnulae orientated in
all directions, more rarely scabrid-barbellate, occasionally reduced or absent. Except for Hypochaeris
(also in South America) and Picris (also in Aleutian
Islands), an exclusively Old World subtribe (Lack
1979).
Key to the Genera
1. Pappus absent in all achenes
2
– Pappus present at least in central achenes
4
2. Outer achenes stellately patent, inner achenes straight
381. Rhagadiolus
– Achenes not stellately patent
3
196
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
3. Perennial, rosette of deeply dentate leaves
372. Aposeris
– Annual, rosette of spathulate leaves
373. Arnoseris
4. Pappus of central achenes often reduced to a crownlike structure
374. Garhadiolus
– Pappus of central achenes well developed
5
5. Median and central achenes winged
377. Hyoseris
– Median and central achenes not winged
6
6. Pappus of central achenes consisting of setaceous rays
with few or no pinnulae
375. Hedypnois
– Pappus of central achenes consisting of plumose rays
with many pinnulae
7
7. Receptacle paleaceous
378. Hypochaeris
– Receptacle naked
8
8. Achene bent, with a diaphragm, involucral bracts in
a single row
382. Urospermum
– Achene not bent, without a diaphragm, involucral
bracts in two or several rows
9
9. Anchor-shaped hairs consistently present
10
– Glabrous or various hair types present, but no anchorshaped hairs
379. Leontodon
10. Outer involucral bracts conspicuously cordate
376. Helminthotheca
– Outer involucral bracts inconspicuous, never cordate
380. Picris
outer incurved, subulate, persistent and partly enclosed by the persistent involucral bracts, central
incurved, beaked, caducous, apically either with
a crown-like structure or with pappus bristles. x =
5, diploids. Two species, Caucasus, Near East, central Asia, eastern Asia.
375. Hedypnois Mill.
Hedypnois Mill., Gard. Dict. abr. ed. 4 (1754); Nordenstam,
Bot. Notiser 124: 483–489 (1971), chrom. nos.
Annual herbs, involucral bracts in two rows, receptacle naked, florets yellow, achenes incurved,
ribbed, dimorphic, outer persistent, partly or completely enclosed by the persistent involucral bracts,
inner caducous, pappus of central achenes of few
coarse bristles, sometimes with few pinnulae, pappus of outer achenes a crown-like structure. x =
9, 8, 7, 6, 5, 4, 3, diploids and tetraploids. Three
species, Mediterranean region east to Iran.
376. Helminthotheca Vaill.
Genera of Hypochaeridinae
372. Aposeris Neck. ex Cass.
Aposeris Neck. ex Cass., Dict. Sci. Nat. 48: 427 (1827).
Perennial scapose herb, leaves pinnate with
triangular-subrhombic lobes in basal rosette,
involucral bracts in two rows, receptacle naked,
ligules yellow, achenes obovoid, pappus absent.
x = 8, diploid. Monotypic, Europe; A. foetida (L.)
Less.
Helminthotheca Vaill., Königl. Akad. Wissensch. Paris
Phys. Abh. 5: 731 (1754); Holzapfel, Willdenowia 24: 97–218
(1994), reg. rev.
Perennial to annual herbs, with rigid anchorshaped hairs, involucral bracts in two rows, outer
ovate to cordate, inner lanceolate, receptacle naked,
florets yellow, achenes somewhat heteromorphic,
provided with a long beak, pappus of plumose
bristles with stiff pinnulae. x = 5, diploids. Four
species, Mediterranean region.
373. Arnoseris Gaertn.
377. Hyoseris L.
Arnoseris Gaertn., Fruct. Sem. Pl. 2: 355 (1791).
Hyoseris L., Sp. Pl. 808 (1753).
Annual scapose herb, stems leafless, branched, peduncles expanded upwards, involucral bracts in
a single row, receptacle naked, flowers yellow, style
branches short, achenes 3–5 angled, pappus absent.
x = 9, diploid. Monotypic, Europe; A. minima (L.)
Schweigger & Koerte.
The unconventional placement of this genus
in Hypochaeridinae is based on molecular data
(Whitton et al. 1995) and on achene characters
(Sennikov and Illarionova 2001).
Perennial to annual scapose herbs, involucral
bracts in two rows, receptacle naked, florets yellow,
outer achenes compressed, median compressed
and winged, inner terete and winged, pappus of
scales or of scales and scabrid bristles. x = 8,
diploids. Two species, Mediterranean region.
378. Hypochaeris L.
Garhadiolus Jaub. & Spach, Ill. Pl. Orient. 3: 119 (1849).
Hypochaeris L., Sp. Pl. 810 (1753); Samuel et al., Amer. J.
Bot. 90: 496–507 (2003), mol. phylog.; Weiss-Schneeweiss
et al., Pl. Syst. Evol. 241: 171–184 (2003), karyol.
Trommsdorffia Bernh. (1800).
Fabera Sch. Bip. (1845).
Annual herbs, involucral bracts in two rows, receptacle naked, florets yellow, achenes dimorphic,
Perennial to annual herbs with coarse, multicellular, unbranched hairs, involucral bracts in several
374. Garhadiolus Jaub. & Spach
Compositae
197
rows, receptacle scaly, florets yellow, white or pink,
achenes beaked, sometimes dimorphic, pappus of
plumose bristles with stiff pinnulae. x = 6, 5, 4, 3,
diploids and tetraploids. Circa 50 species, Eurasia,
northern Africa, Columbia and western Venezuela
to Chile and Argentina.
Testa characters indicate that this genus may
not be as homogenous as previously thought (Tegel
2002).
379. Leontodon L.
Leontodon L., Sp. Pl. 798 (1753); Widder, Phyton (Horn) 17:
23–29 (1975), rev.
Perennial to annual scapose herbs, rarely tuberous, variously hairy but never with anchor-shaped
hairs, leaves usually in basal rosette, involucral
bracts in 2 to several rows, receptacle naked, florets
yellow, pappus of plumose bristles with stiff pinnulae. x = 7, 6, 4, diploids and tetraploids. Circa 40
species, Europe, Mediterranean region to Iran. The
genus is probably not monophyletic.
380. Picris L.
Fig. 48
Picris L., Sp. Pl. 792 (1753); Lack, Diss. Univ. Wien 116:
1–184, cvi (1975), reg. rev.; Holzapfel, Willdenowia 24:
97–218 (1994), reg. rev.
Perennial to annual herbs, with rigid anchorshaped hairs, stems branched, involucral bracts
in 2 to several rows, receptacle naked, ligules
yellow, pappus of plumose bristles with stiff
pinnulae, rarely reduced to crown-like structures
in outer achenes. x = 5, diploids, tetraploids and
hexaploids. Circa 50 species, Eurasia, Aleutian
Islands, south to tropical Africa, east to Australia
and New Zealand.
Fig. 48.
Compositae-Cichorieae. Picris hieracioides.
A Habit. B Middle cauline leaf. C Flowering stems. D Hairs
on leaf margin. E Floret, part of pappus removed. F Achene
with and without pappus. G Transverse section of achene.
(Ross-Craig 1962)
382. Urospermum Scop.
381. Rhagadiolus Vaill.
Rhagadiolus Vaill., Königl. Akad. Wissensch. Paris Phys.
Abh. 5: 737 (1754); Meikle, Taxon 28: 133–141 (1979),
rev.; Voytenko, Bot. Zhurn. (Moscow & Leningrad) 74:
1241–1257 (1989), fruit morph.
Annual herbs, involucral bracts in 2 rows, florets
yellow, achenes subulate or narrowly fusiform,
straight or curved, outer persistent, stellately
patent, smooth, completely enclosed by the persistent involucral bracts, inner strongly curved,
caducous, often hispidulous, pappus absent. x = 5,
diploids. Two species, Mediterranean area, Near
East.
Urospermum Scop., Intr. Hist. Nat. 122 (1777), nom. cons.
prop.; Lack & Leuenberger, Pollen Spores 21: 415–425
(1979), pollen.
Tragopogonoides Vaill. (1754), nom. rej. prop.
Perennial or annual herbs, capitula solitary at
the end of branches, involucral bracts in a single
row, basally connate, receptacle naked, florets pale
yellow, achenes with a basally swollen, hollow beak
separated by a diaphragm from the proximal, compressed, conspicuously rugose embryo-containing
part, pappus of plumose bristles with stiff pinnulae. x = 7, 5, diploids. Two species, Macaronesia,
Mediterranean region, Near East.
198
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
VIII.12. Subtribe Scorzonerinae
Cass. ex Dumort. (1827).
Leaves frequently entire, linear-lanceolate and
parallel-veined, glabrous or tomentose, never hirsute or hispid. Pappus rays at least in basal part and
in central flowers plumose, pinnulae intertwined,
cottony, soft; when pappus absent, achenes with
many hooked projections. Exclusively Old World
subtribe.
Key to the Genera
1. Achenes with hooks or glochids, strongly incurved
385. Koelpinia
– Achenes without hooks or glochids, never strongly
incurved
2
2. Achenes with distinct wings
3
– Achenes without wings
4
3. Achenes columnar to fusiform, with three broad wings
386. Pterachaenia
– Achenes compressed, with two thin wings
388. Tourneuxia
4. Involucral bracts in a single row
5
– Involucral bracts in two to several rows
6
5. Pappus of all achenes of many woolly-plumose rays
389. Tragopogon
– Pappus of marginal achenes of five scabrid rays, pappus of inner achenes with many woolly-plumose rays
384. Geropogon
6. Distal part of achenes densely lanate
383. Epilasia
– Achenes completely glabrous or completely hairy
387. Scorzonera
x = 7, diploid. Monotypic, Mediterranean region
to Iran; G. hybridus (L.) Sch. Bip.
385. Koelpinia Pall.
Koelpinia Pall., Reise Russ. Reich 3: 755 (1776); Nazarova,
Bot. Zhurn. (Moscow & Leningrad) 66: 1755–1758 (1981),
chrom. nos.
Annual herbs, involucral bracts often in a single
row, basally somewhat connate, florets yellow,
achenes incurved, with hooked projections or
glochids, pappus absent. x = 7, 6, diploids,
hexaploids, octoploids. Five species, Mediterranean region, Near East, central Asia.
386. Pterachaenia (Benth.) Lipsch.
Pterachaenia (Benth.) Lipsch., Fragm. Monogr. Roda Scorz.
2: 31 (1939); Lipschitz, Bot. Zhurn. (Moscow & Leningrad)
56: 1150–1152 (1971), rev.; Safavi, Iran. J. Bot. 8: 241–243
(2000), rev.
Annual scapose herbs, florets pale yellow, achenes
with three broad whitish wings, spinulose between
wings, pappus of plumose bristles, pinnulae
intertwined. x = 6, diploid. Monotypic, restricted
to Iran, Afghanistan and Pakistan; P. stewartii
(Hook. f.) R.R. Stewart.
387. Scorzonera L.
Genera of Scorzonerinae
383. Epilasia (Bunge) Benth.
Epilasia (Bunge) Benth. in Benth. & Hook. f., Gen. Pl. 2:
532 (1873).
Annual herbs, leaves parallel-veined, involucral
bracts in a single row, florets pale yellow or violetpink, achenes without wings, ribbed, hispid, upper
portion of achene hairy, pappus of many plumose
bristles, pinnulae intertwined. x = 6, diploids.
Three species, Caucasus, Near East, central Asia.
384. Geropogon L.
Geropogon L., Sp. Pl. ed. 2, 1109 (1763); Díaz de la Guardia
& Blanca, Blancoa 9: 31–44 (1986), rev.
Annual herb, leaves parallel-veined, involucral
bracts in a single row, basally somewhat connate,
florets violet or purple, in outer achenes pappus
of five thick scabrous bristles, in inner achenes
pappus of plumose bristles, pinnulae intertwined.
Scorzonera L., Sp. Pl. 790 (1753); Lipschitz, Fragmenty k
monografii roda Scorzonera L. 1–2 (Moskva 1935, 1939),
rev.; Kamelin & Tagaev, Bot. Zhurn. (Moscow & Leningrad)
71:1672–1682 (1986), rev.; Mavrodiev et al., Taxon 53:
699–712 (2004), mol. phylog.
Podospermum DC. (1805).
Gelasia Cass. (1818).
Lasiospora Cass. (1822).
Achyroseris Sch. Bip. (1845).
Avellara Blanca & Díaz de la Guardia (1985).
Takhtajaniantha Nazarova (1990).
Perennial to annual herbs, often with massive
rootstock, heads often large, involucral bracts
in two or several rows, florets yellow, violet or
purple, achenes fusiform, glabrous or hairy, never
winged, pappus of plumose bristles, sometimes
apically scabrid, pinnulae intertwined. x = 7,
6, diploids and tetraploids. Circa 180 species,
Eurasia, northern Africa. Scorzonera hispanica L.
(scorzonera, Spanish salsify) cultivated for edible
roots.
Compositae
388. Tourneuxia Coss.
Tourneuxia Coss., Bull. Soc. Bot. France 6: 395 (1859).
Annual herb, involucral bracts in two rows, florets
yellow, achenes compressed, winged, pappus
of plumose bristles, pinnulae intertwined, pappus somewhat laterally positioned. Monotypic,
Morocco and Algeria; T. variifolia Coss.
389. Tragopogon L.
Fig. 49
Tragopogon L., Sp. Pl. 789 (1753); Mavrodiev et al., Taxon
53:699–712 (2004), mol. phylog.
Perennial to annual herbs, leaves parallel-veined,
heads large, involucral bracts in a single row, basally
somewhat connate, receptacle naked, ligules yellow, violet or purple, in all achenes pappus with
many plumose bristles, sometimes apically scabrid,
Fig. 49. Compositae-Cichorieae. Tragopogon pratensis. A
Habit. B Flowering stem. C Floret, part of pappus removed.
D Fruiting head. E Achene. F Upper part of achene. (RossCraig 1963)
199
pinnulae intertwined. x = 6, diploids, tetraploids
and hexaploids. Circa 110 species, Eurasia, introduced into other parts of the world, there sometimes forming allopolyploids, e.g. T. mirus Ownbey
of western North America, often precisely datable.
Tragopogon porrifolius L. (salsify) cultivated for edible roots and for young flowering shoots (‘chards’).
IX. Tribe Gundelieae DC. ex Lecoq &
Juillet (1831).
C. Jeffrey
Lacticiferous perennial herbs, shrubs or small
trees. Leaves alternate, lamina dentate-lobulate
or lobulate-pinnatisect, unarmed or very spiny.
Capitula homogamous, discoid, many-flowered,
monomorphic and solitary or 1-flowered, dimorphic and aggregated into syncephalia; phyllaries
multiseriate, imbricate and free or uniseriate and
connate; receptacle epaleaceous. Florets all perfect
or perfect and functionally staminate; corolla regular, with broadened, deeply 5-lobed limb, lobes
narrow; stamens 5; filaments glabrous; anthers
calcarate, basally sagittate, caudate or ecaudate;
endothecial cell wall thickenings polarized. Pollen
spiny. Style slender, arms short to long, flattened,
obtuse to acute, hirtellous or papillose dorsally.
Achenes oblong or ovoid, glabrous or sericeousvillous, shed free or enclosed in syncephalium;
pappus of multiseriate bristles or coroniform.
Two genera and two species, north-western
Africa to central Asia.
The re-establishment of the tribe Gundelieae,
its present circumscription, and the clarification of
its systematic position have resulted mainly from
recent molecular systematic studies (Karis et al.
2001; Funk and Chan 2003). Prior to these, the constituent genera had been variously placed systematically. Gundelia had usually been included, albeit
with some reservation, in Arctotideae, from which
it differs in its vernonioid style and pollen grain
structure (Robinson 1994) – liabioid, similar to that
of the basal Cichorieae genus Scolymus L. Warionia has generally been regarded as a member of
Mutisieae by, for example, Cabrera (1977) but was
excluded from that tribe by Hansen (1991b) on the
basis of its rugose, not mutisioid, petal epidermis
pattern and vernonioid style. The similarity of its
pollen to that of Hesperomannia A. Gray of Vernonieae, noted by Marticorena and Parra (1975),
200
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
had already suggested its position as a member
of Cichorioideae, as did the morphological cladistic study of Karis et al. (1992). The sister-group
relationship of Gundelia to Cichorieae was demonstrated by Karis et al. (2001) and confirmed by Funk
and Chan (2003), who also established the sistergroup relationship of Warionia to Gundelia. The
nature of the unique synflorescence of Gundelia
was established by Claßen-Bockhoff et al. (1989).
shaft hirtellous in upper part, arms rather short,
thick, rather obtuse, dorsally hirtellous. Achenes
oblong, glabrous, each enclosed by the accrescent, lignified, apically spiny phyllaries of the
syncalathium, forming a 1-seeded disseminule;
pappus a denticulate-fimbriate corona. n = 9. One
species, G. tournefortii L., Middle East and Turkey
to central Asia. Other described species are here
considered synonymous.
Key to the Genera
X. Tribe Arctotideae Cass. (1819).
1. Leaves spiny; perennial herb; capitula 1-flowered, aggregated into syncephalia
391. Gundelia
– Leaves not spiny; shrub or small tree; capitula large,
solitary
390. Warionia
390. Warionia Benth. & Coss.
Warionia Benth. & Coss., Bull. Soc. Bot. France 19: 165
(1872); Audissou, Succulentes 21: 30–32 (1998) & Brit. Cact.
Succ. J. 17: 124–126 (1999), col. illustr.
Shrub or small tree, aromatic; leaves coarsely
dentate-lobulate, unarmed. Capitula solitary,
terminal, large, discoid, many-flowered; phyllaries
multiseriate, imbricate, lanceolate, acute, free.
Florets all perfect. Corolla yellow, tube slender;
anthers caudate, tails short, basally slightly fringed;
apical appendage rather long, adaxially somewhat
keeled. Style base swollen, arms long, slender,
acute, dorsally hirtellous with narrow acute hairs
extending to below bifurcation. Achenes obovoid,
densely sericeous with long twin hairs; pappus of
long, multiseriate, scabrid bristles. One species,
W. saharae Benth. & Coss., North Africa (Sahara).
391. Gundelia L.
Gundelia L., Sp. Pl.: 814 (1753); Kamelin, Bot. Zhurn.
(Moscow & Leningrad) 72: 974–978 (1987), distrib.;
Nersesyan & Mekhakyan, Fl. Rastit. Rast. Arm. S.S.R. 12:
43–45 (1999), palynol.
Robust perennial herb; leaves coarsely lobulatepinnatisect, very spiny. Capitula 1-flowered,
aggregated into 5–7-capitulate syncalathia; floret
of central capitulum perfect, florets of the 4–6
outer capitula functionally staminate; syncalathia further aggregated into a large, somewhat
elongated terminal head; phyllaries uniseriate,
connate, 5 in central capitulum, 2 in outer capitula. Corolla greenish, yellow, white, pink or
red-purple, tube short; anthers ecaudate. Pollen
with broad columellae, caveate. Styles slender,
P.O. Karis
Leaves entire or often lobed to pinnatisect,
spinulose to spiny or unarmed, commonly with
woolly hairs at least beneath, sometimes also
with multiseptate, glandular or eglandular hairs,
or longitudinally striate hairs. Latex present in
Gorteriinae. Capitula frequently radiate and heterogamous, sometimes discoid and homogamous,
generally solitary, rarely corymbose or axillary;
involucral bracts usually in several rows, imbricate,
free or ± connate; ray florets 3- or 4-lobed, fertile,
sterile or neuter, sometimes with staminodes,
often yellow; disc florets deeply to more shallowly
5-lobed, perfect or sometimes functionally male,
generally yellow, lobe veins generally continuous;
anthers calcarate, caudate (most Gorteriinae)
or ecaudate (Arctotidinae), endothecial tissue
with outer periclinal and both anticlinal walls
thickened, polarised (some Gorteriinae) or radial
(all Arctotidinae) or without such patterns (most
Gorteriinae), apical appendage firm (Gorteriinae,
Hoplophyllum, Platycarpha) or soft (Arctotidinae,
Eremothamnus, Heterolepis), usually cordate-ovate
and rather short; pollen caveate, echinate, echinolophate or psilolophate; style vernonioid or
± thickened apically, with short or long style
branches, often with longer style hairs in a ring
well below the bifurcation (arctotoid); cypselae generally ± obovoid, rarely oblong-elliptic,
prismatic or obconical, frequently distinctly
ribbed, often sericeous-villous with twin hairs,
or glabrous. Pappus of scales, rarely of bristles or
absent.
Arctotideae comprise 17 genera and about 215
species.
Most recent interpretations of tribal interrelationships of Asteraceae comprise a monophyletic
subfamily Cichorioideae s. str., where the two tribes
Vernonieae and Liabeae appear as sisters (Bremer
Compositae
1996; Karis et al. 2001; Panero and Funk 2002; Funk
et al. 2004). Apart from the latter grouping, the relations within subfamily Cichorioideae s. str. remain
unclear and it is consequently not possible to judge
if Arctotideae are sister to Cichorieae, or if any of
these tribes are more closely related to Vernonieae plus Liabeae. Hoplophyllum and Eremothamnus were placed as subfam. Cichorioideae incertae
sedis by Bremer (1994). Subsequent studies clearly
position Eremothamnus (reviewed by Bremer 1996)
and its sister group Hoplophyllum (Karis et al. 2001;
Funk et al. 2004) with Arctotideae. Apart from the
strong support by molecular data (Karis et al. 2001),
these genera share two unique synapomorphies in
their anther endothecial cells with wall thickenings
in transverse bands confined to the lower part of
each cell, and the 2- to 3-celled sweeping hairs. On
the other hand, the thistle-like Gundelia, placed
by Bremer (1994) in Arctotideae-Gorteriinae, has
been shown to be most closely related to Cichorieae
(Karis et al. 2001), and it is hence not treated here.
Platycarpha, another thistle-like plant from South
Africa, is tentatively accepted in the tribe, since it
shares the scaly pappus, similar pericarp development (Reese 1989) as well as the arctotoid style
with Arctotideae. However, molecular data suggest
that Platycarpha may be closer to the VernonieaeLiabeae clade (Funk et al. 2004). Styles with a ring of
sweeping hairs below the bifurcation are found also
in Cynareae. This previously was used as evidence
for close relationship between the tribes (Lessing
1832; Bremer 1987). This view was strengthened by
the common thistleoid habit. Heterolepis has arctotoid disc floret styles, and shares features with
both the subtribes recognised here. At this point,
it is not evident which features should be regarded
as synapomorphies, and the possibility that Heterolepis belongs elsewhere in Cichorioideae s. str.
must also be considered (Funk et al. 2004). The
3-lobed ray florets (as in subtribe Arctotidinae)
possibly are a plesiomorphy, and this is also the
case with the rather long, but not markedly soft
anther apical appendages. Heterolepis has a long
filament collar, unlike Arctotidinae, but the cells
are inconspicuously reinforced, as in Arctotidinae.
The endothecial cells often have no lateral pattern
on the walls (on anticlinal walls) or some are polarised (as in Gorteriinae), and the involucral bracts
become partly connate (as in Gorteriinae), but are
obtuse and apically scarious (as in Arctotidinae).
As circumscribed here, Arctotideae do not possess
a single morphological synapomorphy, but are diagnosed by a combination of morphological fea-
201
tures which are not present in all included genera,
as well as partly by molecular data (Karis et al. 2001;
Funk et al. 2004). Members of Arctotideae can be
recognised as often radiate species with arctotoid
disc floret styles, and often with a scaly pappus.
Eremothamnus and Hoplophyllum differ by their
bristly pappus and vernonioid disc florets styles,
whereas both Platycarpha and Hoplophyllum have
discoid capitula, although those of Platycarpha are
arranged in secondary heads. Eremothamnus and
subtribe Arctotidinae share the soft anther apical
appendages, but the appendages are longer and
less soft in the former. The pollen morphology of
Arctotideae has been investigated by several authors (Stix 1960; Leins 1970; Leins and Thyret 1971;
Skvarla et al. 1977; Robinson 1994). Interpretations,
however, differ (Leins and Thyret 1971; Skvarla
et al. 1977), and the significance of pollen characters for infratribal and intratribal relationships
are difficult to ascertain (Bremer 1987). For example, the pollen wall is often described as caveate,
i.e. the foot layer is separated by cavities (Skvarla
et al. 1977), but the cavities in Arctotideae pollen
walls may not be homologous with the strictly interapertural cavities prevailing in subfamily Asteroideae (Skvarla et al. 1977; Bremer 1987). It can
be noted, however, that at least the conspicuous
spine channels occur in Hoplophyllum (Robinson
1994), Eremothamnus (Leins 1970) as well as in all
other Arctotideae taxa investigated by Skvarla et al.
(1977).
The basic chromosome number varies to some
extent among genera, and at least Haplocarpha and
Gazania contain polyploid species.
All genera occur in Africa, especially in South
Africa, except Cymbonotus which is native to South
Australia. Subtribal division is not entirely clear,
although most genera can quite readily be placed
in either of the commonly accepted and putatively
monophyletic subtribes Arctotidinae and Gorteriinae. With their slightly aberrant morphology, Eremothamnus, Hoplophyllum, Platycarpha and Heterolepis are left as Arctotideae incertae sedis. All
these taxa need further study in order to elucidate their exact relationships, despite the elaborate molecular study by Funk et al. (2004). Various morphological characters diagnose the two
subtribes Arctotidinae and Gorteriinae, and some
of these can be assumed to be synapomorphies.
Some species of Arctotideae are used as ornamentals. Most common are hybrids originating from
Gazania krebsiana, G. linearis and G. rigens, which
are cultivated all over the world.
202
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Key to the Subtribes
and Unplaced Genera
1. Pappus of barbellate bristles, or of barbellatesubplumose bristle-like, subulate scales
2
– Pappus of scales, never bristle-like, or occasionally
pappus absent
4
2. Capitula discoid
394. Hoplophyllum
– Capitula radiate
3
3. Capitula sessile; involucral bracts spine-tipped; pappus of scabrid-barbellate bristles 392. Eremothamnus
– Capitula pedunculate; involucral bracts unarmed;
pappus of barbellate-subplumose bristle-like,
subulate scales
393. Heterolepis
4. Capitula discoid, in secondary heads ± sessile in a leaf
rosette; florets purple
395. Platycarpha
– Capitula generally radiate, not in secondary heads; if
discoid, then florets yellow
5
5. Involucral bracts free, often obtuse, innermost often
apically scarious; ray florets 3-lobed, mostly 4-veined,
female (rarely neuter), disc floret lobes generally < 4
times as long as wide, without sclerified margins
1. Subtribe Arctotidinae (p. 202)
– Involucral bracts more conspicuously connate, acute,
sometimes spine-tipped, ray florets 4(–2)-lobed, sterile or neuter, disc floret lobes generally 4–10 times as
long as wide, mostly with sclerified margins
2. Subtribe Gorteriinae (p. 204)
Arctotideae Incertae Sedis
lobe ventrally in the sinus of the tube, 3-lobed. Anthers caudate, appendage somewhat soft, endothecium unreinforced laterally or polarised. Style slender, with rather short style branches, apical portion not or only slightly thickened and with short
hairs mainly at the border to the shaft. Cypselae densely sericeous. Pappus of stout, subulate,
bristle-like, marginally barbellate or subplumose
scales in 2 rows. 2n = 20. Three species, South
Africa.
394. Hoplophyllum DC.
Hoplophyllum DC., Prodr. 5: 73 (1836); Karis, Taxon 41:
193–198 (1992), rev.; Karis et al., Taxon 50: 105–114 (2001),
phylog.; Funk et al., Taxon 53: 637–655 (2004), phylog.
Shrubs with linear, entire, terete or flattened, sparsely dentate-spiny, spine-tipped, longitudinally striate, grooved hard leaves. Capitula sessile, discoid.
Involucral bracts chartaceous, spine-tipped. Anthers caudate, endothecial cells with transversal
bands in lower half of cells. Styles vernonioid, with
short 2–3-celled hairs. Cypselae densely sericeous.
Pappus of stout, scabrid-barbellate bristles. 2n =
18. Two species, South Africa.
392. Eremothamnus O. Hoffm.
395. Platycarpha Less.
Eremothamnus O. Hoffm., Bot. Jahrb. 10: 278 (1889); Karis,
Taxon 41: 193–198 (1992), rev.; Karis et al., Taxon 50: 105–
114 (2001), phylog.; Funk et al., Taxon 53: 637–655 (2004),
phylog.
Platycarpha Less., Linnaea 6: 688 (1831); Funk et al., Taxon
53: 637–655 (2004), phylog.
Small woolly shrub with obovate, entire or apically 3–5-dentate-spiny, spine-tipped, fleshy leaves.
Capitula sessile, radiate. Involucral bracts chartaceous, spine-tipped. Ray florets female, fertile,
with staminodes, 3-lobed. Anthers caudate, appendage ± soft, endothecial cells with transverse
bands in lower half of cells. Style vernonioid, with
short 2–3-celled hairs. Cypselae densely sericeous.
Pappus of stout, scabrid-barbellate bristles. One
species, E. marlothianus O. Hoffm., Namibia.
393. Heterolepis Cass.
Heterolepis Cass., Bull. Soc. Philom.: 26 (1820), nom. cons.
Shrublets, sometimes scented, with linear-oblong,
ericoid, entire or remotely dentate, unarmed leaves.
Capitula pedunculate, radiate. Involucral bracts in
2–3 rows, somewhat connate at base, outer lanceolate, foliaceous and acute, inner rounded-truncate,
apically scarious and laciniate. Ray florets female,
fertile, generally with staminodes, with a filiform
Perennial herbs with rosulate, entire, obovateoblanceolate or dentate to pinnatisect, unarmed
or spinulose leaves. Capitula crowded in a large,
sessile secondary head in the centre of the leaf
rosette; each capitulum few-flowered, discoid.
Involucral bracts herbaceous to chartaceous,
innermost resembling paleae. Florets purple.
Anthers often apically emarginate, endothecium
polarised or sometimes not reinforced laterally.
Style slender, with rather short style branches,
apical portion not or only slightly thickened and
with short hairs mainly at the border to the shaft.
Cypselae oblong, prismatic, transversely rugose,
with some apically curled or hooked twin hairs
at the base. Pappus of scales in 1–2 rows. Three
species, South Africa.
X.1. Subtribe Arctotidinae (Cass.) Dumort.
(1819).
Leaves unarmed. Scapes woolly, sometimes also
with reddish-septated multiseptate hairs. Capitula
radiate. Involucral bracts free, outer with foliaceous
Compositae
tips, inner obtuse with scarious tips, ray florets 3lobed, disc florets generally shallowly lobed, styles
often markedly thickened apically. Anthers distinctly ecaudate, apical appendages usually obtuse,
rounded, soft, ± wrinkled, endothecium radial,
collar generally inconspicuous, cells not reinforced.
The supposed monophyly of subtribe Arctotidinae
is supported by the short, soft and wrinkled anther
apical appendages, the rather inconspicuous filament collar, the radial endothecial tissue, as well as
by the apically conspicuously thickened disc floret
styles. The disc corollas are usually more shallowly
lobed than in the taxa outside Arctotidinae, and the
lobes are generally slightly less than four times as
long as wide. This might be an additional synapomorphy for the subtribe but requires investigation
of a larger sample of material. The largest genus
Arctotis is in need of revision, but it should also be
studied in relation to the other genera of the subtribe, where especially the limit against Arctotheca,
Cymbonotus, Dymondia and Haplocarpha must be
considered (McKenzie et al. 2005).
203
brous. Pappus of small scales or absent. 2n = 18.
Four species, South Africa, Mozambique.
397. Arctotis L.
Fig. 50
Arctotis L., Sp. Pl. 922 (1753); Lewin, Feddes Repert. Beih.
11: 1–95 (1922), rev.
Venidium Less. (1831).
Annual or perennial herbs, shrublets or shrubs
with entire or lobed to pinnatisect leaves. Capitula
pedunculate. Ray florets yellow, orange, cream,
white, pink, purple, violet or blue; inner disc
florets sometimes female-sterile, lobes sometimes
apically with an abaxial, thick, deltoid, usually
vividly coloured, almost blackish projection.
Cypselae ventrally smooth or rugose and without
ribs, dorsally with 3–5 strong ribs or wings
sometimes forming 2 dorsal furrows or concavities
at maturity, sericeous or pilose or glabrous, often
with a basal tuft of long hairs. Pappus of 1–2 rows
of large or small scales, sometimes absent. 2n = 18.
Around 60 species, South Africa, Namibia, Angola.
Key to the Genera
1.
–
2.
–
3.
Ray florets neuter
396. Arctotheca
Ray florets female
2
Mat-forming; capitula sessile
399. Dymondia
Not mat-forming; capitula pedunculate or scapose 3
Capitula c. 10 mm wide; disc floret lobe venation discontinuous; South Australia
398. Cymbonotus
– Capitula >> 10 mm wide; disc floret lobe venation
continuous; Africa
4
4. Leaves not conspicuously dentate, cypselae strongly
ribbed, often with concavities between three dorsal
ribs
397. Arctotis
– Leaves conspicuously dentate with broad teeth, cypselae thinly ribbed, without concavities
400. Haplocarpha
Genera of Arctotidinae
396. Arctotheca Vaill.
Arctotheca Vaill., Königl. Akad. Wissensch. Paris Phys. Abh.
5: 604 (1754); Lewin, Feddes Repert. Beih. 11: 1–95 (1922),
rev.
Perennial or sometimes annual herbs with rosulate,
entire or mostly lobed to pinnatifid leaves. Capitula pedunculate. Ray florets neuter, yellow or pale
yellow, sometimes darker towards the base of the
lamina; disc floret lobes apically often with an abaxial, thick, deltoid, usually vividly coloured, almost
blackish projection. Cypselae rather thinly ribbed
mainly dorsally, densely to sparsely villous or gla-
Fig. 50. Compositae-Arctotideae. Arctotis stoechadifolia.
A Habit. B Involucral bract. C Cypsela. (Walsh and Entwisle
1999)
204
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
398. Cymbonotus Cass.
Cymbonotus Cass., Dict. Sci. Nat. 35: 397 (1825).
Annual or perennial herbs with rosulate, dentate
to lobed or pinnatifid leaves. Capitula pedunculate
or scapose, up to c. 10 mm wide. Ray florets with
staminodes; disc florets with discontinuous lobe
veins. Cypselae with 2 lateral and 2 dorsal ribs,
transversely rugose ventrally and between the ribs,
glabrous. Pappus absent. Three species, South Australia.
399. Dymondia Compton
Dymondia Compton, J. S. African Bot. 19: 110 (1953).
Mat-forming perennial with rosulate, linearoblanceolate, entire or sinuate-dentate leaves.
Capitula sessile. Involucral bracts without scarious appendages. Ray florets sometimes with
staminodes; disc floret lobes apically often irregularly 3-lobed. Style branches apically shallowly
3-lobed. Cypselae glabrous. Pappus of 2 rows of
ovate-acuminate, laciniate scales. One species,
D. margaretae Compton, South Africa.
appendages usually acute and firm, endothecium
usually without lateral wall thickenings or with
some polarised cells, collar conspicuous, with
reinforced cells; styles slightly thickened above
or not. Subtribe Gorteriinae is most certainly
a monophyletic group, as evidenced by the often
connate involucral bracts, the deeply alveolate
receptacles, the often 4-lobed ray florets, and the
sclerified disc floret lobe margins. A comparison
of the length/width ratio of the disc corolla lobes
reveals that the corollas are more deeply lobed
than those of Arctotidinae, but this condition
must be considered plesiomorphic. The subtribe
comprises two sister groups, a Berkheya clade
which is supported by molecular data (Funk et al.
2004) and by spiny leaves and mamillate style hairs
(Karis 2006). In this clade, Berkheya is paraphyletic
in relation to Cullumia, Cuspidia, Didelta and
Heterorhachis. The sister group includes Gazania,
Gorteria and Hirpicium, and is supported by
molecular data as well as by longitudinally striate
leaf/scape hairs, fringed apical anther appendages,
"Gazania-type" pollen, and subulate-ensiform
style hairs (Karis 2006). Hirpicium is paraphyletic
in relation to Gazania and Gorteria.
400. Haplocarpha Less.
Haplocarpha Less., Linnaea 6: 90 (1831); Lewin, Feddes
Repert. Beih. 11: 1–95 (1922), rev.
Landtia Less. (1832).
Tufted or mat-forming perennials with ± rosulate, entire or lobed to pinnatifid, pinnately or
sometimes palmately veined leaves; leaf teeth
often broad. Capitula scapose. Ray florets dorsally
often reddish or greenish; inner disc florets
sometimes female-sterile. Cypselae rather thinly
ribbed, sometimes transversely rugose, sericeous
or glabrous, although mostly with a tuft of long
hairs at the base. Pappus of 2 to many rows of
subulate-linear, hyaline scales or absent. 2n = 10,
12, 18. Eight species, southern and eastern Africa
north to Ethiopia.
X.2. Subtribe Gorteriinae Benth. (1873).
Leaves spiny, or unarmed and then often with
longitudinally striate hairs; receptacle deeply alveolate, sometimes becoming lignified at anthesis;
involucral bracts connate at least at the base (free
in Didelta), spiny or mucronate, often acute; ray
florets sterile or neuter, 4(–2)-lobed, disc corolla
lobes generally sclerified marginally along the
veins; anthers caudate or ecaudate, apical anther
Key to the Genera
1. Outer part of receptacle with thick-walled cavities,
becoming lignified, inner part membranous
2
– Receptacle uniform, becoming entirely lignified or not
3
2. Plants unarmed; leaves entire; outer involucral bracts
3–5, large, broadly ovate to ovate, entire; receptacle
breaking into parts each adnate to one outer bract at
maturity
404. Didelta
– Plants armed; leaves pinnatisect; outer involucral
bracts lanceolate to narrowly triangular; receptacle
remaining unbroken at maturity 407. Heterorhachis
3. Leaves and/or involucral bracts spiny, involucral bracts
connate at the base; apical anther appendages with
entire margins
4
– Leaves and involucral bracts unarmed, but sometimes
hispid and/or mucronate; involucral bracts connate
into a lignified cup; apical anther appendages with ±
fringed margins
6
4. Annual herb
403. Cuspidia
– Perennial herbs, shrublets or shrubs
5
5. Pappus absent or rarely present but very inconspicuous
402. Cullumia
– Pappus present
401. Berkheya
6. Seeds germinating from within old capitula; cypselae
without swollen cells; pappus of minute scales hidden
among the twin hairs
406. Gorteria
– Seeds not germinating within old capitula; cypselae
with longitudinal rows of swollen cells; pappus conspicuous
7
Compositae
7. Latex not conspicuous; pappus scales in two distinctly
unequal rows, outer much longer than inner, overlapping, or sometimes in a single row
408. Hirpicium
– Latex often copious; pappus scales in two subequal
rows, outer row not overlapping
405. Gazania
205
a crateriform cup, becoming lignified and enclosing the cypselae after anthesis. Cypselae sericeous.
Pappus of ciliate to barbellate, subulate scales. One
species, C. cernua (L. f.) B.L. Burtt, South Africa.
Genera of Gorteriinae
401. Berkheya Ehrh.
Fig. 51
Berkheya Ehrh., Beitr. 3: 137 (1788), nom. cons.; Roessler,
Mitt. Bot. Staatssamml. München 3: 71–500 (1959), 11: 91–
99 (1973), rev.
Carlinoides Vaill. (1754), nom. rej. prop.
Perennial herbs or shrubs with entire or mostly
dentate to pinnatisect, spiny or spinulose leaves.
Capitula generally radiate, solitary or corymbose,
rarely axillary or umbellate. Involucral bracts connate at base, entire or lobed, spiny. Outer periclinal wall of endothecial cells sometimes with large,
irregularly rounded pores apically, or periclinal
wall rarely ± displaced towards the connectivefacing side of the cell. Cypselae more or less distinctly ribbed, densely to sparsely sericeous or glabrous, entirely enclosed in the receptacle. Pappus
of subequal or unequal scales generally in 1–2 rows.
2n = 14, 16. Around 80 species, southern and tropical Africa north to Nigeria and Ethiopia.
402. Cullumia R. Br.
Cullumia R. Br., Ait. Hort. Kew ed. II. 5: 137 (1813); Roessler,
Mitt. Bot. Staatssamml. München 3: 71–500 (1959), rev.
Shrublets or shrubs with densely arranged,
rigid, entire and often rather small, marginally
conspicuously ciliate to spinulose leaves, often
with a conspicuous sclerified margin, sometimes
decurrent. Capitula sessile, solitary, terminal.
Involucral bracts connate at base, outermost foliaceous, inner serrulate, with herbaceous patches
at the base. Endothecial cells with thickening
on outer periclinal wall, but often ± displaced
towards the connective-facing side of the cell.
Cypselae oblong-ellipsoid, smooth, glabrous or
very rarely minutely pilose, entirely enclosed
in the receptacle. Pappus absent or rarely very
inconspicuous. Fifteen species, South Africa.
403. Cuspidia Gaertn.
Cuspidia Gaertn., Fruct. 2: 454 (1791); Roessler, Mitt. Bot.
Staatssamml. München 3: 71–500 (1959), rev.
Annual herb with dentate to lobed, spiny leaves. Capitula mostly sessile. Involucral bracts connate into
Fig. 51.
Compositae-Arctotideae. Berkheya horrida.
A Habit. B Ray floret with receptacular bract. C Disc floret
and cypsela. (Muschler 1911)
206
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
404. Didelta L’Hér.
Didelta L’Hér., Stirp. Nov.: 55. (1785), nom. cons.; Roessler,
Mitt. Bot. Staatssamml. München 3: 71–500 (1959), rev.
Shrublets, shrubs, or rarely small trees with
entire, often opposite, mostly unarmed leaves.
Capitula sessile or pedunculate. Involucral bracts
dimorphic, free, outer row of 3–5 large, broadly
ovate-triangular bracts, inner row of several
ovate-lanceolate bracts. Receptacle at maturity
breaking into 3–5 outer parts and 1 central part
enclosing the cypselae, outer parts adnate to
1 of the outer involucral bracts and becoming
thickened and lignified, central part membranous.
Cypselae sparsely sericeous or glabrous. Pappus of
lanceolate-subulate, serrulate to ciliate scales. Two
species, South Africa, Namibia.
405. Gazania Gaertn.
Gazania Gaertn., Fruct. 2: 451 (1791), nom. cons.; Roessler,
Mitt. Bot. Staatssamml. München 3: 71–500 (1959), 11:
91–99 (1973), rev.
Perennial or rarely annual herbs or subshrubs
with often rosulate, unarmed, entire to pinnatisect
and then often long-petiolate leaves. Leaf hairs
small to large, longitudinally striate, often only
marginal. Capitula pedunculate or scapose. Scape
± glabrous, rarely with small, longitudinally striate
hairs or floccose. Involucral bracts connate into
a campanulate-cylindrical cup. Ray florets yellow,
orange or reddish with a blackish basal spot and
a blackish dorsal stripe. Apical anther appendages
± fringed marginally, endothecial cells with inner
anticlinal wall thickened, with no thickenings on
outer periclinal wall. Cypselae with longitudinal
rows of swollen cells, sericeous, not enclosed in
the receptacle. Pappus of linear-subulate, subequal
scales in 2 rows. 2n = 10, 12, 14. Seventeen species,
South Africa, Namibia, one species extending into
Tanzania and another into Mozambique.
406. Gorteria L.
Gorteria L., Syst. ed. 10: 1229 (1759); Roessler, Mitt. Bot.
Staatssamml. München 3: 71–500 (1959), 11: 91–99 (1973),
rev.
Erect or sprawling annual herbs germinating
from within old capitula, with unarmed, entire
or dentate to pinnatifid leaves. Stems and leaves
with coarse, rigid, longitudinally striate hairs with
a multicellular base. Capitula solitary or axillary,
pedunculate. Involucral bracts connate into a more
or less urceolate cup, becoming lignified and
enclosing the cypselae after anthesis. Ray florets
yellow or reddish, sometimes with a blackish basal
spot (bulging in G. diffusa Thunb.) and a blackish
dorsal stripe; most disc florets female-sterile, with
large or smaller spine-like hairs on the lobes.
Apical anther appendages ± fringed marginally,
endothecial cells with inner anticlinal wall thickened, with no thickenings on outer periclinal wall.
Cypselae apically sericeous. Pappus of minute
scales, hidden among the cypsela hairs. Three
species, South Africa, Namibia.
407. Heterorhachis Sch. Bip. ex Walp.
Heterorhachis Sch. Bip. ex Walp., Rep. 6: 278 (1847);
Roessler, Mitt. Bot. Staatssamml. München 3: 71–500
(1959), rev.
Shrub with pinnatisect, spiny leaves. Capitula
in terminal racemose clusters. Involucral bracts
somewhat connate at the base, in about 3 rows,
outer and median foliaceous, median row much
longer than outer and inner, ± spreading, innermost ciliate-margined, scarious. Receptacle with
the outer parts becoming thickened and lignified
at maturity, central part membranous. Outer
periclinal wall of endothecial cells sometimes
with large, irregularly rounded pores apically.
Cypselae obconical, sparsely sericeous or glabrous,
entirely enclosed in the receptacle. Pappus of
lanceolate-subulate, serrulate to ciliate scales. One
species, H. aculeata (Burm. f.) Roessler, South
Africa.
408. Hirpicium Cass.
Hirpicium Cass., Bull. Soc. Philom.: 27 (1820); Roessler,
Mitt. Bot. Staatssamml. München 3: 71–500 (1959), 11:
91–99 (1973), rev.
Berkheyopsis O. Hoffm. (1892).
Annual or perennial herbs, subshrubs or shrubs
with entire or dentate to pinnatisect, sometimes
mucronate leaves. Stems and leaves with coarse,
rigid, longitudinally striate hairs with or without
a multicellular base. Involucral bracts connate into
a cupuliform to broadly campanulate-obconical
cup, generally foliaceous, often ciliate-margined.
Disc floret lobes sometimes with large or small
spine-like hairs. Apical anther appendages ±
fringed marginally, endothecial cells with inner
anticlinal wall thickened, with no thickenings on
outer periclinal wall. Cypselae obconical, more
or less ribbed, with longitudinal rows of swollen
Compositae
cells, sericeous, partly enclosed in the receptacle.
Pappus generally of 2 distinct rows of 10+10
broad scales, outer much longer than inner; inner
row sometimes absent. 2n = 10. Twelve species,
southern and eastern Africa north to Ethiopia.
XI. Tribe Corymbieae Panero &
V.A. Funk (2002).
B. Nordenstam
Scapose perennial herbs with a stout, silky-hairy
rhizome. Leaves alternate, mainly rosulate, sessile,
entire, linear-lanceolate to narrowly ellipticoblong, flat or conduplicate, parallel-veined, ±
coriaceous, sometimes cartilaginous or herbaceous, acute to acuminate, narrowed to base,
glabrous or pubescent, sometimes glandular;
cauline leaves gradually smaller. Capitula pedunculate or rarely sessile, several to many in corymbs
to panicles terminating a stout erect bracteate
scape (‘corymbophore’), discoid, single-flowered.
Involucre cylindrical, calyculate, involucral bracts
2, enclosing the floret, narrowly oblong to lanceolate, flat or keeled, 3-nerved, glabrous or glandular
(viscid when fresh), sometimes scabrid, often with
a purplish tinge, apically 2–3-fid or fimbriate;
outer (calyculus) bracts 2 or 3, short. Receptacle
flat, nude. Floret hermaphrodite; corolla 5-lobed,
pink to purplish or white; corolla lobes linear to
oblong, spreading, apically cucullate and dorsally
papillate. Stamens 5; anthers tetrasporangiate with
blackish thecae, shortly sagittate; apical appendage
reduced. Pollen grains caveate. Style bifurcate with
linear branches; style branches and uppermost
part of shaft hairy. Cypselae narrowly oblong,
somewhat compressed, densely pubescent. Pappus
of basally connate short scales and/or discrete fine
bristles. x = 8.
207
of the traditional tribes in the family. Usually
tribe Vernonieae has been suggested, but the
genus is aberrant there on account of its strange
vegetative and floral morphology, caveate pollen
(Bolick 1978) and different phytochemistry. For
example, it lacks sesquiterpene lactones typical
of Vernonieae but contains diterpenes, not occurring in that tribe (Zdero and Bohlmann 1988;
Bohlmann and Jakupovic 1990). Bremer (1994)
accepted the genus in subfam. Cichorioideae
among genera unassigned to tribe. Molecular
evidence suggests a position of Corymbium basal
to Senecioneae in subfam. Asteroideae or as
sister to the subfamily, which led to the proposal
of subfam. Corymbioideae Panero & V.A. Funk
Only one genus:
409. Corymbium L.
Fig. 52
Corymbium L., Sp. Pl.: 928 (1753) & Gen. Pl. ed. 5: 400 (1754);
Markötter, Bot. Jahrb. 70: 354–372 (1939), rev.; Weitz, S.
African J. Bot. 55: 598–629 (1989), rev.; Weitz, Mitt. Inst.
Allg. Bot. Hamburg 23b: 631–642 (1990), rev.
Characters of the tribe. Nine species, Cape region
of South Africa.
The singular South African genus Corymbium
L. has been notoriously difficult to place in any
Fig. 52. Compositae-Corymbieae. Corymbium villosum.
A Habit. B Single-flowered capitulum. C Involucre of two
bracts. D Ovary and style. (Weitz 1989)
208
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
(2002). Many features, such as pollination biology
and cytology, are still practically unknown in this
unique and isolated group within the family. The
flowers produce copious nectar and are frequently
visited by bees, beetles, wasps and ants (Weitz
1989). A single report on chromosome number
has been published, viz. 2n = 16 in C. congestum
E. Mey. ex DC. (Weitz 1989).
XII. Tribe Senecioneae Cass. (1819).
B. Nordenstam
Herbs, shrubs, lianas, epiphytes, treelets or
trees. Leaves alternate, cauline or rosulate, more
rarely opposite, sessile or petiolate, entire or
variously lobed or dissected. Capitula heterogamous or homogamous, radiate, disciform or
discoid, often yellow-flowered but sometimes
white, orange, pink, purple, red or rarely blue,
solitary or corymbose, paniculate or thyrsoid
in terminal or lateral synflorescences. Involucre
commonly uniseriate, sometimes biseriate or
rarely pluriseriate, with or without a calyculus of
smaller bracts. Involucral bracts free or connate
to various degrees. Receptacle flat or convex to
conical, naked or fimbrillate to denticulate with
scale-like projections, glabrous or hairy, solid or
fistulose. Ray florets female, sometimes sterile or
absent, corolla radiate or tubular-filiform; style bifid or simple, fertile or sterile. Disc florets bisexual,
perfect or functionally male, rarely female; corolla
tubular to funnel-shaped or with a campanulate
limb, 4- or 5-lobed. Anthers 4 or 5, tetrasporangiate
or rarely bisporangiate (Gynura; Pullaiah 1983)
with an apical flat appendage, basally obtuse,
sagittate or caudate. Endothecial tissue radial
or polarized, rarely transitional. Filament collar
straight and uniform or basally dilated with larger
cells (balusterform). Pollen caveate, in most genera
Senecioid (columellae solid) but in some genera
Helianthoid (columellae with internal foramina;
Skvarla et al. 1977). Style bifid or simple, fertile
or sterile, apically truncate or convex, rounded
or conical, sometimes with elongate appendage,
papillate or hirsute or only apically with short
sweeping hairs or minutely papillate, sometimes
with a distinct central tuft or pencil of hairs or
fused papillae (Fig. 53); stigmatic areas continuous
or separated. Cypselae terete, elliptic-oblong or
obovoid, sometimes flattened, winged or angled,
often ribbed, glabrous or variously pubescent,
sometimes heteromorphic; carpopodium ringlike, distinct (up to 12 cell layers or more) or
indistinct. Ovary wall crystals often present, often
prismatic, plate-like, elongate or isodiametric,
sometimes drusiform (Nordenstam 1978; Jeffrey
1986). Pappus of few to many bristles or a single
scale or absent, persistent or caducous, uni- to
multiseriate, fine and slender to coarse and rigid,
barbellate to subplumose, white or sometimes
straw-, brick- or red- to purple-coloured.
Senecioneae are one of the largest tribes of
Compositae, with 150 genera presently recognized
and about 3,500 species, and with a worldwide distribution.
The limits of Senecioneae have become more
clear since the 1970s after transfer of several
genera to Heliantheae and Helenieae, and the
recognition of Liabeae and Corymbieae as distinct
tribes (Rydberg 1927; Robinson and Brettell 1973a;
Nordenstam 1977; Panero and Funk 2002). The
tribe is now reasonably well defined, although
the relationships and tribal position of a few
Fig. 53. Compositae-Senecioneae. Style branches from disc
florets. A, B Dendrosenecio keniensis. C, D Lordhowea insularis. E Cacaliopsis nardosmia. F Odontocline tercentenariae. G Othonna brandbergensis. H Jacmaia incana. I Kleinia
longiflora. J Senecio eligulatus. K Io ambondrombeensis.
L Psednotrichia xyridopsis. (Drawings by B. Nordenstam)
Compositae
genera such as Abrotanella and Doronicum are still
problematic or unresolved. A recent addition to the
tribe is Haastia, which was unassigned as to tribe
(Bremer 1994) but fits well in the Brachyglottis
group (Wagstaff and Breitwieser 2002, 2004). The
number of segregates from the core genus Senecio
has increased dramatically in the last decade.
Senecio itself (with over 3,000 binomials) will
eventually be defined as a monophyletic but still
very large genus by continued removal of discordant elements including several sections (e.g. sect.
Jacobaea). On the tribal level, the phylogeny and
relationships within subfamily Asteroideae are not
sufficiently known, and it is difficult to discern the
closest affinities of Senecioneae among the tribes
Astereae, Anthemideae, Calenduleae, Inuleae and
Gnaphalieae. Phytochemically, the tribe is rather
well characterized by the presence of pyrrolizidine
alkaloids in many, but not all genera, and groups
of sesquiterpene lactones known as eremophilanes
and furanoeremophilanes, and by the absence
of polyacetylenes in most genera (Robins 1977).
Doronicum differs from the rest of the tribe by the
presence of acetylenes and in its sesquiterpenoid
constituents (Bohlmann et al. 1973).
A number of subtribes have been suggested but the attempts made so far have not
been all-inclusive. Subtribes proposed include
Blennospermatinae,
Abrotanellinae,
Tussilagininae, Tephroseridinae, Senecioninae and
Adenostylinae (under Eupatorieae). Usually only
two subtribes are recognized, viz. Blennospermatinae and Senecioninae, and within the latter
a ‘senecioid’ and a ‘tussilaginoid’ (initially called
‘cacalioid’) complex have been loosely characterized (‘tussilaginoid’ is a better term than ‘cacalioid’
because of the ambiguity of the rejected name
Cacalia L.; cf. Jeffrey 1992; Barkley 1999). No
formal subtribal classification is proposed here,
pending the outcome of an ongoing molecular
phylogenetic study of the entire tribe (Pelser,
Kadereit, Nordenstam, Breitwieser, Wagstaff,
Watson, in progress).
Key to the Genera
1. Disc-florets 4-lobed
2
– Disc-florets 5-lobed
10
2. Dwarf herbs, sometimes tufted. Leaves sessile, linear,
up to 5 cm long. Capitula small (3–5 mm broad and
long), few-flowered
3
– Herbs, subshrubs or shrubs. Leaves larger, 5–30 cm
long, sessile or often petiolate. Capitula larger (7–
25 mm broad), many-flowered
4
209
3. Capitula disciform with tubular marginal florets
410. Abrotanella
– Capitula radiate; disc floret style undivided; pappus
absent
413. Ischnea p.p.
4. Florets yellow
5
– Florets purple or whitish
8
5. Leaves sessile (Australia, Oceania) 499. Senecio p.p.
– Leaves petiolate (Eurasia)
6
6. Rhizome slender. Capitula solitary or few. Anther base
obtuse. Style branches dorsally glabrous
464. Dolichorrhiza
– Rhizome thick. Capitula few to many. Anther base
caudate or at least sagittate. Style branches dorsally
papillate
7
7. Leaves entire, cordate or deltoid to ovate
557. Caucasalia
– Leaves lobed or dissected
559. Iranecio
8. Leaves ovate or cordate
9
– Leaves lanceolate, cuneate
556. Pojarkovia
9. Perennial herbs with rhizome. Calyculus present. Style
branches subulate
555. Adenostyles
– Shrubs or lianas. Calyculus absent. Style branches linear
543. Faujasiopsis p.p.
10. Plant forming dense cushions; branches and leaves
concealed by dense tomentum. Capitula sessile, disciform (New Zealand)
461. Haastia p.p.
– Habit various but not as above
11
11. Involucral bracts pluriseriate to imbricate
12
– Involucral bracts 1–2-seriate
17
12. Capitula radiate, yellow-flowered
13
– Capitula discoid or disciform
14
13. Leaves cauline, entire, woolly beneath (S Africa)
465. Capelio p.p.
– Basal leaves rosulate, dissected, large, glabrous (Peru)
513. Caxamarca
14. Plants glabrous or somewhat woolly (mostly on young
parts). Capitula few-flowered (3–17 florets)
15
– Plants densely woolly. Capitula many-flowered, discoid
16
15. Leaves sessile, linear or filiform to oblanceolate or reduced to scales. Capitula discoid, yellow-flowered (SW
USA, Mexico)
451. Lepidospartum
– Leaves petiolate. Capitula disciform with tubular
marginal florets, purplish or white to cream-coloured
(Indonesia, New Guinea, Australia)
488. Arrhenechthites
16. Shrub. Capitula several, medium-sized, corymbose
(Cuba)
471. Shafera
– Herbs. Capitula large, solitary or few, nodding (South
America)
521. Culcitium
17. Plants dioecious; capitula radiate or discoid
18
– Plants monoecious or polygamous
23
18. Perennial herbs with rosulate leaves from a rhizome
19
– Shrubs or shrublets, or trees, rarely perennial herbs
with cauline leaves and a woody caudex
20
19. Rhizome thin. Leaves present at flowering time. Capitula radiate
417. Endocellion
– Rhizome thick. Leaves developing postflorally. Capitula discoid or disciform
418. Petasites
20. Leaves crowded at branch ends. Styles undivided in
male capitula (Juan Fernandez)
498. Robinsonia
– Leaves evenly distributed. Styles bifurcate
21
210
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
21. Perennial herbs or shrublets with mostly tomentose
leaves; leaf blades up to 5 cm long. Capitula discoid,
yellow-flowered
415. Chersodoma
– Shrubs with larger glabrous leaves
22
22. Leaves entire. Capitula discoid, pink- to purplishflowered (Bonin Islands)
422. Dendrocacalia
– Leaves serrate. Capitula radiate, yellow-flowered (New
Guinea)
467. Brachionostylum
23. Indumentum of stellate, T-shaped or pseudo-stellate
hairs
24
– Trichomes, if present, not stellate or branched
26
24. Ray florets long, narrowly linear, recurved, reddish
519. Dresslerothamnus
– Ray florets oblong, patent, white or yellow
25
25. Ray florets white. Trichomes peltate and stellate. Style
branches obtuse or subtruncate
456. Aequatorium
– Ray florets yellow. Trichomes substellate and irregularly branched. Style branches acute to acuminate
455. Nordenstamia
26. Styles of disc florets undivided (sterile)
27
– Styles of disc florets bilobed (fertile or sterile)
32
27. Aquatic floating herb. Rays white (South Africa)
535. Cadiscus
– Terrestrial plants. Rays yellow, pink or purple (seldom
white in Othonna spp.)
28
28. Small herbs with solitary pedunculate capitula. Pappus absent
29
– Herbs, subshrubs or shrubs. Pappus present, at least
in ray florets
30
29. Annual. Leaves alternate, mostly pinnatifid. Cypselae
papillate
411. Blennosperma
– Perennial herbs. Leaves rosulate, entire or serrate.
Cypselae glabrous
413. Ischnea p.p.
30. Herb with postflorally developing, large basal leaves.
Ray florets in many series, with linear lamina
419. Tussilago
– Leaves and flowers at the same time. Ray florets narrowly oblong to elliptic-oblong
31
31. Perennial herbs, subshrubs or shrubs 531. Othonna
– Annual herbs
532. Gymnodiscus
32. Filament collar straight, uniform. Endothecial tissue
usually polarized. (‘Tussilaginoid’ genera, usually
with uniseriate ecalyculate involucre, florets often
non-yellow, stigmatic areas of disc floret styles mostly
confluent, disc floret corollas frequently deeply
5-lobed)
33
– Filament collar ‘balusterform’ or subcylindric with
enlarged basal or marginal cells. Endothecial tissue
radial (except in Graphistylis). (‘Senecioid’ genera, involucre 1–2-seriate and mostly calyculate, florets often
yellow, stigmatic areas of disc floret styles often separated, disc floret corolla lobes mostly triangular-ovate)
96
33. Small annual herb (NW USA)
512. Crocidium
– Perennials
34
34. Leaves parallel-veined with 3 or more veins
35
– Leaves pinnately or palmately veined or with a single
midrib
38
35. Leaves petiolate, three-veined
36
– Leaves sessile (New Zealand)
37
36. Involucre calyculate, green, with c. 10 phyllaries
(China)
424. Dicercoclados
– Involucre ecalyculate, yellow, with 5 phyllaries (western USA)
446. Yermo
37. Leaves basal and cauline, lanceolate. Capitula pedunculate, radiate
464. Dolichoglottis
– Leaves cauline, ovate or obovate. Capitula sessile, disciform
461. Haastia p.p.
38. Petioles, if present, with vaginate sheaths
39
– Petioles, if present, not vaginately sheathing
41
39. Capitula solitary or rarely few, campanulate or hemispherical, nodding, usually ecalyculate
429. Cremanthodium
– Capitula several to many in racemose-paniculate or
corymbose synflorescences, calyculate
40
40. Leaves rosulate with involute vernation. Cypselae setose
420. Farfugium
p.p.
– Leaves cauline or rosulate, not involute in bud. Cypselae glabrous
427. Ligularia
41. Leaves pinnately compound, large (Mexico)
443. Villasenoria
– Leaves not pinnately compound
42
42. Stem abruptly contracted below synflorescence
43
– Stems and branches gradually tapering
45
43. Leaves palmately veined, often deciduous before anthesis. Resiniferous subsucculent shrubs
444. Pittocaulon
– Leaves pinnately veined, persistent at anthesis
44
44. Epiphytes or lax suffrutescent herbs. Endothecium polarized
441. Nelsonianthus
– Trees, shrubs or shrublets. Endothecium radial
442. Telanthophora
45. Capitula discoid or disciform
46
– Capitula radiate
76
46. Leaves 3-nerved, elliptic-ovate or lanceolate
47
– Leaves pinnately or palmately veined or just midribbed
48
47. Capitula axillary, solitary or in pairs, calyculate. Involucral bracts c. 10, green (China) 424. Dicercoclados
– Capitula corymbose, ecalyculate. Involucral bracts 5,
yellow (western USA)
446. Yermo
48. Leaves sessile or subsessile
49
– Leaves petiolate
52
49. Shrubs or small trees. Leaf venation pinnate-reticulate
from a prominent midrib. Capitula disciform or sometimes discoid; florets white or cream to pink or purplish (New Guinea)
466. Papuacalia p.p.
– Shrubs, shrublets or herbs. Leaves pinnately to nearly
parallel-veined or one-nerved, sometimes forming
spines. Capitula discoid, bright or creamy yellow
(western USA)
50
50. Shrubs or shrublets. Leaves small (0.5–5 cm long), often fascicled, or transformed into spines
450. Tetradymia
– Herbs or subshrubs. Leaves usually larger, ellipticovate to lanceolate, not fascicled
51
51. Leaves mostly basal and 15–35 cm long, glabrous. Capitula in elongate thyrsoid-racemiform synflorescence
447. Rainiera
– Leaves cauline, 4–12 cm long, white-tomentose beneath. Capitula in short corymbose synflorescence
449. Luina
52. Subshrub with white cortex on old stems. Involucral
bracts 5 in 2 series (Namibia)
554. Dauresia
– Herbs or shrubs, not with white cortex. Involucral
bracts uniseriate
53
53. Florets yellow or orange-coloured
54
Compositae
–
54.
–
55.
–
56.
–
57.
–
58.
–
59.
–
60.
–
61.
–
62.
–
63.
–
64.
–
65.
–
66.
–
67.
–
68.
–
69.
–
70.
Florets white to cream or pink to purplish
60
Shrubs with opposite leaves
55
Herbs with alternate or rosulate leaves
56
Scandent shrub. Involucre ecalyculate (Hispaniola)
474. Herodotia
Erect shrub or shrublet. Involucre calyculate (Colombia, Venezuela)
470. Scrobicaria
Leaves pinnately veined, with broadly winged and amplexicaul petiole. Synflorescence elongate, spike-like
(China)
428. Ligulariopsis
Leaves palmately veined, at least basally; petiole
winged or not
57
Capitula thyrsoid-racemose in elongate synflorescence; involucre ecalyculate (Eurasia)
423. Parasenecio p.p.
Capitula corymbose-paniculate or racemose; involucre calyculate
58
Villous herb with few, palmately lobed leaves. Capitula
racemose. Style branches attenuate, papillate (Japan)
421. Miricacalia
Plants glabrous or sparsely pubescent to floccosetomentose. Capitula corymbose-paniculate
59
Leaves mostly basal, tomentose beneath, palmately
lobed. Capitula campanulate. Style branches attenuate, obtuse to rounded. Endothecium polarized (western USA)
448. Cacaliopsis
Leaves cauline, glabrous or pubescent, entire or variously lobed. Capitula cylindrical to narrowly campanulate. Style branches linear, ± truncate. Endothecium
radial (SW USA, Central America) 440. Roldana p.p.
Shrubs or trees
61
Perennial herbs
67
Capitula disciform; marginal florets with reduced
ligules (New Guinea)
466. Papuacalia p.p.
Capitula discoid
62
Scandent shrublets (Bolivia, Peru)
452. Paracalia
Erect shrubs or trees
63
Plants densely tomentose especially on lower leaf surface
64
Plants glabrous or thinly tomentose in parts
65
Leaves linear to lanceolate (or narrowly ovate); young
stems and leaves glandular (Australia) 462. Bedfordia
Leaves elliptic-oblong to obovate, not distinctly glandular (New Zealand)
458. Brachyglottis p.p.
Glutinous shrub with scattered leaves 5–8 cm long
(New Zealand)
463. Traversia
Trees with large leaves crowded towards branch ends
66
Divaricately much-branched tree. Synflorescences lateral, axillary, pendulous (St. Helena) 469. Lachanodes
Unbranched or little-branched trees or treelets. Synflorescences terminal, erect (Venezuela, Colombia)
453. Paragynoxys
Leaves pinnately veined. Plant pubescent in most parts
(southern USA)
437. Rugelia
Leaves palmately veined
68
Leaves peltate
69
Leaves with marginal petiole attachment
71
Leaves few, cauline, palmatisect (E Asia) 426. Syneilesis
Leaves several to many, entire or lobed
70
Scapigerous herbs with rosulate leaves. Involucre calyculate. Corolla shortly lobed (Central America)
433. Psacaliopsis p.p.
211
– Leaves basal and cauline. Involucre ecalyculate with 5
phyllaries and 5 florets. Corolla deeply lobed (E & SE
USA)
438. Arnoglossum
71. Capitula disciform
72
– Capitula discoid
73
72. Capitula solitary on a long scape, disciform with purple (rarely white) shortly ligulate marginal florets (Europe)
416. Homogyne
– Capitula corymbose. Floret colour various but not purple (Central America)
440. Roldana p.p.
73. Leaves rosulate or subrosulate. Corollas deeply lobed
434. Psacalium
– Leaves cauline, alternate
74
74. Corollas deeply lobed. Endothecium polarized (Mexico)
436. Digitacalia
– Corollas lobed to about the middle of the limb
75
75. Capitula in elongate thyrsoid-racemose synflorescence. Endothecium polarized (Eurasia)
423. Parasenecio p.p.
– Capitula corymbose-paniculate. Endothecium radial
(SW USA, Central America)
440. Roldana p.p.
76. Rays yellow
77
– Rays white or cream
83
77. Trees or shrubs
78
– Herbs
86
78. Leaves opposite, tomentose beneath
79
– Leaves alternate or rosulate
80
79. Erect trees or shrubs. Leaves entire. Capitula campanulate or cup-shaped, many-flowered (South America)
454. Gynoxys
– Scandent shrubs or shrublets. Leaves serrate. Capitula
turbinate-obconical, few-flowered (Hispaniola)
473. Ekmaniopappus p.p.
80. Leaves linear to lanceolate, sessile or subsessile, clustered terminally on branches
81
– Leaves elliptic-oblong to obovate, usually petiolate 82
81. Leaves linear, single-veined, with glossy exudates (Tasmania)
459. Centropappus
– Leaves linear-lanceolate to lanceolate, pinnately
veined (SW USA, Central America)
445. Barkleyanthus
82. Leaves elliptic-ovate or obovate, entire with denticulate margins, ± herbaceous (Chile)
457. Acrisione
– Leaves elliptic-oblong, entire or serrate-lobed, coriaceous (New Zealand)
458. Brachyglottis p.p.
83. Ray florets reduced, cream (New Guinea)
466. Papuacalia p.p.
– Capitula distinctly radiate with white ligules
84
84. Leaves ± tomentose below (New Zealand)
458. Brachyglottis p.p.
– Leaves glabrous or glabrescent
85
85. Leaves up to 40 cm long. Ray florets short, c. 0.5 cm.
Anthers ecaudate (St. Helena)
468. Pladaroxylon
– Leaves up to 10 cm long. Ray florets 1–2 cm long. Anthers conspicuously caudate
460. Urostemon
86. Leaves peltate, petiolate, with hairy base
433. Psacaliopsis p.p.
– Leaves not peltate
87
87. Leaves pinnatisect, cauline (China, Taiwan, Japan)
430. Nemosenecio
– Leaves entire or dentate-lobate
88
88. Involucre biseriate
89
– Involucre uniseriate
90
212
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
89. Stout and somewhat suffrutescent herbs. Leaves
densely woolly beneath
465. Capelio p.p.
– Glabrous or sparsely pubescent herbs
414. Doronicum
90. Leaves basal and cauline, tapering to the base like
a winged petiole
91
– Leaves rosulate or cauline, distinctly petiolate, at least
the basal ones
92
91. Involucre ecalyculate
432. Tephroseris
– Involucre calyculate (Mexico, Guatemala)
439. Robinsonecio
92. Rhizome thick or roots tuberous
93
– Rhizome thin or roots fibrous
94
93. Leaves rosulate with involute vernation. Cypselae
hairy
420. Farfugium p.p.
– Leaves cauline; vernation not involute. Cypselae glabrous
425. Sinacalia
94. Leaves palmately laciniate. Capitula large, showy. Pappus absent (Mexico)
435. Pippenalia
– Leaves entire to lobed. Capitula small or mediumsized. Pappus usually present
95
95. Leaves rosulate and/or cauline, palmately veined
(China, Korea, Myanmar)
431. Sinosenecio
– Leaves rosulate, entire, pinnately veined (New
Zealand)
458. Brachyglottis p.p.
96. Involucral bracts connate (to various extents)
97
– Involucral bracts free
111
97. Annual herbs (often delicate, glabrous)
98
– Perennial herbs, subshrubs or shrubs
105
98. Leaves rosulate
99
– Leaves cauline
100
99. Leaves linear-filiform (Angola)
540. Psednotrichia
– Leaves pinnatipartite (South Africa)
530. Euryops annuus
100. Capitula discoid or disciform
101
– Capitula radiate
102
101. Leaves sessile. Capitula discoid. Cypselae ellipsoid (W
Africa)
539. Bafutia
– Leaves petiolate, auriculate. Capitula disciform.
Cypselae compressed (S Africa)
494. Stilpnogyne
102. Pappus absent
103
– Pappus present
104
103. Ray florets yellow (S Africa)
538. Steirodiscus
– Ray florets white or pink (tropical Africa) 536. Stenops
104. Leaves amplexicaul, serrulate. Pappus of few short caducous bristles (South Africa)
537. Oligothrix
– Leaves not amplexicaul, entire. Pappus of many persistent bristles (Australia)
499. Senecio gregorii, S. calcicola
105. Compact low perennials, forming mats, cushions or
rosettes. Leaves densely set, entire or apically lobed.
Ray florets mostly white (yellow in 2 species of Werneria) (South America)
106
– Perennial herbs, subshrubs or shrubs. Ray florets (if
present) yellow
108
106. Mat- or cushion-forming plants. Leaves very closely
set, those covering rhizomes brown, blackish or
whitish; angular-terete, entire or apically few-lobed
524. Xenophyllum
– Rosette-forming or solitary small perennials. Leaves
assembled near rhizome tips and below capitula, green
107
107. Plants glabrous. Style branch tips papillate or glabrous
525. Werneria
– Plants densely strigose. Style branch tips with elongated hair tuft
523. Misbrookea
108. Leaves succulent, entire or with serrate margins. Style
branches of disc floret papillate-hairy
109
– Leaves coriaceous or herbaceous, entire or variously
lobed. Style branches of disc floret truncate to obtuse
with apical papillae, otherwise glabrous
110
109. Capitula discoid, florets white or pink to purplish.
Style branches with a long papillate appendage
534. Lopholaena
– Capitula radiate or disciform, yellow-flowered. Style
branches papillate-hairy outside
533. Hertia
110. Capitula borne on unbranched axillary or pseudoterminal naked peduncles (Africa, SW Arabia)
530. Euryops
– Capitula many in terminal bracteate corymbs. Leaves
large, herbaceous, pinnatilobate (Cuba) 481. Oldfeltia
111. Style branches of disc floret apically acute, conical or
acuminate, with an appendage, tuft or pencil of fused
hairs or papillae
112
– Style of disc floret apically truncate or rounded-obtuse
134
112. Capitula discoid or disciform
113
– Capitula radiate
122
113. Capitula disciform; marginal female florets numerous,
filiform
503. Erechtites
– Capitula discoid, homogamous
114
114. Style appendage triangular-conical or elongate, made
up of cellular tissue, papillate
115
– Style appendage a central tuft or pencil of fused hairs
or papillae
117
115. Truly succulent herbs or shrubs
501. Kleinia p.p.
– Plants herbaceous, at most subsucculent, or shrubs
with coriaceous leaves
116
116. Herbs or subshrubs. Capitula corymbose-paniculate
or solitary, calyculate
502. Gynura p.p.
– Shrublet with small coriaceous leaves. Capitula ecalyculate, solitary on long peduncles
527. Lamprocephalus
117. Involucre ecalyculate
541. Emilia
– Involucre calyculate
118
118. Florets pure yellow
500. Solanecio p.p.
– Floret colour various but not pure yellow
119
119. Leaves distinctly petiolate. Capitula nodding
517. Aetheolaena
– Leaves subsessile or shortly petiolate. Capitula erect
or nodding
120
120. Leaves imbricate with revolute margins, tomentose beneath
516. Lasiocephalus
– Leaves not imbricate, flat
121
121. Leaves herbaceous, entire or lobed to dissected. Florets
white, yellowish, red, blue, etc. Involucre cylindrical
(Africa to Yemen)
506. Crassocephalum p.p.
– Leaves coriaceous, entire, serrate. Florets greenish yellow. Involucre campanulate (Colombia)
518. Arbelaezaster
122. Herbs
123
– Shrubs or subshrubs
128
123. Leaves rosulate, pinnatipartite or laciniate. Involucre
calyculate or conspicuously calyculate (Peru, Ecuador)
496. Dorobaea
– Leaves not rosulate. Involucre calyculate or ecalyculate
124
124. Rays white (Chile, Argentina)
504. Iocenes
Compositae
– Rays variously coloured but not white (except in
Graphistylis organensis f. albiflora from Brazil) 125
125. Leaves distinctly petiolate
126
– Leaves subsessile or shortly petiolate
127
126. Petiole conspicuously auriculate and amplexicaul.
Leaves strongly nerved
515. Garcibarrigoa
– Petiole without or with small auricles. Leaves normally
veined
514. Pseudogynoxys p.p.
127. Coarse herbs (or subshrubs). Leaves ellipticlanceolate, serrate, subcoriaceous. Capitula many;
involucre campanulate or cup-shaped (Brazil)
511. Graphistylis
– Soft herbs. Leaves entire or lobed, herbaceous. Capitula few to several; involucre cylindrical (tropical
Africa, Yemen)
506. Crassocephalum p.p.
128. Scandent subshrubs or shrubs. Rays conspicuous, red
or orange to yellow, fragrant (Central & South America)
514. Pseudogynoxys p.p.
– Erect shrubs or small trees
129
129. Leaves linear-lanceolate, < 3 cm wide. Receptacle
naked (Costa Rica)
522. Charadranaetes
– Leaves elliptic-lanceolate to oblong-ovate, > 3 cm
wide. Receptacle denticulate (i.e. with tooth-like
processes)
130
130. Leaves ± hairy. Ray florets numerous (8–21), lamina
distinctly longer than tube, yellow or orange (South &
Central America)
512. Talamancalia
– Leaves glabrous at least when mature. Ray florets few
(2–8 or rarely more in Lundinia), short; lamina about
equalling the tube in length
131
131. Low shrubs. Leaf-blades up to 4 cm long. Rays narrow,
pale yellow (Lord Howe Island)
487. Lordhowea
– Tall shrubs or small trees. Leaves larger. Rays bright
yellow
132
132. Style branches apically with an acuminate papillate
appendage; stigmatic areas continuous (Jamaica)
484. Jacmaia
– Style branches with apical hair tuft; stigmatic areas
separated
133
133. Style branches apically rounded with hair pencil. Anthers ecaudate (Costa Rica)
486. Jessea
– Style branches truncate with minute hair tuft. Anthers
caudate (Cuba, Hispaniola)
483. Lundinia
134. Leaves opposite, sessile or subsessile, entire with dentate or denticulate margins
135
– Leaves alternate or rosulate
136
135. Erect subshrub with radiate capitula (Madagascar)
505. Io
– Scandent shrubs with discoid or radiate capitula
(Colombia)
520. Cabreriella
136. Anthers distinctly caudate (‘Synotoids’ a.o.)
137
– Anthers basally obtuse to sagittate
153
137. Erect shrubs or shrublets with narrowly linear or acicular leaves
138
– Herbs or shrubs, sometimes scandent. Leaves flat, pinnately or sometimes palmately veined
139
138. Leaves closely set, patent. Capitula several, pedunculate
548. Parafaujasia
– Leaves imbricate, appressed. Capitula solitary, sessile
545. Eriotrix
139. Pappus bristles few, rigid, basally flattened
544. Faujasia
– Pappus bristles numerous, soft or slender, terete 140
140. Scandent herbs or subshrubs
141
–
141.
–
142.
–
143.
–
144.
–
145.
–
146.
–
147.
–
148.
–
149.
–
150.
–
151.
–
152.
–
153.
–
154.
–
155.
–
156.
–
157.
–
158.
–
159.
213
Erect shrubs or sometimes subshrubs
149
Leaves pinnately veined
142
Leaves palmately veined
144
Florets white or pink to lilac (Mascarene Islands)
543. Faujasiopsis p.p.
Florets yellow
143
Style branches truncate (Madagascar)
546. Hubertia faujasioides
Style branches obtuse-rounded (Jamaica)
482. Odontocline p.p.
Leaves not auriculate
145
Leaves auriculate
147
Petioles not prehensile (Madagascar, La Réunion)
547. Humbertacalia p.p.
Petioles prehensile
146
Capitula radiate or discoid (Indomalesia)
552. Cissampelopsis
Capitula disciform (Africa)
550. Mikaniopsis
Florets whitish. Petioles not prehensile (Madagascar,
La Réunion)
547. Humbertacalia p.p.
Florets yellow. Petioles prehensile
148
Plant subsucculent. Capitula discoid (South Africa)
551. Delairea
Plant herbaceous. Capitula radiate (tropical Africa)
553. Austrosynotis
Capitula small, with 1–2 ray florets and 4–5 disc florets,
ecalyculate (Canary Isl.)
489. Bethencourtia
Capitula > 5-flowered, calyculate
150
Styles with clearly separated stigmatic areas
151
Style with continuous or barely separated stigmatic
areas
152
Diffuse subshrubs or shrubs with leaves often caulirosulate (S & SE Asia)
549. Synotis
Leaves evenly distributed (Cuba, Hispaniola)
483. Lundinia
Style branch apex blunt with only very short papillae
(Madagascar, La Réunion)
546. Hubertia
Style branch apex rounded-conical with 1–2 lateral
bunches of sweeping hairs (Jamaica)
482. Odontocline p.p.
Cypselae compressed. Herbs or subshrubs. Leaves
mostly lobed to dissected, palmately veined
154
Cypselae not compressed. Leaves mostly pinnately
veined, or linear, midveined
155
Florets yellow. Cypselae distinctly compressed with
thickened margins (Africa & Madagascar)
491. Cineraria
Florets non-yellow (white, pink, purple, etc.). Cypselae
not much compressed, margins not thickened (Macaronesian islands)
490. Pericallis
Ray florets bilabiate with a short limb and two small
ventral lobes. Scandent shrub with three-nerved leaves
(Hispaniola)
472. Mattfeldia
Ray florets ligulate or absent. Non-scandent plants 156
Capitula ecalyculate
157
Capitula calyculate (sometimes with a single calyculus
bract)
162
Shrubs or subshrubs
158
Herbs
160
Capitula borne singly on unbranched terminal or axillary peduncles
159
Capitula several, corymbose
499. Senecio p.p.
Peduncles terminal, scapose, bracteate. Ray florets
white or cream. Cypselae dimorphic: in ray florets
214
–
160.
–
161.
–
162.
–
163.
–
164.
–
165.
–
166.
–
167.
–
168.
–
169.
–
170.
–
171.
–
172.
–
173.
–
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
glabrous with caducous pappus, in disc florets with
myxogenic hairs and persistent pappus
528. Phaneroglossa
Peduncles axillary, naked. Ray florets yellow. Cypselae
homomorphic
530. Euryops spp.
Pappus a single scale or absent. Capitula few-flowered,
pink to purplish
542. Emiliella
Pappus of slender bristles. Florets several to many,
colour various
161
Cypselae dimorphic: outer adaxially midribbed and
glabrous, inner smooth and hairy. Capitula radiate,
yellow
492. Bolandia
Cypselae homomorphic. Capitula discoid or sometimes radiate; floret colour various 541. Emilia p.p.
Style branches of disc florets short, erect, apically
dilated, dorsally papillate-hairy. Capitula disciform
with purplish-creamy white florets (Indonesia, New
Guinea, Australia)
488. Arrhenechthites p.p.
Style branches of disc florets linear or oblong, spreading
163
Pachycaul trees or tree-like suffrutescent herbs (‘dendrophorbs’) with large leaves assembled terminally on
stems and branches
164
Herbs, shrubs or small trees with more slender stems
and more evenly distributed (or rosulate) leaves 165
Megaphytic rosette-trees. Styles of disc florets with
continuous stigmatic areas. Cypselae 5-ribbed or 5angled (tropical African mountains)
529. Dendrosenecio
Subshrubs, shrubs or treelets. Styles of disc florets
with separated stigmatic areas. Cypselae 8–10-ribbed
(South America)
510. Dendrophorbium
Herbs
166
Subshrubs, shrubs or shrublets, or small trees
172
Annual herbs
167
Perennial herbs
168
Involucral bracts c. biseriate; inner with 2 dark resin
ducts. Cypselae subtriquetrous, black, papillose-hairy
apically and basally and along three longitudinal lines
(southern Africa)
493. Mesogramma
Involucral bracts ± uniseriate, without dark stripes.
Cypselae ± terete
499. Senecio p.p.
Roots fibrous (rhizome present in some Old World and
South American spp.)
169
Rhizomatous herbs with most leaves basal, often rosulate or nearly so
171
Styles of disc florets with separate stigmatic areas,
apically ± truncate and with sweeping hairs
170
Styles with continuous stigmatic areas, apically obtuse
and without distinct sweeping hairs. Capitula discoid,
orange-flowered (Hispaniola)
585. Ignurbia
Cypselae dimorphic: outer 4-winged, glabrous; inner
6-ribbed, with mucilaginous hairs
526. Oresbia
Cypselae homomorphic
499. Senecio p.p.
Capitula campanulate to cup-shaped, radiate or rarely
discoid; florets yellow, orange or red. Corolla lobes
deltoid-ovate
495. Packera
Capitula cylindrical-turbinate, discoid; florets white
or pink. Corolla lobes lanceolate
497. Hasteola
Scandent shrubs or shrublets
173
Erect subshrubs, shrubs or small trees
176
Styles of disc florets with continuous stigmatic areas
(Cuba, Hispaniola)
174
Styles of disc florets with separated stigmatic areas 175
174. Leaves 3-nerved from the base, glabrous 475. Leonis
– Leaves pinnately veined, tomentose beneath
476. Nesampelos
175. Leaves tomentose beneath. Cypselae 8–10-ribbed
(Hispaniola)
479. Elekmania p.p.
– Leaves glabrous or variously pubescent. Cypselae 5ribbed or 5-angled (South & Central America)
508. Pentacalia p.p.
176. Styles of disc florets apically rounded and subglabrous,
dorsally puberulous. Capitula discoid, few-flowered
(Brazil)
507. Hoehneophytum
– Style of disc florets truncate or obtuse, penicillate. Capitula radiate or discoid, usually many-flowered 177
177. Cypselae 5-ribbed or 5-angled (South & Central America)
178
– Cypselae 8–10-ribbed
179
178. Erect shrubs or shrublets. Leaves closely set, subsessile
or shortly petiolate, often small, ericoid to lanceolate
or elliptic-ovate
509. Monticalia
– Epiphytic shrubs. Leaves more laxly distributed, petiolate
508. Pentacalia p.p.
179. Styles of disc florets with continuous stigmatic areas
(Cuba)
180
– Styles of disc florets with separated stigmatic areas 181
180. Involucral bracts biseriate. Style branches with stigmatic areas extending to the adaxial side, which is
papillate in the middle
480. Herreranthus
– Involucral bracts uniseriate. Style branches stigmatic
inside, glabrous outside
478. Antillanthus p.p.
181. Style of disc florets obtuse, with short and few sweeping hairs (Caribbean)
182
– Style of disc florets ± truncate with distinct sweeping
hairs. Capitula yellow-flowered
183
182. Capitula radiate, white-flowered (Jamaica)
477. Zemisia
– Capitula radiate or discoid, yellow-flowered (Hispaniola)
479. Elekmania p.p.
183. Ovary wall crystals prismatic, plate- or needle-like.
Capitula radiate or sometimes discoid or disciform
499. Senecio p.p.
– Ovary wall crystals drusiform. Capitula always discoid
(Africa, Madagascar, Yemen)
500. Solanecio p.p.
Genera of Senecioneae
410. Abrotanella Cass.
Fig. 54
Abrotanella Cass., Dict. Sci. Nat. 36: 27 (1825); Swenson,
Pl. Syst. Evol. 197: 149–193 (1995), rev.; Swenson & Bremer,
Syst. Bot. 22: 493–508 (1997), biogeogr.; Heads, Biol. J. Linn.
Soc. 67: 39–432 (1999), biogeogr.
Dwarf or prostrate perennial herbs, often forming mats or cushions. Leaves alternate, sessile with
sheathing base, entire, coriaceous or subcarnose,
glabrous with sunken glands. Capitula mostly sessile, small, disciform; florets few, whitish, greenishyellow or red to purplish. Marginal florets female,
tubular. Disc florets perfect or functionally male,
4-lobed; corolla lobes with central vascular strand
and without lateral strands.
Compositae
215
lobed. Capitula solitary, radiate, yellow-flowered.
Receptacle conical. Ray florets female, with distinct
tube, with or without pappus bristles. Disc florets
perfect. Anther base obtuse, ecaudate; endothecium polarized. Style branches with conical hair
tip; stigmatic areas separated. Cypselae obovoid,
with short myxogenic twin hairs. Pappus bristles
slender, densely barbellate, caducous. n = 9. One
species, C. multicaule Hook., north-western USA.
413. Ischnea F. Muell.
Ischnea F. Muell., Trans. Roy. Soc. Victoria 1, 2: 13 (1889);
Koster, Blumea 22: 207–217 (1975), rev.; Royen, Alpine Fl.
New Guinea 4: 3352–3363 (1983), rev.; Swenson, Pl. Syst.
Evol. 191: 247–263 (1994), rev.
Fig. 54. Compositae-Senecioneae. Abrotanella linearis.
A Habit. B Leaf base. C Capitulum. D Involucral bracts.
E Outer floret. F Central floret. G Mature cypsela. (Swenson
1995)
Anthers shortly caudate; endothecium polarized.
Style simple or shortly bilobed, truncate-obtuse,
papillose. Cypselae oblong-obovoid, glabrous or
papillose-puberulous with twin hairs, without pappus, but sometimes with an apical rim of small
teeth. n = 9, 18. Twenty species, Australasia, southern South America.
411. Blennosperma Less.
Blennosperma Less., Syn. Gen. Comp.: 267 (1832); Ornduff,
Brittonia 16: 289–295 (1964), rev.; Swenson, Doct. Diss. Uppsala Univ.: 10–12 (1995).
Small annual herbs. Leaves alternate, sessile, mostly
pinnatifid. Capitula solitary on naked peduncles,
radiate, yellow- or purplish-flowered. Receptacle
conical. Ray florets female, tube reduced. Disc florets 5-lobed, functionally male. Anthers ecaudate;
endothecium polarized. Style simple, sterile, apically enlarged. Cypsela obovoid, with short myxogenic twin hairs. Pappus absent. n = 7, 8, 9, 16.
Three species, Chile, California.
412. Crocidium Hook.
Crocidium Hook., Fl. Bor.-Am. 1: 335, t. 112 (1834); St.
John, Torreya 28: 73–77 (1928), rev.; Morton, N. Amer. Fl.
ser. 2, 10: 147 (1978), rev.
Small annual herb. Leaves basal, rosulate, and
cauline and clasping, entire or somewhat dentate to
Small perennial herbs, subrosulate, tufted or solitary. Leaves linear-oblanceolate, entire or sparsely
dentate. Capitula solitary, small and few-flowered,
shortly radiate, yellow- or purplish-flowered.
Involucral bracts biseriate. Receptacle slightly convex, naked. Ray florets with ± reduced tube. Disc
florets functionally male; corolla 4- or 5-lobed.
Anthers ecaudate; endothecium polarized. Style
simple, sterile. Cypselae elliptic-oblong, glabrous,
without pappus. n = 9. Four species, New Guinea.
414. Doronicum L.
Doronicum L., Sp. Pl.: 855 (1753) & Gen. Pl. ed. 5: 377
(1754); Cavillier, Ann. Cons. Jard. Bot. Genève 13: 195–367
(1911), rev.; Gorshkova, Fl. U.S.S.R. 26: 669–682 (1961), reg.
rev.; Edmondson, Fl. Turkey 5: 137–145 (1975), reg. rev.;
Ferguson, Fl. Eur. 4: 190–191 (1976), reg. rev.; Edmondson,
Notes Roy. Bot. Gard. Edinburgh 37: 67–74 (1978), reg.
rev.; Rechinger, Fl. Iran. 164: 44–48 (1989), reg. rev.; Chen,
Fl. Reip. Pop. Sin. 77, 1: 4–13 (1999), reg. rev.; Alvarez
Fernandez, Ann. Missouri Bot. Gard. 90: 319–389 (2003),
rev.
Perennial herbs. Leaves basal and cauline,
rounded-cordate or ovate-oblong; basal leaves
petiolate; cauline leaves sessile, often amplexicaul.
Capitula solitary or few to several, corymbose,
large, radiate, yellow-flowered. Involucral bracts
biseriate. Anther base obtuse or auriculate; endothecium polarized. Style branches obtuse with
few sweeping hairs; stigmatic areas continuous.
Cypselae oblong, ribbed, glabrous or pubescent.
Pappus bristles numerous, often lacking in ray
florets, slender, sometimes dimorphic. Pollen
helianthoid. n = 15, 20, 30, 45, 60. Circa 40 species,
Eurasia, northern Africa.
216
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
415. Chersodoma Phil.
Chersodoma Phil., Anales Mus. Nac. Chile 1891: 33 (1891);
Cabrera, Revista Mus. La Plata, Bot. 6: 343–355 (1946),
re-establ., rev.; Nordenstam, Opera Bot. 44: 22–23 (1978),
notes, subg. nov.; Dillon & Sagástegui, Brittonia 48: 582–604
(1997), rev.
Dioecious shrubs, subshrubs or perennial herbs.
Leaves petiolate or subsessile, often tomentose
beneath, entire; margins sometimes dentate, often
revolute. Capitula solitary, pedunculate, or few
and laxly paniculate, homogamous; florets either
pistillate with aborted anthers or functionally
male with sterile ovary. Anthers caudate; endothecial tissue polarized; filament collar cylindrical,
uniform. Style branches apically rounded to
subtruncate without sweeping hairs, dorsally
puberulous-papillose; stigmatic areas separated
at least in the basal half. Cypselae oblong, ribbed,
glabrous or hairy. Pappus bristles slender, white
or purplish-brown, persistent. n = 10. Nine
species, Andean South America from Argentina to
Peru.
416. Homogyne Cass.
Homogyne Cass., Bull. Soc. Philom. Paris 1816: 198 (1816);
Vierhapper, Oesterr. Bot. Z. 1923(6–8): 150–164 (1923),
syst. position; Tutin, Fl. Eur. 4: 188–189 (1976), rev.
Perennial herbs. Leaves mostly basal, petiolate,
rounded to cordate, entire with dentate to lobulate
margins, palmately veined; cauline leaves smaller,
few. Capitula solitary on long peduncles, disciform,
purple- or white-flowered. Marginal florets female,
tubular and very shortly radiate. Disc florets
perfect. Anthers exserted, ecaudate. Style branches
truncate, penicillate. Cypselae oblong, ribbed,
glabrous. Pappus bristles numerous, slender,
white or dirty white. n = c. 28, 29, 30, 60, 65,
65–70, 70, 80. Three species, central and southern
Europe.
417. Endocellion Turcz. ex Herder
Endocellion Turcz. ex Herder, Bull. Soc. Imp. Naturalistes
Moscou 38: 345 (1865); Toman, Folia Geobot. Phytotax.
(Prague) 7: 381–406 (1972), rev.
Perennial dioecious herbs with a slender rhizome.
Leaves rosulate, developing after anthesis, petiolate, ± ovate, palmately veined; cauline leaves sessile, scale-like. Capitula solitary or rarely few together, yellow-flowered, calyculate. Male capitula
discoid or disciform with functionally male disc
florets. Female capitula radiate, lacking disc florets;
florets radiate. Cypselae oblong, glabrous. Pappus
bristles numerous, slender. n = 28, 29, 30, c. 50+,
56. Two species, Siberia, eastern Asia.
418. Petasites Mill.
Petasites Mill., Gard. Dict. abr. ed.: 4 (1754); Toman,
Folia Geobot. Phytotax. (Prague) 7: 381–406 (1972), rev.;
Cherniawski & Bayer, Canad. J. Bot. 76: 2061–2075 (1999),
reg. rev.; Chen, Fl. Reip. Pop. Sin. 77, 1: 94–101 (1999), reg.
rev.; Kupriyanova, Flora U.S.S.R. 26: 645–654 (1961), reg.
rev. (sub Nardosmia).
Nardosmia Cass. (1825).
Perennial dioecious or subdioecious herbs with
a thick rhizome. Leaves rosulate, developing
after anthesis, petiolate, rounded-cordate or
triangular, entire to palmately lobed, palmately
veined. Cauline leaves scale-like. Capitula several,
paniculate-racemose, large, calyculate, radiate,
disciform or discoid; ‘male’ capitula with or
without marginal florets and with numerous
tubular functionally male florets. Female capitula
with sterile tubular or shortly radiate florets and
numerous filiform female florets. Styles apically
clavate to cylindrical, shortly bilobed. Cypselae
cylindric-oblong, glabrous, ribbed. n = 10, 14, 16,
26, 28, 29, 30, 40, c. 44, 45, 60. Circa 20 species,
Eurasia, North America.
419. Tussilago L.
Tussilago L., Sp. Pl.: 865 (1753) & Gen. Pl. ed. 5: 372 (1754).
Perennial rhizomatous herb. Leaves rosulate, petiolate, large, developing after anthesis, roundedcordate, entire with dentate margins. Capitula solitary on a bracteate scapiform stem, radiate, yellow. Ray florets numerous, female, with narrow
rays. Disc florets functionally male, tubular. Anthers sagittate. Style branches apically clavate, sterile. Cypselae oblong, ribbed, glabrous. Pappus bristles numerous, slender, persistent. n = 30, 36. One
species, T. farfara L., Eurasia.
420. Farfugium Lindl.
Farfugium Lindl., Gard. Chron. 1857: 4 (1857); Kitamura,
Acta Phytotax. Geobot. 8: 77–89 (1939), rev.; Koyama, Mem.
Fac. Sci. Kyoto Univ. ser. Biol. 2: 38 (1968), reg. rev.
Perennial herbs with a thick rhizome. Leaves rosulate, petiolate, with involute vernation, ovatecordate or reniform, entire with dentate margins,
palmately veined. Capitula few to several, corym-
Compositae
bose, radiate, calyculate, yellow-flowered. Disc florets perfect. Anthers caudate. Style branches tapering towards the obtuse apex, papillate throughout.
Cypselae oblong, hairy. Pappus bristles numerous,
slender. n = 30, 31. Three species, Japan, southeastern China, Korea, Taiwan.
421. Miricacalia Kitam.
Miricacalia Kitam., Acta Phytotax. Geobot. 5: 214 (1936);
Koyama, Mem. Fac. Sci. Kyoto Univ. ser. Biol. 2: 163 (1969),
rev.; Robinson & Brettell, Phytologia 27: 265–276 (1973),
notes.
Perennial villous herb. Leaves few, petiolate, palmately lobed and veined. Capitula several, racemose, discoid, calyculate, yellow-flowered. Anthers
ecaudate, auriculate. Style branches long, papillate,
apically truncate, penicillate. Cypselae narrowly
elliptic-oblong, glabrous, beaked and with a long
carpopodium. Pappus bristles numerous, slender.
n = 26, 27, 30. One species, M. makineana (Yatabe)
Kitam., Japan.
422. Dendrocacalia (Nakai) Nakai ex Tuyama
Dendrocacalia (Nakai) Nakai ex Tuyama, Bot. Mag. (Tokyo)
50: 129 (1936); Koyama, Mem. Fac. Sci. Kyoto Univ. ser. Biol.
2: 157–158 (1969), rev.; Kato & Nagamasu, J. Pl. Res. 108:
443–450 (1995), morph.
Erect branched dioecious shrub with soft wood.
Leaves ovate to oblanceolate, entire, glabrous, pinnately veined. Capitula several, corymbose, discoid, calyculate, pink to purplish-flowered. Florets
5, female or functionally male. Anther base obtuse.
Style branches papillate, apically truncate, penicillate; stigmatic areas continuous. Cypselae oblong, pubescent. Pappus bristles numerous, slender. n = 30. One species, D. crepidifolia (Nakai)
Nakai ex Tuyama, Bonin Island.
423. Parasenecio W.W. Smith & Small
Parasenecio W.W. Smith & Small, Trans. Proc. Bot. Soc.
Edinburgh 28: 93 (1923); Chen, Fl. Reip. Pop. Sin. 77, 1:
19–87 (1999), reg. rev.; sub Cacalia auct. non L.: Pojarkova,
Fl. U.S.S.R. 26: 683–697 (1961), reg. rev.; Koyama, Mem.
Fac. Sci. Kyoto Univ. ser. Biol. 2: 163–181 (1969), reg. rev.;
Koyama, Acta Phytotax. Geobot. 29: 65–84, 171–178 (1978)
& 30: 65–84 (1979), reg. rev.
Koyamacalia H. Robins. & Brettell (1973).
Perennial herbs. Leaves basal and cauline,
petiolate, or cauline leaves sometimes sessile,
rounded-cordate to deltoid or ovate, entire or pal-
217
mately lobed, usually palmately veined; margins
usually denticulate to serrate. Capitula numerous,
discoid, paniculate-racemose or in spike-like
synflorescences, small, ecalyculate, white-, creamor yellow-flowered. Anthers sagittate to shortly
caudate. Style branches with continuous stigmatic
areas, apically obtuse, papillate. Cypselae ellipticoblong, glabrous, with distinct carpopodium.
Pappus bristles numerous, slender, white or tawny
to reddish. n = 20, 26, 29, 30, 31, 45, 60. Circa 70
species, Eurasia from Russia eastwards with the
majority in eastern Asia, one species reaching the
Aleutian Isl.
424. Dicercoclados C. Jeffrey & Y.L. Chen
Dicercoclados C. Jeffrey & Y.L. Chen, Kew Bull. 39: 213
(1984).
Perennial rhizomatous herb. Leaves cauline,
petiolate, entire, lanceolate, three-veined; margins
sparsely denticulate. Capitula solitary or few
together, axillary, pedunculate, discoid, calyculate,
yellow-flowered. Anthers caudate with long firm
tails. Style branches with continuous stigmatic
areas, apically with two hair tufts. Cypselae narrowly oblong, glabrous. Pappus bristles numerous,
slender. One species, D. triplinervis C. Jeffrey &
Y.L. Chen, China.
425. Sinacalia H. Rob. & Brettell
Sinacalia H. Rob. & Brettell, Phytologia 27: 274 (1973); Nordenstam, Opera Bot. 44: 15 (1978), syst., nomencl.; Jeffrey
& Chen, Kew Bull. 39: 215–222 (1984), rev.; Chen, Fl. Reip.
Pop. Sin. 77, 1: 13–18 (1999), rev.
Perennial herbs with an underground tuber. Leaves
cauline, petiolate, ovate to rounded-cordate or deltoid, entire or palmately lobed, palmately veined or
nearly so. Capitula several to many, rarely solitary,
thyrsoid or paniculate-corymbose, narrow, radiate, ecalyculate, yellow-flowered. Anthers auriculate or sagittate, apical appendage dark-coloured,
midlined. Style branches ± truncate, apically papillate, stigmatic areas continuous but basally discrete. Cypselae oblong, ribbed, glabrous, with distinct cylindrical carpopodium. Pappus bristles numerous, slender. n = 30. Four species, China.
426. Syneilesis Maxim.
Syneilesis Maxim., Prim. Fl. Amur.: 165 (1859); Koyama,
Mem. Fac. Sci. Kyoto Univ. ser. Biol. 2: 159–162 (1969), rev.;
Robinson & Brettell, Phytologia 27: 265–276 (1973), notes;
Chen, Fl. Reip. Pop. Sin. 77, 1: 89–93 (1999), reg. rev.
218
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Perennial rhizomatous herbs. Seedling with
a single cotyledon. Leaves cauline, few, petiolate, peltate, palmately lobed or dissected and
palmately veined. Capitula several to many,
racemose-paniculate or corymbose, discoid,
calyculate, white-flowered. Anthers caudate. Style
branches elongate, papillate, apically truncate and
penicillate. Cypselae narrowly elliptic to oblong,
glabrous. Pappus bristles numerous, slender.
n = 26, 39. Seven species, eastern Asia.
427. Ligularia Cass.
Ligularia Cass., Bull. Soc. Philom. Paris 1816: 198 (1816),
nom. cons.; Handel-Mazzetti, Bot. Jahrb. 69: 95–142 (1938),
reg. rev.; Pojarkova, Notul. Syst. Herb. Inst. Bot. Komarov.
Acad. Sci. U.R.S.S. 12: 293–318 (1950), syst.; Pojarkova, Fl.
U.S.S.R. 26: 788–857 (1961), reg. rev.; Koyama, Mem. Fac.
Sci. Kyoto Univ. ser. Biol. 2: 39–52 (1968), reg. rev.; Liu, Fl.
Reip. Pop. Sin. 77, 2: 4–115 (1989), reg. rev.; Rechinger, Fl.
Iran. 164: 48–51 (1989), reg. rev.
Perennial herbs with a creeping rhizome. Leaves
basal and cauline, petiolate with strongly sheathing leaf base, oblong to oblanceolate or cordate,
entire with often serrate-dentate margins. Capitula several to many, racemose-paniculate, radiate
or sometimes discoid, yellow-flowered. Involucre
narrowly cup-shaped to cylindrical, calyculate. Anthers auriculate to shortly caudate. Style branches
subtruncate to obtuse with short lateral sweeping hairs, papillate apically or throughout. Cypselae narrowly oblong to elliptic-oblong, ribbed, glabrous. Pappus bristles numerous, slender, white or
tawny to rufous. n = 15–16, 16, 24, 25–26, 27, 29,
30, 31. Circa 125 or more species, Eurasia. Genus
under revision (I. Illarionova).
428. Ligulariopsis Y.L. Chen
429. Cremanthodium Benth.
Cremanthodium Benth., Hooker’s Icon. Pl. 12: 37, t. 1141
(1873); d’O. Good, J. Linn. Soc., Bot. 48: 259–316 (1929),
rev.; Koyama, Mem. Fac. Sci. Kyoto Univ. ser. Biol. 2: 52–60
(1968), reg. rev.; Liu, Fl. Reip. Pop. Sin. 77, 2: 115–171
(1989), reg. rev.
Perennial herbs. Leaves rosulate, sometimes also
a few cauline, petiolate, ovate-cordate to reniform
or elliptic-oblong to lanceolate, rarely pinnatifid;
leaf base sheathing. Capitula solitary or few,
racemose, broad, nodding, radiate or discoid,
ecalyculate. Involucre broadly cup-shaped or
campanulate. Florets white, pink, purple, yellow
or orange-coloured. Anthers ecaudate. Style
branches obtuse, penicillate, dorsally papillate.
Cypselae elliptic-oblong, glabrous. Pappus bristles
numerous, slender, white or reddish. n = 29. Circa
75 species, China, Tibet and Himalaya.
430. Nemosenecio (Kitam.) B. Nord.
Fig. 55
Nemosenecio (Kitam.) B. Nord., Opera Bot. 44: 45 (1978),
45–46, new comb., new sp.; Jeffrey & Chen, Kew Bull. 39:
262–266 (1984), reg. rev.; Chen, Fl. Reip. Pop. Sin. 77, 1:
161–166 (1999), reg. rev.
Senecio sect. Nemosenecio Kitam. (1937).
Perennial herbs. Leaves cauline, shortly petiolate,
pinnately lobed or dissected. Capitula several,
corymbose, radiate, yellow-flowered, ecalyculate.
Anthers short, ecaudate, with narrow apical appendage; endothecium transitional. Style branches
truncate, stigmatic areas partly confluent but
discrete at least basally. Cypselae narrowly oblong,
glabrous or sparsely pubescent. Pappus bristles
numerous, slender, persistent. n = 5, 10, 20, 24 .
Six species, Japan, China.
431. Sinosenecio B. Nord.
Ligulariopsis Y.L. Chen, Acta Phytotax. Sin. 34: 631 (1996);
Chen, Fl. Reip. Pop. Sin. 77, 1: 87–89 (1999), rev.
Sinosenecio B. Nord., Opera Bot. 44: 48 (1978), 48–51, new
comb.; Jeffrey & Chen, Kew Bull. 39: 222–261 (1984), rev.;
Chen, Fl. Reip. Pop. Sin. 77, 1: 101–141 (1999), rev.
Perennial herb. Leaves cauline, petiolate, ovate,
pinnately veined; petiole winged and with amplexicaul base. Capitula numerous in elongated
spike-like thyrsoid-paniculate synflorescence,
discoid, few-flowered. Disc florets perfect, yellow.
Anther base obtuse, ecaudate. Style branches
obtuse, penicillate, papillate to below bifurcation.
Cypselae cylindrical-oblong, ribbed, glabrous.
Pappus bristles numerous, slender, purplishbrown. n = 29. One species, L. shichuana Y.L.
Chen, China.
Perennial herbs with fibrous roots or a rhizome.
Leaves rosulate and/or cauline, petiolate, rounded
to cordate, reniform or triangular-ovate, entire or
margins dentate or sinuate-lobate. Capitula solitary
or several, corymbose-paniculate, radiate, yellowflowered, ecalyculate. Anthers ecaudate; endothecium polarized. Style branches apically truncate
to obtuse with short or long sweeping hairs, dorsally papillate, stigmatic areas confluent or narrowly separated. Cypselae elliptic-oblong, ribbed,
glabrous or pubescent, sometimes heteromorphic
Compositae
219
Perennial or biennial herbs. Leaves radical and
cauline, sessile with often petioliform base, entire
with often serrate or dentate margins, pinnately
veined. Capitula few or several, rarely solitary, radiate or occasionally discoid, ecalyculate, yellow-,
orange- or red-flowered. Anthers with radial
endothecium, filament collar cylindrical. Style
branches truncate to obtuse, with continuous
stigmatic areas. Cypselae oblong, ribbed, glabrous
or pubescent. Pappus bristles numerous, slender.
n = 9, 13, 14, 17, 18, 19(?), 20, 22, 23, 24, 25, 32,
36, 40, c. 40, 48, 45–52. Circa 50 species, northern
Eurasia, one species in North America.
433. Psacaliopsis H. Rob. & Brettell
Psacaliopsis H. Rob. & Brettell, Phytologia 27: 408 (1974).
Perennial scapigerous herbs. Leaves rosulate,
petiolate, peltate, entire or lobed, palmately
veined, with hairy leaf base. Capitula solitary
or few, radiate, yellow-flowered, or discoid
and reddish-flowered, calyculate. Disc floret
corolla shortly lobed, with resin canals. Anthers with radial endothecium; filament collar
cylindrical. Style branches with continuous
stigmatic areas. Cypselae oblong, glabrous.
Pappus bristles numerous, slender, caducous.
n = 30. Circa 6 species, Mexico, Guatemala.
434. Psacalium Cass.
Fig. 55. Compositae-Senecioneae. Nemosenecio yunnanensis. A Habit. B Ray floret. C Disc floret. D Corolla of disc
floret, opened. E Stamens. F Style branches of disc floret.
(Drawing by B. Nordenstam)
Psacalium Cass., Dict. Sci. Nat. 43: 461 (1826); Rydberg,
Bull. Torrey Bot. Club 51: 370–376 (1924), reg. rev.; Pippen,
Contr. U.S. Natl Herb. 34: 415–436 (1968), 384–410 (1968),
rev. (sub Odontotricho); Robinson & Brettell, Phytologia
27: 254–264 (1973), rev.
Odontotrichum Zucc. (1832).
(glabrous and epappose in ray florets, hairy and
pappose in disc florets). Pappus bristles numerous,
slender, persistent or caducous, sometimes absent
in ray florets. n = 23, 24. Circa 40 species, southeastern Asia, mainly China, one species in Canada
(prob. not congeneric).
Scapigerous perennial herbs. Leaves rosulate
or at least mainly basal, petiolate, peltate or
with marginal petiole, rounded-ovate, entire or
dentate-lobate to pinnatisect; leaf bases hairy;
cauline leaves reduced. Capitula several to many,
paniculate to corymbose or racemose, discoid,
calyculate. Corolla white or yellowish or purplish,
deeply lobed. Anthers ecaudate; endothecium
radial. Style branches with continuous stigmatic
areas. Cypselae elliptic-obovoid, somewhat compressed, ribbed, glabrous or pubescent. Pappus
bristles numerous, slender, or absent. n = c. 25,
c. 29, 30. Circa 40 species, south-western USA
(Arizona), Mexico, Guatemala.
432. Tephroseris (Reichenb.) Reichenb.
Tephroseris (Reichenb.) Reichenb., Deutsche Bot. Fl. Sax.:
146 (1842); Cufodontis, Feddes Repert. Beih. 70: 1–266
(1933), rev.; Nordenstam, Opera Bot. 44: 43–45 (1978), new
comb.; Jeffrey & Chen, Kew Bull. 39: 267–285 (1984), reg.
rev.; Chen, Fl. Reip. Pop. Sin. 77, 1: 141–161 (1999), reg. rev.
220
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
435. Pippenalia McVaugh
Pippenalia McVaugh, Contr. Univ. Michigan Herb. 9: 470
(1972).
Perennial scapigerous herb. Leaves rosulate, longpetiolate, peltate, laciniate, palmately veined; leaf
base hairy. Capitula solitary, long-pedunculate, radiate, yellow, ecalyculate. Corolla lobes of disc floret
short. Anther base ecaudate, obtuse. Style branches
apically obtuse, shortly penicillate, dorsally papillate. Cypselae oblong-obovoid, ribbed, glabrous.
Pappus absent. n = 15, 30. One species, P. delphiniifolia (Rydb.) McVaugh, Mexico.
ser. 2, 10: 151–159 (1978), rev. (sub Cacalia).
Mesadenia Raf. (1832).
Perennial herbs. Leaves basal and cauline, petiolate, entire or somewhat lobed, ovate-cordate
to elliptic, palmately veined. Capitula few, small
and few-flowered, corymbose, discoid, ecalyculate. Receptacle conical. Corolla deeply lobed,
white to yellowish or pink. Anthers with radial
endothecium. Style branches truncate, penicillate,
with continuous stigmatic areas. Cypselae oblong,
ribbed, glabrous. Pappus bristles numerous, slender. n = 25, 26, 27, 28, 55. Eight species, central,
eastern and south-eastern USA.
436. Digitacalia Pippen
439. Robinsonecio T.M. Barkley & J.P. Janovec
Digitacalia Pippen, Contr. U.S. Natl Herb. 34: 378 (1968),
378–383, rev.; Robinson & Brettell, Phytologia 27: 407
(1974), notes; Turner, Phytologia 69: 150–159 (1990), rev.
Robinsonecio T.M. Barkley & J.P. Janovec, Sida 17: 77–81
(1996).
Erect perennial herbs. Leaves evenly distributed
on stem, petiolate, ± palmately lobed and veined.
Capitula several, corymbose, discoid, calyculate.
Corolla white, cream or purplish, deeply lobed.
Anthers ecaudate; endothecium polarized. Style
branches truncate to obtuse with continuous
stigmatic areas. Cypselae narrowly cylindrical,
oblong, ribbed, glabrous or pubescent. Pappus
bristles numerous, slender. n = 30. Five species,
Mexico.
Perennial herbs with a thick rhizome. Stem subscapose. Leaves rosulate, petiolate, ovate-lanceolate
or spathulate, entire with denticulate margins.
Capitula solitary or up to 8, corymbose, radiate,
yellow-flowered. Involucral bracts biseriate, with
a distinct or reduced calyculus. Disc florets perfect.
Anther collars cylindrical. Style branches with
continuous stigmatic areas. Cypselae ellipticoblong, glabrous or appressed-hirsute. Pappus
bristles numerous, slender. Pollen helianthoid.
n = 30. Two species, Mexico, Guatemala.
437. Rugelia Shuttlew. ex Chapm.
440. Roldana La Llave & Lex.
Rugelia Shuttlew. ex Chapm., Fl. South. U.S.: 246 (1860);
Pippen, N. Amer. Fl. ser. 2, 10: 153–154 (1978), rev. (as
Cacalia rugelia).
Roldana La Llave & Lex., Nov. Veg. Descr. 2: 10 (1825);
Robinson & Brettell, Phytologia 27: 408–424 (1974), review;
Turner, Phytologia 80: 276–279 (1996), new taxa; Panero &
Villaseñor, Brittonia 48: 79–90 (1996), new taxa; Rydberg,
Bull. Torrey Bot. Club 51: 376–377 (1924), rev. (sub Pericalia); Pippen, Contr. U.S. Natl Herb. 34: 410–415 (1968),
rev. (sub Pericalia). Revision ongoing (A.M. Funston).
Pericalia Cass. (1827).
Perennial rhizomatous herb, pubescent in most
parts. Radical leaves large, petiolate, ovate, denticulate, pinnately veined; cauline leaves smaller,
subsessile, entire, elliptic-ovate, palmately veined.
Capitula few to several, long-pedunculate, cymose,
many-flowered, discoid, calyculate; florets white
or ochroleucous. Anthers ecaudate, basally obtuse.
Style branches apically truncate and penicillate.
Cypselae narrowly cylindrical-oblong, striate,
glabrous. Pappus bristles numerous, rather rigid,
scabrid. n = 28. One species, R. nudicaulis
Shuttlew. ex Chapm., southern USA.
438. Arnoglossum Raf.
Arnoglossum Raf., Fl. Ludov.: 64 (1817); Vuilleumier,
J. Arnold Arbor. 50: 104–123 (1969), rev., notes; Robinson,
Phytologia 28: 294–295 (1974), rev.; Pippen, N. Amer. Fl.
Herbs, subshrubs or treelets, mostly with a hairy
tuberous caudex and fleshy roots; stems with
large pith. Leaves alternate, petiolate, often
peltate, palmately or sometimes pinnately veined,
rounded-ovate, entire and dentate to lobate or
pinnatisect. Capitula several to numerous, paniculate to corymbose, radiate, disciform or discoid,
yellow-, white- or greenish-flowered. Involucre
cylindrical to narrowly campanulate, calyculate.
Anthers with transitional endothecium; filament
collars cylindrical. Style branches with continuous
stigmatic areas. Cypselae elliptic-obovoid, ribbed,
glabrous or pubescent. Pappus bristles numerous,
Compositae
slender. n = 30, 60. Circa 65 species, south-western
USA (Arizona), Mexico to Panama.
441. Nelsonianthus H. Rob. & Brettell
Nelsonianthus H. Rob. & Brettell, Phytologia 27: 54 (1973).
Epiphytes or lax suffrutescent herbs. Stems
abruptly contracted below synflorescence. Leaves
clustered towards stem ends, petiolate, 3–5-veined
from the base, entire with serrulate margins.
Capitula few to several, radiate or discoid, yellowflowered, calyculate. Anthers sagittate to shortly
caudate; endothecium polarized. Style branches
obtuse; stigmatic areas continuous except basally.
Cypselae oblong, ribbed, glabrous. Pappus bristles
numerous, slender. Two or three species, Mexico,
Guatemala.
442. Telanthophora H. Rob. & Brettell
Telanthophora H. Rob. & Brettell, Phytologia 27: 424 (1974).
Treelets, shrubs or shrublets. Leaves alternate,
mostly terminal on abruptly contracted stem
ends, petiolate, persistent, entire or dentate-lobed,
pinnately veined. Capitula several or numerous,
small, paniculate-corymbose, radiate or discoid,
± distinctly calyculate, yellow-flowered. Anthers
with radial endothecium; filament collar cylindrical. Style branches obtuse with continuous
stigmatic areas. Cypselae elliptic-oblong, ribbed,
mostly glabrous. Pappus bristles numerous, slender. Pollen helianthoid. n = 30, c. 38, c. 65. Twelve
to 14 species, Panama to Mexico.
443. Villasenoria B.L. Clark
Villasenoria B.L. Clark, Sida 18: 631–634 (1999).
Tall perennial suffruticose herb. Stem simple,
erect, tapering to synflorescence. Leaves cauline,
petiolate, large, pinnately compound. Capitula numerous in elongate paniculate-racemose clusters,
radiate, yellow-flowered. Involucre cylindrical
or narrowly campanulate, biseriate, calyculate.
Anthers with radial endothecium, filament collar
basally somewhat enlarged. Style branches apically
truncate or subconical with short sweeping hairs,
stigmatic areas continuous. Cypselae oblongcylindrical, glabrous. Pappus bristles numerous,
slender. One species, V. orcuttii (Greenm.) B.L.
Clark, Mexico.
444. Pittocaulon H. Rob. & Brettell
Pittocaulon H. Rob. & Brettell, Phytologia 26: 451 (1973).
221
Erect shrubs or small trees, resiniferous, subsucculent. Leaves cauline, mostly assembled terminally
at branch ends, petiolate, often deciduous before
anthesis, palmately lobed and veined. Capitula several, corymbose, radiate, yellow-flowered, calyculate. Anthers with radial endothecium; filament collar uniformly cylindrical. Style branches with continuous stigmatic areas. Cypselae elliptic-oblong,
ribbed, glabrous. Pappus bristles numerous, slender. n = 30. Six species, Mexico.
445. Barkleyanthus H. Rob. & Brettell
Barkleyanthus H. Rob. & Brettell, Phytologia 27: 407 (1974).
Erect branching glabrous shrub, with a weak taproot and many lateral roots; stems with chambered
pith. Leaves cauline, assembled terminally on
branches, subsessile, entire, lanceolate or linearlanceolate, pinnately veined. Capitula several,
corymbose, radiate, yellow-flowered, ecalyculate.
Anthers with radial endothecium. Style branches
with continuous stigmatic areas. Cypselae narrowly elliptic-oblong, ribbed, sparsely pubescent.
n = 30. One species, B. salicifolius (Kunth) H. Rob.
& Brettell, south-western USA, Mexico, Central
America.
446. Yermo R.D. Dorn
Yermo R.D. Dorn, Madroño 38: 199 (1991).
Perennial herb with a thick taproot. Leaves
basal and cauline, entire, petiolate, lanceolate to
elliptic-obovate, 3-nerved. Capitula numerous,
corymbosely crowded, discoid. Involucre with
5 phyllaries, yellow, ecalyculate. Corolla deeply
lobed, yellow. Anthers auriculate. Style branches
obtuse to truncate, penicillate, with continuous stigmatic areas. Cypselae elliptic-oblong,
c. 10-veined, mostly puberulous. Pappus bristles numerous, slender, caducous. One species,
Y. xanthocephalus R.D. Dorn, western USA.
447. Rainiera Greene
Rainiera Greene, Pittonia 3: 291 (1898); Strother, N. Amer.
Fl. ser. 2, 10: 162–163 (1978), rev.
Perennial herb with a short rhizome. Leaves basal
and cauline, sessile but tapering into a petioliform
base, entire, broadly oblanceolate, upper leaves
gradually smaller, pinnately veined. Capitula
several in elongated thyrsoid panicles, narrow and
few-flowered, discoid. Corolla lobes lanceolate,
pale yellowish. Anther base obtuse or minutely
222
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
sagittate. Style branches truncate, papillate. Cypselae cylindrical-oblong, many-nerved, glabrous.
Pappus bristles numerous, slender. One species,
R. stricta (Greene) Greene, north-western USA.
448. Cacaliopsis A. Gray
Cacaliopsis A. Gray, Proc. Amer. Acad. Arts 19: 50 (1883);
Nordenstam, Opera Bot. 44: 21–22 (1978), notes; Strother,
N. Amer. Fl. ser. 2, 10: 160–161 (1978), rev.
Perennial rhizomatous herb. Leaves mostly basal,
petiolate, palmately lobed and veined. Capitula
solitary or few, corymbose to racemose or subumbellate, discoid, yellow- to orange-flowered,
calyculate. Corolla lobes short. Anthers ecaudate. Style branches apically obtuse to rounded
to subconical, dorsally papillate, stigmatic
areas discrete but apically confluent. Cypselae
cylindrical-oblong, many-nerved, glabrous. Pappus bristles numerous, slender. n = 30. One
species, C. nardosmia (A. Gray) A. Gray, western
North America.
449. Luina Benth.
Luina Benth., Hooker’s Icon. Pl. 12: 36, t. 1139 (1873);
Nordenstam, Opera Bot. 44: 21–22 (1978), notes; Strother,
N. Amer. Fl. ser. 2, 10: 161–162 (1978), rev.
Perennial tomentose or lanate herbs or subshrubs.
Leaves cauline, sessile or petiolate, entire, lanceolate or oblanceolate to elliptic-ovate. Capitula
several, corymbose to racemose, discoid, yellow- or
cream-flowered. Anthers shortly caudate; filament
collar narrow, elongate. Style branches obtuse or
subtruncate, dorsally finely papillate, stigmatic
areas discrete except towards the apex. Cypselae
narrowly oblong to fusiform, nerved, glabrous
or pubescent. Pappus bristles numerous, slender.
n = 30. Two species, western North America.
papillate-puberulous. Cypselae oblong to narrowly
elliptic-obovate or turbinate, with glandular ribs,
otherwise glabrous or pubescent. Pappus bristles
numerous, slender or scale-like, sometimes lacking. n = 30, 31, 45, 60, 62, 90. Ten species, western
North America.
451. Lepidospartum (A. Gray) A. Gray
Lepidospartum (A. Gray) A. Gray, Proc. Amer. Acad. Arts
19: 50 (1883); Strother, N. Amer. Fl. ser. 2, 10: 171–173
(1978), reg. rev.
Broom-like shrubs or treelets. Leaves sessile,
linear-oblanceolate to filiform or scale-like, entire.
Capitula solitary to several or many, racemose,
axillary or terminal, discoid, small and mostly
few-flowered, yellow-flowered. Involucral bracts
pluriseriate. Anthers sagittate or caudate. Style
branches slender, papillate-puberulous, apically
with a conical penicillate appendage. Cypselae
narrowly elliptic-oblong or fusiform, glabrous or
pubescent. Pappus bristles numerous, slender.
n = 30, c. 45. Three species, south-western USA,
Mexico.
452. Paracalia Cuatrec.
Paracalia Cuatrec., Brittonia 12: 183 (1960).
Scandent shrublets. Leaves cauline, petiolate,
elliptic-ovate, entire. Capitula several, corymbose,
small and few-flowered, discoid, ecalyculate.
Corolla white, deeply lobed. Anther base acute;
endothecium polarized. Style branches apically
with a pointed tuft of hairs; stylopodium distinct. Cypselae oblong, glabrous. Pappus bristles
numerous, slender. Two species, Bolivia, Peru.
453. Paragynoxys (Cuatrec.) Cuatrec.
450. Tetradymia DC.
Tetradymia DC., Prodr. 6: 440 (1838); Strother, Brittonia 26:
177–202 (1974), rev.; N. Amer. Fl. ser. 2, 10: 163–170 (1978),
rev.
Shrubs, spinescent or unarmed, often canescent.
Leaves cauline, sessile, alternate, sometimes fascicled, small and narrow, filiform or linear-lanceolate
to oblanceolate (some transformed into spines).
Capitula solitary or few together, axillary or corymbose, small and few-flowered, discoid, ecalyculate,
cream- or yellow-flowered. Anthers sagittate, ecaudate. Style branches truncate to obtusely conical,
Paragynoxys (Cuatrec.) Cuatrec., Brittonia 8: 153 (1955);
Correa, Brittonia 55: 157–168 (2003), rev.
Erect single-stemmed or little-branched trees
or treelets. Leaves alternate, crowded towards
branch ends, petiolate, large, entire, elliptic-ovate
or obovate, coriaceous. Capitula numerous, fewflowered, paniculate, discoid, calyculate. Corolla
white, deeply lobed. Anthers sagittate or shortly
caudate. Style branches apically obtuse-conical,
papillate. Cypselae oblong, ribbed, glabrous.
Pappus bristles numerous, slender. n = c. 40.
Twelve species, Andes of Colombia and Venezuela.
Compositae
223
454. Gynoxys Cass.
Gynoxys Cass., Dict. Sci. Nat. 48: 455 (1827).
Shrubs or trees. Leaves opposite, petiolate or
subsessile, entire, elliptic-oblong to ovate or
obovate, coriaceous, densely tomentose beneath.
Capitula few to many, corymbose-paniculate, radiate or discoid, yellow-flowered. Anthers ecaudate.
Style branches with an apical conical-pointed
hair pencil, stigmatic areas confluent. Cypselae
elliptic-oblong, ribbed, glabrous or pubescent.
Pappus bristles numerous, slender. n = c. 12, c. 36,
c. 40, 40. Circa 120 species, Andes from Argentina
and Bolivia to Venezuela.
455. Nordenstamia Lundin
Nordenstamia Lundin, Comp. Newslett. 44: 15 (2006);
Nordenstam, Comp. Newslett. 44: 19–23 (2006); new comb.
Senecio L. sect. Praegynoxys Cuatrec. (1951).
Gynoxys Cass. sect. Praegynoxys (Cuatrec.) Cuatrec. (1955).
Aequatorium B. Nord. subg. Praegynoxys (Cuatrec.)
B. Nord. (1997).
Shrubs or trees (up to 18 m tall), pubescent
with branching, globular-echinate or substellate
trichomes especially on leaves and young parts.
Leaves alternate (rarely opposite). Capitula several,
paniculate-corymbose, radiate, yellow-flowered,
calyculate. Ray florets few (2–5). Anthers sagittate
or shortly caudate. Style branches with an apical
pointed appendage; stigmatic areas continuous
or barely separated. Cypselae elliptic-oblong,
8–10-ribbed, glabrous or glandular-puberulous.
Pappus bristles numerous, slender but often
dilated apically, white to fulvous. Circa 20 species,
Argentina, Bolivia, Peru, southern Ecuador.
456. Aequatorium B. Nord.
Fig. 56
Aequatorium B. Nord., Opera Bot. 44: 59 (1978); DíazPiedrahita & Cuatrecasas, Revista Acad. Colomb. Ci. 17, 67:
659–666 (1990), reg. rev.; Díaz-Piedrahita & Cuatrecasas,
Revista Acad. Colomb. Ci. 19, 73: 247–252 (1994), new taxa,
key; Nordenstam, Comp. Newslett. 31: 1–16 (1997), reg. rev.
Erect shrubs or trees, stellate-tomentose in young
parts. Leaves alternate, petiolate, entire, lanceolate
to elliptic-ovate, with margins often dentate or
denticulate, coriaceous; tomentum mainly on
lower side, in two layers; a white or greyish
inner layer overlain by a scurfy rusty-brownish
looser layer peeling off in patches; trichomes
peltate-stellate. Capitula several to numerous,
paniculate-corymbose, radiate, calyculate. Ray
Fig. 56. Compositae-Senecioneae. Aequatorium jamesonii.
A Habit. B Capitulum. C Ray floret. D Disc floret. E Corolla
of disc floret, opened. F Stamens. G Style branches of disc
floret. (Drawing by B. Nordenstam)
florets white or cream. Disc florets perfect; corolla
deeply lobed, white or pale yellowish. Anthers
ecaudate; endothecium polarized; filament collar
cylindrical. Style branches obtuse or subtruncate,
apically and dorsally papillate, stigmatic areas
continuous. Cypselae oblong, ribbed, glabrous.
Pappus bristles numerous, somewhat coarse,
persistent, white or brownish to fulvous. Circa 12
species, Ecuador, Colombia, south Venezuela.
457. Acrisione B. Nord.
Acrisione B. Nord., Bot. Jahrb. Syst. 107: 582 (1985).
Shrubs or small trees with large soft pith. Leaves
alternate, cauline, petiolate, entire with denticulate
margins, elliptic-ovate to obovate. Capitula several
to many, corymbose, radiate, ecalyculate, yellowflowered. Anthers sagittate, becoming exserted.
Style branches of disc florets with ± continuous
stigmatic areas, apically obtuse, papillate. Cypselae elliptic-oblong, ribbed, puberulous. Pappus
bristles numerous, slender. Two species, Chile.
458. Brachyglottis J.R. Forst. & G. Forst.
Brachyglottis J.R. Forst. & G. Forst., Char. Gen. Pl.: 91
(1776); Allan, Fl. N. Z. 1: 736–758 (1961), rev. (mostly as
224
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Senecio); Drury, N. Z. J. Bot. 11: 731–784 (1977), key, rev.
(as Senecio); Nordenstam, Opera Bot. 44: 25–31 (1978),
emend., new comb.; Webb, N. Z. J. Bot. 25: 148–152 (1987),
notes; Wagstaff & Breitwieser, Syst. Bot. 29: 1003–1010
(2004), phylog.
Shrubs or trees or perennial herbs with a ± thick
rhizome. Leaves cauline or rosulate, sessile or
petiolate, entire or dentate to lobed, pinnately
veined. Capitula solitary and scapose, or several
and corymbose, radiate or disciform, yellow-,
cream- or white-flowered. Anthers caudate or
sagittate. Style branches with ± continuous stigmatic areas, basally somewhat discrete. Cypselae
oblong, ribbed, glabrous or pubescent. Pappus
bristles numerous, slender, usually persistent,
white. n = 30. Circa 30 species, New Zealand.
459. Centropappus Hook. f.
Centropappus Hook. f., London J. Bot. (Hooker) 6: 124
(1847).
Glabrous shrub or small tree. Leaves alternate,
bunched terminally, sessile, linear, one-nerved,
entire, obtuse, with glossy exudate. Capitula in
short terminal corymbs, radiate, yellow-flowered.
Involucral bracts uniseriate. Corolla lobes of disc
florets deeply lobed, without midvein. Anthers
basally rounded, ecaudate. Style branches apically
truncate, papillate; stigmatic areas continuous.
Cypselae oblong, glabrous, smooth. Pappus bristles apically subplumose. One species, C. brunonis
Hook. f., Tasmania.
460. Urostemon B. Nord.
Urostemon B. Nord., Opera Bot. 44: 31 (1978); Drury, N. Z.
J. Bot. 13: 769–780 (1975, as Senecio), morph.
Shrub or treelet, sometimes epiphytic, glabrous except for glands and T-shaped trichomes on young
parts. Leaves cauline, petiolate, somewhat sinuatedentate to subentire. Capitula several, laxly corymbose, radiate, ecalyculate. Ray florets white. Disc
florets perfect; corolla yellow, deeply lobed. Anthers
caudate with long papillate tails. Style branches
apically obtuse to convex, papillate also dorsally,
stigmatic areas continuous. Cypselae narrowly oblong, glabrous, striate with 10 veins and 10 resin
canals. Pappus bristles numerous, stiff and rather
coarse, dirty white, persistent. n = 30. One species,
U. kirkii (Hook. f. ex Kirk) B. Nord., New Zealand
(North Island).
461. Haastia Hook. f.
Haastia Hook. f., Handb. N. Z. Flora: 156 (1864); Allan, Fl.
N. Z. 1: 675–676 (1961), rev.; Bailes, Q. Bull. Alp. Gard. Soc.
55: 118–122 (1987), notes.
Compact shrub forming dense cushions or mats, or
procumbent to suberect subshrubs. Leaves densely
imbricate, spathulate, apically crenulated, covered
with long hairs, or alternate, patent to recurved,
oblong-obovate, tomentose. Capitula solitary, sessile, terminal, heterogamous and disciform. Involucral bracts subuniseriate to biseriate, glabrous
or hairy. Marginal florets pistillate with exserted
style branches. Disc florets hermaphroditic, tubular, white. Anthers ecaudate. Style branches linear
with apical conical tuft of obtuse hairs, or slender and dorsally papillate-hairy; stigmatic areas
separated or continuous. Cypselae linear-oblong,
compressed or subterete, glabrous. Pappus bristles
numerous, coarse and basally flattened, or slender
throughout, persistent. n = 30. Three species, New
Zealand (not monophyletic, perhaps better treated
as two separate genera).
462. Bedfordia DC.
Bedfordia DC. in Guill., Arch. Bot. (Paris) 2: 332 (1833);
Gray, Muelleria 3: 64–66 (1974), rev.; Orchard, Muelleria
19: 81–94 (2004), rev.
Erect shrubs or treelets. Leaves sessile, alternate,
entire, linear-lanceolate to elliptic, margins serrulate to denticulate, lower side tomentose. Capitula
solitary and axillary or several and corymbose, discoid, ecalyculate, yellow-flowered. Corolla deeply
lobed with lanceolate lobes. Anther base shortly
auriculate; filament collar short. Style branches obtuse, dorsally papillose. Cypselae oblong, ribbed,
glabrous. Pappus bristles numerous, slender.
n = 30. Two species, south-eastern Australia,
Tasmania.
463. Traversia Hook. f.
Traversia Hook. f., Handb. N. Z. Fl.: 163 (1864); Allan, Fl. N.
Z. 1: 758 (1961), rev.
Erect branching glutinous shrub. Leaves cauline,
sessile, elliptic to obovate, entire with serrate margins, pinnately to reticulately veined. Capitula few
to several, corymbose, discoid, calyculate. Corolla
white, deeply lobed. Anther base ecaudate, auriculate. Style branches apically truncate and papillose.
Cypselae elliptic-oblong, ribbed, glabrous. Pappus
bristles coarse and rigid, basally connate, scabrid,
Compositae
unequal, off-white, persistent. n = 30. One species,
T. baccharoides Hook. f., New Zealand.
Pappus bristles numerous, slender but apically
clavate. n = 36. Fourteen species, New Guinea.
464. Dolichoglottis B. Nord.
467. Brachionostylum Mattf.
Dolichoglottis B. Nord., Opera Bot. 44: 33 (1978); Allan, Fl.
N. Z. 1: 741–742 (1961, as Senecio), rev.
Brachionostylum Mattf., Nova Guinea 14: 527 (1932).
Perennial herbs with a thick woolly caudex. Stem
erect, silky-woolly and glandular. Leaves basal and
cauline, sessile, lanceolate or linear-lanceolate,
parallel-veined, entire with margins sometimes
toothed. Capitula few to several, corymbose,
radiate. Involucral bracts biseriate. Ray florets
yellow, white or pinkish. Disc florets perfect,
yellow, 5-, occasionally 4-lobed. Anthers ecaudate.
Style branches obtuse or subtruncate, stigmatic
areas continuous or only basally discrete. Cypselae
oblong, ribbed, with mucilaginous twin hairs.
Pappus bristles stiff and rather coarse, persistent.
n = 30. Two species, New Zealand.
465. Capelio B. Nord.
Capelio B. Nord., Comp. Newslett. 38: 72 (2002), 71–78, rev.,
key, new sp.; Comp. Newslett. 39: 48–51 (2003), nomencl.
Celmisia Cass. (1817), nom. rej., non Cass. (1825), nom.
cons.
Alciope DC. in Lindley (1836) & DC. (1836), nom. illegit.
Perennial woolly herbs. Leaves cauline, petiolate,
elliptic-ovate, entire, densely woolly beneath.
Capitula solitary or few to several, corymbose,
long-pedunculate, radiate, yellow-flowered. Involucre bi- to triseriate, ecalyculate. Anthers
ecaudate; endothecium polarized with few thickenings. Style branches apically obtuse, papillate;
stigmatic areas continuous. Cypselae oblong,
pubescent. Pappus bristles numerous, slender.
Three species, southern Africa.
225
Dioecious shrub. Leaves alternate, petiolate,
elliptic-ovate, entire with serrulate or denticulate
margins. Capitula several, axillary, corymbosely
paniculate, radiate, calyculate. Ray florets female
or sterile, yellow; disc florets functionally male.
Stamens with cylindrical uniform filament collar.
Style branches apically rounded, glabrous. Cypselae narrowly oblong, ribbed, glabrous. Pappus
bristles numerous, slender, caducous. One species,
B. pullei Mattf., New Guinea.
468. Pladaroxylon (Endl.) Hook. f.
Pladaroxylon (Endl.) Hook. f., Hooker’s Icon. Pl. 11: 47,
t. 1055 (1870); Mabberley, Kew Bull. 30: 413–420 (1975, as
Senecio), morph., notes; Drury, N. Z. J. Bot. 13: 769–780
(1975, as Senecio), morph.; Nordenstam, Opera Bot. 44: 36–
38 (1978), re-establ.
Erect tree to 5 m tall, upwards branching dichotomously or trichotomously. Leaves shortly petiolate,
assembled towards branch ends, elliptic-obovate,
entire with somewhat dentate margins. Capitula
numerous, corymbose in terminal synflorescences,
radiate, narrowly campanulate, calyculate, whiteflowered. Disc florets perfect; corolla deeply lobed,
slightly zygomorphic. Anthers ecaudate, with subcylindrical filament collar. Style branches linear,
with narrowly separated stigmatic areas, apically
truncate, papillate. Cypselae oblong, ribbed, glabrous. Pappus bristles numerous, slender, persistent. One species, P. leucadendron (G. Forst.) Hook.
f., St. Helena.
469. Lachanodes DC.
466. Papuacalia Veldk.
Papuacalia Veldk., Blumea 36: 168 (1991).
Erect shrubs or trees, often pubescent and
glandular. Leaves sessile or petiolate, entire, linearlanceolate to elliptic-oblong. Capitula several to
numerous in lateral synflorescences, disciform or
discoid, with creamy-white or pinkish-purple florets. Ray florets, if present, with reduced ray. Disc
florets perfect or functionally male. Anthers ecaudate with polarized endothecium. Style branches
truncate-obtuse, apically papillate, sometimes also
dorsally. Cypselae oblong, glabrous or pubescent.
Lachanodes DC. in Guill., Arch. Bot. (Paris) 2: 332 (1833);
Mabberley, Kew Bull. 30: 413–420 (1975, as Senecio),
morph., notes; Drury, N. Z. J. Bot. 13: 769–780 (1975,
as Senecio), morph.; Nordenstam, Opera Bot. 44: 36–38
(1978), re-establ., nomencl.
Branching tree to 5 m tall. Leaves cauline, large,
petiolate, oblong-obovate to elliptic-ovate, entire with denticulate margins, pinnately veined.
Capitula several, paniculate in lateral pendulous
synflorescences, discoid, few-flowered. Corolla
white, deeply lobed. Stamens with narrowly
cylindrical filament collar. Style branches long
226
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
and much exserted, apically obtuse, papillate also
dorsally towards the apex, with mostly continuous
stigmatic areas. Cypselae oblong, glabrous, ribbed.
Pappus bristles pluriseriate, slender, persistent.
One species, L. arborea (Roxb.) B. Nord., St.
Helena.
470. Scrobicaria Cass.
Scrobicaria Cass., Dict. Sci. Nat. 48: 456 (1827); Nordenstam, Opera Bot. 44: 63–64 (1978), re-establ.
Small erect shrubs. Leaves cauline, opposite,
shortly petiolate or subsessile, entire with dentate
margins. Capitula few to several, rather densely
corymbose, discoid, calyculate, yellow-flowered.
Involucre ± biseriate. Anthers caudate; endothecial
tissue transitional; filament collar subcylindrical
or slightly balusterform. Style branches apically
obtuse to conical, papillate; stigmatic areas continuous. Cypselae oblong, ribbed, glabrous. Pappus
bristles numerous, slender, white. Two species,
Colombia, Venezuela.
471. Shafera Greenm.
Shafera Greenm., Publ. Field Mus. Nat. Hist., Bot. Ser. 2:
327 (1912).
Erect or somewhat scandent tomentose shrub or
subshrub. Leaves cauline, petiolate, entire, ellipticovate. Capitula several, corymbose, discoid,
yellow-flowered, calyculate. Involucre pluriseriate
with imbricate phyllaries. Anthers sagittate. Style
branches apically obtuse. Cypselae elliptic-oblong,
pubescent with twin hairs; carpopodium distinct,
annular. Pappus bristles numerous, slender. One
species, S. platyphylla Greenm., Cuba.
472. Mattfeldia Urb.
Mattfeldia Urb., Ark. Bot. 23, A(11): 90 (1931).
Scandent shrub. Leaves cauline, petiolate, entire, elliptic-ovate, three-nerved. Capitula few
or several, paniculately corymbose, small and
few-flowered, radiate, yellow-flowered. Ray florets
bilabiate with a short lamina and two small ventral
lobes. Disc florets perfect. Style branches truncate,
penicillate. Cypselae narrowly elliptic-oblong,
glabrous. Pappus bristles numerous, slender. One
species, M. triplinervis Urb., Hispaniola.
473. Ekmaniopappus A. Borhidi
Ekmaniopappus A. Borhidi, Acta Bot. Hung. 37: 109 (1992).
Scandent shrubs or shrublets. Leaves opposite or
alternate, petiolate, ± ovate with serrate to dentate
margins, tomentose beneath. Capitula several
to many, paniculate in axillary and terminal
synflorescences, small and few-flowered, radiate,
yellow-flowered. Involucre cylindrical, calyculate.
Anthers sagittate or auriculate. Style branches
apically truncate and shortly penicillate. Cypselae
elliptic-oblong, 8-ribbed, glabrous. Pappus bristles
numerous, slender. Two species, Hispaniola.
474. Herodotia Urb. & Ekm.
Herodotia Urb. & Ekm., Ark. Bot. 20 A, 5: 63 (1926).
Scandent shrub. Leaves opposite, petiolate, entire,
elliptic-ovate, glabrous, with sinuate-lobulate margins. Capitula several, paniculately corymbose, discoid, 2–3-flowered, yellow. Involucre biseriate with
8 phyllaries, ecalyculate. Anthers ecaudate with obtuse base. Style branches dorsally papillate, apically
truncate, penicillate. Cypselae elliptic-oblong, subglabrous with scattered small hairs, with thick carpopodium. Pappus bristles numerous, slender. One
species, H. haitiensis Urb. & Ekm., Hispaniola.
475. Leonis B. Nord.
Leonis B. Nord., Comp. Newslett. 44: 55 (2006).
Almost glabrous scandent shrublet (vine). Leaves
alternate, ovate to elliptic, entire or shallowly trilobate, 3-nerved, petiolate. Capitula in short axillary corymbs along the stems and branches, radiate, calyculate. Involucral bracts uniseriate. Resin
ducts present in involucre, corollas, styles etc. Anthers sagittate. Style branches of disc florets obtuse
with few short sweeping hairs; stigmatic areas continuous. Cypselae glabrous or sparsely setose or
papillate, 10-ribbed. Pappus bristles pluriseriate,
basally connate, persistent. One species, L. trineura
(Griseb.) B. Nord., Cuba, Hispaniola.
476. Nesampelos B. Nord.
Nesampelos B. Nord., Comp. Newslett. 44: 58 (2006).
Lianas with tomentose glabrescent stems. Leaves
alternate, petiolate, elliptic-ovate, entire with dentate to denticulate margins, coriaceous, upper
side glabrescent and glossy, lower side brownishtomentose. Capitula in terminal or axillary lateral
corymbs, radiate, calyculate. Involucre tomentose.
Ray florets yellow, creamy or white. Anthers
basally obtuse, ecaudate. Style branches apically
subtruncate with short sweeping hairs; stigmatic
Compositae
areas separated. Cypselae 10-striate, ciliate in
upper half. Pappus bristles pluriseriate, basally
connate, persistent. Three species, Hispaniola.
477. Zemisia B. Nord.
Zemisia B. Nord., Comp. Newslett. 44: 71 (2006).
Erect, diffuse or divaricate shrub, tomentose
throughout, but leaves glabrescent adaxially.
Leaves alternate, petiolate, ovate to lanceolate, with
denticulate to subentire margins, midribbed and
pinnately veined. Capitula numerous in terminal
corymbose synflorescence, radiate, calyculate.
Involucral bracts uniseriate, 8–13. Receptacle
alveolate. Ray florets 3–6, white. Disc florets
hermaphrodite; corolla cream; lobes distinctly
midlined, apically thickened. Anthers distinctly
caudate. Style branches subtruncate with few short
pili abaxially; stigmatic areas separated. Cypsela
8-nerved, villous-papillate with white obtuse
mucilaginous hairs; carpopodium distinct. Pappus
bristles numerous, c. biseriate, persistent. One
species, Z. discolor (Sw.) B. Nord., Jamaica.
478. Antillanthus B. Nord.
Antillanthus B. Nord., Comp. Newslett. 44: 51 (2006).
Erect or scandent shrubs. Leaves cauline, linear
or lanceolate to oblong-ovate, entire, glabrous or
pubescent. Capitula few to several, corymbose or
paniculate, radiate or discoid, white-, cream- or
yellow-flowered. Receptacle alveolate. Resin canals
often present in corolla and style. Anthers sagittate
or caudate. Style branches obtuse with rather few
and short sweeping hairs; stigmatic areas continuous. Cypselae elliptic-oblong, 10-ribbed, glabrous
or pubescent. Pappus bristles numerous, slender,
basally connate. Seventeen species, Cuba.
479. Elekmania B. Nord.
Elekmania B. Nord., Comp. Newslett. 44: 66 (2006).
Erect or scandent shrubs or subshrubs, sometimes
strongly resiniferous. Leaves alternate, petiolate,
entire or with dentate-lobulate margins, usually
tomentose below, glabrous above. Capitula several to numerous in lateral or terminal corymbose
or cymose synflorescences, radiate or sometimes
discoid, yellow-flowered. Corolla lobes midveined,
apically papillate. Anthers ecaudate or shortly caudate; apical appendage lanceolate. Style branches
truncate or somewhat obtuse with short sweeping
227
hairs; stigmatic areas separated. Cypselae glabrous
or hirsute, 8–10-ribbed. Pappus bristles numerous,
slender. Circa ten species, Hispaniola.
480. Herreranthus B. Nord.
Herreranthus B. Nord., Comp. Newslett. 44: 62 (2006).
Shrub or treelet; stem with large pith; young
branches tomentose, ± glabrescent. Leaves
alternate, petiolate, entire, elliptic-oblong to
lanceolate or oblanceolate, glabrous above, densely
tomentose beneath, midribbed. Capitula terminal,
cymose or corymbose, discoid. Involucral bracts
biseriate. Anthers shortly or distinctly caudate.
Style branches apically shortly papillate; stigmatic
areas continuous and extending to the dorsal
side. Cypselae 10-ribbed, hairy. Pappus bristles
pluriseriate, persistent. One species, H. rivalis
(Greenm.) B. Nord., Cuba.
481. Oldfeltia B. Nord. & Lundin
Oldfeltia B. Nord. & Lundin, Comp. Newslett. 38: 66 (2002).
Erect glabrous shrub. Leaves cauline, shortly
petiolate, elliptic-lanceolate, large, entire with
serrate to denticulate margins. Capitula numerous,
corymbosely paniculate, discoid, yellow-flowered.
Involucre cylindrical, with 5 connate bracts, pale
yellowish-green, calyculus bracts few. Receptacle
convex, nude. Anthers basally sagittate to shortly
caudate. Style branches apically triangular-conical
with many short clavate sweeping hairs; stigmatic areas discrete. Cypselae oblong, glabrous.
Pappus bristles numerous, slender. One species,
O. polyphlebia (Griseb.) B. Nord. & Lundin,
Cuba.
482. Odontocline B. Nord.
Odontocline B. Nord., Opera Bot. 44: 23 (1978).
Erect or scandent shrubs. Leaves alternate, petiolate, entire or pinnatifid, margins sometimes
serrate. Capitula several to many, corymbose,
radiate, yellow- or orange-flowered, fragrant.
Involucre narrowly campanulate to subcylindrical,
uniseriate, minutely calyculate. Receptacle denticulate. Anthers caudate. Style branches apically
rounded-obtuse with subterminal sweeping hairs;
stigmatic areas continuous. Cypselae oblong,
ribbed, glabrous or pubescent. Pappus bristles
numerous, slender, persistent. n = 30. Six species,
Jamaica.
228
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
483. Lundinia B. Nord.
486. Jessea H. Rob. & Cuatrec.
Lundinia B. Nord., Comp. Newslett. 44: 64 (2006).
Jessea H. Rob. & Cuatrec., Novon 4: 49 (1994); Nordenstam,
Bot. Jahrb. Syst. 118: 147–152 (1996), rev.
Erect shrub or small tree. Leaves petiolate, lanceolate or oblong-lanceolate, entire with ± denticulate
margins. Capitula several, corymbose, radiate,
yellow-flowered. Receptacle denticulate. Anthers
caudate. Style branches apically truncate, often
with a minute short hair tuft; stigmatic areas
separated. Cypselae elliptic-oblong, pubescent.
Pappus bristles numerous, slender, short. One
species, L. plumbea (Griseb.) B. Nord., Cuba,
Hispaniola.
484. Jacmaia B. Nord.
Jacmaia B. Nord., Opera Bot. 44: 64 (1978).
Erect shrub. Leaves sessile or subsessile, lanceolate
or oblanceolate, with sinuate-dentate margins,
pinnatilobate towards the base. Capitula numerous, corymbose, radiate, yellow-flowered.
Involucre cylindrical or narrowly campanulate, tomentose, basally incrassate, uniseriate, calyculate.
Receptacle denticulate. Ray florets few and short.
Disc florets hermaphrodite; corolla tubular and
widening above, slightly zygomorphic. Anthers
shortly caudate; endothecial tissue transitional.
Style branches apically with an acuminate distinct
papillate appendage; stigmatic areas continuous.
Cypselae narrowly oblong, ribbed, setose apically, with distinct carpopodium. Pappus bristles
numerous, very slender and almost smooth,
persistent. One species, J. incana (Sw.) B. Nord.,
Jamaica.
485. Ignurbia B. Nord.
Ignurbia B. Nord., Willdenowia spec. vol. 31, 1: 464 (2006).
Erect perennial herb, shortly hairy or glandular
on stems, leaves and peduncles. Leaves alternate,
herbaceous, irregularly incised-lobed, petiolate.
Capitula numerous in terminal corymbs, discoid,
calyculate, orange-flowered. Corolla tubular,
gradually widening above. Anthers ecaudate;
apical appendage narrow; endothecial tissue radial
with short ± isodiametric cells; filament collar
long, uniformly wide, with larger cells along the
margins. Pollen grains minutely spinulose. Style
branches apically obtuse, papillate, without distinct sweeping hairs; stigmatic areas continuous.
Cypselae 10-ribbed, glabrous. Pappus bristles
pluriseriate, basally connate, persistent. Two
species, Hispaniola.
Erect shrubs or subshrubs, with large pith. Leaves
sessile or petiolate, large, oblong-ovate or ellipticlanceolate, entire or lobed to laciniate, with dentate
margins, closely arcuately pinnativeined; leaf base
± clasping. Capitula numerous, densely corymbosely paniculate, radiate, yellow-flowered. Involucre uniseriate, cylindrical or narrowly cup-shaped,
calyculate. Receptacle with scale-like projections.
Corolla of disc florets with lanceolate to narrowly
ovate lobes. Anthers ecaudate. Style branches apically rounded-obtuse with a central hair tuft; stigmatic areas separated. Cypselae oblong, ribbed,
glabrous or pubescent. Pappus bristles numerous,
slender, caducous. n = c. 50. Four species, Costa
Rica, Panama.
487. Lordhowea B. Nord.
Lordhowea B. Nord., Opera Bot. 44: 38 (1978); Drury, N. Z.
J. Bot. 13: 769–780 (1975, as Senecio), morph.
Erect glabrous shrub. Leaves cauline, alternate,
petiolate, elliptic-ovate, coarsely dentate or lobate.
Capitula many, corymbose, overtopped by leaves,
radiate with few and short pale yellow ray florets,
paucicalyculate. Receptacle denticulate. Disc
florets hermaphrodite, corolla lobes lanceolate.
Anthers ecaudate. Style branches apically conical
and acute, dorsally papillate especially towards the
apex, stigmatic areas separated. Cypselae narrowly
oblong, 10-ribbed, glabrous. Pappus bristles
numerous, slender, persistent. n = 19. One species,
L. insularis (Benth.) B. Nord., Lord Howe Isl.
488. Arrhenechthites Mattf.
Arrhenechthites Mattf., Bot. Jahrb. Syst. 69: 288 (1938);
Belcher, Ann. Missouri Bot. Gard. 43: 1–85 (1956), rev.
Shrubs or robust perennial herbs. Leaves cauline,
petiolate, elliptic-ovate, entire with dentate to
serrate margins. Capitula numerous, in terminal
cymose synflorescences, disciform. Involucre
cylindrical, biseriate to pluriseriate, paucicalyculate. Marginal florets few, female, tubular,
purplish. Disc florets few, perfect or functionally
male, white or cream-coloured, sometimes with
purplish tube, deeply lobed. Anthers with broadly
balusterform collars. Style branches short, apically
dilated and truncate, papillate. Cypselae oblong,
glabrous, with distinct carpopodium. Pappus
Compositae
bristles numerous, slender. n = c. 50. Six species,
Indonesia, New Guinea, south-eastern Australia.
489. Bethencourtia Choisy
Bethencourtia Choisy in Buch, Phys. Beschr. Canar. Ins.: 148
(1825); Nordentstam, Comp. Newslett. 44: 24–31 (2006), rev.
(as Canariothamnus)
Canariothamnus B. Nord. (2006).
Senecio sect. Bethencourtii DC. (1838).
Glabrous shrublets. Leaves alternate, sessile, linearoblanceolate, entire, few-toothed or lobed. Capitula small, corymbose, ecalyculate, yellow-flowered.
Involucral bracts 4–7. Ray florets 1–3, short. Disc
florets 3–5. Anthers caudate; endothecium radial;
filament collar balusterform. Style branches apically obtuse, with short papilliform sweeping hairs;
stigmatic areas confluent or barely discrete. Cypselae oblong, shortly hirsute. Pappus semi-persistent.
n = 10. Three species, Canary Islands.
490. Pericallis D. Don in Sweet
Pericallis D. Don in Sweet, Brit. Fl. Gard. ser. 2: pl. 228
(1834); Nordenstam, Opera Bot. 44: 15–21 (1978), re-establ.,
new comb.; Swenson & Manns, Taxon 52: 533–546 (2003),
phylog.; Swenson, Svensk Bot. Tidskr. 98: 193–206 (2004),
popular review.
Perennial herbs or subshrubs. Leaves cauline, petiolate, cordate to reniform or ovate, entire or dentate to lobed, palmately veined. Capitula several or
rarely solitary, or many, corymbose, radiate, ecalyculate. Florets white, pink, purple or blue. Anthers ecaudate; filament collar much dilated basally.
Style branches truncate to obtuse, stigmatic areas discrete. Cypselae terete or somewhat compressed, elliptic-oblong, ribbed, glabrous. Pappus
bristles numerous, sometimes absent in ray florets, caducous. Pollen helianthoid. n = 30. Fifteen
species, Canary Islands, Madeira, Azores (P. hybrida B. Nord., the florists’ so-called Cineraria, is
widely used in horticulture).
or rarely discoid, paucicalyculate, yellow-flowered.
Disc florets perfect or rarely functionally male.
Anthers slightly sagittate; filament collar balusterform. Style branches truncate to obtuse with short
sweeping hairs; stigmatic areas separated. Cypselae compressed with winged or thickened margins,
brown or blackish. Pappus bristles numerous, slender, caducous. n = 10, 20. Circa 35 species, Africa,
Madagascar, Arabia.
492. Bolandia Cron
Bolandia Cron, Novon 16:224 (2006).
Annual or perennial herbs. Leaves alternate,
petiolate, entire or lobed to dissected, araneose
or tomentose, glabrescent. Capitula solitary on
bracteate peduncles, radiate, ecalyculate, yellowflowered. Involucral bracts uniseriate. Anthers
minutely sagittate. Style branches apically with
a central hair pencil; stigmatic areas separated.
Cypselae dimorphic: outer abaxially convex and
white-villous, adaxially midribbed and glabrous;
inner weakly ribbed, uniformly hairy. Pappus
bristles pluriseriate, caducous. Two species, South
Africa, Lesotho.
493. Mesogramma DC.
Mesogramma DC., Prodr. 6: 304 (1838); Nordenstam &
Pelser, Comp. Newslett. 42: 74–88 (2005), re-establ.
Glabrous annual herb. Leaves alternate, petiolate,
ovate-lanceolate, grossly dentate or lobate. Capitula solitary or few to several, laxly corymbose,
radiate, yellow-flowered, minutely calyculate.
Involucral bracts biseriate, inner phyllaries with
two distinct dark resin ducts. Corolla of disc florets
black-veined. Anthers sagittate. Style branches
truncate; stigmatic areas separated. Cypselae
subtriquetrous, black, with white papilliform hairs
apically and basally and in three longitudinal lines.
Pappus bristles white, persistent. One species,
M. apiifolium DC., southern Africa.
491. Cineraria L.
494. Stilpnogyne DC.
Cineraria L., Sp. Pl. ed. 2: 1242 (1763) & Gen. Pl. ed. 6: 426
(1764); Hilliard, Compositae in Natal: 372–387 (1977), reg.
rev. Genus under revision (G. Cron).
Stilpnogyne DC., Prodr. 6: 293 (1838).
Perennial herbs or subshrubs. Leaves cauline or
radical, petiolate, cordate or rounded to ovate, entire or dentate, lobed or pinnatilobate or pinnatisect, glabrous or tomentose, palmately veined. Capitula few or several to many, corymbose, radiate
229
Small annual glabrous herb. Leaves alternate, petiolate, ovate-cordate, entire with distally dentate or
lobulate margins. Capitula few to several, corymbose, disciform, yellow-flowered. Involucral bracts
uniseriate, basally connate, ecalyculate. Marginal
florets usually 3, female, tubular or with a short
limb. Disc florets few, hermaphrodite. Anthers
230
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
ecaudate with rounded base. Style branches apically subtruncate to somewhat convex, penicillate.
Cypselae oblong-obovate, compressed, puberulous. Pappus bristles uniseriate, slender, lacking in
marginal florets. One species, S. bellidioides DC.,
southern Africa.
discoid, distinctly calyculate. Corolla white or
pinkish, with narrow lobes. Anthers ecaudate.
Style branches obtuse. Cypselae narrowly oblong,
ribbed, glabrous. Pappus bristles numerous,
slender. n = 18, 20. One species, H. suaveolens
(Raf.) Pojark., eastern USA.
495. Packera A. Löve & D. Löve
498. Robinsonia DC.
Packera A. Löve & D. Löve, Bot. Notiser 128: 520 (1976);
Barkley, Trans. Kansas Acad. Sci. 65: 318–408 (1962, as
Senecio), rev.; Freeman & Barkley, Sida Contr. Bot. 16:
699–709 (1995), reg. syn.; Trock, Sida Contr. Bot. 20:
1023–1041 (2003), reg. rev.
Robinsonia DC. in Guill., Arch. Bot. (Paris) 2: 333 (1833),
nom. cons.; Sanders et al., Opera Bot. 92: 195–215 (1987),
biogeogr., evol.; Sang et al., Syst. Bot. 20: 55–64 (1995),
phylog.
Rhetinodendron Meisn. (1839).
Symphyochaeta (DC.) Skottsb. (1951).
Perennial herbs with a creeping rhizome and
fibrous roots. Leaves basal, often rosulate, and
cauline, reduced upwards. Capitula solitary to
many, corymbose, radiate or sometimes discoid,
yellow- or orange- to red-flowered. Corolla lobes
deltoid to triangular-ovate. Anthers ecaudate.
Style branches truncate, apically penicillate.
Cypselae oblong, ribbed, glabrous or sometimes
pubescent. Pappus bristles numerous, slender.
Pollen helianthoid. n = mostly 22 and 23; also
reported: 20 (doubtful), 24, c. 30, 35, 40, 46, 60, 69.
Circa 75 species, North America, Mexico, Siberia.
496. Dorobaea Cass.
Dorobaea Cass., Dict. Sci. Nat. 48: 453 (1827); Nordenstam, Opera Bot. 44: 51–53 (1978), re-establ.; Nordenstam
& Pruski, Comp. Newslett. 27: 31–42 (1995), reg. rev.
Perennial glabrous herbs with fibrous roots. Leaves
rosulate, petiolate, lanceolate to ovate, pinnatipartite or laciniate. Capitula solitary on long, bracteate
peduncles, large, radiate, conspicuously calyculate,
yellow- or orange-flowered. Corolla lobes lanceolate with a median resin duct. Anthers ecaudate
with radial endothecium and balusterform filament collar. Style branches apically convex to conical, stigmatic areas separated. Cypselae oblong,
ribbed, glabrous or shortly hirsute. Pappus bristles
numerous, slender. n = c. 50. Three species, Peru,
Ecuador.
497. Hasteola Raf.
Hasteola Raf., New Fl. 4: 79 (1837).
Synosma Raf. ex Britton & A. Br. (1898).
Tall perennial herb with rhizome. Leaves basal,
long-petiolate, hastate, deltoid with serrate margins; cauline leaves progressively smaller. Capitula
numerous, corymbose, cylindrical or turbinate,
Dioecious trees or tall shrubs. Leaves cauline,
sessile, entire, narrowly elliptic to elliptic-ovate,
mostly assembled towards branch ends. Capitula
many, corymbose, radiate or discoid, calyculate.
Florets yellow to greenish or purplish. Anthers
aborted in female heads. Styles undivided in male
heads. Cypselae elliptic-oblong, ribbed, glabrous
or pubescent. Pappus bristles numerous, often
connate basally. n = 20. Seven species, Juan
Fernandez Islands.
499. Senecio L.
Fig. 57
Senecio L., Sp. Pl.: 866 (1753) & Gen. Pl. ed. 5: 373
(1754); Hooker, Fl. Brit. India 3: 338–356 (1881), reg. rev.;
Greenman, Bot. Jahrb. 32: 1–33 (1902), reg. rev.; Ann.
Missouri Bot. Gard. 25: 795–822 (1938), reg. rev.; Cabrera,
Lilloa 5: 65–120 (1939), reg. rev.; Lilloa 15: 27–501 (1949),
reg. rev.; Brittonia 7, 2: 53–74 (1950), reg. rev.; Arquiv. Jard.
Bot. Rio de Janeiro 15: 163–269 (1957), reg. rev.; Schischkin,
Fl. U.S.S.R. 26: 699–788 (1961), reg. rev.; Cabrera, Fl. Prov.
Buenos Aires 4, 6: 282–319 (1963), reg. rev.; Humbert, Fl.
Madag. 189: 677–804 (1963), reg. rev.; Aristeguieta, Fl.
Venez. 10: 765–815 (1964), reg. rev.; Merxmüller, Prodr.
Fl. Südwestafr. 139: 162–173 (1967), reg. rev.; Cabrera, Fl.
Patagonica 7: 176–275 (1971), reg. rev.; Cabrera, Fl. Ilustr.
Entre Ríos Argent. 6: 433–451(1974), reg. rev.; Cabrera &
Klein, Fl. Ilustr. Catarinense, Compostas: 135–217 (1975),
reg. rev.; Matthews, Fl. Turkey 5: 145–168 (1975), reg. rev.;
Chater & Walters, Fl. Eur. 4: 191–205 (1976), reg. rev.;
Hilliard, Compositae in Natal: 387–502 (1977), reg. rev.;
Barkley, N. Amer. Fl. ser. 2, 10: 50–139 (1978), reg. rev.;
Cabrera, Fl. Prov. De Jujuy 10: 489–586 (1978), reg. rev.;
Cabrera & Zardini, Darwiniana 22: 427–492 (1980), reg.
syn., key; Jeffrey & Chen, Kew Bull. 39: 351–432 (1984),
reg. rev.; Cabrera, Darwiniana 26: 79–217 (1985), reg. rev.;
Jeffrey, Kew Bull. 41: 879–904 (1986), reg. rev.; Nordenstam,
Fl. Iran. 164: 59–95, pl. 38–63 (1989), reg. rev.; Chen, Fl.
Compositae
Reip. Pop. Sin. 77, 1: 225–304 (1999), reg. rev.; Thompson,
Muelleria 19: 101–214 (2004), part. reg. rev.
Madaractis DC. (1838).
Crociseris (Reichenb.) Fourr. (1868).
Curio P.V. Heath (1999).
Annual or perennial herbs, subshrubs or shrubs.
Leaves alternate, cauline or rosulate, sessile or petiolate, rarely peltate, entire or variously lobed or
dissected. Capitula solitary or few to numerous,
corymbose, radiate, disciform or discoid, usually
231
calyculate. Florets often yellow, sometimes pink to
purple, rarely blue or white. Anthers usually ecaudate; filament collar balusterform. Style branches
truncate with short sweeping hairs; stigmatic areas separated. Cypselae elliptic-ovate to narrowly
oblong, often 8–12-ribbed, glabrous or pubescent.
Pappus bristles numerous, slender, persistent or caducous. n = mostly 10 or 20 but also reported are
9, 16, 18, 19, 21, 22, 30, 32, 35, 40, c. 45, 46, 49, 50,
52, 60, c. 70, c. 80, 90, c. 91. Circa 1,250 species, cosmopolitan, many in South America and southern
Africa.
Several discordant elements have recently been
segregated (Nordenstam 2006c, e, f; Nordenstam &
Pelser 2005) and some more need to be removed
from Senecio in order to achieve monophyly. One
of these segregates is the Eurasian genus Jacobaea
Mill. (1754) with c. 35 species (Nordenstam 2006a;
Pelser et al. 2006).
500. Solanecio (Sch. Bip.) Walp.
Solanecio (Sch. Bip.) Walp., Rep. 6: 273 (1846); Jeffrey, Kew
Bull. 41: 920–923 (1986), rev.
Perennial herbs, shrubs or small trees. Leaves
cauline, sessile or petiolate, entire to serrate or
lobed to laciniate. Capitula few to many, corymbose, discoid, calyculate, yellow-flowered. Anthers
ecaudate. Style branches truncate, occasionally
with a central hair tuft. Cypselae oblong, ribbed,
glabrous or pubescent; carpopodium distinct,
annular. Ovary wall crystals drusiform. Pappus
bristles numerous, slender. n = 10, c. 90. Sixteen
species, tropical Africa, Madagascar, Yemen.
501. Kleinia Mill.
Kleinia Mill., Gard. Dict. abr. ed.: 4 (1754); Halliday, Kew
Bull. 39: 817–827 (1984), reg. rev.; Jeffrey, Kew Bull. 41:
923–929 (1986), reg. rev.; Halliday, Hooker’s Icon. Pl. 39,
4: t. 3876–3900 (1988), part. rev.; Thulin, Nordic J. Bot. 22:
419–426 (2003), new spp.
Notonia DC. (1833).
Notoniopsis B. Nord. (1978).
Fig. 57. Compositae-Senecioneae. Senecio vernalis. A Habit.
B Ray floret. C Disc floret. D Corolla of disc floret, opened.
E Stamens. F Style branches of disc floret. G Cypsela (pappus
removed). (Drawing by B. Nordenstam)
Succulent herbs or shrubs. Leaves cauline, sessile
or shortly petiolate, entire to incised-lobed, sometimes reduced to scales or absent. Capitula solitary or few to numerous, corymbose, discoid, with
or without calyculus. Florets white, yellow, greenish, orange, red or purplish; corolla tubular, gradually widened upwards. Anthers ecaudate, with long,
slightly balusterform filament collar. Style branches
linear, apically with a convex or conical to long-
232
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
acuminate, papillose-hairy appendage; stigmatic
areas separated. Cypselae oblong, ribbed, glabrous.
Ovary wall crystals drusiform. Pappus bristles numerous, slender, persistent. n = 9, 10, 20, c. 50.
Circa 50 species, Africa, Madagascar, Canary Islands, Arabia, east to India and Sri Lanka.
502. Gynura Cass.
Gynura Cass., Dict. Sci. Nat. 34: 392 (1825), nom. cons.;
Davies, Kew Bull. 33: 335–342 (1978), Kew Bull. 33: 629–640
(1979), Kew Bull. 35: 363–367 (1980), Kew Bull. 35: 711–734
(1981), rev.; Jeffrey, Kew Bull. 41: 929–930 (1986), reg. rev.;
Chen, Fl. Reip. Pop. Sin. 77, 1: 309–322 (1999), reg. rev.
Perennial herbs or subshrubs, erect or scrambling
or climbing, sometimes subsucculent; roots fibrous
or tuberous. Leaves cauline, sessile or petiolate, entire or lobed to pinnatifid. Capitula solitary or few
to several, corymbosely paniculate, discoid. Involucre cylindrical or campanulate, calyculate. Florets
white, greenish, yellow, orange, red or purplish. Anthers ecaudate, basally obtuse or sometimes auriculate. Style branches apically with elongate vascularized papillate appendage. Cypselae oblong, ribbed,
glabrous or pubescent; carpopodium annular or
indistinct. Ovary wall crystals drusiform. Pappus
bristles numerous, slender. n = 5, 10, 11, 17, 20.
Circa 40 species, Palaeotropics.
503. Erechtites Raf.
Erechtites Raf., Fl. Ludov.: 65 (1817); Belcher, Ann. Missouri Bot. Gard. 43: 1–85 (1956), rev.; Barkley & Cronquist,
N. Amer. Fl. ser. 2, 10: 139–142 (1978), reg. rev.
Annual or perennial herbs. Leaves cauline, sessile, pinnatilobate or entire with serrate margins.
Capitula several, corymbose, disciform. Involucre
cylindrical, calyculate. Marginal florets female, filiform. Disc florets perfect, white to yellowish. Style
branches with apical appendage of fused hairs.
Cypselae oblong, ribbed; carpopodium indistinct.
Pappus bristles numerous, slender. n = 20. Five
species, North and South America, West Indies.
504. Iocenes B. Nord.
Iocenes B. Nord., Opera Bot. 44: 58 (1978).
Erect perennial herb with a short rhizome. Leaves
basal and cauline, lower leaves petiolate, lyratopinnatifid or incised-lobed, upper leaves sessile,
half-clasping. Capitula several, corymbose, radiate, paucicalyculate. Ray florets white. Disc florets
perfect, yellow. Anthers ecaudate. Style branches
apically with an elongated hair pencil. Cypselae oblong, 10-ribbed, glabrous; carpopodium distinct.
Pappus bristles numerous, slender, persistent. n =
20. One species, I. acanthifolius (Hombr. & Jacq.)
B. Nord., with two subspecies, Argentina, Chile.
505. Io B. Nord.
Io B. Nord., Comp. Newslett. 40: 47 (2003).
Glabrous half-shrub. Leaves opposite, subsessile,
ovate-lanceolate, entire with denticulate margins.
Capitula laxly corymbose, radiate, yellow-flowered,
calyculate. Anthers sagittate. Style branches apically truncate without sweeping hairs, dorsally
papillate; stigmatic areas continuous. Cypselae
oblong, glabrous. Pappus bristles uniseriate, slender. One species, I. ambondrombeensis (Humb.)
B. Nord., Madagascar.
506. Crassocephalum Moench
Crassocephalum Moench, Methodus: 516 (1794); Humbert,
Fl. Madag. 189: 829–843 (1963), reg. rev.; Jeffrey, Kew Bull.
41: 904–908 (1986), reg. rev.
Perennial herbs. Leaves cauline, sessile or petiolate, sometimes amplexicaul, entire or lobed, margins often serrate. Capitula few or several, corymbose, radiate or discoid, cylindrical, calyculate. Florets creamy white or yellow, orange, pink to red
or mauve, purple or blue. Anthers ecaudate; filament collar narrow, elongate. Style branches apically rounded and shortly appendiculate, dorsally
papillate. Cypselae oblong, ribbed, glabrous or with
thickened twin hairs; carpopodium indistinct. Pappus bristles numerous, slender. n = 5, 10, 12, 16,
20, 24, c. 45. Twenty-four species, tropical Africa,
Madagascar, Yemen (two weedy species introduced
in many tropical countries).
507. Hoehneophytum Cabrera
Hoehneophytum Cabrera (‘Hoehnephytum’), Brittonia 7: 53
(1950); Hind, Kew Bull. 48: 257–259 (1993), notes; Hind,
Kew Bull. 54: 897–904 (1999, publ. 2000), notes.
Suffrutescent herbs or shrubs. Leaves cauline, petiolate, elliptic, entire. Capitula several, corymbose,
discoid, few-flowered, yellow, calyculate. Anther
base obtuse. Style branches apically rounded with
short sweeping hairs, dorsally puberulous; stigmatic areas separated. Cypselae oblong, ribbed,
pubescent. Pappus bristles numerous, slender.
Three species, Brazil.
Compositae
508. Pentacalia Cass.
Pentacalia Cass., Dict. Sci. Nat. 48: 461 (1827); Robinson
& Cuatrecasas, Phytologia 40: 37–50 (1978), reg. rev.;
Cuatrecasas, Phytologia 49: 241–260 (1981), reg. rev.;
Díaz-Piedrahita & Cuatrecasas, Asteraceas de la Flora de
Colombia, Senecioneae-I: 25–174 (1999), reg. rev.
Scandent shrubs or epiphytes, glabrous or
pubescent with simple hairs. Leaves petiolate,
oblong to elliptic-ovate, entire with margins often
dentate to serrate, often coriaceous. Capitula
several to many, corymbosely paniculate in
terminal or lateral synflorescences, radiate or disciform, yellow- or rarely white-flowered. Anthers
sagittate to caudate. Style branches truncate or
obtuse, with separated stigmatic areas. Cypselae
elliptic-oblong, 5-angled or -ribbed, glabrous or
sometimes pubescent. Pappus bristles numerous,
slender, white or pink to rufous. n = 20, c. 40,
45–50, 50, c. 51. Circa 200 species, Andes of South
America, Central America north to Mexico.
509. Monticalia C. Jeffrey
Monticalia C. Jeffrey, Kew Bull. 47: 69 (1992), 69–74,
notes, new comb.; Robinson & Cuatrecasas, Phytologia 40:
37–50 (1978, as Pentacalia subg. Microchaete), reg. rev.;
Díaz-Piedrahita & Cuatrecasas, Asteraceas de la Flora de
Colombia, Senecioneae-I: 175–345 (1999, as Pentacalia
subg. Microchaete), reg. rev.
Microchaete Benth. (1845), nom. illegit. rej., non Thuret ex
Bornat & Flahault (1886), nom. cons.
Erect shrubs or shrublets. Leaves closely set,
shortly petiolate to subsessile, often small, ericoid
or linear-lanceolate to elliptic-ovate. Capitula
several to numerous, corymbose, radiate or discoid, mostly yellow-flowered. Anthers sagittate to
shortly caudate. Style branches truncate to obtuse
with discrete stigmatic areas, papillate apically
and sometimes dorsally. Cypselae 5-angled or
-ribbed, with distinct carpopodium. Pappus
bristles numerous, slender, white. n = c. 10, 20,
c. 40. Circa 70 species, Andes of South America,
Central America north to Costa Rica.
510. Dendrophorbium (Cuatrec.) C. Jeffrey
Dendrophorbium (Cuatrec.) C. Jeffrey, Kew Bull. 47: 65
(1992), 65–69, notes, new comb.; Díaz-Piedrahita & Cuatrecasas, Asteraceas de la Flora de Colombia, Senecioneae-I:
8–25 (1999), reg. rev.
Senecio L. sect. Dendrophorbium Cuatrec. (1951).
Erect suffrutescent herbs (‘dendrophorbs’), shrubs
233
or trees. Leaves large, cauline, often assembled
apically on branches, entire, petiolate, margins
dentate or denticulate. Capitula numerous in
± dense terminal corymbose or paniculate synflorescences. Anther base ecaudate or rarely caudate,
rounded-obtuse or minutely auriculate. Style
branches apically truncate to obtusely convex;
stigmatic areas narrowly separated. Cypselae
elliptic-oblong, 5-ribbed; carpopodium distinct.
Pappus bristles numerous, slender. n = c. 40,
49–50, c. 50. Circa 75 species, Andes of South
America from Bolivia to Venezuela, Argentina,
Brazil, Paraguay.
511. Graphistylis B. Nord.
Graphistylis B. Nord., Opera Bot. 44: 56 (1978); Hind, Kew
Bull. 48: 285 (1993), notes; Nordenstam, Comp. Newslett.
24: 51 (1994), new comb.
Erect subshrubs or somewhat lignescent herbs,
usually pubescent and glabrescent. Leaves shortly
petiolate or subsessile, entire, elliptic to lanceolate,
with serrate or denticulate margins. Capitula
several to numerous, corymbose-paniculate, radiate, yellow- or rarely white-flowered, calyculate.
Disc florets perfect, corolla lobes triangular-ovate
with a median resin duct. Anther base obtuse,
auriculate; endothecial tissue mainly polarized.
Style branches apically with an elongated hair
pencil and lateral shorter hair tufts; stigmatic areas
narrowly separated. Cypselae narrowly oblong,
ribbed, glabrous. Pappus bristles numerous,
slender, persistent. n = 50. Eight species, Brazil.
512. Talamancalia H. Rob. & Cuatrec.
Talamancalia H. Rob. & Cuatrec., Novon 4: 50 (1994); Nordenstam & Pruski, Comp. Newslett. 27: 31–42 (1995), key,
new comb.
Herbs, subshrubs or shrubs, erect or ascending,
hirsute or lanate, glabrescent. Leaves cauline, petiolate with a ± winged petiole and half-clasping base,
pinnatilobate at least proximally, otherwise entire
with serrate margins. Capitula few to several, laxly
corymbose, radiate, orange- or yellow-flowered;
involucre broadly campanulate, calyculate. Receptacle naked. Disc florets numerous; corolla lobes
narrowly oblong. Anthers ecaudate. Style branches
apically rounded with a short central hair tuft; stigmatic areas separated. Cypselae 10-ribbed, with
short mucilaginous hairs. Pappus bristles pluriseriate, slender, caducous. Four species, Peru, Ecuador,
Panama, Costa Rica.
234
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
513. Caxamarca Dillon & Sagástegui
516. Lasiocephalus Schlecht.
Caxamarca Dillon & Sagástegui, Novon 9: 156–161 (1999).
Lasiocephalus Schlecht., Gesell. Naturf. Freunde Berlin
Mag. Neusten Entdeck. Gesamt. Naturk. 8: 308 (1818); Nordenstam, Opera Bot. 44: 53–56 (1978, under Aetheolaena).
Perennial herb with foetid tuberous fasciculate
roots and unbranched erect stem. Basal leaves
rosulate, large, dissected; cauline leaves smaller,
sessile, entire with serrate to dentate margins, with
decurrent base forming wings. Capitula several,
corymbose-cymose, radiate, yellow-flowered.
Involucre hemispherical, basally swollen, pluriseriate, calyculate. Disc florets numerous, with
deeply lobed corolla. Anthers ecaudate. Style
branches apically with a conical hair tuft; stigmatic areas separated. Cypselae 10-ribbed, pilose.
Pappus bristles biseriate, slender, persistent.
One species, C. sanchezii Dillon & Sagástegui,
Peru.
514. Pseudogynoxys (Greenm.) Cabrera
Pseudogynoxys (Greenm.) Cabrera, Brittonia 7: 54 (1950);
Robinson & Cuatrecasas, Phytologia 36: 177–192 (1977),
rev.; Pruski, Syst. Bot. 21: 101–105 (1996), new sp.
Scandent herbs or subshrubs. Leaves petiolate, entire or rarely lobate, margins often serrate. Capitula few to several, occasionally solitary, corymbose, radiate, orange-red- or orange- to yellowflowered, fragrant, calyculate. Anther base obtuse
to sagittate, ecaudate. Style branches apically with
an elongate acuminate appendage of fused hairs.
Cypselae oblong, ribbed, glabrous or puberulous
to hirsute; carpopodium indistinct. Pappus bristles
numerous, slender. n = 10 (questionable), 45, 47–
48. Fourteen species, South America from Bolivia
and Brazil to Central America and Mexico. Pseudogynoxys chenopodioides (Kunth) Cabrera is cultivated as an ornamental and occurs as a garden escapee.
515. Garcibarrigoa Cuatrec.
Garcibarrigoa Cuatrec., Caldasia 15: 6 (1986).
Perennial herbs. Leaves petiolate, entire with
serrulate margins, elliptic-lanceolate, basally
auriculate and stem-clasping, strongly pinnately
veined. Capitula solitary or few, corymbose,
radiate, orange- or yellow-flowered. Anthers
ecaudate. Style branches apically with a triangularacuminate hair tuft. Cypselae oblong, ribbed,
glabrous; carpopodium indistinct. Pappus bristles numerous, slender. Two species, Ecuador,
Colombia.
Erect or ascending little-branched shrublets.
Leaves small and closely set, subsessile, entire,
elliptic or obovate to lanceolate, tomentose,
margins revolute. Capitula solitary or few together,
nodding, discoid, yellow-flowered, calyculate.
Anther base ecaudate, obtuse. Style branches
apically obtuse or truncate with a median hair
pencil. Cypselae oblong, ribbed, glabrous. Pappus
bristles numerous, slender. Two or three species,
Andes of Peru, Ecuador, Colombia.
517. Aetheolaena Cass.
Aetheolaena Cass., Dict. Sci. Nat. 48: 453 (1827); Nordenstam, Opera Bot. 44: 53–56 (1978), re-establ., new comb.;
Cuatrecasas, Phytologia 40: 307–312 (1978), rev. (sub
Lasiocephalo).
Lasiocephalus sensu auct. plur., non Schlecht. (1818).
Scandent or ascending half-shrubs. Leaves scattered, petiolate but upper cauline leaves often
sessile, triangular-ovate, entire, often dentate
or serrulate. Capitula few to many, corymbosepaniculate, discoid, nodding, calyculate; florets
dirty white to greenish-yellow. Anther base ecaudate, obtuse. Style branches apically convex or
conical with a central elongate hair pencil and
shorter lateral hairs or papillae; stigmatic areas
separated. Cypselae oblong, ribbed, glabrous.
Pappus bristles numerous, slender, caducous.
n = 20. Circa 15 species, Andes of South America
from Bolivia to Venezuela.
518. Arbelaezaster Cuatrec.
Arbelaezaster Cuatrec., Caldasia 15: 1 (1986).
Erect perennial herb with rhizome. Leaves cauline,
shortly petiolate, ovate, entire with denticulate
or serrate margins. Capitula several, paniculatecorymbose, discoid with greenish-yellow florets,
calyculate. Anthers sagittate. Style branches with
an apical tuft of fused long hairs; stigmatic areas
separated. Cypselae oblong, glabrous, ribbed.
Pappus bristles numerous, slender. One species,
A. ellesworthii (Cuatrec.) Cuatrec., Colombia.
519. Dresslerothamnus H. Rob.
Dresslerothamnus H. Rob., Phytologia 40: 493 (1978);
Robinson, Syst. Bot. 14: 380–388 (1989), rev.
Compositae
Scandent shrubs with T-shaped or substellate
trichomes. Leaves cauline, petiolate, elliptic-ovate,
entire. Capitula several, corymbose-paniculate,
radiate, calyculate. Rays flagelliform, twisted and
recurved, reddish. Disc florets perfect, yellow.
Anthers caudate with long and often papillose
tails. Style branches apically truncate, papillate
and with a short median hair tuft; stigmatic areas
separated. Cypselae oblong, ribbed or angled,
glabrous. Pappus bristles numerous, slender with
enlarged tips, caducous. Four or five species,
Colombia, Panama, Costa Rica.
520. Cabreriella Cuatrec.
Cabreriella Cuatrec., Bol. Soc. Argent. Bot. 119: 15 (1980).
Scandent glabrous shrubs. Leaves opposite, sessile
or subsessile, ovate-cordate, entire with dentate
margins. Capitula several, corymbose, radiate or
discoid, yellow-flowered, calyculate. Disc florets
perfect. Anthers basally sagittate or shortly caudate. Style branches of disc florets apically truncate
or obtuse, papillate; stigmatic areas separated.
Cypselae elliptic-oblong, 5-ribbed, glabrous.
Pappus bristles numerous, slender. Two species,
Colombia.
521. Culcitium Humb. & Bonpl.
235
Style branches apically rounded with subterminal
short sweeping hairs. Achenes oblong, ribbed,
glabrous. Pappus bristles many, slender, caducous.
One species, C. durandii (Klatt) J. Janovec &
H. Rob., Costa Rica.
523. Misbrookea V.A. Funk
Misbrookea V.A. Funk, Brittonia 49: 111 (1997).
Perennial rhizomatous herb forming mats or growing solitary, densely hairy with long strigose trichomes. Leaves closely set and appressed, sessile,
oblong-obovate, greyish-green. Capitula solitary,
sessile or subsessile, radiate, ecalyculate. Involucral bracts connate. Ray florets white. Disc florets perfect, yellow. Anthers shortly sagittate. Style
branches apically obtuse, papillate and centrally
penicillate; stigmatic areas separated. Cypselae oblong, ribbed, hairy. Pappus bristles numerous, slender, persistent. n = c. 58, c. 58 + 2B. One species,
M. strigosissima (A. Gray) V.A. Funk, high Andes
of Bolivia and Peru.
524. Xenophyllum V.A. Funk
Xenophyllum V.A. Funk, Novon 7: 235 (1997); Funk,
Estudios sobre diversidad y ecologia de plantas, Univ.
Catol. Quito: 25–35 (1997), reg. rev., key.
Perennial herbs, densely tomentose. Leaves
rosulate, lanceolate to elliptic, entire, margins
sometimes serrulate. Capitula large, solitary or
few, corymbose, nodding, discoid, many-flowered,
yellow- or cream-flowered. Involucre pluriseriate,
calyculate. Anthers ecaudate. Style branches truncate to obtuse. Cypselae oblong, ribbed, glabrous.
Pappus bristles numerous, very slender. Circa 15
species, Andes of South America, from Chile to
Venezuela.
Perennial herbs forming hummocks or mats;
rhizomes covered with leaves or leaf bases. Leaves
closely spirally set, entire and ± thick or small
with lobed tips, bright green at least distally,
becoming brown or blackish. Capitula solitary,
sessile, ecalyculate, radiate. Involucral bracts
connate at least basally. Ray florets white or
purplish. Disc florets perfect, white or yellow. Style
branches apically truncate to rounded, papillate.
Cypselae obovate, ribbed, glabrous. Pappus bristles numerous, slender. n = 52, c. 54. Twenty-one
species, high Andes from Argentina and Chile to
Colombia.
522. Charadranaetes J. Janovec & H. Rob.
525. Werneria Kunth
Charadranaetes J. Janovec & H. Rob., Novon 7: 162–168
(1997).
Werneria Kunth in HBK., Nov. Gen. Sp. Pl. ed. folio 4: 148
(1818); Rockhausen, Bot. Jahrb. Syst. 70: 273–342 (1940),
rev.; Cabrera, Not. Mus. La Plata 13 Bot. 60: 49–61 (1948),
reg. rev.; Funk, Estudios sobre diversidad y ecologia de
plantas, Univ. Catol. Quito: 25–35 (1997), reg. rev., key.
Culcitium Humb. & Bonpl., Pl. Aequin. 2: 1 (1808);
Cuatrecasas, Fieldiana Bot. 27: 40–51 (1950), rev.
Semi-woody glabrous shrub. Leaves cauline,
petiolate, entire with serrate margins, foetid
when crushed. Capitula 2 or 3, radiate, narrow;
involucre uniseriate, calyculate. Ray florets yellow
or orange-coloured. Disc florets perfect, yellow;
corolla lobes linear-lanceolate with resin ducts.
Anther base ecaudate, obtuse or slightly sagittate.
Perennial rosulate herbs growing solitary or in
small clumps. Leaves sessile, closely set at rhizome
tips or below capitula, entire, glabrous. Capitula
solitary, sessile or pedunculate, ecalyculate. In-
236
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
volucral bracts connate to various degrees. Ray
florets white or rarely yellow. Disc florets perfect,
white or yellow. Anthers sagittate. Style branches
apically obtuse, papillate. Cypselae oblong, ribbed,
glabrous. Pappus bristles numerous, slender.
n = c. 21, c. 48, 50, c. 58, c. 75, c. 77, c. 103, 111.
Twenty-five to 30 species, high Andes of South
America.
continuous. Cypselae dimorphic, oblong with
wing-like ridges: those of ray florets glabrous,
brown to black; those of disc florets hirsute on
the curved ridges with mucilaginous twin hairs,
brown. Pappus bristles numerous, pluriseriate,
slender, caducous in ray florets, persistent in disc
florets. One species, P. bolusii (Oliv.) B. Nord.,
South Africa.
526. Oresbia Cron & B. Nord.
529. Dendrosenecio (Hauman ex Hedberg) B.
Nord.
Oresbia Cron & B. Nord., Novon 16:216 (2006).
Perennial tomentose-araneose herb. Leaves alternate, sessile, elliptic-ovate or spathulate with
denticulate to serrate margins, glabrescent above,
base half-clasping. Capitula few to several, laxly
corymbose, rarely solitary on long peduncle,
radiate, yellow-flowered, calyculate. Anthers
sagittate or shortly caudate. Style branches apically
truncate, papillate. Cypselae dimorphic: outer
4-winged, glabrous; inner 6-ribbed with mucilaginous hairs. Pappus bristles white, caducous. One
species, O. heterocarpa Cron & B. Nord., South
Africa.
527. Lamprocephalus B. Nord.
Lamprocephalus B. Nord., Bot. Notiser 125: 323 (1976).
Shrublet. Leaves cauline, sessile, linear, coriaceous.
Capitula solitary on long, bracteate peduncles,
discoid, red-flowered. Involucre uniseriate, ecalyculate. Corolla tubular, shortly lobed. Anthers
slightly sagittate. Style branches apically with large
oblong-triangular obtuse appendage, glabrous but
minutely papillate; stigmatic areas continuous.
Cypselae oblong, 5-angular, with short myxogenic
twin hairs. Pappus bristles copious, pluriseriate,
slender, persistent. One species, L. montanus
B. Nord., South Africa.
528. Phaneroglossa B. Nord.
Phaneroglossa B. Nord., Opera Bot. 44: 66 (1978).
Ascending subshrub, glabrous except for woolly
leaf-axils. Leaves cauline, sessile, closely set, linearlanceolate or oblanceolate, entire, midribbed,
basally half-clasping; margins denticulate and
somewhat revolute. Capitula solitary, longpedunculate, radiate, ecalyculate, white- or
cream-flowered. Involucral bracts uniseriate,
basally connate. Tube of ray florets shortly hirsute.
Disc florets perfect. Anther base auriculate.
Style branches apically conical; stigmatic areas
Dendrosenecio (Hauman ex Hedberg) B. Nord., Opera Bot.
44: 40 (1978); Mabberley, Kew Bull. 28: 61–96 (1973, as
Senecio subg. Dendrosenecio), morph.; Jeffrey, Kew Bull.
41: 887–892 (1986, as Senecio ser. Arborei), rev.; Mabberley
in Vuilleumier & Monasterio (eds) High altitude tropical
biogeography: 81–102 (1986), morph.; Knox, Contr. Univ.
Michigan Herb. 19: 241–257 (1993), rev.; Knox & Palmer,
Amer. J. Bot. 82: 1567–1573 (1995), phylog.
Pachycaul trees or tree-like herbs (‘dendrophorbs’).
Leaves very large, assembled towards ends of stem
and branches, sessile or petiolate, ovate-elliptic or
panduriform, pinnately and reticulately veined.
Capitula numerous in terminal panicles, large,
nodding, radiate or discoid, yellow-flowered.
Involucre ± biseriate, calyculate. Anther base
auriculate; filament collar somewhat balusterform
to almost straight. Style branches truncate and
shortly penicillate, with continuous stigmatic
areas. Cypselae oblong, ribbed, glabrous. Pappus
bristles numerous, slender. n = 10, c. 40, 50. Eleven
species, mountains of tropical Africa.
530. Euryops (Cass.) Cass.
Euryops (Cass.) Cass., Dict. Sci. Nat. 16: 49 (1820), nom.
cons. prop.; Nordenstam, Opera Bot. 20: 1–409 (1968),
rev.; Nordenstam, Bot. Notiser 121: 209–232 (1968), cyt.,
morph.; Nordenstam, Opera Bot. 23: 1–77 (1969), biogeogr.
Jacobaeastrum Vaill. (1754), nom. rej. prop.
Gamolepis sensu auct. plur., non Less. (1832).
Ruckeria DC. (1838).
Lasiocoma Bol. (1906).
Thodaya Compt. (1931).
Lysichlamys Compt. (1943).
Shrubs or shrublets or perennial (one annual)
herbs. Leaves cauline or rarely subrosulate, sessile,
entire to variously lobed or pinnatisect. Capitula
axillary on naked peduncles, often numerous, or
pseudo-terminal and solitary, radiate or rarely
discoid, yellow-flowered. Involucre uniseriate, ecalyculate; bracts connate or rarely free. Disc florets
Compositae
perfect or rarely functionally male; corolla lobes
deltoid-ovate. Anthers ecaudate. Style branches
truncate-obtuse, with discrete stigmatic areas.
Cypselae elliptic-oblong, glabrous or pubescent
with often myxogenic twin hairs. Pappus bristles
uniseriate, flexuous, caducous, or absent. n = 10,
20, 30. One hundred species, Africa, Arabia,
Socotra.
531. Othonna L.
Othonna L., Sp. Pl.: 924 (1753) & Gen. Pl. ed. 5: 396 (1754);
Merxmüller, Prodr. Fl. Südwestafr. 139: 129–135 (1967), reg.
rev.; Rowley, The succulent Compositae (1994), horticult.
review.
Perennial succulent shrubs or herbs with underground tubers or corms. Leaves cauline or
rosulate, sessile or shortly petiolate, entire or
dentate to serrate or lobed, terete to flat, mostly
glabrous. Capitula solitary and pedunculate to
many and corymbose, radiate or discoid, ecalyculate. Involucral bracts uniseriate, connate
at least basally. Florets yellow, white, pink or
purple. Anthers ecaudate. Style branches of disc
florets simple, sterile, apically conical or obtuse.
Cypselae elliptic-oblong, glabrous or pubescent.
Pappus bristles numerous, slender, persistent
or caducous, white, straw-coloured or pink to
rufous or purplish. n = 10, 20, 30, c. 40. Circa
120 species, southern Africa north to Angola and
Zimbabwe. Genus biphyletic and under revision
(B. Nordenstam).
532. Gymnodiscus Less.
Gymnodiscus Less., Linnaea 6: 95 (1831).
Annual glabrous herbs. Leaves rosulate or some
cauline, pinnately lobed or dissected. Capitula few
to several, corymbose, small and few-flowered,
radiate, yellow-flowered, ecalyculate. Involucral
bracts connate. Disc florets functionally male with
a simple sterile style. Anthers ecaudate. Cypselae
oblong-ovoid, glabrous. Pappus bristles numerous,
slender, lacking in disc florets. n = 9. Two species,
southern Africa.
533. Hertia Less.
Hertia Less., Syn. Gen. Comp.: 88 (1832).
Othonnopsis Jaub. & Spach (1852).
Shrubs or subshrubs or perennial herbs with
a thick rhizome. Leaves cauline, sessile, entire
with margins sometimes serrate, linear to lance-
237
olate or obovate, glabrous, ± succulent. Capitula
solitary, axillary or terminal, or several paniculatecorymbose, radiate or disciform, yellow-flowered.
Involucral bracts uniseriate, often connate at least
basally. Receptacle flat, naked. Disc florets functionally male. Anthers ecaudate. Style branches
obtuse, dorsally hirsute or papillate. Cypselae
elliptic-oblong, ribbed, glabrous or pubescent.
Pappus bristles numerous, slender, persistent.
n = 10. Circa 10 species, southern and northern
Africa, south-western Asia east to Iran.
534. Lopholaena DC.
Lopholaena DC., Prodr. 6: 335 (1838); Phillips & Smith,
Trans. Roy. Soc. S. Afr. 21: 221–238 (1934), rev.; Jeffrey, Kew
Bull. 41: 932 (1986), reg. rev.
Perennial herbs, subshrubs or shrubs, with a thick
rhizome. Leaves cauline, sessile, entire, fleshy
or coriaceous, margins often serrate. Capitula
solitary or several, paniculate-corymbose, discoid,
ecalyculate. Involucral bracts ± connate, dorsally
keeled. Florets white or pink to purple. Style
branches with elongate subulate appendage.
Cypselae elliptic-oblong, glabrous or pubescent.
Pappus bristles numerous, slender. Eighteen
species, southern Africa.
535. Cadiscus E. Mey. ex DC.
Cadiscus E. Mey. ex DC., Prodr. 7: 254 (1838).
Aquatic herb. Leaves floating, alternate, linear, amplexicaul. Capitula solitary, axillary, radiate, ecalyculate. Involucral bracts uniseriate, connate. Ray
florets white. Disc florets functionally male, yellow;
style simple, sterile. Cypselae oblong, hirsute. Pappus bristles few, coarse, subpaleaceous. n = 10. One
species, C. aquaticus E. Mey. ex DC., South Africa.
536. Stenops B. Nord.
Stenops B. Nord., Opera Bot. 44: 73 (1978); Nordenstam,
Comp. Newslett. 23: 1–2 (1993), syn., new comb.
Pseudocadiscus Lisowski (1987).
Glabrous annual subcarnose sub-aquatic herbs.
Leaves cauline, sessile, simple, linear to narrowly
elliptic-oblong, margins slightly denticulate.
Capitula few, corymbose-paniculate, radiate,
ecalyculate. Involucral bracts uniseriate, connate
to about the middle. Receptacle conical. Ray florets
white or pink to mauve. Disc florets perfect, yellow.
Anthers ecaudate; filament collar much dilated
basally. Style branches apically truncate with
238
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
short sweeping hairs; stigmatic areas separated.
Cypselae 5-angled, glabrous. Pappus absent. Two
species, Tanzania, Zambia.
537. Oligothrix DC.
Oligothrix DC., Prodr. 6: 293 (1838).
Small annual herb. Leaves cauline, sessile and
amplexicaul, entire with somewhat serrate margins. Capitula few, laxly corymbose, small and
few-flowered, radiate. Involucral bracts connate.
Ray florets short, white or yellow. Disc florets
perfect, yellow. Anthers ecaudate; filament collar
basally much enlarged. Style branches truncate,
penicillate. Cypselae elliptic-oblong, ribbed, puberulous. Pappus bristles few, slender, caducous.
One species, O. gracilis DC., South Africa.
538. Steirodiscus Less.
Steirodiscus Less., Syn. Gen. Comp.: 251 (1832).
Gamolepis Less. (1832).
Psilothonna E. Mey. ex DC. (1838).
Small annual glabrous herbs. Leaves cauline, sessile, entire and linear or pinnately lobed. Capitula solitary or few, corymbose, radiate, yellowflowered, ecalyculate. Involucral bracts uniseriate,
connate at least basally. Receptacle convex. Disc
florets perfect or functionally male. Anthers ecaudate with rounded base. Style branches truncate to
conical-obtuse, penicillate. Cypselae oblong-ovate,
glabrous or puberulous. Pappus absent. n = 8, 10.
Five species, southern Africa.
539. Bafutia C.D. Adams
Bafutia C.D. Adams, Kew Bull. 15: 439 (1962).
Annual herb. Leaves cauline, sessile, oblanceolate,
entire with sometimes serrulate margins. Capitula few, corymbose, small, discoid, ecalyculate.
Involucral bracts uniseriate, connate. Florets
hermaphrodite, pink to purplish; corolla lobes
oblong-ovate. Anthers ecaudate. Style branches
short, apically obtuse-clavate, penicillate. Cypselae
oblong, glabrous. Pappus bristles few, short,
caducous. One species, B. tenuicaulis C.D. Adams,
Cameroun Mts., western tropical Africa.
540. Psednotrichia Hiern
Psednotrichia Hiern, J. Bot. 36: 289 (1898); Anderberg &
Karis, Nordic J. Bot. 15: 375–379 (1995), re-establ.
Xyridopsis B. Nord. (1978).
Small annual herbs, ± glabrous. Leaves mostly
basal, entire, linear-filiform. Capitula solitary
on slender naked peduncles, small, discoid,
yellow-flowered, ecalyculate. Involucral bracts
uniseriate, connate to about the middle. Corolla
lobes unequal, narrowly ovate, with a median resin
duct. Anthers ecaudate; filament collar gradually
widening towards the base. Style branches apically
with a short conical hair tuft, dorsally minutely
papillate; stigmatic areas separated. Cypselae
elliptic-oblong, shortly hirsute with obtuse mucilaginous twin hairs. Pappus bristles several,
flexuous, distinctly barbellate, caducous. Two
species, Angola.
541. Emilia (Cass.) Cass.
Emilia (Cass.) Cass., Dict. Sci. Nat. 34: 393 (1825); Humbert, Fl. Madag. 189: 804–823 (1963), reg. rev.; Barkley &
Cronquist, N. Amer. Fl. ser. 2, 10: 147–150 (1978), reg. rev.;
Nicolson, Syst. Bot. 5: 391–407 (1980), reg. rev.; Jeffrey, Kew
Bull. 41: 908–920 (1986), reg. rev.
Pithosillum Cass. (1826).
Pseudactis S. Moore (1919).
Annual or perennial herbs with fibrous roots,
mostly glabrous. Leaves cauline, sessile or petiolate, sometimes amplexicaul. Capitula solitary or
few to several, corymbose, radiate or discoid, ecalyculate. Florets white, pink, red, purple, orange or
yellow. Involucral bracts uniseriate. Style branches
apically truncate or obtuse, minutely papillate,
sometimes with a small central hair tuft. Cypselae
elliptic-oblong; carpopodium indistinct. Pappus
bristles numerous, slender, persistent. n = 5, 8,
10, 11, 15, 20. Circa 100 species, tropical regions,
mostly in Africa (some tropical weeds).
542. Emiliella S. Moore
Emiliella S. Moore, J. Bot. 56: 225 (1918); Torre, Garcia de
Orta, sér. Bot. 2: 85–88 (1975), rev.
Annual herbs, mostly glabrous. Leaves cauline,
petiolate or sessile, entire or lobed, sometimes
basally auriculate and/or amplexicaul. Capitula
solitary or few, corymbose, discoid. Involucre
cylindrical, ecalyculate. Florets few, pink to
purplish blue. Style branches truncate. Cypselae
narrowly oblong, ribbed, glabrous. Pappus a single scale or absent. Five species, Angola, Zaire,
Zambia.
Compositae
543. Faujasiopsis C. Jeffrey
Faujasiopsis C. Jeffrey, Kew Bull. 47: 77 (1992), rev.; Jeffrey,
Fl. Mascar. 109: 134–139 (1993), rev.
Scandent or erect shrubs. Leaves cauline, entire,
broad, glabrous. Capitula several, corymbose,
disciform or discoid, white- or lilac-flowered,
ecalyculate. Female florets tubular-campanulate,
4- or 5-lobed. Anthers caudate; filament collar
short. Style branches apically obtuse. Cypselae
oblong, glabrous; carpopodium distinct. Pappus
bristles numerous, slender. n = 10. Three species,
Mauritius, La Réunion.
544. Faujasia Cass.
Faujasia Cass., Bull. Soc. Philom. Paris 1819: 20 (1819);
Jeffrey, Kew Bull. 47: 78 (1992), rev.; Jeffrey, Fl. Mascar. 109:
139–142 (1993), rev.
Erect shrubs or shrublets. Leaves cauline, sessile, linear to ovate, entire with often serrate
or dentate margins. Capitula many in terminal
corymbiform cymes, radiate or discoid, yellow- or
cream- to white-flowered, calyculate; peduncles
stiff, bracteolate. Anthers caudate. Style branches
apically obtuse. Cypselae oblong, ribbed, glabrous;
carpopodium indistinct, annular. Pappus bristles
numerous, basally flattened. Four species, La
Réunion.
545. Eriotrix Cass.
Eriotrix Cass., Bull. Soc. Philom. Paris 1817: 32 (1817);
Jeffrey, Fl. Mascar. 109: 142–144 (1993), rev.
Erect glabrous shrublets. Leaves cauline, small,
closely appressed, narrowly linear-filiform or
subulate. Capitula solitary, sessile or some subsessile, disciform, white- or yellowish-flowered.
Marginal florets female, tubular. Disc florets
perfect. Anthers sagittate. Style branches truncate
or obtuse, penicillate. Cypselae oblong, ribbed,
glabrous. Pappus bristles several or numerous,
flexuous and entangled. Two species, La Réunion.
546. Hubertia Bory
Hubertia Bory, Voy. Iles Afrique 1: 334, t. 14 (1804); Jeffrey,
Kew Bull. 47: 79–81 (1992), emend., new comb.; Jeffrey, Fl.
Mascar. 109: 147–150 (1993), reg. rev.
Shrubs or suffrutescent herbs, erect or rarely scandent. Leaves cauline, sessile or petiolate, unlobed
or lobed with margins often serrate to dentate. Capitula several to many, corymbose, radiate, disci-
239
form or discoid, yellow- or cream-flowered, calyculate. Anthers caudate; filament collars cylindrical
or only slightly balusterform. Style branches apically obtuse or rounded; stigmatic areas separated
or partly confluent. Cypselae oblong, glabrous or
sometimes pubescent; carpopodium distinct. Pappus bristles numerous, slender. Circa 25 species,
Madagascar, La Réunion, Comores.
547. Humbertacalia C. Jeffrey
Humbertacalia C. Jeffrey, Kew Bull. 47: 82 (1992), 82–83,
rev.; Jeffrey, Fl. Mascar. 109: 132–134 (1993), rev.
Scandent subshrubs. Leaves cauline, petiolate,
pinnately or palmately veined, unlobed with
margins often serrate or dentate. Capitula numerous in axillary or terminal synflorescences,
discoid, few-flowered, calyculate; florets white or
sometimes greenish-yellowish. Anthers caudate.
Style branches apically obtuse or rounded, sometimes with a pointed hair tuft. Cypselae oblong,
glabrous or pubescent; carpopodium distinct.
Pappus bristles numerous, slender. Eight species,
Madagascar, La Réunion.
548. Parafaujasia C. Jeffrey
Parafaujasia C. Jeffrey, Kew Bull. 47: 79 (1992); Jeffrey, Fl.
Mascar. 109: 146 (1993), rev.
Erect shrubs. Leaves cauline, closely set, sessile,
spreading, linear, one-nerved. Capitula numerous
in terminal pyramidal thyrsoid synflorescences,
discoid, yellow-flowered. Anthers caudate. Style
branches apically truncate with short papillae.
Cypselae oblong, glabrous; carpopodium indistinct. Pappus bristles ± thick, straight, scabrid,
free from the base. Two species, Mauritius, La
Réunion.
549. Synotis (C.B. Clarke) C. Jeffrey & Y.L. Chen
Synotis (C.B. Clarke) C. Jeffrey & Y.L. Chen, Kew Bull. 39:
285 (1984); 211, 285–339, reg. rev.; Chen, Fl. Reip. Pop. Sin.
77, 1: 167–217 (1999), reg. rev.
Erect or scrambling herbs or subshrubs with
woody rhizome. Leaves cauline, often assembled
on branch ends, petiolate or sometimes sessile,
pinnately veined, entire or somewhat lobed,
rarely pinnatisect, lanceolate to elliptic-oblong
or ovate, sometimes cordate; margins usually
dentate or serrate. Capitula several to numerous,
corymbose or racemose, radiate, disciform or
discoid, yellow- or cream-flowered, calyculate.
240
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Disc florets perfect; corolla lobes short. Anthers
caudate. Style branches truncate to obtuse, penicillate, sometimes with a tuft of longer papillae.
Cypselae narrowly cylindrical-oblong, glabrous or
sometimes pubescent. Pappus bristles numerous,
slender, white or fulvous to rufous. n = 10, 18,
20. Circa 55 species, eastern Asia, especially
Sinohimalayan region.
550. Mikaniopsis Milne-Redh.
Mikaniopsis Milne-Redh. in Exell, suppl. Cat. Vasc. Pl. S.
Thomé: 27 (1956); Jeffrey, Kew Bull. 41: 878–879 (1986),
reg. rev.; Lisowski, Fragm. Flor. Geobot. 35: 43–53 (1991),
reg. rev.
Scandent perennial herbs or subshrubs. Leaves
cauline, with prehensile petioles, exauriculate,
ovate-cordate, palmately veined. Capitula several,
paniculate-corymbose, disciform, yellow- or
white-flowered, calyculate. Marginal florets female
with tubular-campanulate corolla. Disc florets
perfect. Anthers caudate. Style branches apically
obtuse and appendiculate with a central hair tuft.
Cypselae oblong, ribbed, glabrous. Pappus bristles
numerous, slender. Fifteen species, tropical and
subtropical Africa.
551. Delairea Lem.
Delairea Lem., Ann. Sci. Nat. Bot. sér. 3, 1: 329 (1844).
Scandent-climbing succulent herb. Leaves petiolate, ovate-cordate, palmately veined; petioles auriculate at base. Capitula several, corymbose, discoid, yellow-flowered, calyculate. Anthers caudate.
Style branches obtuse, penicillate. Cypselae oblong,
ribbed, glabrous; carpopodium distinct. Pappus
bristles numerous, slender. n = 10. One species,
D. odorata Lem., South Africa.
552. Cissampelopsis (DC.) Miq.
Cissampelopsis (DC.) Miq., Fl. Ned. Ind. 2: 102 (1856);
Jeffrey & Chen, Kew Bull. 39: 339–351 (1984), reg. rev.;
Koyama, Acta Phytotax. Geobot. 39: 151–163 (1988), reg.
rev.; Chen, Fl. Reip. Pop. Sin. 77, 1: 217–225 (1999), reg. rev.
Scandent perennial herbs or subshrubs. Leaves
cauline, with prehensile petioles, exauriculate,
ovate or cordate to deltoid, palmately veined.
Capitula many, corymbose-paniculate, discoid
or radiate, calyculate. Ray florets yellow. Disc
florets perfect, white to pink or yellow; corolla
lobes lanceolate. Anthers caudate. Style branches
truncate or obtuse, penicillate. Cypselae narrowly
oblong, ribbed, glabrous. Pappus bristles numerous, slender, white or rufescent. Ten species,
eastern and south-eastern Asia (Indomalesia).
553. Austrosynotis C. Jeffrey
Austrosynotis C. Jeffrey, Kew Bull. 41: 828 (1986).
Scandent perennial herb. Leaves cauline, petiolate
with prehensile petioles, auriculate, ovate-cordate,
palmately veined. Capitula several, paniculatecorymbose, radiate, yellow-flowered, calyculate.
Anthers caudate. Style branches truncate, penicillate. Cypselae oblong, glabrous. Pappus bristles
numerus, slender. One species, A. rectirama (Bak.)
C. Jeffrey, Tanzania, Malawi.
554. Dauresia B. Nord. & Pelser
Dauresia B. Nord. & Pelser, Comp. Newslett. 42: 76 (2005).
Glabrous subshrubs with white cortex. Leaves alternate, petiolate, rounded-cordate, dentate, palmately veined. Capitula densely corymbose, discoid, white- or yellow-flowered, ecalyculate. Involucral bracts 5, biseriate. Corolla deeply 5-lobed.
Anthers shortly caudate; endothecial tissue polarized. Style branches apically obtuse with hair tufts.
Cypselae papillate-hirsute on ribs, mucilaginous
when soaked. Pappus bristles numerous, white.
Two species, Namibia.
555. Adenostyles Cass.
Adenostyles Cass., Dict. Sci. Nat. 1, suppl. 59 (1816);
Vierhapper, Oesterr. Bot. Z. 1923 (6–8): 150–164 (1928),
syst. position; Toman et al., Preslia (Praha) 40: 122–132
(1968), syst. position; Tutin, Fl. Eur. 4: 189 (1976), rev.;
Wagenitz, Phyton (Horn) 23: 141–159 (1983), rev.
Cacalia L. (1753 & 1754), nom. rej.
Perennial rhizomatous herbs. Leaves basal and
cauline, petiolate, upper cauline leaves sometimes
sessile and half-clasping, rounded-cordate, entire
with dentate margins, pinnately veined. Capitula
numerous, corymbose, narrow, discoid, calyculate.
Florets 4-lobed, white or purplish. Anther base
ecaudate, obtuse. Style branches elongate, subulate, papillose; stigmatic areas largely continuous.
Cypselae oblong, ribbed, glabrous. Pappus bristles
numerous, slender, persistent. n = 19. Three
species, central and southern Europe.
556. Pojarkovia Askerova
Pojarkovia Askerova, Nov. Syst. Vyssh. Rast. 21: 186 (1984);
Nordenstam, Pl. Syst. Evol. 206: 19–32 (1997), syst.
Compositae
Perennial herb with short thick rhizome. Leaves
cauline, shortly petiolate, lanceolate, entire,
cuneate. Capitula numerous, corymbose, narrow,
discoid, calyculate. Corolla deeply 4-lobed, whitish.
Anther base ecaudate, auriculate; endothecium
transitional (partly polarized). Style branches
long, linear, puberulous, apically truncate, penicillate; stigmatic areas separated. Cypselae oblong,
glabrous. Pappus bristles numerous, slender,
persistent. n = 20. One species, P. stenocephala
(Boiss.) Askerova, Caucasus.
241
Leaves alternate, petiolate, entire to pinnatisect.
Capitula few to several, corymbose-paniculate,
broadly campanulate to hemispherical, radiate,
yellow-flowered. Disc floret corollas 4-lobed.
Anthers caudate. Style branches with continuous
or partially separated stigmatic areas, apically
truncate to obtuse and papillate. Cypselae oblong,
glabrous or shortly villous, ribbed, with a distinct
carpopodium. Pappus bristles numerous, slender,
white to sordid or straw-coloured, persistent.
n = 12, 15–17, 20. Sixteen species, south-western
Asia.
557. Caucasalia B. Nord.
Caucasalia B. Nord., Pl. Syst. Evol. 206: 22 (1997), 19–32,
rev.
XIII. Tribe Calenduleae Cass. (1819).
Perennial rhizomatous herbs, glabrous or
glandular-puberulous. Leaves cauline, petiolate,
cordate or oblong-ovate, with dentate or denticulate margins. Capitula several, corymbose, radiate
or discoid, yellow-flowered, shortly calyculate.
Disc floret corolla 4-lobed. Anthers with radial
endothecium, filament collar basally dilated. Style
branches obtuse to truncate with short sweeping
hairs, stigmatic areas confluent or barely separated. Cypselae oblong, glabrous, ribbed. Pappus
bristles numerous, persistent. n = 19, 38. Four
species, Caucasus, south-western Asia.
B. Nordenstam
558. Dolichorrhiza (Pojark.) Galushko
Dolichorrhiza (Pojark.) Galushko, Nov. Sist. Vyssh. Rast. 6:
210 (1970); Nordenstam, Fl. Iran. 164: 51–53 (1989), reg.
rev.; Nordenstam, Pl. Syst. Evol. 206: 19–32 (1997), syst.
Perennial herbs with a long thin rhizome. Leaves
mostly basal and some cauline, petiolate, roundedcordate or reniform, palmately veined; margins
dentate or lobulate. Capitula solitary or several,
corymbose, radiate or discoid, yellow-flowered, calyculate. Disc florets 4-lobed. Anther base ecaudate,
obtuse. Style branches apically truncate and penicillate, otherwise glabrous; stigmatic areas separated at least basally. Cypselae oblong, ribbed, glabrous; carpopodium distinct. Pappus bristles numerous, slender, caducous. n = 15–16, 20, 22. Four
species, Caucasus, Iran.
559. Iranecio B. Nord.
Iranecio B. Nord., in Rech. f. (ed.) Fl. Iran. 164: 53 (1989),
53–59, reg. rev.; Jeffrey, Kew Bull. 47: 102–103 (1992),
emend., new comb.
Perennial herbs with a short stout rhizome.
Herbs, subshrubs, shrubs or small trees; unarmed
or sometimes spinescent. Leaves alternate or
opposite, sessile or petiolate, entire or variously
lobed or dissected. Capitula solitary or corymbose,
pedunculate or rarely sessile, heterogamous,
radiate. Involucre 1–3-seriate or imbricate; receptacle naked. Ray florets female, fertile or sterile,
rarely neuter, yellow to orange or white, pink to
purple or blue. Disc florets hermaphrodite, perfect
or functionally male, 5-lobed, yellow to orange
or reddish. Anthers caudate; apical appendage
triangular-ovate, flat; endothecial tissue polarized.
Pollen grains spinulose, exine without bacula
(Praglowski and Grafström 1980). Style fertile
or sterile, entire or shortly bilobed to bifurcate;
stigmatic areas divided at least basally; sweeping
hairs in a subapical collar or extending down
the style-branches (Garuleum). Cypselae homomorphic to polymorphic, terete, triquetrous or
flattened, winged or wingless, straight or curved,
sometimes rostrate, glabrous, in some taxa with
a fleshy coloured or whitish exocarp (Fig. 58).
Pappus absent. Chromosome number known
from seven genera, with x = 7, 8, 9, 10, 11 or 15.
Calendula has the most complex karyology, with
all basic numbers represented except 10.
This small tribe currently comprises 12 genera
and about 120 species.
Taxonomically, the tribe is fairly well understood at the species level, based on a lifelong
study by Norlindh (e.g. 1943, 1960, 1977) and
ongoing work by Nordenstam and colleagues
(Nordenstam 1994a, b, 1996, 2004, 2006d; Nordenstam et al. 2006). The generic delimitation still
poses problems, however, the most problematic
242
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Fig. 58. Compositae-Calenduleae. Cypsela morphology.
A Calendula arvensis. Annulate cypsela. B–E Calendula
maderensis. B Falcate, C rostrate, D trialate, E alaterostrate cypsela. F, G Nephrotheca ilicifolia. F Radial side.
G Longitudinal section, showing cavities. H Gibbaria
scabra. Reniform cypsela, radial side. I–N Oligocarpus calendulaceus. I Wingless, J, K trialate, L aculeate with enclosed
apical cavity, M subterete, N rostrate cypsela. O Norlindhia
amplectens. Cypsela trialate with efenestrate small cavity.
P Norlindhia breviradiata. Cypsela trialate with efenestrate
small cavity. Q Norlindhia aptera. Exalate cypsela. R,
S Garuleum bipinnatum. R Triangular-obpyramidate ray
cypsela. S Outer disc cypsela, 2-winged with thickened
margin. T Inuloides tomentosa. Cypsela trialate with large
efenestrate apical cavity. U Tripteris nervosa. Cypsela
trialate trifenestrate. V Osteospermum grandidentatum.
Cypsela cylindrical. W Osteospermum herbaceum. Cypsela
ellipsoid triangulate. X Osteospermum ciliatum. Cypsela
cylindrical, basally sulcate. Y Monoculus monstrosus.
Cypsela trialate unifenestrate. (A–H Norlindh 1962, I–P,
T–Y Norlindh 1943, Q original, R, S Norlindh 1977)
genus being Osteospermum. Although some of its
elements have been recently transferred to other
genera, it remains heterogeneous and further
taxonomic changes can be forseen. Osteospermum
sect. Homocarpa is closely related to the genus
Chrysanthemoides, which is defined by a single
character, the drupaceous fruits. A thin fleshy
and sometimes coloured exocarp is found on the
cypselae of members of sect. Homocarpa and sect.
Coriacea as well, and future changes in generic
taxonomy are inevitable. Pending further studies,
a conservative view on the limits of Osteospermum
and Chrysanthemoides is taken here (cf. Wood
and Nordenstam 2004). Secondary compounds
found in only Calenduleae include two fatty acids,
viz. calendic acid in one group of genera (Calendula, Chrysanthemoides, Osteospermum) and
dimorphecolic acid in another (Dimorphotheca)
(Smith et al. 1960; Barclay and Earle 1965).
Cyanoglycosides, mainly linamarin, also serve
to characterize the genus Dimorphotheca in its
present circumscription, i.e. including Castalis
and Osteospermum sect. Blaxium (Valadon 1977).
Calenduleae are also noteworthy for the high
occurrence of diterpenes with a pimarane skeleton
(Alvarenga et al. 2005). The centre of the tribe
is clearly southern Africa, where all genera but
one occur. Of these, Osteospermum and Tripteris
extend into the northern hemisphere. Calendula is confined to the northern hemisphere,
with a mainly Mediterranean distribution range
extending eastwards as far as Iran.
The habitats of many Calenduleae are the South
African fynbos and karoo vegetation types, also
mountains and deserts. Some of the annuals, such
as Dimorphotheca pluvialis, contribute to the spectacular spring flower displays after good rains in
South Africa. In horticulture, plants of the tribe
are well known since ancient times, such as the
common marigold (Calendula), and more recently
several cultivars of Dimorphotheca are becoming
increasingly popular. The tribe is traditionally regarded as related to Senecioneae (Bremer 1987)
but molecular evidence also indicates affinities to
Astereae, Anthemideae and Gnaphalieae (Panero
and Funk 2002).
Key to the Genera
1. Ray florets female-sterile or neuter
569. Dimorphotheca
– Ray florets female, fertile
2
2. Cypselae polymorphic, some curved and/or rostrate 3
Compositae
– Cypselae homo- or dimorphic, seldom rostrate
4
3. Outer cypselae cymbiform, straight or curved, inner
strongly curved and sometimes annular. Style undivided
560. Calendula
– Cypselae straight or slightly curved, but not annular,
winged or wingless or 6-angled, some rostrate, surface
smooth or rugose to aculeate. Style bifid
561. Oligocarpus
4. Style of disc florets deeply bifurcate and papillatehairy outside
570. Garuleum
– Style of disc florets shallowly bifid with a subapical
collar of sweeping hairs
5
5. Cypselae drupaceous with a distinct black-blue or
orange-red fleshy layer
567. Chrysanthemoides
– Cypselae dry or rarely with a thin whitish or blueblackish fleshy layer
6
6. Disc florets perfect, with flattened cypselae
569. Dimorphotheca
– Disc florets functionally male, with cypselae undeveloped and not flattened
7
7. Cypselae reniform with a ventral cavity
8
– Cypselae straight or slightly curved without ventral
cavity
9
8. Leaves linear-subulate. Involucral bracts imbricate.
Rays white or cream
568. Gibbaria
– Leaves ovate-oblong with dentate margins. Involucral
bracts subuniseriate. Rays yellow 568a.Nephrotheca
9. Rays white, pink or purple
569. Dimorphotheca
– Rays yellow-orange
10
10. Cypselae 3-winged (rarely wings reduced) with apical
cavity
11
– Cypselae winged or not, without apical cavity
566. Osteospermum
11. Disc corolla lobes with sclerenchymatic strands.
Cypselae fenestrate
12
– Disc corolla without sclerenchymatic strands. Cypselae not fenestrate
13
12. Cypselae 3-winged and 3-fenestrate
564. Tripteris
– Cypselae 3-winged, unifenestrate
563. Monoculus
13. Perennial herb with entire tomentose leaves
565. Inuloides
– Annual herbs, glandular-pubescent, leaves lobate to
dentate
562. Norlindhia
Genera of Calenduleae
243
polymorphic, some curved, rostrate, winged or annular. 2n = 14, 16, 18, 22, 28, 30, 32, 36, 42, 44,
46, 54, c. 85. About 15 species (or 10–25, depending on taxonomic opinion), Macaronesia, northern
Africa, Mediterranean region, southern and central
Europe, Anatolia, Yemen, Iraq to Iran.
561. Oligocarpus Less.
Oligocarpus Less., Syn. Gen. Comp.: 90 (1832); Norlindh,
Studies in the Calenduleae I: 346–352 (1943), rev. (as
Osteospermum sect. Oligocarpus); Nordenstam, Comp.
Newslett. 44: 45 (2006).
Annual herbs, glandular-puberulous. Leaves alternate, entire or dentate to lobate. Capitula solitary,
small, radiate. Involucre ± uniseriate. Ray florets
female-fertile, short, yellow. Cypselae polymorphic, curved or straight, winged or wingless, some
rostrate, surface smooth or aculeate. Disc florets
functionally male, yellow. Style shortly bilobed.
Two species, South Africa, St. Helena.
562. Norlindhia B. Nord.
Norlindhia B. Nord., Comp. Newslett. 44: 41 (2006).
Annual herbs, glandular-pubescent with blacktipped glands. Leaves alternate, petiolate or sessile,
sinuato-dentate or denticulate, base half-clasping.
Capitula corymbose, radiate. Involucral bracts
uniseriate or nearly so. Ray florets female, fertile,
yellow to orange. Cypselae homo- or dimorphic
with apical cavity (sometimes small or absent),
without fenestra, 3-winged or wingless, some
often rostrate. Disc florets functionally male,
yellow; corolla without sclerenchymatic strands.
Style apex minutely bifid. Three species, northwestern Cape Province in South Africa, southern
Namibia.
560. Calendula L.
Calendula L., Sp. Pl.: 921 (1753); Lanza, Atti R. Accad. Sci.
Palermo 11: 1–166 (1919), rev.; Meusel & Ohle, Oesterr. Bot.
Z. 113: 191–210 (1966), part. rev.; Heyn et al., Israel J. Bot.
23: 169–201 (1974), part. rev.; Ohle, Feddes Repert. 85: 245–
283 (1974), reg. rev.; Ohle, Feddes Repert. 86: 1–17 (1975),
reg. rev.
Herbs or subshrubs, often glandular and aromatic.
Leaves alternate, sessile, entire or dentate. Capitula pedunculate, radiate. Involucre 1–2-seriate. Ray
florets female-fertile, yellow to orange. Disc florets
functionally male with undivided style. Cypselae
563. Monoculus B. Nord.
Monoculus B. Nord., Comp. Newslett. 44: 39 (2006).
Annual herbs, glandular-pubescent. Leaves alternate, dentate or sinuately lobed. Capitula corymbose, radiate. Involucral bracts biseriate with broad
scarious margins. Ray florets female, fertile, yellow
or orange. Cypselae 3-winged with a unifenestrate
apical cavity. Disc florets functionally male with
blackish-purplish corolla lobes. Style shortly bifid.
2n = 16. Two species, western Cape Province in
South Africa, southern Namibia.
244
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
564. Tripteris Less.
Tripteris Less., Linnaea 6: 95 (1831), nom. cons.; Norlindh,
Studies in the Calenduleae I: 263–346 (1943), rev. (as
Osteospermum subg. Tripteris).
Annual or perennial herbs, subshrubs or shrubs.
Leaves alternate or opposite, entire or variously
lobed. Capitula solitary or corymbose, radiate.
Involucre biseriate to subuniseriate. Ray florets
female-fertile, yellow to orange. Cypselae 3-winged
with an apical trifenestrate cavity, homomorphic.
Disc florets functionally male. Style shortly bifid.
2n = 18, 32. About 20 species, southern and
tropical Africa north to Egypt, Arabia and Jordan.
565. Inuloides B. Nord.
Inuloides B. Nord., Comp. Newslett. 44: 44 (2006).
Tomentose shrublet or small subshrub. Leaves
mainly rosulate, petiolate, spathulate-orbiculate,
entire. Capitula solitary, scapose, radiate. Involucral bracts subuniseriate, inner ones with
scarious-fimbriate margins. Ray florets femalefertile, yellow but turning blue upon drying.
Cypselae 3-winged, with apical cavity but lacking
fenestra, rugose between the wings and apically
glandular-hairy. Disc florets functionally male,
yellow. Style bilobed. Anther appendage margined,
blackish. One species, I. tomentosa (L. fil.) B. Nord.,
south-western Cape Province in South Africa.
566. Osteospermum L.
Fig. 59
Osteospermum L., Sp. Pl.: 923 (1753); Norlindh, Studies in
the Calenduleae I: 98–263 (1943), rev.
Annual or perennial herbs, subshrubs or shrubs.
Leaves alternate or opposite, entire or divided. Capitula radiate. Involucre 1–3-seriate. Ray florets
female-fertile, yellow or orange. Cypselae homoor heteromorphic, straight or slightly curved, terete
or triquetrous, sometimes winged, with or without
apical cavities. Disc florets functionally male. Style
minutely bifid. 2n = 16, 18, 36. About 45 species,
southern and tropical Africa, Somalia and southwestern Arabia.
Fig. 59. Compositae-Calenduleae. Osteospermum burttianum. A Habit. B Ray floret. C Disc floret. D Stamens.
E Style of disc floret. F Cypsela. (Drawing by B. Nordenstam)
dentate. Capitula solitary or corymbose, radiate,
shortly pedunculate. Ray florets female-fertile, yellow. Cypsela drupaceous with a fleshy red, orange,
blue, purple or black exocarp, globose to ovoid.
Disc florets functionally male. Style minutely bifid.
2n = 18, 20, 36. Two, or perhaps up to six species,
southern and eastern tropical Africa, introduced in
St. Helena, Australia, etc.
568. Gibbaria Cass.
567. Chrysanthemoides Fabr.
Chrysanthemoides Fabr., Enum.: 79 (1759); Norlindh, Studies in the Calenduleae I: 367–403 (1943), rev.; Griffoen, M.Sc.
Thesis, Univ. Cape Town (1995), rev.
Shrubs or small trees. Leaves alternate, shortly petiolate, glabrous (pubescent when young), entire or
Gibbaria Cass., Bull. Sci. Soc. Philom. Paris 1817: 139
(1817); Norlindh, Studies in the Calenduleae I: 358–366
(1943), rev.
Erect shrublet. Leaves alternate, linear-subulate,
entire, scabrid. Capitula terminal, 1–3, pedunculate, radiate. Involucre 2–3-seriate. Ray florets
Compositae
245
Glandular smelly shrub or shrublet. Leaves
alternate, ovate-oblong, flat; margins dentate,
scabrid. Capitula terminal, solitary, shortly pedunculate. Involucre subuniseriate. Ray florets
female-fertile, yellow, sometimes reddish-purple
beneath, obliquely laterally attached to the ovary.
Cypselae homomorphic, reniform, with ventral
cavity. Disc florets functionally male, yellow. Style
minutely bilobed. 2n = 20. One species, N. ilicifolia
(L.) B. Nord. & Källersjö, south-western Cape
Province in South Africa.
569. Dimorphotheca Vaill.
Fig. 60
Dimorphotheca Vaill., Königl. Akad. Wissensch. Paris Phys.
Abh. 5: 547 (1754), nom. cons.; Norlindh, Studies in the Calenduleae I: 38–76 (1943), rev.
Blaxium Cass. (1824).
Castalis Cass. (1824).
Xenismia DC. (1836).
Acanthotheca DC. (1838).
Osteospermum sect. Blaxium (Cass.) T. Norl. (1943).
Fig. 60. Compositae-Calenduleae. Cypsela forms in Dimorphotheca. A, B Dimorphotheca polyptera. A Ray cypsela.
B Disc cypsela. C, D D. zeyheri. C Ray cypsela. D Disc
cypsela. E, F D. cuneata. E Ray cypsela. F Disc cypsela. G, H
D. montana. G Ray cypsela. H Disc cypsela. I D. nudicaulis
var. graminifolia. Disc cypsela suborbicular, flattened with
thickened margins. J D. fruticosa. Cypsela triquetrous. K
D. ecklonis. Cypsela acutely triangulate. L D. scabra. Cypsela
triquetrous with incrassate angles. M D. dregei. Cypsela triangulate tuberculate. N D. caulescens. Cypsela obtusely 5–
6-angled. O D. jucunda. Cypsela acutely triangulate with
additional weak rib(s). P D. walliana. Cypsela globose and
basally sulcate. (Norlindh 1943)
female-fertile, white or cream with yellow to
orange reverse. Cypselae homomorphic, reniform with a ventral cavity, glabrous. Disc florets
functionally male, yellow or orange. Style shortly
bilobed. One species, G. scabra (Thunb.) T. Norl.,
southern Cape Province in South Africa.
568a. Nephrotheca B. Nord. & Källersjö
Nephrotheca B. Nord. & Källersjö, Comp. Newslett. 44: 33
(2006).
Annual or perennial herbs, subshrubs or shrubs.
Leaves alternate, entire or divided. Capitula solitary or corymbose, pedunculate, radiate. Involucre uniseriate. Ray florets female-fertile or femalesterile or neuter, yellow or orange, purple or white.
Ray floret cypsela (when present) triquetrous or
subterete, sometimes winged or tuberculate. Disc
florets perfect or functionally male. Style shortly
bilobed. Disc floret cypsela (when present) flattened with thickened margin. 2n = 18, 20. Twenty
species, southern Africa, Zimbabwe, Angola.
570. Garuleum Cass.
Garuleum Cass., Bull. Sci. Soc. Philom. Paris 1819: 172
(1819); Norlindh, Bot. Notiser 130: 377–380 (1977), part.
rev.
Perennial herbs or shrubs. Leaves alternate, sessile,
dentate to pinnatisect. Capitula solitary or corymbose, pedunculate, radiate. Involucre 2–3-seriate,
± imbricate. Ray florets female-fertile, blue or purple to mauve or white or yellow. Cypselae straight,
triquetrous or angular, glabrous. Disc florets perfect or functionally male, yellow. Style deeply bifurcate with linear papillate-hirsute branches. Cypselae (when present) flattened, winged with a thickened rim, glabrous. Eight species, South Africa,
Namibia.
246
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
XIV. Tribe Gnaphalieae (Cass.)
Lecoq & Juillet (1831).
R.J. Bayer, I. Breitwieser, J. Ward
and C. Puttock
Herbs, subshrubs or shrubs (Fig. 61), generally unarmed. Stems generally with fibres in
phloem, without resin canals. Leaves alternate,
rarely opposite, generally entire, and generally
tomentose at least on the lower surface. Capitula
heterogamous disciform or homogamous discoid,
rarely heterogamous radiate, solitary or variously
grouped, sometimes forming dense secondary
heads. Involucral bracts (Figs. 62, 63) generally
papery, generally brightly coloured or hyaline
and with a thickened, cartilaginous basal portion
(stereome) composed of compact sclerenchyma.
Receptacles generally epaleate. Female outer florets
generally filiform or often absent. Central florets
generally perfect, sometimes functionally male.
Anthers ecalcarate, with tails (Fig. 62); endothecial
tissue almost always polarised. Pollen (Fig. 62)
with two-layered ectexine comprising an outer
columellate layer and an irregularly interlaced
basal layer (‘gnaphalioid’ type). Style branches
(Fig. 62) with hairs apically, sometimes apically
and dorsally, or rarely dorsally only; stigmatic rows
generally separated (Fig. 62). Cypselae (Fig. 63)
generally small and oblong to obovoid, usually
hairy; pericarp generally with 2–3 or 5 vascular
bundles. Pappus (Fig. 63) generally of plumose or
barbellate to scabrid capillary bristles, occasionally of bristles and scales, only scales, or absent.
Base chromosome number variable but often
n = 7 (in some groups, 8 or 9). Phytochemistry:
sesquiterpene lactones usually absent.
Comprising 185 genera and about 1,240
species, cosmopolitan, but most diverse in
Australia and South Africa.
The work of Anderberg (1989, 1991a, b, c)
and Karis (1993b) suggests that the tribe Inuleae
(Asteraceae: Asteroideae) sensu Merxmüller et al.
(1977) should be considered as three separate
tribal lineages: Inuleae s.s., Gnaphalieae and
Plucheeae. More recently, however, Anderberg
(in this series) returned Plucheeae to Inuleae
s.s. These recircumscriptions, based in part on
a cladistic analysis of morphological characters,
are supported by two molecular analyses (Kim
and Jansen 1995; Bayer and Starr 1998), both
molecular studies indicating that Inuleae s.l. are
not a monophyletic lineage. In the trnL intron and
trnL/trnF intergenic spacer analysis by Bayer and
Starr (1998), Inuleae s.s. and Plucheeae together
form a clade sister to the remainder of Asteroideae. Kim and Jansen (1995), using ndhF, also
suggest a strong sister relationship of Plucheeae
and Inuleae within Asteroideae. The topological
relationships identified by Bayer and Starr (1998)
were almost identical to those established by Karis
(1993b) based on morphology. Therefore, the
segregation of Gnaphalieae from Inuleae s.l. is
warranted. The sister relationships of Gnaphalieae
proposed by these studies remain controversial.
Karis (1993b), using morphology, revealed them as
sister to a clade containing the tribes Astereae and
Fig. 61. Compositae-Gnaphalieae. Habit. A, B Raoulia eximia, cushion shrub, New Zealand. C Leucogenes grandiceps,
decumbent herb, dense clusters of capitula surrounded by
ring of showy modified leaves, New Zealand. D Bryomorphe aretioides, ericoid shrub, capitula with radiate outer
florets, South Africa. E Xerochrysum bracteatum, upright
herb, colourful papery involucral bracts, Australia. F Leucochrysum albicans, mat-forming herb, white papery involucral bracts, Australia. G Mniodes pulvinulata, suffruticose cushion, Peru. H Gnaphalium supinum, stoloniferous rosette herb, inconspicuous involucral bracts, Eurasia.
(Photographs A–C, F John Lovis, D Marinda Koekemoer,
E Randall Bayer, G Michael Dillon, H Ilse Breitwieser)
Compositae
Anthemideae. Bayer and Starr (1998), based on
trnL intron and trnL/trnF spacer data, proposed
that Gnaphalieae are sister to Senecioneae. The
RFLP (restriction fragment length polymorphism)
analysis of Jansen et al. (1991) placed Gnaphalieae
as sister to Inuleae (represented by Inula L.).
Keeley and Jansen (1991) showed them as sister
to a clade consisting of Inuleae and Plucheeae.
Finally, the ndhF analysis of Kim and Jansen (1995)
had Gnaphalieae in an unresolved clade containing
Calenduleae, Astereae and Anthemideae. Until
more information is at hand, the tribal circumscription and sister relationships of Gnaphalieae
will remain controversial. Some genera which are
currently part of Gnaphalieae (sensu Anderberg
in Bremer 1994) will need to be relocated to other
tribes but, until more data are presented, these
transfers would be premature. Additionally, our
current molecular work on the tribe (Bayer and
Starr 1998; Bayer et al. 2000) indicates that the
subtribes of Gnaphalieae, as delimited by Anderberg (1991a), are non-monophyletic and will
need recircumscription once the final analyses are
completed. It is for this reason that we have chosen
not to place the genera within any subtribal or
informal classification system, as there is no strong
phylogenetic evidence yet to erect such groups. As
more phylogenetic information becomes available,
it will be possible to establish such groups but,
at this stage, this would be premature and would
lead to instability. Therefore, we decided to list the
genera alphabetically. The descriptions of genera
are based largely on Anderberg (1991a), except for
numerous amendments made by us, especially to
Australian, New Zealand and South African taxa.
We recognize 185 genera, in contrast to the 167
which Anderberg (1991a) accepted. Some of the 18
additional genera are ones which were resurrected
by us and some are new genera, described since
Anderberg’s monograph. Also, in light of recent
molecular and reassessed morphological evidence,
Printzia and Isoetopsis have been transferred to
Astereae (Bayer and Cross 2002). A cautionary
note should be expressed concerning the key
to genera. It is very difficult to write a key to
genera of a large and taxonomically problematical
group such as Gnaphalieae. This key should be
considered tentative at this point as, in many cases,
it considers mainly the typical character states in
each genus. Therefore, anomalous species may
prove impossible to identify correctly to genus
with this key.
247
Key to the Genera
1.
–
2.
–
3.
–
4.
–
5.
–
6.
–
7.
–
8.
–
9.
–
10.
–
11.
–
12.
–
13.
–
14.
–
15.
–
16.
–
17.
–
18.
–
19.
–
20.
–
21.
–
22.
–
23.
–
24.
–
25.
–
26.
–
27.
–
Plants perennial
2
Plants annual or biennial
125
Plants woody
3
Plants herbaceous
61
Plants forming alpine cushions
4
Plants shrubby
10
Plants dioecious
5
Plants synoecious
6
Outer florets yellow; Asia
734. Sinoleontopodium
Outer florets purple to blue; South America
692. Mniodes
Habit ericoid
604. Bryomorphe
Habit not ericoid
7
Sweeping hairs on style branches apically and dorsally
or dorsally only
8
Sweeping hairs on style branches apically only
9
Central florets yellow
725. Raouliopsis
Central florets purple
668. Jalcophila
Leaf apex apiculate; Tasmania
719. Pterygopappus
Leaf apex truncate or obtuse; New Zealand
724. Raoulia
Habit ericoid
11
Habit not ericoid
22
Capitulum not narrowly cylindric
12
Capitulum narrowly cylindric
17
Leaves extremely densely set, imbricate
13
Leaves not extremely densely set, not imbricate
14
Central florets purple
704. Phaenocoma
Central florets yellow
631. Dolichothrix
Involucral bracts cartilaginous 669. Lachnospermum
Involucral bracts papery
15
Pappus scabrid or barbellate
664. Hydroidea
Pappus distinctly plumose, at least apically
16
Stereome undivided
718. Pterothrix
Stereome divided
750. Trichogyne
Outer radiate florets present
18
Outer radiate florets absent
19
Involucral bracts with brown papery lamina
578. Amphiglossa
Involucral bracts without lamina
629. Disparago
Involucral bracts cartilaginous
20
Involucral bracts papery
21
Capitulum with one floret
738. Stoebe
Capitulum with two or more florets 633. Elytropappus
Capitula many in corymbs
687. Metalasia
Capitula solitary or only a few together 708. Planea
Capitula heterogamous, radiate or disciform
23
Capitula homogamous, discoid
44
Outer florets filiform or tubular
24
Outer florets radiate
34
Central florets perfect
25
Central florets functionally male
29
Sweeping hairs on style branches apically only
26
Sweeping hairs on style branches dorsally only or apically and dorsally
27
Branches terminating in a thorn
571. Acanthocladium
Branches not terminating in a thorn
700. Ozothamnus
Outer florets purple to blue 650. Gnaphaliothamnus
Outer florets white or yellow
28
248
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
28. Capitula solitary or only a few together; North Africa
576. Aliella
– Capitula many in terminal panicles; Australia
623. Cremnothamnus
29. Leaf lamina flat
30
– Leaf lamina not flat
31
30. Receptacle paleate
703. Petalacte
– Receptacle epaleate
670. Langebergia
31. Leaf margin concave to involute
683. Loricaria
– Leaf margin revolute
32
32. Outer florets purple to blue
33
– Outer florets white or yellow
583. Anderbergia
33. Sweeping hairs on style branches dorsally only; Central and South America
616. Chionolaena
– Sweeping hairs on style branches apically only; South
Africa
581. Anaxeton
34. Outer florets yellow with purple stripes abaxially 35
– Outer florets not discolourous
40
35. Receptacle paleate
36
– Receptacle epaleate
39
36. Central florets with 1–4 pappus bristles 729. Rosenia
– Central florets without pappus bristles
37
37. Subterranean thick woody tubers or rhizomes present
621. Comborhiza
– Subterranean thick woody tubers or rhizomes absent
38
38. Plants glabrous (if hairy, then with capitula congested
in secondary heads)
697. Oedera
– Plants ± hairy or glandular-hairy, capitula generally
solitary
726. Relhania
39. Pappus scales present
681. Leysera
– Pappus scales absent
589. Antithrixia
40. Sweeping hairs on style branches dorsally, not reaching the apex
41
– Sweeping hairs on style branches dorsally and apically
43
41. Pappus scales present
605. Callilepis
– Pappus scales absent
42
42. Leaf margin entire; central florets functionally male
699. Oxylaena
– Leaf margin not entire; central florets perfect
702. Pentatrichia
43. Receptacle paleate
593. Arrowsmithia
– Receptacle epaleate
686. Macowania
44. Leaf lamina flat
45
– Leaf lamina not flat
53
45. Corolla veins reaching ends of lobes; capitula solitary
or only a few together
46
– Corolla veins ending below apex of lobes; capitula
many in corymbs
50
46. Leaf margins entire
47
– Leaf margins not entire
48
47. Receptacle conical, involucral bracts cartilaginous
592. Argyroglottis
– Receptacle flat to convex, involucral bracts papery
700. Ozothamnus
48. Pappus of elongated scale-like awns 586. Anisochaeta
– Pappus of capillary bristles or scales
49
49. Central floret cypsela shorter than corolla
587. Anisothrix
– Central floret cypsela equal to or longer than corolla
702. Pentatrichia
50. Sweeping hairs on style branches dorsally and apically;
style apex acute to conical
610. Catatia
– Sweeping hairs on style branches apically only; style
apex truncate
51
51. Pappus absent
743. Syncephalum
– Pappus present
52
52. Pappus bristles connate
700. Ozothamnus
– Pappus bristles free
736. Stenocline
53. Leaf margin concave to involute
54
– Leaf margin revolute
56
54. Central florets yellow or white
700. Ozothamnus
– Central florets purple
55
55. Capitula 3–6 together, in corymbs 597. Atrichantha
– Capitulum solitary or 2–3 together
608. Calotesta
56. Capitula many in terminal panicles; Madagascar
661. Humeocline
– Capitula many in corymbs; Australia
57
57. Receptacle flat or convex
58
– Receptacle conical
60
58. Pappus absent
655. Haeckeria
– Pappus present
59
59. Anthers with distinct tails
700. Ozothamnus
– Anthers without distinct tails
609. Cassinia
60. Florets 20–30 per capitulum; inner involucral bracts
with claw
667. Ixodia
– Florets 4–6 per capitulum; inner involucral bracts
without claw
696. Odixia
61. Capitula homogamous
62
– Capitula heterogamous
80
62. Stereome divided
63
– Stereome undivided
69
63. Involucral bracts brownish or hyaline
64
– Involucral bracts coloured
66
64. Receptacle flat or convex
573. Acomis
– Receptacle conical
65
65. Pappus of bristles with flattened axis; cypsela without
reddish knobs
680. Leucophyta
– Pappus of rudimentary bristles or of scales; cypsela
with conspicuous reddish knobs
730. Rutidosis
66. Anther apex narrower than thecae
591. Argentipallium
– Anther apex as wide as thecae
67
67. Basal claw on involucral bracts distinct
678. Leucochrysum
– Basal claw on involucral bracts absent
68
68. Receptacle fimbriliferous; cypsela trichomes present
742. Syncarpha
– Receptacle not fimbriliferous; cypsela trichomes absent
737. Stenophalium
69. Plants dioecious, stolons usually present
70
– Plants synoecious, stolons absent
71
70. Leaves tomentose abaxially; pappus bristles connate
in a ring
588. Antennaria
– Leaves glabrous abaxially; pappus bristles free
701. Parantennaria
71. Capitula solitary or only a few together
72
– Capitula many
76
72. Involucral bracts papery
73
– Involucral bracts cartilaginous
586. Anisochaeta
73. Involucral bracts brownish or hyaline; Asia
620. Cladochaeta
– Involucral bracts coloured; Australia
74
74. Basal claw on involucral bract distinct; receptacle
epaleate
75
– Basal claw on involucral bract absent; receptacle
paleate
577. Ammobium
Compositae
75.
–
76.
–
77.
–
78.
–
79.
–
80.
–
81.
–
82.
–
83.
–
84.
–
85.
–
86.
–
87.
–
88.
–
89.
–
90.
–
91.
–
92.
–
93.
–
94.
–
95.
–
96.
–
97.
–
98.
–
99.
Pappus bristles barbellate
584. Anemocarpa
Pappus bristles plumose
678. Leucochrysum
Capitula in glomerules
77
Capitula in panicles
78
Capitula pedicellate
622. Craspedia
Capitula sessile
720. Pycnosorus
Receptacle conical, paleate
590. Apalochlamys
Receptacle flat or convex, epaleate
79
Involucral bracts brownish or hyaline; Australia
607. Calomeria
Involucral bracts coloured; Madagascar
661. Humeocline
Outer florets blue to purple
81
Outer florets yellow, white or pale green
100
Central florets perfect
82
Central florets functionally male or female
97
Sweeping hairs on style branches dorsally and apically
83
Sweeping hairs on style branches apically only or dorsally only
90
Distinct outer florets radiate or absent
84
Distinct outer florets filiform
85
Distinct outer florets radiate
596. Athrixia
Distinct outer florets absent
730. Rutidosis
Style branches acute to conical
86
Style branches obtuse to truncate
88
Pappus bristles scabrid or barbellate without shorter
teeth apically
87
Pappus bristles distinctly plumose but barbellate towards apex with shorter teeth apically
601. Berroa
Cypsela with globose hairs
600. Belloa
Cypsela with elongated twin hairs
684. Lucilia
Pappus bristles free or cohering by patent cilia; Australasia
637. Euchiton
Pappus bristles connate in a ring or in groups; South
America
89
Anther tails distinct; elongated or globose twin hairs
present on cypsela
685. Luciliocline
Anther tails absent; short clavate twin hairs present on
cypsela
614. Chevreulia
Style apex obtuse
91
Style apex truncate
92
Leaf margin entire; receptacle paleate 605. Callilepis
Leaf margin denticulate; receptacle epaleate
675. Lepidostephium
Stereome divided
632. Edmondia
Stereome undivided
93
Cypsela hairs short, clavate
651. Gnaphalium
Cypsela hairs globose, elongated, or absent
94
Capitula solitary in axils of uppermost leaves; receptacle conical
723. Rachelia
Capitula clustered; receptacle flat
95
Receptacle paleate; leaves tomentose abaxially only
745. Tenrhynea
Receptacle epaleate; leaves tomentose on both sides 96
Involucral bracts white; pappus bristles free; New
Zealand
640. Ewartiothamnus
Involucral bracts brown; pappus bristles connate in
a ring; mainly South America
646. Gamochaeta
Leaf margin flat
98
Leaf margin concave to involute
99
Plants synoecious
688. Mexerion
Plants dioecious
624. Cuatrecasasiella
Plants not ericoid; Australia
639. Ewartia
249
– Plants ericoid; South Africa
750. Trichogyne
100. Sweeping hairs on style branches apically and dorsally
101
– Sweeping hairs on style branches only apically or only
dorsally
110
101. Capitula many
102
– Capitula solitary or only few
106
102. Pappus bristles free or cohering by patent cilia 103
– Pappus bristles connate in a ring or in groups, in Euchiton apparently so
104
103. Involucral bracts white
580. Anaphalis
– Involucral bracts brownish or hyaline 637. Euchiton
104. Stereome divided; South America
647. Gamochaetopsis
– Stereome undivided; Eurasia, Australasia
105
105. Central florets functionally male
674. Leontopodium
– Central florets perfect
637. Euchiton
106. Involucral bracts coloured
755. Xerochrysum
– Involucral bracts brownish or hyaline
107
107. Style apex obtuse; stereome undivided
108
– Style apex much prolonged; stereome divided
730. Rutidosis
108. Outer florets filiform
109
– Outer florets radiate
706. Philyrophyllum
109. Leaf margin entire
637. Euchiton
– Leaf margin not entire
705. Phagnalon
110. Sweeping hairs on style branches dorsally only 111
– Sweeping hairs on style branches apically only
112
111. Outer florets filiform
576. Aliella
– Outer florets radiate
605. Callilepis
112. Involucral bracts brownish or hyaline
113
– Involucral bracts coloured
116
113. Capitula surrounded by whorl of white-lanate leaves
679. Leucogenes
– Capitula not surrounded by whorl of white-lanate
leaves
114
114. Outer florets filiform or tubular
115
– Outer florets radiate
698. Oreoleysera
115. Plant forming mats
724. Raoulia
– Plant not forming mats
666. Ixiolaena
116. Stereome divided
117
– Stereome undivided
121
117. Distinct outer florets fewer than central florets or absent
118
– Distinct outer florets more numerous than central florets
120
118. Anther appendages flat, cypselae oblong
119
– Anther appendages concave, cypselae ellipsoid to
turbinate
618. Chrysocephalum
119. Anther apex narrower than thecae
591. Argentipallium
– Anther apex as wide as thecae
659. Helichrysum
120. Capitula with < 20 florets
572. Achyrocline
or
659. Helichrysum
– Capitula with > 20 florets
715. Pseudognaphalium
or
659. Helichrysum
121. Receptacle conical
579. Anaphalioides
– Receptacle flat or convex
122
122. Capitula solitary or only a few together
123
– Capitula many together
124
123. Capitula solitary; New Zealand
724. Raoulia
– Capitula only a few together; South Africa
658. Helichrysopsis
250
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
124. Pappus bristles capillary; South Africa
709. Plecostachys
– Pappus bristles rudimentary or of scales; NE tropical
Africa
615. Chiliocephalum
125. Capitula homogamous
126
– Capitula heterogamous
181
126. Cypsela ellipsoid or turbinate or long and rod-like 127
– Cypsela oblong or obovoid
143
127. Cypsela long and rod-like
617. Chondropyxis
– Cypsela ellipsoid or turbinate
128
128. Involucral bracts in one row
129
– Involucral bracts in more than one row
131
129. Pappus bristles capillary
691. Millotia
– Pappus bristles rudimentary or of scales
130
130. Distinct leaf glands present
748. Toxanthes
– Distinct leaf glands absent
732. Scyphocoronis
131. Involucral bracts with distinct claw
132
– Involucral bracts without claw
135
132. Centremost florets functionally male
697. Schoenia (S. ayersii)
– Centremost florets bisexual
133
133. Style apex obtuse or truncate
134
– Style apex acute to conical
754. Waitzia
134. Style apex obtuse
678. Leucochrysum
– Style apex truncate
657. Haptotrichion
135. Involucral bracts herbaceous
136
– Involucral bracts papery or cartilaginous
137
136. Receptacle epaleate
712. Podotheca
– Receptacle paleate
694. Neotysonia
137. Involucral bracts cartilaginous
138
– Involucral bracts papery
139
138. Pappus bristles capillary
744. Taplinia
– Pappus with a few scale-like bristles
721. Quinetia
139. Cypsela longer than or equal to corolla
140
– Cypsela shorter than corolla
142
140. Receptacle conical
599. Bellida
– Receptacle flat or convex
141
141. Receptacle paleate
694. Neotysonia
– Receptacle epaleate
722. Quinqueremulus
142. Stereome divided; receptacle conical 730. Rutidosis
– Stereome undivided; receptacle flat
648. Gilberta
143. Central floret cypsela oblong
144
– Central floret cypsela obovoid
145
144. Involucral bracts brownish or hyaline
645. Galeomma
– Involucral bracts coloured
753. Vellereophyton
145. Involucral bracts brownish or hyaline
146
– Involucral bracts coloured
168
146. Receptacle paleate
147
– Receptacle epaleate
151
147. Receptacle flat or convex
148
– Receptacle conical or peg-like
150
148. Involucral bracts cartilaginous; pappus of connate
scales
749. Trichanthodium
– Involucral bracts papery; pappus of scales absent 149
149. Capitula surrounded by whorl of leaves
619. Chthonocephalus
– Capitula not surrounded by whorl of leaves
626. Decazesia
150. Receptacle conical
733. Siloxerus
– Receptacle peg-like
622. Craspedia
151. Cypsela trichomes present
152
– Cypsela trichomes absent
162
152. Cypsela densely villous
153
–
153.
–
154.
–
155.
–
156.
–
157.
–
158.
–
159.
–
160.
–
161.
–
162.
–
163.
–
164.
–
165.
–
166.
–
167.
–
168.
–
169.
–
170.
–
171.
–
172.
–
173.
–
174.
–
Cypsela not densely villous
156
Receptacle peg-like
644. Fitzwillia
Receptacle conical
154
Leaf glands distinct
714. Polycalymma
Leaf glands absent
155
Pappus of basally connate subplumose to plumose
bristles
642. Feldstonia
Pappus of short reduced flattened basally connate bristles
630. Dithyrostegia
Receptacle flat or convex
157
Receptacle conical or peg-like
159
Involucral bracts cartilaginous 749. Trichanthodium
Involucral bracts papery
158
Leaf glands distinct
635. Eriochlamys
Leaf glands absent
713. Pogonolepis
Receptacle peg-like
602. Blennospora
Receptacle conical
160
Leaf glands distinct; pappus bristles scabrid or barbellate
161
Leaf glands absent; pappus distinctly plumose, at least
apically
574. Actinobole
Capitula surrounded by whorl of leaves
693. Myriocephalus
Capitula not surrounded by whorl of leaves
652. Gnephosis
Involucral bracts cartilaginous
163
Involucral bracts papery
164
Bracts in more than one row
628. Dielitzia
Bracts in one row
735. Sondottia
Leaves glabrous
673. Lemooria
Leaves tomentose
165
Leaf glands distinct; receptacle conical
662. Hyalochlamys
Leaf glands absent; receptacle flat or convex
166
Leaves alternate throughout; capitula not surrounded
by whorl of leaves
677. Leptotriche
Leaves opposite, at least basally; capitula surrounded
by whorl of leaves
167
Pappus basally connate into a jagged scale-like cup
612. Cephalosorus
Pappus absent
634. Epitriche
Involucral bracts herbaceous
712. Podotheca
Involucral bracts papery
169
Inner involucral bracts conspicuously longer than
outer bracts
170
Inner involucral bracts not conspicuously longer than
outer bracts
176
Pappus present
171
Pappus absent
707. Pithocarpa
Pappus scabrid or barbellate
172
Pappus distinctly plumose, at least apically
174
Leaves filiform, alternate throughout
636. Erymophyllum
Leaves flat, opposite, at least basally
173
Cypsela of outer florets dorsiventrally compressed; 2
vascular bundles medial in relation to cotyledons
731. Schoenia
Cypsela of outer florets not dorsiventrally compressed;
2 vascular bundles lateral in relation to cotyledons
672. Lawrencella
Receptacle flat or convex ; pappus bristles free or cohering by patent cilia
611. Cephalipterum
Receptacle conical; pappus bristles connate in a ring
175
Compositae
175. Cypsela densely villous with elongated twin hairs
727. Rhodanthe
– Cypsela not densely villous, but with globose hairs
without basal cell
663. Hyalosperma
176. Pappus absent
746. Thiseltonia
– Pappus present
177
177. Pappus of jagged scales
710. Pleuropappus
– Pappus of flattened plumose bristles
178
178. Cypsela trichomes absent
649. Gilruthia
– Cypsela trichomes present
179
179. Receptacle paleate
720. Pycnosorus
– Receptacle epaleate
180
180. Involucral bracts usually 4, with 2 concave bracts (always present) surrounding usually 2 (sometimes 0 or
1) flat bracts
585. Angianthus
– Involucral bracts 7–c.22, variously flat to somewhat
concave
606. Calocephalus
181. Central floret cypsela long, rod-like
182
– Central floret cypsela ellipsoid, turbinate, oblong,
obovoid or abortive
183
182. Biennial
695. Nestlera
– Annual
728. Rhynchopsidium
183. Central floret cypsela obovoid
184
– Central floret cypsela ellipsoid, turbinate, or oblong
186
184. Capitula arranged in glomerules or spikes
747. Tietkensia
– Capitula solitary or only a few together
185
185. Inner bracts conspicuously longer than outer bracts,
coloured
672. Lawrencella
– Inner bracts not conspicuously longer than outer
bracts, hyaline
656. Haegiela
186. Central floret cypsela ellipsoid or turbinate
187
– Central floret cypsela oblong
202
187. Outer florets bilabiate
672. Denekia
– Outer florets radiate, filiform, tubular or corolla absent
188
188. Outer florets radiate
189
– Outer florets filiform or tubular, or corolla absent 192
189. Outer florets purple to blue; receptacle paleate
575. Alatoseta
– Outer florets yellow; receptacle epaleate
190
190. Outer florets yellow with purple stripes abaxially;
South Africa
728. Rhynchopsidium
– Outer florets without purple stripes abaxially; Australia
191
191. Bract laminae filiform-subulate, densely plumose
595. Asteridea
– Bract laminae flat, entire to long-ciliate on margins
711. Podolepis
192. Central florets functionally male
598. Basedowia
– Central florets perfect
193
193. Leaf glands distinct
194
– Leaf glands absent
197
194. Involucral bracts brownish or hyaline
195
– Involucral bracts coloured
196
195. Involucral bracts with distinct basal claw
595. Asteridea
– Involucral bracts without distinct basal claw
594. Artemisiopsis
196. Pappus bristle one
654. Gratwickia
– Pappus bristles more than one 618. Chrysocephalum
197. Capitula surrounded by a whorl of leaves
198
– Capitula not surrounded by a whorl of leaves
200
198.
–
199.
–
200.
–
201.
–
202.
–
203.
–
204.
–
205.
–
206.
–
207.
–
208.
–
209.
–
210.
–
211.
–
212.
–
213.
–
214.
–
215.
–
216.
–
217.
–
218.
–
219.
–
220.
–
221.
–
222.
–
223.
–
224.
251
Involucral bracts brownish or hyaline
199
Involucral bracts coloured
717. Pterochaeta
Receptacle conical
751. Triptilodiscus
Receptacle flat or convex
637. Euchiton
Leaves opposite, at least basally
656. Haegiela
Leaves alternate throughout
201
Outer florets purple to blue
641. Facelis
Outer florets yellow
676. Leptorhynchos
Central florets perfect
203
Central florets functionally male
215
Stereome divided
204
Stereome undivided
209
Involucral bracts brownish or hyaline
205
Involucral bracts coloured
207
Outer florets scattered in axils of outer involucral
bracts
665. Ifloga
Outer florets all inside a common involucre
206
Florets purple
671. Lasiopogon
Florets yellow
653. Gnomophalium
Florets yellow
715. Pseudognaphalium
Florets purple
208
Capitula not surrounded by a whorl of leaves
752. Troglophyton
Capitula surrounded by a whorl of leaves
753. Vellereophyton
Receptacle flat or convex
210
Receptacle conical
213
Receptacle paleate
689. Micropsis
Receptacle epaleate
211
Pappus bristles connate in a ring 646. Gamochaeta
Pappus bristles free or cohering by patent cilia 212
Style branches truncate
651. Gnaphalium
Style branches obtuse
637. Euchiton
Inner involucral bracts folded around florets
682. Logfia
Inner involucral bracts not enclosing florets
214
Pappus present
643. Filago
Pappus absent
739. Stuartina
Receptacle peg-like
216
Receptacle flat to conical
222
Inner involucral bracts folded around florets
217
Inner involucral bracts not enclosing florets
218
Corolla of outer florets attached obliquely to cypsela
613. Chamaepus
Corolla of outer florets attached straight to cypsela
638. Evacidium
Anther appendage concave; apex narrower than thecae
219
Anther appendage flat; apex as wide as thecae
221
Leaves opposite, at least basally; cypsela with
trichomes
716. Psilocarphus
Leaves alternate throughout; cypsela without
trichomes
220
Pappus present
741. Stylocline
Pappus absent
582. Ancistrocarphus
Pappus present
625. Cymbolaena
Pappus absent
603. Bombycilaena
Receptacle flat or convex
223
Receptacle conical
226
Outer florets scattered in axils of the outer involucral
bracts
224
Outer florets all inside a common involucre
225
Pappus bristles without patent cilia at base
750. Trichogyne
252
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
– Pappus bristles with patent cilia at base
665. Ifloga
225. Inner involucral bracts enclosing florets
690. Micropus
– Inner involucral bracts not enclosing florets
740. Stuckertiella
226. Receptacle paleate; pappus absent 660. Hesperevax
– Receptacle epaleate; pappus present
643. Filago
Genera of Gnaphalieae
571. Acanthocladium F. Muell.
Acanthocladium F. Muell., Fragm. 2: 155 (1861); Burbidge,
Austral. J. Bot. 6: 229–284 (1958), rev.
Spiny shrub. Leaves alternate, flat with entire margins, tomentose on both surfaces. Capitula solitary.
Involucral bracts papery, brownish, stereome undivided. Receptacle flat. Outer florets filiform, yellow.
Central florets perfect, yellow. Anthers with flat appendages. Style branches truncate, with hairs apically. Cypselae ellipsoid, with globose twin hairs.
Pappus bristles capillary, scabrid or barbellate, connate in groups. One species, A. dockeri F. Muell.,
Australia.
sally. Cypselae ellipsoid, with globose hairs without a basal cell. Pappus absent. Three species,
Australia.
574. Actinobole Fenzl ex Endl.
Fig. 63Q
Actinobole Fenzl ex Endl., Gen. Pl. suppl. 3: 70 (1843);
Short, Muelleria 6: 9–22 (1985), rev.
Gnaphalodes A. Gray (1852), nom. illegit., non Mill. (1754).
Annual herbs. Leaves alternate to subopposite, flat
with entire margins, tomentose on both surfaces.
Capitula many, crowded together among basal
leaves. Involucral bracts papery, hyaline, stereome
undivided. Receptacle conical, epaleate. All florets
perfect, yellow. Anthers with concave appendages.
572. Achyrocline (Less.) DC.
Achyrocline (Less.) DC., Prodr. 6: 219 (1838); Dillon &
Sagástegui-Alva, Fieldiana, Bot. n.s. 26: 1–70 (1991), reg.
rev.
Gnaphalium subg. Achyrocline Less. (1832).
Perennial herbs. Leaves alternate, flat with entire
margins, tomentose on both surfaces. Capitula
many in corymbs, rarely solitary. Involucral bracts
papery, coloured, stereome divided. Receptacle flat, epaleate. Outer florets filiform, yellow.
Central florets perfect, yellow. Anthers with flat
appendages. Style branches truncate, with hairs
apically. Cypselae oblong, with short clavate twin
hairs. Pappus bristles capillary, barbellate, free.
n = 14. Thirty-two species, Africa, Madagascar,
Central and South America.
573. Acomis F. Muell.
Acomis F. Muell., Fragm. 4: 145 (1864); Wilson, Nuytsia 8:
479–483 (1992), reg. rev.
Perennial herbs. Leaves alternate, flat with entire
margins, tomentose on both surfaces. Capitula
solitary or only a few together. Involucral bracts
papery, brownish, stereome divided. Receptacle
flat, epaleate. All florets perfect, yellow. Anthers
with concave appendages. Style branches with
much prolonged apex, with sweeping hairs dor-
Fig. 62.
Compositae-Gnaphalieae. Tribal characters.
A Ozothamnus aggregatus. Transverse section of stem, fibres (arrow) within phloem (×185). B, C Papery involucral
bracts. B Pseudognaphalium luteoalbum. Hyaline lamina,
divided stereome (×25). C Anaphalioides bellidioides.
Showy lamina, undivided stereome (×13). D Ozothamnus
depressus. Stamen with ecalcarate, tailed anther (×67).
E Anaphalis margaritacea. Pollen with 2-layered ectexine,
TEM section (×2,600). F, G Style branches from central
florets. F Ozothamnus leptophyllus. Separate stigmatic
rows, hairs apically (SEM ×115). G Ewartia catipes. Hairs
apically and dorsally (SEM ×112. 5)
Compositae
Style branches truncate, with hairs apically.
Cypselae obovoid, glabrous or with globose hairs
without a basal cell. Pappus bristles with flattened
axis, distinctly plumose, at least apically, connate
in a ring. n = 10. Four species, Australia.
575. Alatoseta Compton
Alatoseta Compton, Trans. Roy. Soc. S. Afr. 19: 314 (1931);
Dyer, The genera of southern African flowering plants
(1975), gen. consp.
Annual herb. Leaves alternate, subterete with entire
margins, tomentose on both surfaces. Capitula
solitary. Involucral bracts cartilaginous, stereome
undivided. Receptacle flat, paleate. Outer florets
radiate, purple. Central florets perfect, yellow.
Anthers with flat appendages. Style branches
obtuse, with hairs dorsally. Cypselae ellipsoid,
glabrous. Pappus a short rim. One species, A. tenuis
Compton, Africa.
576. Aliella Qaiser & Lack
Aliella Qaiser & Lack, Bot. Jahrb. Syst. 106: 488 (1986);
Qaiser & Lack, Bot. Jahrb. Syst. 106: 487–498 (1986), rev.
Perennial subshrubs and herbs. Leaves alternate,
revolute or flat with dentate margins, sparsely tomentose on both surfaces. Capitula solitary. Involucral bracts cartilaginous, brown, stereome undivided. Receptacle flat, epaleate. Outer florets filiform, yellow. Central florets perfect, yellow. Anthers with flat appendages. Style branches obtuse,
with hairs dorsally. Cypselae ellipsoid, with a few
elongated twin hairs. Pappus bristles capillary, barbellate, basally connate. n = 7, 9. Three species,
Africa.
578. Amphiglossa DC.
Amphiglossa DC., Prodr. 6: 258 (1838); Dyer, The genera
of southern African flowering plants (1975), gen. consp.;
Koekemoer, Bothalia 29: 65–75 (1999), rev.
Shrubs. Leaves alternate, flat to twisted with entire margins, sparsely tomentose on both surfaces.
Capitula only a few together, in dense corymbs.
Involucral bracts papery, brown, stereome undivided. Receptacle flat, epaleate. Outer florets radiate, purple. Central florets perfect, purple. Anthers
with flat appendages. Style branches truncate, with
hairs apically. Cypselae ellipsoid, glabrous. Pappus
bristles capillary, plumose, basally connate. Nine
species, Africa.
579. Anaphalioides (Benth.) Kirp.
Fig. 63L
Ammobium R. Br. in J. Sims, Bot. Mag. ad t. 2459 (1824);
Anderberg, Telopea 4: 129–135 (1990), rev.; Orchard, Telopea 5: 1–12 (1992), rev.
Nablonium Cass. (1825).
Perennial herbs. Leaves alternate, flat with entire
margins, tomentose on both surfaces. Capitula solitary or only a few together. Involucral bracts papery, white, stereome undivided. Receptacle conical, paleate. All florets perfect, yellow. Anthers with
concave appendages. Style branches truncate, with
hairs apically. Cypselae ellipsoid, glabrous. Pappus
of two spines. n = 12, 13. Three species, Australia.
Fig. 62C
Anaphalioides (Benth.) Kirp., in Kirpichnikov & Kuprijanova, Trudy Bot. Inst. Akad. Nauk S.S.S.R. ser. 1, Fl. Sist.
Vyssh. Rast. 9: 33 (1950); Glenny, N. Z. J. Bot. 35: 451–477
(1997), rev.; Glenny & Wagstaff, N. Z. J. Bot. 35: 441–449
(1997), phylog., Ward & Breitwieser, N. Z. J. Bot. 36: 165–
171 (1998), reg. rev.
Perennial herbs or subshrubs. Leaves alternate, flat
or revolute with entire margins, tomentose abaxially or on both surfaces. Capitula solitary, only a few
together, or many in corymbs. Involucral bracts papery, white, stereome undivided. Receptacle conical, epaleate. Outer florets filiform, pale green.
Central florets perfect, pale green. Anthers with
flat appendages. Style branches truncate, with hairs
apically. Cypselae oblong to narrowly oblong, generally glabrous. Pappus bristles capillary, scabrid,
free. n = 14, 28, 42. Seven species, New Zealand and
New Guinea.
580. Anaphalis DC.
577. Ammobium R. Br.
253
Fig. 62E
Anaphalis DC., Prodr. 6: 271 (1838); Beauverd, Bull. Soc.
Bot. Genève sér. 2, 5: 146–147 (1913), gen. consp.; Koster,
Blumea 20: 193–226 (1972), reg. rev.; Grierson, Notes Roy.
Bot. Gard. Edinburgh 31: 389–392 (1972), reg. rev.
Subdioecious perennial herbs or shrubs. Leaves alternate, flat or revolute with entire margins, tomentose on both surfaces. Capitula many in corymbs.
Involucral bracts papery, white, stereome divided.
Receptacle flat, epaleate. Outer florets filiform, yellow. Central florets functionally male, yellow. Anthers with flat appendages. Style branches truncate,
with hairs dorsally and apically. Cypselae oblong,
with short clavate twin hairs. Pappus bristles cap-
254
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Fig. 63. Compositae-Gnaphalieae. A Rhodanthe moschata.
Acute pappus apex. B Hyalosperma cotula. Clavate pappus
apex. C Helichrysum calvertianum. Obtuse pappus apex.
D Ozothamnus bidwillii. Scabrid pappus bristle. E Haptotrichion conicum. Barbellate pappus bristle. F Rhodanthe
frenchii. Plumose pappus bristle. G Relhania fruticosa. Pappus scale. H Rutidosis leptorrhynchoides. Fimbriate pappus
scale. I Xerochrysum bracteatum. Flat, honeycombed receptacle. J Triptilodiscus pygmaeus. Peg-like receptacle. K
Helichrysum miconiifolium. Fimbriate receptacle. L Ammobium craspedioides. Paleaceous receptacle. M Anemocarpa
saxatilis. Laminar (clawless) involucral bract. N Helichrysum milfordiae. Clawed involucral bract. O Leucochrysum
stipitatum. Stipitate involucral bract. P Lucilia acutifolia.
Elongated twin cypsela hair. Q Actinobole oldfieldiana. Globose twin cypsela hair. R Rhodanthe anthemoides. Pappus
bristles fused into groups. S Gilberta tenuifolia. Pappus bristles fused into a ring. T Pseudognaphalium luteoalbum.
Pappus bristles coherent by patent cilia. U Helichrysum
adenophorum. Pappus bristles weakly coherent. V Helichrysum aureum. ‘Helichrysum-type’, oblong cypsela with attached floret. W Macowania hamata. ‘Relhania-type’, rodlike cypsela with attached floret. X Gnephosis eriocarpa.
‘Gnephosis-type’, obovoid cypsela with attached floret. Y
Lawrencella davenportii. Triangular style apex. Z Bellida
graminea. Acute style apex. AA Ozothamnus lepidophyllus.
Truncate style apex. BB Chrysocephalum semicalvum. Obtuse style apex. (P, Q modified after Anderberg 1991a; Y–BB
after Wilson 1992; drawings by William Murray)
illary, barbellate, free. n = 7, 13, 14, 21, 24. About
110 species, Asia, North and South America.
581. Anaxeton Gaertn.
Anaxeton Gaertn., Fruct. Sem. Pl. 2: 406 (1791); Lundgren,
Opera Bot. 31 (1972), rev.
Shrubs. Leaves alternate, revolute with entire margins, sparsely tomentose on both surfaces. Capitula many in dense corymbs. Involucral bracts papery, white, stereome undivided. Receptacle flat,
epaleate. Outer florets filiform, purple. Central florets functionally male, purple. Anthers with flat appendages. Style branches truncate, with hairs apically. Cypselae ellipsoid, glabrous. Pappus bristles
capillary, barbellate, basally connate. Ten species,
Africa.
582. Ancistrocarphus A. Gray
Ancistrocarphus A. Gray, Proc. Amer. Acad. Arts 7: 355
(1868).
Compositae
Annual herb. Leaves alternate, flat with entire
margins, tomentose on both surfaces. Capitula
only a few together. Involucral bracts cartilaginous,
hyaline, stereome undivided. Receptacle peg-like,
epaleate. Outer florets filiform, purple. Central
florets functionally male, purple. Style branches
with much prolonged apex, with hairs dorsally.
Anthers with concave appendages. Cypselae
oblong, glabrous. Pappus absent. One species,
A. filagineus A. Gray, North America.
583. Anderbergia B. Nord.
Anderbergia B. Nord., Ann. Wiener Mus. Naturgesch. 98:
407 (1996); Nordenstam, Ann. Wiener Mus. Naturgesch. 98,
B: 403–418 (1996), rev.
Shrubs. Leaves alternate, revolute with entire margins, sparsely tomentose on both surfaces. Capitula many in dense corymbs. Involucral bracts papery, white, stereome undivided. Receptacle flat,
epaleate. Outer florets filiform, purple. Central florets functionally male, purple. Anthers with flat appendages. Style branches truncate, with hairs apically. Cypselae ellipsoid, glabrous. Pappus bristles
capillary, barbellate, free. Six species, Africa.
584. Anemocarpa Paul G. Wilson
Fig. 63M
Anemocarpa Paul G. Wilson, Nuytsia 8: 452 (1992); Wilson,
Nuytsia 8: 447–460 (1992), rev.
Perennial herbs. Leaves alternate, flat with entire
margins, tomentose on both surfaces. Capitula
solitary or only a few together. Involucral bracts
papery, coloured, stereome undivided. Receptacle
convex, epaleate. All florets perfect, yellow. Style
branches obtuse. Cypselae ellipsoid. Pappus
bristles capillary, barbellate, free. Two species,
Australia.
585. Angianthus J.C. Wendl.
Angianthus J.C. Wendl., Col. Pl. 2: 31. t. 48 (1808?), nom.
cons.; Short, Muelleria 5: 143–183 (1983), rev.; Short,
Muelleria 5: 185–214 (1983), rev.
Phyllocalymma Benth. (1837).
Skirrhophorus DC. (1838).
Annual herbs and a single shrub. Leaves alternate or opposite at least basally, flat with entire
margins, tomentose on both surfaces. Capitula
many, forming secondary heads. Involucral bracts
papery, coloured, stereome undivided. General
receptacle peg-like, epaleate. All florets perfect,
yellow. Anthers with concave appendages. Style
255
branches truncate, with hairs apically. Cypselae
obovoid, with globose hairs without basal cells.
Pappus bristles with flattened axis, barbellate or
distinctly plumose apically, connate in a ring.
n = 6, 12, 13. Fifteen species, Australia.
586. Anisochaeta DC.
Anisochaeta DC., Prodr. 5: 109 (1836); Dyer, The genera of
southern African flowering plants (1975), gen. consp.
Perennial subshrub. Leaves alternate, flat with
deeply lobed margins, sparsely tomentose on
both surfaces. Capitula only a few together, in
lax corymbs. Involucral bracts cartilaginous,
stereome undivided. Receptacle flat, epaleate. All
florets perfect, whitish-yellow. Anthers with flat
appendages. Style branches obtuse, with hairs
dorsally. Cypselae ellipsoid, triquetrous, with few
elongated twin hairs. Pappus of several elongated
awns. One species, A. mikanioides DC., southern
Africa.
587. Anisothrix O. Hoffm.
Anisothrix O. Hoffm. in Kuntze, Revis. Gen. Pl. 3: 129
(1898); Anderberg, Bot. Jahrb. Syst. 109: 363–372 (1988),
rev.
Perennial subshrubs. Leaves alternate, flat with
dentate margins, sparsely tomentose on both
surfaces. Capitula solitary. Involucral bracts
cartilaginous, stereome undivided. Receptacle flat,
epaleate. All florets perfect, yellow. Anthers with
flat appendages. Style branches obtuse, with hairs
dorsally. Cypselae ellipsoid, with few elongated
twin hairs. Pappus of scales and capillary bristles,
few, barbellate, free. n = 7. Two species, Africa.
588. Antennaria J. Gaertn.
Fig. 64A
Antennaria J. Gaertn., Fruct. Sem. Pl. 2: 410 (1791), nom.
cons.; Bayer & Stebbins, Syst. Bot. 7: 300–313 (1982), reg.
rev.; Bayer, Syst. Bot. 9: 74–83 (1984), reg. rev.; Bayer, Biol.
Zentralbl. 106: 683–698 (1987), agamic complexes; Bayer,
Taxon 37: 292–298 (1988), nomencl.; Bayer, Brittonia 41:
53–60 (1989), rev.; Dillon & Sagástegui-Alva, Fieldiana, Bot.
n.s. 26: 1–70 (1991), reg. rev.; Bayer, Arctic Alpine Res. 25:
150–159 (1993), reg. rev.; Bayer & Stebbins, Canad. J. Bot.
71: 1589–1604 (1993), rev., syn.
Antennaria Link ex Fr. (1821), non J. Gaertn. (1791).
Dioecious perennial herbs or subshrubs. Leaves alternate, flat with entire margins, tomentose on both
surfaces, abaxially only, or rarely glabrous. Capitula
solitary, only a few together, or many in corymbs.
256
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Involucral bracts papery, coloured, stereome undivided. Receptacle flat, epaleate. Female florets filiform, white or purplish. Male florets tubular, white
or purplish. Anthers with flat appendages. Style
branches truncate, with hairs dorsally and apically.
Cypselae oblong, with short clavate twin hairs. Pappus bristles capillary, barbellate, connate in a ring.
n =14, 21, 28, 42, 49, 56, 70, 84. About 40 species,
Asia, Europe, North and South America.
589. Antithrixia DC.
Antithrixia DC., Prodr. 6: 277 (1838); Bremer, Bot. Notiser
131: 449–453 (1978), rev.
Shrub. Leaves alternate or opposite, revolute or flat
with entire margins, densely tomentose adaxially.
Capitula solitary. Involucral bracts papery, brown,
stereome undivided. Receptacle flat, epaleate.
Outer florets radiate, yellow with purple bands.
Central florets perfect, yellow. Anthers with
flat appendages. Style branches truncate, with
hairs apically. Cypselae cylindrical, glabrous or
with few elongated twin hairs. Pappus bristles
capillary, barbellate, basally connate. One species,
A. flavicoma DC., Africa.
590. Apalochlamys Cass.
Apalochlamys Cass., Dict. Sci. Nat. 56: 223 (1828).
Perennial herb. Leaves alternate, flat, broad with
entire margins, tomentose on both surfaces. Capitula many in terminal panicles. Involucral bracts
papery, brownish, stereome undivided. Receptacle conical, paleate. All florets perfect, yellow. Anthers with flat appendages. Style branches truncate,
with hairs apically. Cypselae ellipsoid, glandularhairy. Pappus bristles capillary, barbellate, connate
in a ring. One species, A. spectabilis (Labill.) J.H.
Willis, Australia.
591. Argentipallium Paul G. Wilson
Argentipallium Paul G. Wilson, Nuytsia 8: 455 (1992); Wilson, Nuytsia 8: 447–460 (1992), rev.
Perennial herbs. Leaves alternate, flat with entire
margins, tomentose on both surfaces or only abaxially. Capitula solitary or many in terminal panicles. Involucral bracts papery, coloured, stereome
divided. Receptacle convex, epaleate. Outer filiform
florets yellow or absent. Central florets perfect, yellow. Anthers with flat appendages. Style branches
obtuse or truncate. Cypselae obovoid, with short
clavate twin hairs. Pappus bristles capillary, barbellate, shortly connate in a ring. n =12. Six species,
Australia.
592. Argyroglottis Turcz.
Argyroglottis Turcz., Bull. Soc. Imp. Naturalistes Moscou 24:
83 (1851); Burbidge, Austral. J. Bot. 6: 229–284 (1958), rev.
Fig. 64. Compositae-Gnaphalieae. A Antennaria pulchella.
Habit (pistillate plant). B Logfia gallica. Habit. C Leontopodium alpinum. Flowering branch. D Gnaphalium uliginosum. Flowering branch. (Drawings by William Murray)
Shrub. Leaves alternate, flat with entire margins,
tomentose on both surfaces. Capitula solitary. Involucral bracts cartilaginous, white, stereome un-
Compositae
divided. Receptacle conical, epaleate. All florets
perfect, yellow. Anthers with flat appendages. Style
branches conical, with hairs dorsally. Cypselae ellipsoid, with elongated twin hairs. Pappus bristles
capillary, barbellate, connate in groups. n =12. One
species, A. turbinata Turcz., Australia.
593. Arrowsmithia DC.
Arrowsmithia DC., Prodr. 7: 254 (1838); Dyer, The genera
of southern African flowering plants (1975), gen. consp.;
Hilliard & Burtt, Notes Roy. Bot. Gard. Edinburgh 42:
227–260 (1985), rev.
Perennial subshrub. Leaves alternate, revolute to
flat with entire margins, sparsely glandular-hairy
on both surfaces. Capitula solitary. Involucral
bracts cartilaginous, stereome undivided. Receptacle flat, epaleate. Outer florets radiate, yellow.
Central florets perfect or functionally male, yellow.
Anthers with flat appendages. Style branches
obtuse, with hairs dorsally. Cypselae ellipsoid,
glabrous or with few elongated twin hairs. Pappus
bristles capillary, few, barbellate, free. One species,
A. styphelioides DC., Africa.
594. Artemisiopsis S. Moore
Artemisiopsis S. Moore, J. Linn. Soc., Bot. 35: 331 (1902);
Merxmüller, Prodromus einer Flora von Südwestafrika 139
(1967), reg. rev.
Annual herb. Leaves alternate, flat with entire margins, sparsely tomentose on both surfaces. Capitula
only a few together, in dense corymbs. Involucral
bracts cartilaginous, stereome divided. Receptacle
flat, epaleate. Outer florets filiform, yellow. Central florets perfect, yellow. Anthers with flat appendages. Style branches truncate, with hairs apically. Cypselae ellipsoid, with few elongated twin
hairs. Pappus bristles capillary, few, barbellate, free.
One species, A. villosa (O. Hoffm.) Schweickerdt,
Africa.
257
appendages. Style branches truncate, with hairs
apically. Cypselae ellipsoid or turbinate, with
globose twin hairs. Pappus bristles capillary,
barbellate or apically plumose, free. n = 7, 9. Nine
species, Australia.
596. Athrixia Ker Gawl.
Athrixia Ker Gawl., Bot. Reg. 8, t. 681 (1823); Kroner, Mitt.
Bot. Staatssamml. München 16: 1–268 (1980), rev.
Klenzea Sch. Bip. ex Walp. (1840).
Perennial herbs. Leaves alternate, revolute with
entire or dentate margins, sparsely tomentose
on both surfaces. Capitula solitary or only a few
together, in lax corymbs. Involucral bracts cartilaginous, stereome undivided. Receptacle flat,
epaleate. Outer florets filiform, purple or white.
Central florets perfect, yellow. Anthers with flat
appendages. Style branches obtuse, with hairs
dorsally. Cypselae ellipsoid, with few elongated
twin hairs. Pappus bristles capillary, barbellate,
free. Fourteen species, Africa.
597. Atrichantha Hilliard & B.L. Burtt
Atrichantha Hilliard & B.L. Burtt, Bot. J. Linn. Soc. 82:
219 (1981); Hilliard & Burtt, Bot. J. Linn. Soc. 82: 181–232
(1981), gen. consp.; Karis, Bot. J. Linn. Soc. 102: 23–36
(1990), rev.
Shrub. Leaves alternate, flat to twisted with
entire margins, sparsely tomentose adaxially.
Capitula three to six in corymbs. Involucral bracts
papery, white, stereome undivided. Receptacle
flat, epaleate. Outer florets absent. Central florets
perfect, purple. Anthers with flat appendages. Style
branches truncate, with hairs apically. Cypselae
ellipsoid, glabrous. Pappus bristles capillary,
barbellate, free. One species, A. gemmifera (Bol.)
Hilliard & B.L. Burtt, Africa.
598. Basedowia Pritzel
595. Asteridea Lindl.
Asteridea Lindl., Sketch Veg. Swan R. 24 (1839); Kroner,
Mitt. Bot. Staatssamml. München 16: 1–268 (1980), rev.;
Short, Austral. Syst. Bot 13: 739–744 (2000), reg. rev.
Annual herbs. Leaves alternate, flat with entire
margins, tomentose on both surfaces. Capitula
solitary. Involucral bracts papery, hyaline, stereome forming two ribs. Receptacle flat, epaleate.
Outer florets radiate, yellow to white. Central
florets perfect, yellow. Anthers with concave
Basedowia Pritzel, Ber. Deutsch. Bot. Gesell. 36: 332 (1918).
Annual herb. Leaves alternate, flat with entire
margins, very sparsely tomentose on both surfaces.
Capitula only a few together, unisexual. Involucral
bracts papery, white, stereome undivided. Receptacle convex, epaleate. Female florets without
corolla. Male florets tubular, yellow. Style branches
truncate, with hairs apically. Cypselae ellipsoid
or turbinate, glabrous. Pappus usually absent, if
present of capillary bristles, barbellate, connate at
258
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
base. One species, B. tenerrima (F. Muell. & Tate)
J.M. Black, Australia.
599. Bellida Ewart
Fig. 63Z
Bellida Ewart, Proc. Roy. Soc. Victoria ser. 2: 34, t. 10, 11
(1907); Wilson, Nuytsia 8: 361–377 (1992), rev.
Annual herb. Leaves alternate, filiform with entire
margins, weakly tomentose on both surfaces. Capitula solitary. Involucral bracts papery, brownish,
stereome undivided. Receptacle conical, epaleate.
All florets perfect, except very innermost ones
functionally male, yellow. Anthers with concave
appendages. Style branches conical, with hairs
dorsally. Cypselae turbinate, with elongated twin
hairs. Pappus bristles capillary, barbellate, violetcoloured, free. n = 9. One species, B. graminea
Ewart, Australia.
towards the apex, connate in groups. One species,
B. gnaphalioides (Less.) Beauverd, South America.
602. Blennospora A. Gray
Blennospora A. Gray, J. Bot. Kew Gard. Misc. 3: 98: 172
(1851); Short, Muelleria 6: 349–358 (1987), rev.
Annual herbs. Leaves mainly alternate but often opposite, at least basally, filiform with entire margins,
tomentose on both surfaces. Capitula many, forming terminal secondary heads. Involucral bracts papery, hyaline, stereome undivided. Receptacle peglike, epaleate. All florets perfect, yellow. Anthers
with concave appendages. Style branches truncate,
with hairs apically. Cypselae obovoid, with globose
hairs without a basal cell. Pappus bristles with flattened axis, distinctly plumose, connate in a ring.
n = 11. Two species, Australia.
600. Belloa Remy
Belloa Remy in Gay, Fl. Chile 3: 336 (1848); Cabrera, Bol.
Soc. Argent. Bot. 7: 79–85 (1958), rev.; Sagástegui-Alva &
Dillon, Phytologia 58: 392–400 (1985), reg. rev.; Freire, Darwiniana 27: 431–490 (1986), rev.; Dillon & Sagástegui-Alva,
Fieldiana, Bot. n.s. 26: 1–70 (1991), reg. rev.
Perennial herbs. Leaves alternate, flat with entire
margins, tomentose on both surfaces. Capitula
solitary or only a few together. Involucral bracts
papery, brownish, stereome usually undivided. Receptacle flat, epaleate. Outer florets filiform, purple.
Central florets perfect, purple. Anthers with flat appendages. Style branches acute, with hairs dorsally.
Cypselae ellipsoid to turbinate, with globose twin
hairs. Pappus bristles capillary, barbellate, connate
in groups. n = 12. About nine species, South
America.
603. Bombycilaena (DC.) Smoljan.
Bombycilaena (DC.) Smoljan., Bot. Mater. Gerb. Bot. Inst.
Komarova Akad. Nauk S.S.S.R. 17: 448 (1955).
Micropus sect. Bombycilaena DC. (1836).
Annual herbs. Leaves alternate, flat with entire
margins, tomentose on both surfaces. Capitula
only a few together. Involucral bracts cartilaginous, brownish, stereome undivided. Receptacle
peg-like, epaleate. Outer florets filiform, purple.
Central florets functionally male, purple. Anthers
with flat appendages. Style branches with much
prolonged apex, with hairs dorsally. Cypselae
oblong, glabrous. Pappus absent. n = 14. Three
species, Asia, Europe and North America.
604. Bryomorphe Harv.
601. Berroa Beauverd
Berroa Beauverd, Bull. Soc. Bot. Genève sér. 2, 5: 210 (1913);
Beauverd, Bull. Soc. Bot. Genève sér. 2, 5: 210–212 (1913),
rev.
Perennial herb. Leaves alternate, flat with entire
margins, tomentose on both surfaces. Capitula
solitary. Involucral bracts papery, brownish,
stereome undivided. Receptacle flat, epaleate.
Outer florets filiform, purple. Central florets
perfect, purple. Anthers with flat appendages.
Style branches acute, with hairs dorsally. Cypselae
turbinate, with elongated twin hairs. Pappus
bristles capillary, distinctly plumose, barbellate
Fig. 61D
Bryomorphe Harv., Thes. Cap. 2: 33 (1863); Dyer, The
genera of southern African flowering plants (1975), gen.
consp.
Shrub. Leaves alternate, twisted, revolute with
entire margins, sparsely tomentose adaxially.
Capitula solitary. Involucral bracts papery, brown,
stereome undivided. Receptacle flat, epaleate.
Outer florets radiate, white. Central florets perfect
or perhaps functionally male, purple. Anthers
with flat appendages. Style branches truncate,
with hairs apically. Cypselae cylindrical, glabrous.
Pappus bristles capillary, barbellate, free. One
species, B. aretioides (Turcz.) Druce, Africa.
Compositae
605. Callilepis DC.
Callilepis DC., Prodr. 5: 671 (1836); Dyer, The genera of
southern African flowering plants (1975), gen. consp.
Zoutpansbergia Hutch. (1946).
Perennial herbs and a shrub. Leaves alternate or
opposite, flat with entire margins, sparsely tomentose on both surfaces. Capitula solitary. Involucral
bracts cartilaginous, stereome undivided. Receptacle flat, paleate. Outer florets radiate, blue or white.
Central florets perfect, purple. Anthers with flat
appendages. Style branches obtuse, with hairs dorsally. Cypselae ellipsoid, triquetrous, with few elongated twin hairs. Pappus of several elongated awns.
Five species, southern Africa.
606. Calocephalus R. Br.
Calocephalus R. Br., Trans. Linn. Soc. London 12: 106
(1817).
Perennial or annual herbs. Leaves opposite, at
least basally, or alternate, flat with entire margins,
tomentose on both surfaces. Capitula many,
forming terminal secondary heads. Involucral
bracts papery, coloured, stereome undivided.
Receptacle peg-like, epaleate. All florets perfect,
yellow. Anthers with concave appendages. Style
branches truncate, with hairs apically. Cypselae
obovoid, with globose hairs without a basal cell.
Pappus bristles with flattened axis, distinctly
plumose apically, connate in a ring. n = 12, 14, 28.
Eleven species, Australia.
607. Calomeria Vent.
Calomeria Vent., Jard. Malmaison ad t. 73 (1804); Heine,
Adansonia sér. 2, 7: 115–140 (1967), rev.
Humea J.E. Sm. (1804).
Biennial herb. Leaves alternate, large, flat with entire margins, weakly tomentose on both surfaces.
Capitula many in large, drooping, terminal panicles composed of many small corymbs. Involucral
bracts papery, pink, stereome undivided. Receptacle flat, epaleate. All florets perfect, purplish.
Anthers with concave appendages. Style branches
truncate, with hairs apically. Cypselae obovoid,
with short clavate twin hairs. Pappus absent. One
species, C. amaranthoides Vent., Australia.
608. Calotesta P.O. Karis
Calotesta P.O. Karis, Bot. J. Linn. Soc. 102: 25 (1990); Karis,
Bot. J. Linn. Soc. 102: 23–36 (1990), rev.
259
Shrub. Leaves alternate, flat to twisted with entire margins, sparsely tomentose adaxially. Capitula only a few together, in small clusters. Involucral
bracts papery, white, stereome undivided. Receptacle flat, epaleate. All florets perfect, purple. Anthers
with flat appendages. Style branches truncate, with
hairs apically. Cypselae ellipsoid, glabrous. Pappus bristles capillary, barbellate, free. One species,
C. alba P.O. Karis, southern Africa.
609. Cassinia R. Br.
Cassinia R. Br., Trans. Linn. Soc. London 12: 126, nom.
cons. (1817); Breitwieser & Ward, N. Z. J. Bot. 35: 125–128
(1997), reg. rev.; Orchard, Austral. Syst. Bot. 17: 469–481
(2004), rev.; Orchard, Austral. Syst. Bot. 17: 505–533 (2004),
rev.; Orchard, Austral. Syst. Bot. 17: 535–566 (2004), rev.;
Orchard, Austral. Syst. Bot. 17: 567–570 (2004), nomencl.;
Orchard, Taxon 54: 199–201 (2005), nomencl.
Shrubs. Leaves alternate, revolute with entire margins, tomentose on both surfaces. Capitula many
in corymbs. Involucral bracts papery, coloured,
stereome undivided. Receptacle flat, usually
paleate. Outer filiform florets, when present,
yellow. Central florets perfect, yellow. Anthers with
flat appendages. Style branches truncate, with hairs
apically. Cypselae ellipsoid, with elongated twin
hairs. Pappus bristles capillary, barbellate, connate
in a ring. n =14. About 20 species, Australia.
610. Catatia Humbert
Catatia Humbert, Mém. Soc. Linn. Normandie 25: 67, 288
(1923).
Shrubs. Leaves alternate, flat with entire margins,
tomentose. Capitula many in corymbs. Involucral
bracts papery, hyaline, inner involucral bracts tubelike and enclosing florets, stereome undivided. Receptacle flat, epaleate. All florets perfect, yellow.
Anthers with flat appendages. Style branches conical, with hairs dorsally. Cypselae oblong, glabrous.
Pappus absent or forming a small dentate rim. Two
species, Madagascar.
611. Cephalipterum A. Gray
Cephalipterum A. Gray, J. Bot. Kew Gard. Misc. 4: 271
(1852).
Annual herb. Leaves alternate, flat with entire margins, tomentose on both surfaces. Capitula many
in terminal clusters. Involucral bracts papery,
coloured, stereome undivided. Receptacle flat,
epaleate. All florets perfect, yellow. Anthers with
260
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
concave appendages. Style branches truncate, with
hairs apically. Cypselae obovoid, with elongated
twin hairs. Pappus bristles capillary with flattened
axis, distinctly plumose apically, free. n =12, 14.
One species, C. drummondii A. Gray, Australia.
612. Cephalosorus A. Gray
Cephalosorus A. Gray, J. Bot. Kew Gard. Misc. 3: 98, 152
(1851); Short, Muelleria 5: 143–183 (1983), rev.
Piptostemma Turcz. (1851).
Annual herb. Leaves opposite throughout, flat with
entire margins, tomentose on both surfaces. Capitula many, forming terminal secondary heads.
Involucral bracts papery, hyaline, stereome undivided. Receptacle flat, epaleate. All florets perfect,
yellow. Anthers with concave appendages. Style
branches truncate, with hairs apically. Cypselae
obovoid, glabrous. Pappus scale-like, connate in
a ring. n =12. One species, C. carpesioides (Turcz.)
P.S. Short, Australia.
613. Chamaepus Wagenitz
Chamaepus Wagenitz in K.H. Rechinger, Fl. Iran. 145: 12
(1980).
Annual herb. Leaves alternate, flat with entire margins, tomentose on both sides. Capitula only a few
together. Involucral bracts cartilaginous and papery at the apex, brownish, stereome undivided.
Receptacle peg-like, epaleate. Outer florets filiform,
purple. Central florets functionally male, purple.
Anthers with flat appendages. Style branches with
hairs dorsally. Cypselae oblong, glabrous. Pappus
absent. One species, C. afghanicus Wagenitz, Middle East.
614. Chevreulia Cass.
Chevreulia Cass., Bull. Sci. Soc. Philom. Paris 1817: 69
(1817); Dillon & Sagástegui-Alva, Fieldiana, Bot. n.s. 26:
1–70 (1991), reg. rev.
Perennial herbs. Leaves opposite, flat with entire
margins, tomentose abaxially. Capitula solitary. Involucral bracts papery, brownish, stereome undivided. Receptacle flat, epaleate. Outer florets filiform, purple. Central florets perfect, purple. Anthers with flat appendages. Style branches obtuse,
with hairs dorsally. Cypselae ellipsoid, with short
clavate twin hairs. Pappus bristles capillary, barbellate, connate in a ring. About six species, South
America.
615. Chiliocephalum Benth.
Chiliocephalum Benth., Icon. Pl. ser. 3, 2: 34, pl. 1137 (1873);
Bentham, Icon. Pl. ser. 3, 2: 34, pl. 1137 (1873), reg. rev.;
Anderberg, Bot. Jahrb. Syst. 110: 1–6 (1988), rev.
Perennial herb. Leaves alternate, flat with entire
margins, sparsely tomentose on both surfaces. Capitula only a few together, in dense corymbs. Involucral bracts papery, yellow, stereome undivided.
Receptacle flat, epaleate. Outer florets filiform, yellow. Central florets functionally male, yellow. Anthers with flat appendages. Style branches truncate,
with hairs apically. Cypselae oblong, glabrous. Pappus absent. One species, C. schimperi Benth., northeastern tropical Africa.
616. Chionolaena DC.
Chionolaena DC., Prodr. 5: 397 (1836); Baker, Fl. Brasil. 6
(1882), reg. rev.; Anderberg & Freire, Notes Roy. Bot. Gard.
Edinburgh 46: 37–41 (1989), reg. rev.; Freire, Ann. Missouri
Bot. Gard. 80: 397–438 (1993), rev.
Pseudoligandra M.O. Dillon & Sagást. (1990).
Parachionolaena M.O. Dillon & Sagást. (1992).
Shrubs. Leaves alternate, revolute with entire margins, tomentose on both surfaces. Capitula solitary
or many in corymbs or dense clusters. Involucral
bracts papery, coloured, stereome undivided. Receptacle flat, epaleate. Outer florets filiform, purple.
Central florets functionally male, purple. Anthers
with flat appendages. Style branches truncate, with
hairs dorsally. Cypselae ellipsoid, generally with
elongated twin hairs or glabrous. Pappus bristles
capillary, barbellate, connate in a ring. n = 14.
Eighteen species, Central and South America.
617. Chondropyxis D.A. Cooke
Chondropyxis D.A. Cooke, in Jessop & Toelken, Fl. S. Australia 3: 1612 (1986).
Annual herb. Leaves subopposite, semi-succulent,
flat with entire margins, glabrous. Capitula solitary.
Involucral bracts cartilaginous, hyaline. Receptacle flat. Outer florets filiform. Central florets perfect. Style branches lanceolate-subulate. Cypselae
linear with dilated apex, glabrous. Pappus bristles
capillary, plumose, connate in a ring. One species,
C. halophila D.A. Cooke, Australia.
618. Chrysocephalum Walp.
Chrysocephalum Walp., Linnaea 14: 503 (1840).
Argyrophanes Schltdl. (1847), nom. illegit.
Fig. 63BB
Compositae
Perennial or annual herbs. Leaves alternate, flat
with entire margins, tomentose on both surfaces.
Capitula solitary or only a few together. Involucral bracts papery, coloured, stereome divided. Receptacle flat, epaleate. Outer florets filiform, yellow. Central florets perfect, yellow. Anthers with
concave appendages. Style branches truncate, with
hairs apically. Cypselae narrowly ellipsoid, with
globose hairs without a basal cell, and one cell overtopping the other. Pappus bristles capillary, barbellate, free. n =12, 24. About seven species, Australia.
619. Chthonocephalus Steetz
Chthonocephalus Steetz in Lehm., Pl. Preiss. 1: 444 (1845);
Short, Muelleria 7: 225–238 (1990), rev.
Rosulate annual herbs. Leaves opposite, flat with
entire margins, tomentose on both surfaces. Capitula many in dense secondary heads surrounded
by leaves. Involucral bracts papery, hyaline, stereome undivided. Receptacle flat, paleate. All florets
perfect, yellow. Anthers with concave appendages.
Style branches truncate, with hairs apically. Cypselae obovoid, with globose hairs without a basal cell.
Pappus absent. Four species, Australia.
620. Cladochaeta DC.
Cladochaeta DC., Prodr. 6: 245 (1838).
Perennial herbs. Leaves alternate, flat with entire
margins, tomentose on both surfaces. Capitula only
a few together. Involucral bracts papery, brownish, stereome undivided. Receptacle flat, epaleate.
All florets perfect, yellow. Anthers with flat appendages. Style branches truncate and with hairs
apically. Cypselae oblong, with globose twin hairs.
Pappus bristles capillary, barbellate, basally connate in groups. n =8, 9. Two species, Eurasia.
621. Comborhiza Anderb. & K. Bremer
Comborhiza Anderb. & K. Bremer, Ann. Missouri Bot.
Gard. 78: 1070 (1991); Anderberg & Bremer, Ann. Missouri
Bot. Gard. 78: 1061–1072 (1991), rev.
Shrubs with stems arising from subterranean,
thick, woody tubers or rhizomes. Leaves alternate,
flat with entire margins, sparsely tomentose
abaxially. Capitula solitary. Involucral bracts
papery, brown, stereome undivided. Receptacle
flat to convex, paleate. Outer florets radiate, yellow
with purple bands. Central florets perfect, yellow.
Anthers with flat appendages. Style branches
truncate, with hairs apically. Cypselae ellipsoid,
261
glabrous or with few elongated twin hairs. Pappus
scales and bristles capillary, few, barbellate, basally
connate. Two species, Africa.
622. Craspedia G. Forst.
Craspedia G. Forst., Fl. Ins. Austral. 58 (1786); Ward &
Breitwieser, N. Z. J. Bot. 36: 165–171 (1998), reg. rev.;
Breitwieser et al., N. Z. J. Bot. 37: 399–412 (1999), phylog.
Richea Labill. (1800).
Perennial or annual herbs. Leaves alternate, in
a basal rosette and sometimes cauline, flat with
entire margins, tomentum various. Capitula
many in dense glomerules. Involucral bracts
papery, brownish or hyaline, stereome undivided.
Receptacle peg-like, paleate. All florets perfect,
white, yellow or orange. Anthers with concave
appendages. Style branches truncate, with hairs
apically. Cypselae obovoid, with papillae, glandular hairs, or elongated or globose twin hairs.
Pappus bristles with flattened axis, distinctly
plumose, connate in a ring. n =11, 22, 33, 44, 55.
About 20 species, Australia, New Zealand.
623. Cremnothamnus C.F. Puttock
Cremnothamnus C.F. Puttock, Austral. Syst. Bot. 7: 576
(1994); Puttock, Austral. Syst. Bot. 7: 569–583 (1994), rev.
Perennial subshrub. Leaves alternate, flat with entire margins, weakly tomentose on both surfaces.
Capitula many in terminal panicles composed of
many small corymbs. Involucral bracts papery,
coloured, stereome undivided. Receptacle convex,
epaleate. Outer florets filiform, yellow. Central florets perfect, yellow. Anthers with flat appendages.
Style branches obtuse, with hairs dorsally and
apically. Cypselae turbinate, with elongated twin
hairs. Pappus bristles capillary, barbellate, weakly
cohering in a ring. One species, C. thomsonii
(F. Muell.) C.F. Puttock, Australia.
624. Cuatrecasasiella H. Rob.
Cuatrecasasiella H. Rob., Fl. Neotrop. 2, suppl.: 14 (1985);
Robinson, Fl. Neotrop. 2, suppl.: 13–16 (1985), rev.; Dillon &
Sagástegui-Alva, Fieldiana, Bot. n.s. 26: 1–70 (1991), reg.
rev.
Dioecious perennial herbs. Leaves opposite, flat
with entire margins, tomentose on both surfaces.
Capitula solitary, sessile. Involucral bracts papery,
brownish, stereome undivided. Receptacle flat,
epaleate. Female florets filiform, purple. Male
florets purple. Anthers with flat appendages. Style
262
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
branches acute to conical, with hairs dorsally.
Cypselae turbinate, glabrous. Pappus bristles capillary, barbellate, connate in a ring. Two species,
South America.
625. Cymbolaena Smoljan.
Cymbolaena Smoljan., Bot. Mater. Gerb. Bot. Inst. Komarova Akad. Nauk S.S.S.R. 17: 452 (1955); Wagenitz,
Oesterr. Bot. Z. 119: 399–403 (1971), rev.
Annual herb. Leaves alternate, flat with entire
margins, tomentose on both surfaces. Capitula
only a few together. Involucral bracts cartilaginous, brownish, stereome undivided. Receptacle
peg-like, epaleate. Outer florets filiform, purple.
Central florets functionally male, purple. Anthers
with flat appendages. Style branches with much
prolonged apex, with hairs dorsally and apically.
Cypselae oblong, with short clavate twin hairs.
Pappus bristles capillary, scabrid, cohering by
patent cilia. One species, C. griffithii (A. Gray)
Wagenitz, Middle East, Asia.
626. Decazesia F. Muell.
Decazesia F. Muell., Fragm. 11: 71 (1879).
Annual herb. Leaves alternate, flat with entire margins, tomentose on both surfaces. Capitula many in
loose globose secondary heads. Involucral bracts
papery, hyaline, stereome undivided. Receptacle
flat, paleate. All florets perfect, yellow. Anthers with
concave appendages. Style branches truncate, with
hairs apically. Cypselae obovoid, with globose hairs
without a basal cell. Pappus bristles with flattened
axis, barbellate, connate in a ring. n = 14. One
species, D. hecatocephala F. Muell., Australia.
627. Denekia Thunb.
Denekia Thunb., Prodr. Fl. Cap. 2: 153 (1800); Dyer, The
genera of southern African flowering plants (1975), gen.
consp.
Annual or perennial herb. Leaves alternate, flat with
dentate margins, densely tomentose abaxially. Capitula only a few together, in dense corymbs. Involucral bracts cartilaginous, stereome undivided.
Receptacle flat, epaleate. Outer florets bilabiate,
blue or white. Central florets functionally male,
blue or white. Anthers with concave appendages.
Style branches truncate, with hairs apically. Cypselae ellipsoid, with few elongated twin hairs. Pappus bristle one, capillary, plumose. One species,
D. capensis Thunb., Africa.
628. Dielitzia P.S. Short
Dielitzia P.S. Short, Muelleria 7: 103 (1989); Short, Muelleria
7: 103–116 (1989), rev.
Rosulate annual herb. Leaves opposite, at least
basally, narrowly linear, flat with entire margins,
tomentose on both surfaces. Capitula many in
dense secondary heads among the leaves. Involucral bracts cartilaginous, brown. All florets
perfect, yellow. Style branches truncate, with
hairs apically. Cypselae obovoid. Pappus bristles
capillary, barbellate, connate in a ring. n =13. One
species, D. tysonii P.S. Short, Australia.
629. Disparago Gaertn.
Disparago Gaertn., Fruct. Sem. Pl. 2: 463 (1791); Levyns,
J. S. African Bot. 2: 95–103 (1936), rev.; Koekemoer, Bothalia
21: 158–161 (1991), reg. rev.; Koekemoer, Bothalia 23:
197–206 (1993), rev.
Shrubs. Leaves alternate, flat to twisted, with entire
margins, sparsely tomentose adaxially. Capitula
only a few together, in dense corymbs. Involucral
bracts cartilaginous, stereome undivided. Receptacle flat, epaleate. Outer florets, when present,
radiate, purple to pink or white. Central florets
perfect, purple. Anthers with flat appendages. Style
branches truncate, with hairs apically. Cypselae
ellipsoid, glabrous. Pappus bristles capillary,
plumose, free. n = 9. Seven species, Africa.
630. Dithyrostegia A. Gray
Dithyrostegia A. Gray, J. Bot. Kew Gard. Misc. 3, 97, 100
(1851); Short, Muelleria 5: 185–214 (1983), rev.; Short, Muelleria 7: 103–116 (1989), rev.
Annual herbs. Leaves alternate, flat with entire margins, tomentose on both surfaces. Capitula many in
dense secondary heads surrounded by two basally
connate leaves. Involucral bracts papery, hyaline,
stereome undivided. Receptacle conical, epaleate.
All florets perfect, yellow. Anthers with concave appendages. Style branches truncate, with hairs apically. Cypselae obovoid, with elongated twin hairs.
Pappus bristles short, flat, connate in a ring. Two
species, Australia.
631. Dolichothrix Hilliard & B.L. Burtt
Dolichothrix Hilliard & B.L. Burtt, Bot. J. Linn. Soc. 82:
222 (1981); Hilliard & Burtt, Bot. J. Linn. Soc. 82: 181–232
(1981), rev.
Shrub. Leaves alternate, flat with entire margins,
sparsely tomentose adaxially. Capitula solitary.
Compositae
Involucral bracts papery, white, stereome undivided. Receptacle flat, epaleate. All florets
perfect, yellow. Anthers with flat appendages. Style
branches truncate, with hairs apically. Cypselae
ellipsoid, with elongated twin hairs. Pappus
bristles capillary, barbellate, basally connate. One
species, D. ericoides (Lam.) Hilliard & B.L. Burtt,
Africa.
632. Edmondia Cass.
Edmondia Cass., Bull. Sci. Soc. Philom. Paris 1818: 75 (1818).
Perennial subshrubs. Leaves alternate, involute, adpressed, with entire margins, sparsely tomentose
adaxially. Capitula solitary. Involucral bracts papery, white or pink, stereome divided. Receptacle
flat, epaleate. Outer florets filiform, purple. Central florets perfect, yellow. Anthers with concave
appendages. Style branches truncate, with hairs
apically. Cypselae oblong, with short clavate twin
hairs. Pappus bristles capillary, barbellate, basally
connate. Three species, Africa.
633. Elytropappus Cass.
Elytropappus Cass., Bull. Soc. Sci. Philom. Paris 1816: 199
(1816); Levyns, J. S. African Bot. 1: 89–103 (1935), rev.; Nordenstam, J. S. African Bot. 30: 45–48 (1964), nomencl.
Achyrocome Schrank (1824).
Shrubs. Leaves alternate, flat to twisted with entire margins, sparsely tomentose adaxially. Capitula solitary or many in dense corymbs. Involucral
bracts cartilaginous, stereome undivided. Receptacle flat, epaleate. All florets perfect, purple. Anthers
with flat appendages. Style branches truncate, with
hairs apically. Cypselae ellipsoid, glabrous. Pappus
bristles capillary, plumose, basally connate. Eight
species, southern Africa.
634. Epitriche Turcz.
Epitriche Turcz., Bull. Soc. Imp. Naturalistes Moscou 24: 74
(1851); Short, Muelleria 5: 143–183 (1983), rev.
Pteropogon sect. Pteropogonopsis A. Gray (1852).
Annual herb. Leaves opposite, flat with entire margins, tomentose on both surfaces. Capitula many
in dense secondary heads. Involucral bracts papery, hyaline, stereome undivided. Receptacle flat,
epaleate. All florets perfect, yellow. Anthers with
concave appendages. Style apices truncate, with
hairs apically. Cypselae obovoid, glabrous. Pappus absent. One species, E. demissus (A. Gray)
P.S. Short, Australia.
263
635. Eriochlamys Sond. & F. Muell.
Eriochlamys Sond. & F. Muell. in Sond., Linnaea 25: 488
(1853).
Annual herb. Leaves opposite, flat with entire margins, tomentose on both surfaces. Capitula many
in terminal secondary heads. Involucral bracts papery, hyaline, stereome undivided. Receptacle flat,
epaleate. All florets perfect, yellow. Anthers with
concave appendages. Style branches truncate, with
hairs apically. Cypselae obovoid, with globose hairs
without a basal cell. Pappus absent. n = 14. One
species, E. behrii Sond. & F. Muell., Australia.
636. Erymophyllum Paul G. Wilson
Erymophyllum Paul G. Wilson, Nuytsia 7: 105 (1989); Wilson, Nuytsia 7: 103–116 (1989), rev.
Annual herbs. Leaves alternate, filiform with entire margins, tomentose on both surfaces. Capitula only a few together, in terminal clusters. Involucral bracts papery, coloured, stereome undivided. Receptacle flat, epaleate. All florets perfect,
yellow. Anthers with concave appendages. Style
branches truncate, with hairs apically. Cypselae
obovoid, with elongated twin hairs. Pappus bristles with flattened axis, barbellate, free. n = 11, 14.
Five species, Australia.
637. Euchiton Cass.
Euchiton Cass., Dict. Sci. Nat. 56: 214 (1828); Koster, Blumea
20: 193–226 (1972), reg. rev.; Drury, N.Z.J. Bot. 10: 112–179
(1972), reg. rev.; Breitwieser & Sampson, Grana 36: 65–79
(1997), palynol.; Breitwieser & Sampson, Grana 36: 80–95
(1997), palynol.; Breitwieser & Ward, N. Z. J. Bot. 36: 303–
304 (1998), rev.; Ward & Breitwieser, N. Z. J. Bot. 36: 165–171
(1998), rev.; Breitwieser et al., N. Z. J. Bot. 37: 399–412 (1999),
phylog.
Gnaphalium sect. Euchiton (Cass.) DC. (1837)
Perennial or rarely annual, generally stoloniferous
herbs. Leaves alternate, generally flat with entire
margins, tomentose on both surfaces or abaxially
only. Capitula solitary or few to many in dense
clusters. Involucral bracts papery, brownish or hyaline, stereome undivided. Receptacle flat, epaleate.
Outer florets filiform, purple or colourless. Central florets perfect, purple or colourless. Anthers
with flat appendages. Style branches obtuse, with
hairs dorsally and apically. Cypselae small, oblong or larger and narrowly ellipsoid-oblong, generally with short clavate twin hairs. Pappus bristles
capillary, scabrid, free or cohering by patent cilia.
264
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
n = 14. Twenty species, New Zealand, Australia,
New Guinea, Asia, Oceania.
638. Evacidium Pomel
Evacidium Pomel, Mat. Fl. Atl. 41 (1874).
Annual herb, sometimes perennating. Leaves
alternate, flat with entire margins, tomentose.
Capitula only a few together. Involucral bracts
cartilaginous or apically with a minute papery
portion, brownish, stereome undivided. Receptacle peg-like, epaleate. Outer florets filiform,
purple. Central florets functionally male, purple.
Anthers with flat appendages. Style branches
with a much prolonged sterile apex, with hairs
dorsally. Cypselae oblong, with globose twin hairs.
Pappus absent. One species, E. atlanticum Pomel,
south-eastern Europe and northern Africa.
639. Ewartia Beauverd
Fig. 62G
Ewartia Beauverd, Bull. Soc. Bot. Genève sér. 2, 2: 236
(1910); Beauverd, Bull. Soc. Bot. Genève sér. 2, 3: 253–260
(1911), rev.; Breitwieser & Ward, N. Z. J. Bot. 31: 43–58
(1993), morph.; Breitwieser & Sampson, Grana 36: 65–79
(1997), palynol.; Breitwieser & Sampson, Grana 36: 80–95
(1997), palynol.; Ward & Breitwieser, N. Z. J. Bot. 36:
165–171 (1998), rev.; Breitwieser et al., N. Z. J. Bot. 37:
399–412 (1999), phylog.
Subdioecious or dioecious, perennial, mat-forming
herbs. Leaves alternate, concave with entire margins, tomentose on both surfaces. Capitula solitary
or few together. Involucral bracts papery, white,
stereome divided or undivided. Receptacle flat,
epaleate. Outer florets filiform, purple. Central
florets purple. Anthers with flat appendages. Style
branches obtuse, with hairs dorsally and apically.
Cypselae oblong to obovoid, with elongated or
short clavate twin hairs. Pappus bristles capillary,
scabrid, connate in a ring or in groups. n = 14, 28.
Four species, Australia.
640. Ewartiothamnus Anderb.
Ewartiothamnus Anderb., Opera Bot. 104: 94 (1991);
Anderberg, Opera Bot. 104: 1–195 (1991), rev.; Breitwieser
& Ward, N. Z. J. Bot. 31: 43–58 (1993), morph.; Breitwieser
& Sampson, Grana 36: 65–79 (1997), palynol.; Breitwieser
& Sampson, Grana 36: 80–95 (1997), palynol.; Ward &
Breitwieser, N. Z. J. Bot. 36: 165–171 (1998), rev., Breitwieser
et al., N. Z. J. Bot. 37: 399–412 (1999), phylog.
Perennial herb. Leaves alternate, flat with entire
margins, tomentose on both surfaces. Capitula
many in corymbs. Involucral bracts papery, white,
stereome undivided. Receptacle flat, epaleate.
Outer florets filiform, purple. Central florets perfect, purple. Anthers with flat appendages. Style
branches truncate, with hairs apically. Cypselae
ellipsoid to obovoid, glabrous. Pappus bristles
capillary, scabrid, free. n = 14. One species,
E. sinclairii (Hook. f.) Anderb., New Zealand.
641. Facelis Cass.
Facelis Cass., Bull. Sci. Soc. Philom. Paris 1819: 94 (1819);
Beauverd, Bull. Soc. Bot. Genève sér. 2, 5: 212–220 (1913),
rev.; Robinson, Fl. Neotrop. 2, suppl.: 13–16 (1985), reg.
rev.; Dillon & Sagástegui-Alva, Fieldiana, Bot. n.s. 26: 1–70
(1991), reg. rev.
Annual herbs. Leaves alternate, flat with entire
margins, tomentose on both surfaces. Capitula
solitary or only a few together. Involucral bracts
papery, brownish, stereome undivided. Receptacle
flat, epaleate. Outer florets filiform, purple. Central
florets perfect, purple. Anthers with flat appendages. Style branches acute with hairs dorsally.
Cypselae turbinate, with elongated twin hairs.
Pappus bristles capillary, distinctly plumose below,
barbellate apically, connate in groups. n = 14. Four
species, South America.
642. Feldstonia P.S. Short
Feldstonia P.S. Short, Muelleria 7: 108 (1989); Short,
Muelleria 7: 103–116 (1989), rev.
Annual herb. Leaves opposite basally, alternate
above, flat with entire margins, generally glabrous.
Capitula many in terminal dense secondary
heads. Involucral bracts cartilaginous, hyaline.
Receptacle conical, epaleate. All florets perfect,
yellow. Style branches truncate, with hairs apically.
Cypselae obovoid, hairy. Pappus bristles capillary,
subplumose to plumose, connate in a ring. n = 11.
One species, F. nitens P.S. Short, Australia.
643. Filago L.
Filago L., Sp. Pl. 2: 927 (1753), nom. cons.; Chrtek & Holub,
Preslia 35: 1–17 (1963), rev.; Wagenitz, Willdenowia 4:
37–59 (1965), rev.; Wagenitz, Ber. Deutsch. Bot. Gesell. 79:
336–342 (1966), rev.; Wagenitz, Willdenowia 4: 283–298
(1968a), reg. rev., Wagenitz, Willdenowia 5: 55–66 (1968b),
rev.; Wagenitz, Willdenowia 5: 395–444 (1969), rev.; Wagenitz, Willdenowia 6: 115–139 (1970), reg. rev.; Wagenitz,
Feddes Repert. 81: 107–117 (1970), rev.; Wagenitz, Israel
J. Bot. 19: 260–265 (1970), reg. rev.; Wagenitz, Sida 6:
221–223 (1976), reg. rev.
Compositae
Evax Gaertn. (1791).
Gifola Cass. (1819).
Filaginopsis Torr. & A. Gray (1842).
Evacopsis Pomel (1874).
Annual herbs. Leaves alternate, flat with entire
margins, tomentose on both surfaces. Capitula
only a few together. Involucral bracts cartilaginous,
brownish, stereome undivided. Receptacle conical,
epaleate. Outer florets filiform, purple. Central
florets perfect or functionally male, purple.
Anthers with flat appendages. Style branches
with much prolonged apex, with hairs dorsally.
Cypselae oblong, with short clavate twin hairs.
Pappus bristles capillary, scabrid, free. n = 9, 13,
14. Forty-six species, Asia, Europe, North Africa
and North America.
644. Fitzwillia P.S. Short
Fitzwillia P.S. Short, Muelleria 7: 111 (1989); Short,
Muelleria 7: 103–116 (1989), rev.
Annual herb. Leaves opposite, flat with entire
margins, semi-succulent, glabrous. Capitula many
in flattened secondary heads. Involucral bracts
cartilaginous, hyaline. Receptacle peg-like. All
florets perfect, white. Style branches truncate, with
hairs apically. Cypselae obconic, villous. Pappus
of scales, connate in a ring forming a cup. One
species, F. axilliflora (W.V. Fitzg. ex Ewart & Jean
White) P.S. Short, Australia.
645. Galeomma Rauschert
265
Capitula many in head-like or spike-like clusters.
Involucral bracts papery, brownish, stereome undivided. Receptacle flat, epaleate. Outer florets filiform, purple. Central florets perfect, purple. Anthers with flat appendages. Style branches truncate, with hairs apically. Cypselae oblong, with globose twin hairs. Pappus bristles capillary, barbellate, connate in a ring. n = 14. About 50–80 species,
South America.
647. Gamochaetopsis Anderb. & Freire
Gamochaetopsis Anderb. & Freire, Bot. J. Linn. Soc. 106:
186 (1991); Anderberg & Freire, Bot. J. Linn. Soc. 106:
173–198 (1991), rev.
Perennial herb. Leaves alternate, flat with entire
margins, tomentose on both surfaces. Capitula few
together in small, flat-topped glomerules. Involucral bracts papery, brownish, stereome divided.
Receptacle flat. Outer florets filiform. Central
florets perfect. Style branches truncate, with hairs
dorsally and apically. Cypselae oblong-elliptic,
with short clavate twin hairs. Pappus bristles
capillary, scabrid, connate in a ring. One species,
G. alpina (Poepp. & Endl.) Anderb. & S.E. Freire,
South America.
648. Gilberta Turcz.
Fig. 63S
Gilberta Turcz., Bull. Soc. Imp. Naturalistes Moscou 24: 192
(1851); Wilson, Nuytsia 8: 379–438 (1992), rev.
Antheidosorus A. Gray (1851).
Annual herbs. Leaves alternate, flat with entire margins, sparsely tomentose on both surfaces. Capitula
many in dense corymbs. Involucral bracts papery,
hyaline, stereome divided. Receptacle flat, epaleate.
All florets perfect, yellow. Anthers with flat appendages. Style branches truncate, with hairs apically. Cypselae oblong, with globose hairs. Pappus
bristles capillary, barbellate, free or cohering by
patent cilia. Two species, southern Africa.
Annual herb. Leaves alternate, filiform with entire margins, tomentose on both surfaces. Capitula many in terminal secondary heads. Involucral
bracts papery, hyaline, stereome undivided. Receptacle flat, epaleate. All florets perfect or some central ones functionally male, yellow. Anthers with
concave appendages. Style branches truncate, with
hairs apically. Cypselae ellipsoid, with elongated
twin hairs. Pappus bristles with flattened axis, distinctly plumose, connate in a ring. n = 10. One
species, G. tenuifolia Turcz., Australia.
646. Gamochaeta Wedd.
649. Gilruthia Ewart
Gamochaeta Wedd., Chlor. Andina 1: 151 (1856); Dillon &
Sagástegui-Alva, Fieldiana, Bot. n.s. 26: 1–70 (1991), reg.
rev.
Gnaphalium sect. Gamochaeta (Wedd.) Benth., pro parte et
quoad typum (1873).
Gilruthia Ewart in Ewart, Jean White & B. Rees, Proc. Roy.
Soc. Victoria ser. 2, 22: 13 (1909).
Galeomma Rauschert, Taxon 31: 557 (1981).
Annual or perennial herbs. Leaves alternate, flat
with entire margins, tomentose on both surfaces.
Annual herb. Leaves alternate, flat with entire
margins, tomentose on both surfaces. Capitula
only a few together, in terminal clusters. Involucral bracts papery, green, stereome undivided.
Receptacle flat, epaleate. All florets perfect, yellow.
266
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Anthers with concave appendages. Style branches
truncate, with hairs apically. Cypselae obovoid,
glabrous. Pappus bristles with flattened axis,
distinctly plumose, connate in a ring. n = 13. One
species, G. osbornii Ewart & Jean White in Ewart,
Jean White & B. Rees, Australia.
650. Gnaphaliothamnus Kirp.
Gnaphaliothamnus Kirp. in Kirp. & Kuprijanova, Trudy
Bot. Inst. Akad. Nauk S.S.S.R. ser. 1, Fl. Sist. Vyssh. Rast.
9: 33 (1950), reg. rev.; Nesom, Phytologia 68: 366–381
(1990), rev.; Nesom, Phytologia 69: 1–3 (1990), rev.; Nesom,
Phytologia 76: 185–191 (1994), gen. consp.
Shrub. Leaves alternate, revolute with entire margins, tomentose on both surfaces. Capitula many in
corymbs. Involucral bracts papery, white or pink,
stereome undivided. Receptacle flat, epaleate.
Outer florets filiform, purple. Central florets
perfect, purple. Anthers with flat appendages.
Style branches truncate, with hairs dorsally and
apically. Cypselae oblong, with short clavate twin
hairs. Pappus bristles capillary, barbellate, free.
n = 14. One species, G. rhodanthus (Sch. Bip.)
Kirp., Central America.
651. Gnaphalium L.
Figs. 61H, 64D
Gnaphalium L., Sp. Pl. 2: 850 (1753); Drury, N. Z. J. Bot. 8:
222–248 (1970), rev.; Chang & Tseng, Fl. Reip. Pop. Sin. 75
(1979), reg. rev.; Jeffrey, Taxon 28: 349–351 (1979), nomencl.;
Hilliard, Bot. J. Linn. Soc. 82: 267–292 (1981), gen. consp.;
Hilliard & Burtt, Bot. J. Linn. Soc. 82: 233–265 (1981),
nomencl.; Dillon & Sagástegui-Alva, Fieldiana, Bot. n.s. 26:
1–70 (1991), reg. rev.
Omalotheca Cass. (1828).
Amphidoxa DC. (1838).
Filaginella Opiz (1854).
Gnaphalium sect. Eu-Gnaphalium O. Hoffm., pro parte et
quoad typum (1889).
Homognaphalium Kirp. (1950), quoad typum, excl. descr.
Annual or perennial herbs. Leaves alternate, flat
with entire margins, tomentose on both surfaces.
Capitula solitary or only a few together. Involucral
bracts papery, brownish, stereome undivided. Receptacle flat, epaleate. Outer florets filiform, purple. Central florets perfect, purple. Anthers with
flat appendages. Style branches truncate, with hairs
apically. Cypselae oblong, with short clavate twin
hairs. Pappus bristles capillary, scabrid, free. n = 7,
8, 9, 10, 14, 21, 26, 28, 29. About 80 species, cosmopolitan.
652. Gnephosis Cass.
Figs. 63X, 65D–F
Gnephosis Cass., Bull. Sci. Soc. Philom. Paris 1820: 43
(1820); Short, Muelleria 5: 185–214 (1983), rev.; Short,
Muelleria 6: 317–319 (1987), rev.; Jeffrey, Comp. Newslett.
16 (1989), nomencl.
Crossolepis Benth. (1837).
Chrysocoryne Endl. (1843).
Cyathopappus F. Muell. (1861).
Annual herbs. Leaves opposite basally, alternate
above, flat with entire margins, tomentose on both
surfaces. Capitula many in elongate secondary
heads. Involucral bracts papery, hyaline, stereome
undivided. Receptacle conical, epaleate. Anthers
with concave appendages. All florets perfect,
yellow. Style branches truncate, with hairs apically.
Cypselae obovoid, with globose hairs without
a basal cell. Pappus of small scales connate in
a ring to form a cup or absent. n = 6, 12, 14. Eight
species, Australia.
653. Gnomophalium Greuter
Gnomophalium Greuter, Willdenowia 33: 242 (2003);
Hilliard & Burtt, Bot. J. Linn. Soc. 82: 181–232 (1981), reg.
rev., sub Homognaphalio; Fayed & Zareh, Willdenowia 18:
445–453 (1989), reg. rev., sub Homognaphalio.
Homognaphalium Kirp. (1950), quoad descr., typo excluso.
Annual herb. Leaves alternate, flat with entire margins, tomentose on both surfaces. Capitula many in
dense clusters among a group of leaves. Involucral
bracts papery, hyaline, stereome divided. Receptacle flat, epaleate. Outer florets filiform, yellow.
Central florets perfect, yellow. Anthers with flat
appendages. Style branches truncate, with hairs
apically. Cypselae oblong, with short clavate twin
hairs. Pappus bristles capillary, scabrid, cohering
by patent cilia. One species, G. pulvinatum (Delile)
Greuter, North Africa and Asia.
654. Gratwickia F. Muell.
Gratwickia F. Muell., Bot. Centralbl. 64: 445 (1895); Eichler,
Taxon 12: 295–297 (1963), nomencl.
Annual herb. Leaves alternate, flat with entire
margins, tomentose on both surfaces. Capitula
only a few together, in terminal clusters. Involucral bracts papery, yellowish, stereome divided.
Receptacle flat, epaleate. Outer florets filiform,
yellow. Central florets perfect, yellow. Anthers
with concave appendages. Style branches truncate,
with hairs apically. Cypselae ellipsoid, glabrous.
Pappus bristle capillary, one, plumose, on central
Compositae
florets only. One species, G. monochaeta F. Muell.,
Australia.
655. Haeckeria F. Muell.
Haeckeria F. Muell., Linnaea 25: 406 (1853); Ewart, Flora
of Victoria (1930), reg. rev.; Heine, Adansonia sér. 2,
7: 115–140 (1967), rev.; Orchard, Austral. Syst. Bot. 17:
447–467 (2004), rev.
Humea sect. Haeckeria (F. Muell.) F. Muell. (1858).
Shrubs. Leaves alternate, revolute with entire
margins, tomentose on both surfaces. Capitula
many in corymbs. Involucral bracts papery,
brownish, stereome undivided. Receptacle flat,
epaleate. All florets perfect, yellow. Anthers with
flat appendages. Style branches truncate, with
hairs apically. Cypselae ellipsoid, with short
clavate twin hairs. Pappus absent. Two species,
Australia.
656. Haegiela P.S. Short & Paul G. Wilson
Haegiela P.S. Short & Paul G. Wilson, Muelleria 7: 259 (1990);
Short & Wilson, Muelleria 7: 259–265 (1990), rev.
Annual herb. Leaves opposite, at least basally, flat
with entire margins, tomentose on both surfaces.
Capitula axillary, solitary. Involucral bracts papery,
silvery. Receptacle flat, epaleate. Outer florets filiform, yellow. Central florets perfect, yellow with
purple apices. Anthers with flat appendages. Style
branches truncate. Cypselae ellipsoid, with globose hairs without a basal cell. Pappus absent. One
species, H. tatei (F. Muell.) P.S. Short & Paul G.
Wilson, Australia.
657. Haptotrichion Paul G. Wilson
Fig. 63E
Haptotrichion Paul G. Wilson, Nuytsia 8: 422 (1992);
Wilson, Nuytsia 8: 379–438 (1992), rev.
Annual herbs. Leaves alternate, flat with entire
margins, glandular-hairy on both surfaces. Capitula solitary. Involucral bracts papery, coloured,
clawed, stereome undivided. Receptacle flat or
convex, epaleate. All florets perfect, yellow. Anthers
with flat appendages. Style branches truncate.
Cypselae beaked, ellipsoid. Pappus bristles capillary, barbellate, connate in a ring to form a small
cup. n = 12. Two species, Australia.
658. Helichrysopsis Kirp.
Helichrysopsis Kirp., Trudy Bot. Inst. Akad. Nauk S.S.R. ser.
1: 32 (1950).
267
Perennial subshrub. Leaves alternate, flat with entire margins, sparsely tomentose on both surfaces.
Capitula only a few together, in dense corymbs. Involucral bracts papery, white, stereome undivided.
Receptacle flat, epaleate. Outer florets filiform, yellow. Central florets perfect, yellow. Anthers with
flat appendages. Style branches truncate, with hairs
apically. Cypselae oblong, with globose twin hairs.
Pappus bristles capillary, plumose, basally connate.
One species, H. septentrionalis (Vatke) Hilliard &
B.L. Burtt, Africa.
659. Helichrysum Mill.
Fig. 63C,K,N,U,V
Helichrysum Mill., Gard. Dict. abr. ed. 4: 28 (1754), corr.
Pers., Syn. Pl. 2: 414 (1807), nom. et orth. cons.; Moeser,
Bot. Jahrb. 43: 420–460 (1909), reg. rev.; Moeser, Bot. Jahrb.
44: 239–345 (1910), reg. rev.; Schwartz, Mitt. Inst. Bot. Hamburg 10: 1–393 (1939), reg. rev.; Hedberg, Symb. Bot. Upsal.
15: 1–424 (1957), reg. rev.; Cufodontis, Bull. Jard. Bot. Natl
Belg. 36, suppl. (1966), reg. rev.; Gill, Willdenowia 12: 95–
128 (1982), reg. rev.; Scott, Fl. Mascar. 109: 62–70 (1993),
reg. rev.
Leontonyx Cass. (1822).
Perennial or annual herbs or sometimes shrublets.
Leaves alternate, generally flat with entire margins, often tomentose. Capitula solitary or many
in corymbs. Involucral bracts papery, brown, yellow, pink or white, stereome divided or undivided.
Receptacle flat, epaleate or rarely paleate. Outer
filiform florets yellow, or absent. Central florets
perfect, yellow. Anthers with flat appendages. Style
branches truncate, with hairs apically. Cypselae oblong, glabrous or with elongated or short clavate
twin hairs. Pappus bristles capillary, barbellate or
subplumose, connate or free. n = 7, 8, 9, 14, 21,
24, 28, 42. About 600 species, Africa, Madagascar,
Europe and Asia.
660. Hesperevax (A. Gray) A. Gray
Hesperevax (A. Gray) A. Gray, Proc. Amer. Acad. Arts 7:
356. (1868); Morefield, Syst. Bot. 17: 293–310 (1992), rev.
Evax [sect.] Hesperevax A. Gray (1857).
Annual herbs. Leaves alternate or seemingly
whorled below, flat with entire margins, tomentose
on both surfaces. Capitula only a few together.
Involucral bracts none or vestigial. Receptacle
conical, paleate. Outer florets filiform, white. Central florets functionally male, lobes purple, tubes
white. Cypselae ovoid, dorsiventrally compressed,
glabrous. Pappus absent. Three species, North
America.
268
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
661. Humeocline Anderb.
Humeocline Anderb., Opera Bot. 104: 138 (1991); Anderberg, Opera Bot. 104: 1–195 (1991), rev.
Perennial herb or subshrub. Leaves alternate, revolute with entire margins, tomentose. Capitula many
in terminal pyramidal synflorescence. Involucral
bracts papery, yellowish. Receptacle flat, epaleate.
All florets perfect, yellow. Anthers with flat appendages. Style branches truncate, with hairs apically. Cypselae oblong, glabrous. Pappus absent.
One species, H. madagascariensis (Humbert) Anderb., Madagascar.
662. Hyalochlamys A. Gray
Hyalochlamys A. Gray, J. Bot. Kew Gard. Misc. 3: 98–101
(1851); Short, Muelleria 5: 185–214 (1983), rev.
Rosulate annual herb. Leaves opposite, at least
basally, flat with entire margins, tomentose on both
surfaces. Capitula many, forming dense secondary
heads among the leaves. Involucral bracts papery,
hyaline, stereome undivided. Receptacle conical,
epaleate. All florets perfect, yellow. Anthers with
concave appendages. Style branches truncate,
with hairs apically. Cypselae obovoid, glabrous.
Pappus absent. One species, H. globifera A. Gray,
Australia.
663. Hyalosperma Steetz
Fig. 63B
Hyalosperma Steetz in Lehm., Pl. Preiss. 1: 476 (1845);
Wilson, Nuytsia 7: 75–101 (1989), rev.
Helipterum sect. Pachypterum Steetz in Lehm. (1845), pro
parte et quoad typum.
Annual herbs. Leaves alternate, filiform with entire
margins, tomentose on both surfaces. Capitula
solitary or only a few together. Involucral bracts
papery, white or yellow, stereome undivided.
Receptacle conical, epaleate. All florets perfect,
yellow. Anthers with concave appendages. Style
branches truncate, with hairs apically. Cypselae
obovoid, with globose hairs without a basal cell.
Pappus bristles with flattened axis, plumose,
connate in a ring. n = 8, 11, 12. Nine species,
Australia.
664. Hydroidea P.O. Karis
Hydroidea P.O. Karis, Bot. J. Linn. Soc. 102: 29 (1990);
Hilliard & Burtt, Bot. J. Linn. Soc. 82: 181–232 (1981), reg.
rev. sub Atricantha; Karis, Bot. J. Linn. Soc. 102: 23–36
(1990), rev.
Shrub. Leaves alternate, twisted with revolute
entire margins, sparsely tomentose on both
surfaces. Capitula solitary. Involucral bracts
papery, white, stereome undivided. Receptacle
flat, epaleate. All florets perfect, purple. Anthers
with flat appendages. Style branches truncate, with
hairs apically. Cypselae ellipsoid, glabrous. Pappus
bristles capillary, barbellate, free. One species,
H. elsiae (Hilliard) P.O. Karis, Africa.
665. Ifloga Cass.
Ifloga Cass., Bull. Sci. Soc. Philom. Paris 1819: 142 (1819);
Fayed & Zareh, Willdenowia 17: 115–123 (1988), reg. rev.;
Hilliard, Bot. J. Linn. Soc. 82: 293–312 (1981), rev.
Trichogyne sect. Ifloga (Cass.) DC. (1838).
Comptonanthus B. Nord. (1964), pro parte et quoad typum.
Annual herbs. Leaves alternate, concave to involute
with entire margins, tomentose adaxially. Capitula
only a few together, in spikes. Involucral bracts papery, brownish, stereome divided. Receptacle flat,
epaleate. Outer florets filiform, purple. Central florets perfect, rarely functionally male, yellow. Anthers with flat appendages. Style branches truncate, with hairs apically. Cypselae oblong, with globose twin hairs. Pappus bristles capillary, distinctly
plumose apically, cohering by patent cilia. n = 7.
Six species, Asia, Europe, Middle East and Africa.
666. Ixiolaena Benth.
Ixiolaena Benth. in Endl., Enum. Pl. 66 (1837).
Perennial herbs. Leaves alternate, flat with entire
margins, tomentose on both surfaces. Capitula solitary. Involucral bracts cartilaginous with papery
tips, stereome undivided. Receptacle flat, epaleate.
Outer florets filiform, yellow. Central florets perfect, yellow. Style branches truncate, with hairs apically. Anthers with concave appendages. Cypselae
ellipsoid with elongated twin hairs. Pappus bristles
capillary, barbellate, connate in a ring. n = 20, 30.
Eight species, Australia.
667. Ixodia R. Br.
Ixodia R. Br. in Aiton, Hort. Kew. ed. 2. 4: 517 (1812); Copley, J. Adelaide Bot. Gard. 6: 41–54 (1982), rev.; Orchard,
Brunonia 4: 185–197 (1981), gen. consp.
Shrubs. Leaves alternate, revolute with entire margins, tomentose on both surfaces. Capitula many
in corymbs. Involucral bracts papery, white, stereome undivided. Receptacle conical, paleate. All florets perfect, yellow or white. Anthers with flat appendages. Style branches truncate, with hairs api-
Compositae
cally. Cypselae ellipsoid, with short clavate twin
hairs. Pappus absent. n = 13. Two species, Australia.
269
pus bristles capillary, barbellate, free. One species,
L. canescens (DC.) Anderb., southern Africa.
671. Lasiopogon Cass.
668. Jalcophila M.O. Dillon & Sagást.
Jalcophila M.O. Dillon & Sagást., Brittonia 38: 162 (1986);
Dillon & Sagástegui-Alva, Brittonia 38: 162–167 (1986), rev.;
Anderberg & Freire, Brittonia 42: 138–141 (1990), rev.; Dillon & Sagástegui-Alva, Fieldiana, Bot. n.s. 26: 1–70 (1991),
reg. rev.
Perennial, alpine cushion plants. Leaves alternate,
slightly revolute with entire margins, tomentose
adaxially only. Capitula solitary. Involucral bracts
papery, brownish, stereome undivided. Receptacle
flat, epaleate. Outer florets filiform, purple. Central florets perfect, purple. Style branches truncate,
with hairs dorsally. Cypselae ellipsoid to ovoid, glabrous or with glandular hairs. Pappus bristles capillary, scabrid, connate in a ring. Three species, South
America.
669. Lachnospermum Willd.
Lachnospermum Willd., Sp. Pl. 3: 1787 (1803); Dyer, The
genera of southern African flowering plants (1975), gen.
consp.; Hilliard & Burtt, Bot. J. Linn. Soc. 82: 181–232
(1981), gen. consp.
Ericoid shrubs. Leaves alternate, flat to twisted
with revolute entire margins, sparsely tomentose
adaxially. Capitula solitary or only a few together,
in dense corymbs. Involucral bracts cartilaginous,
stereome undivided. Receptacle flat, epaleate.
All florets perfect, purple. Anthers with flat
appendages. Style branches truncate, with hairs
apically. Cypselae ellipsoid, with few elongated
twin hairs. Pappus bristles capillary, barbellate,
basally connate. Three species, southern Africa.
670. Langebergia Anderb.
Langebergia Anderb., Opera Bot. 104: 93 (1991); Anderberg, Opera Bot. 104: 1–195 (1991), rev.; Nordenstam, Ann.
Wiener Mus. Naturgesch. 98, B: 403–418 (1996), rev.
Petalacte D. Don sect. Ampilasia DC. (1837).
Shrub. Leaves alternate, almost flat with revolute,
entire margins, sparsely tomentose on both surfaces. Capitula many in dense corymbs. Involucral
bracts papery, white, stereome undivided. Receptacle flat, epaleate. Outer florets filiform, purple.
Central florets functionally male, purple. Anthers
with flat appendages. Style branches truncate, with
hairs apically. Cypselae ellipsoid, glabrous. Pap-
Lasiopogon Cass., Bull. Sci. Soc. Philom. Paris 1818: 75
(1818); Nordenstam, J. S. African Bot. 30: 53–65 (1964),
rev.
Annual herbs. Leaves alternate, flat with entire margins, densely tomentose on both surfaces. Capitula
only a few together, in dense corymbs. Involucral
bracts papery, brownish, stereome divided. Receptacle flat, epaleate. Outer florets filiform, purple.
Central florets perfect, purple. Anthers with flat appendages. Style branches truncate, with hairs apically. Cypselae oblong, with globose hairs. Pappus
bristles capillary, plumose or barbellate, free. n = 7.
Eight species, Africa and Middle East.
672. Lawrencella Lindl.
Figs. 63Y, 65G
Lawrencella Lindl., Bot. Reg., Appendix to vols. 1–23: 23
(1839); Wilson, Nuytsia 8: 361–377 (1992), rev.
Helichrysum sect. Lawrencella (Lindl.) Benth. (1867).
Annual or perennial herbs. Leaves opposite, at
least basally, flat with entire margins, tomentose
on both surfaces. Capitula solitary or only a few
together. Involucral bracts papery, white, yellow
or pink, stereome undivided. Receptacle flat,
epaleate. Outer filiform florets, when present,
yellow. Central florets perfect, yellow. Anthers with
concave appendages. Style branches conical, with
hairs dorsally. Cypselae obovoid, with elongated
twin hairs. Pappus bristles capillary, barbellate,
free. n = 8, 11. About 35 species, Australia.
673. Lemooria P.S. Short
Lemooria P.S. Short, Muelleria 7: 112 (1989); Short, Muelleria 7: 103–116 (1989), rev.
Annual herb. Leaves opposite, at least basally, flat
with entire margins, glabrous. Capitula many in
dense secondary heads. Involucral bracts papery,
hyaline. Receptacle flat to convex, epaleate. All florets perfect, yellow. Style branches truncate, with
hairs apically. Cypselae obovoid, with elongated
twin hairs. Pappus bristles capillary, subplumose,
connate in a ring. One species, L. burkittii (Benth.)
P.S. Short, Australia.
674. Leontopodium R. Br. ex Cass.
Fig. 64C
Leontopodium R. Br. ex Cass., Bull. Sci. Soc. Philom. Paris
1819: 144 (1819); Beauverd, Bull. Soc. Bot. Genève sér. 2, 4:
270
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
12–55 (1912), rev.; Handel-Mazzetti, Beih. Bot. Centralb.
44, 2: 1–178 (1927), rev.
Perennial herbs. Leaves alternate, flat with entire
margins, tomentose on both surfaces. Capitula
many in corymbs. Involucral bracts papery,
brownish, stereome undivided. Receptacle flat,
epaleate. Outer florets filiform, yellow. Central
florets functionally male, yellow. Anthers with
flat appendages. Style branches truncate, with
hairs dorsally and apically. Cypselae oblong, with
short clavate twin hairs. Pappus bristles capillary,
barbellate, connate in a ring. n = 7, 12, 13, 14, 22,
24, 25, 26, 52. Fifty-eight species, Asia and Europe.
675. Lepidostephium Oliv.
Lepidostephium Oliv., Icon. Pl. ser. 3: 22 (1868); Kroner,
Mitt. Bot. Staatssamml. München 16: 1–268 (1980), rev.
Perennial herbs. Leaves alternate, revolute with
denticulate margins, sparsely tomentose on both
surfaces. Capitula solitary or only a few together,
in lax corymbs. Involucral bracts cartilaginous
to herbaceous, stereome undivided. Receptacle flat, epaleate. Outer florets radiate, violet.
Central florets perfect, yellow. Anthers with flat
appendages. Style branches obtuse, with hairs
dorsally. Cypselae ellipsoid, with few elongated
twin hairs. Pappus bristles capillary, barbellate,
free. Two species, southern Africa.
676. Leptorhynchos Less.
Leptorhynchos Less., Syn. Gen. Comp. 273 (1832); Wilson,
Nuytsia 12: 303–305 (1998), nomencl.; Wilson, Nuytsia 13:
607–611 (2001), rev.
Rhytidanthe Benth. (1837).
Annual or rarely perennial herbs. Leaves alternate,
flat with entire margins, tomentose on both surfaces or adaxially only. Capitula solitary. Involucral
bracts papery, brownish, stereome forming two
ribs. Receptacle flat, epaleate. Outer florets tubular, yellow. Central florets perfect, yellow. Anthers
with concave appendages. Style branches truncate,
with hairs apically. Cypselae ellipsoid, with globose
hairs without a basal cell, and one cell overtopping
the other. Pappus bristles capillary, barbellate, free.
n = 10, 12. Ten species, Australia.
677. Leptotriche Turcz.
Leptotriche Turcz., Bull. Soc. Imp. Naturalistes Moscou 24:
73 (1851); Munir, Adelaide Bot. Gard. 12: 101–117 (1989),
reg. rev.
Annual herbs. Leaves alternate, flat with entire margins, tomentose on both surfaces. Capitula many
in terminal secondary heads. Involucral bracts papery, hyaline, stereome undivided. Receptacle flat,
epaleate. All florets perfect, yellow. Anthers with
concave appendages. Style branches truncate, with
hairs apically. Cypselae obovoid, glabrous. Pappus
bristles scale-like, connate in a ring, absent in some
species. Twelve species, Australia.
678. Leucochrysum (DC.) Paul G. Wilson Fig. 61F
Leucochrysum (A. Cunn. ex DC.) Paul G. Wilson, Nuytsia 8:
441 (1992); Wilson, Nuytsia 8: 439–446 (1992), rev.
Helipterum sect. Leucochrysum A. Cunn. ex DC. (1838).
Perennial or annual herbs. Leaves alternate, flat or
sometimes filiform with entire margins, tomentose
or glandular. Capitula solitary. Involucral bracts
papery, coloured, inner bracts clawed, stereome
undivided. Receptacle flat, epaleate. All florets perfect, yellow. Style branches obtuse. Cypselae ellipsoid. Pappus bristles capillary, distinctly plumose,
free or shortly connate. n = 9. Five species, Australia.
679. Leucogenes Beauverd
Fig. 61C
Leucogenes Beauverd, Bull. Soc. Bot. Genève sér. 2, 2:
241–242 (1910); Molloy, N. Z. J. Bot. 33: 53–63 (1995), rev.;
Ward & Breitwieser, N. Z. J. Bot. 36: 165–171 (1998), rev.;
Breitwieser et al., N. Z. J. Bot. 37: 399–412 (1999), phylog.
Helichrysum sect. Leontopodioides Benth. (1873).
Perennial herbs. Leaves alternate, flat with entire
margins, tomentose on both surfaces. Capitula in
a dense corymb surrounded by conspicuous whorl
of leafy, white-lanate bracts. Involucral bracts
papery, brownish or hyaline, stereome undivided.
Receptacle conical, epaleate. Outer florets filiform,
generally yellow. Central florets perfect, generally yellow. Anthers with flat appendages. Style
branches truncate, with hairs apically. Cypselae
narrowly oblong to ellipsoid, with elongated twin
hairs. Pappus bristles capillary, scabrid, free.
n = 14, 28, 56. Four species, New Zealand.
680. Leucophyta R. Br.
Leucophyta R. Br., Trans. Linn. Soc. London 12: 106 (1818).
Perennial herb or subshrub. Leaves alternate,
flat with entire margins, densely tomentose on
both surfaces. Capitula many in dense, globose
secondary heads. Involucral bracts papery, hyaline,
stereome divided. Receptacle conical, epaleate.
Compositae
All florets perfect, yellow. Anthers with concave
appendages. Style branches truncate, with hairs
apically. Cypselae obovoid, glabrous. Pappus
bristles with flattened axis, plumose, connate
in a ring. n = 9. One species, L. brownii Cass.,
Australia.
271
volucral bracts papery, brownish, stereome undivided. Receptacle flat, often paleate. Female florets
filiform, yellow. Male florets yellow. Anthers with
flat appendages. Style branches truncate, with hairs
apically. Cypselae ellipsoid, glabrous. Pappus bristles capillary, barbellate, free. n = ca.14. Nineteen
species, South America.
681. Leysera L.
Leysera L., Sp. Pl. ed. 2, 2: 1249 (July 1763); Bremer, Bot.
Notiser 131: 369–383 (1978), rev.; Anderberg & Bremer,
Ann. Missouri Bot. Gard. 78: 1061–1072 (1991), phylog.,
gen. consp.; Fayed, Willdenowia 20: 97–102 (1991), reg. rev.
Leptophytus Cass. (1817).
Annual herbs or subshrubs. Leaves alternate, flat
with entire margins, tomentose on both surfaces.
Capitula solitary. Involucral bracts papery, brown,
stereome undivided. Receptacle flat, paleate. Outer
florets radiate, yellow with purple bands dorsally.
Central florets perfect, yellow. Anthers with flat appendages. Style branches obtuse, with hairs apically. Cypselae cylindrical, glabrous or with few
elongated twin hairs. Pappus of scales and capillary, plumose, free bristles. n = 4, 7, 8. Three
species, Africa.
682. Logfia Cass.
Fig. 64B
Logfia Cass., Bull. Sci. Soc. Philom. Paris 1819: 143 (1819).
Oglifa Cass. (1819).
Annual herbs. Leaves alternate, flat with entire
margins, tomentose on both surfaces. Capitula
only a few together. Involucral bracts cartilaginous, brownish, stereome undivided. Receptacle
conical, epaleate. Outer florets filiform, purple.
Central florets perfect, purple. Anthers with flat
appendages. Style branches with much prolonged
apex, with hairs dorsally and apically. Cypselae
oblong, with short clavate twin hairs. Pappus
bristles capillary, scabrid, cohering by basal cilia.
n = 14. Nine species, Europe, Middle East and
northern Africa.
683. Loricaria Wedd.
Fig. 65A–C
Loricaria Wedd., Chlor. Andina 1: 165 (1856) non J.V. Lamour., nom. rej. (1825); Cuatrecasas, Feddes Repert. 56: 149–
172 (1954), rev.; Dillon & Sagástegui-Alva, Phytologia 59:
227–233 (1986), rev.; Dillon & Sagástegui-Alva, Fieldiana,
Bot. n.s. 26: 1–70 (1991), reg. rev.
Dioecious shrubs. Leaves alternate, concave to involute with entire margins, tomentose adaxially
only. Capitula solitary or only a few together. In-
Fig. 65. Compositae-Gnaphalieae. A Loricaria leptothamna.
Pistillate floret. B, C Loricaria thuyoides. B Staminate floret. C Flowering branch. D–F Gnephosis uniflora. D Habit.
E Flowering branch. F Floret. G Lawrencella davenportii.
Habit. (Drawings by William Murray)
272
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
684. Lucilia Cass.
Fig. 63P
Lucilia Cass., Bull. Sci. Soc. Philom. Paris 1817: 32 (1817);
Freire, Darwiniana 27: 431–490 (1986), rev.; Freire, Bol. Soc.
Argent. Bot. 24: 411–413 (1986), cytol.; Zardini, Ann. Missouri Bot. Gard. 74: 431 (1987), rev.; Freire, Cladistics 3:
254–272 (1987), phylog.; Freire, Darwiniana 28: 409–411
(1987), rev.; Anderberg & Freire, Bot. J. Linn. Soc. 106: 173–
198 (1991), phylog.; Dillon & Sagástegui-Alva, Fieldiana,
Bot. n.s. 26: 1–70 (1991), reg. rev.
Oligandra Less. (1832).
Perennial herbs. Leaves alternate, flat with entire
margins, tomentose on both surfaces. Capitula solitary or only a few together. Involucral bracts papery, brownish, stereome usually undivided. Receptacle flat, epaleate. Outer florets filiform, purple. Central florets perfect, purple. Anthers with
flat appendages. Style branches acute to conical,
with hairs dorsally. Cypselae turbinate, with elongated twin hairs. Pappus bristles capillary, barbellate, connate in groups. n = 9, 14. Eleven species,
South America.
radiate, yellow or rarely white. Central florets
perfect or functionally male, yellow. Anthers with
flat appendages. Style branches obtuse, with hairs
dorsally. Cypselae ellipsoid, with few elongated
twin hairs. Pappus bristles capillary, barbellate,
free. Twelve species, southern Africa.
687. Metalasia R. Br.
Metalasia R. Br., Trans. Linn. Soc. London 12: 124 (1817);
Karis, Opera Bot. 99: 1–150 (1989), rev.
Shrubs. Leaves alternate, twisted with revolute entire margins, sparsely tomentose adaxially. Capitula many in dense corymbs. Involucral bracts papery, white, pink or yellow, stereome undivided.
Receptacle flat, epaleate. All florets perfect, purple.
Anthers with flat appendages. Style branches truncate, with hairs apically. Cypselae ellipsoid, glabrous. Pappus bristles capillary, barbellate, free.
n = 7. Fifty-two species, southern Africa.
688. Mexerion G.L. Nesom
685. Luciliocline Anderb. & Freire
Luciliocline Anderb. & Freire, Bot. J. Linn. Soc. 106: 187
(1991); Anderberg & Freire, Bot. J. Linn. Soc. 106: 173–198
(1991), rev., phylog.
Rosulate perennial herbs. Leaves basal, flat or revolute with entire margins, tomentose on both surfaces. Capitula solitary or many in glomerules arranged in spikes. Involucral bracts brownish, stereome undivided (rarely divided). Receptacle flat.
Outer florets filiform, white with purplish corolla
lobes. Central florets perfect. Style branches truncate or obtuse, with hairs dorsally and apically.
Cypselae ellipsoid or oblong-ellipsoid, with globose twin hairs. Pappus bristles capillary, scabrid,
connate in a ring. Five species, South America.
Mexerion G.L. Nesom, Phytologia 68: 249 (1990); Nesom,
Phytologia 68: 247–254 (1990), rev.
Rosulate, stoloniferous, perennial herbs. Leaves,
flat with entire margins, tomentose on both surfaces. Capitula arranged in glomerules or spikes.
Involucral bracts brownish or hyaline, stereome
undivided. Outer florets filiform, purple. Central
florets functionally male, purple. Style branches
with much prolonged apex, with hairs dorsally and
apically. Cypselae ellipsoid or turbinate-oblong,
with elongated twin hairs. Pappus bristles capillary, barbellate, connate in a ring. Two species,
Mexico.
689. Micropsis DC.
686. Macowania Oliv.
Fig. 63W
Macowania Oliv., Icon. Pl. 11: tab. 1062 (1870); Burtt &
Grau, Notes Roy. Bot. Gard. Edinburgh 31: 373–376 (1972),
rev.; Hilliard & Burtt, Notes Roy. Bot. Gard. Edinburgh 34:
253–286 (1976), reg. rev.; Hilliard & Burtt, Notes Roy. Bot.
Gard. Edinburgh 42: 227–260 (1985), reg. rev.
Homochaete Benth. (1872).
Perennial subshrubs or shrubs. Leaves alternate,
revolute or rarely flat with entire margins, sparsely
tomentose adaxially, densely abaxially. Capitula
solitary. Involucral bracts cartilaginous, stereome
undivided. Receptacle flat, epaleate. Outer florets
Micropsis DC., Prodr. 5: 459 (1836); Beauverd, Bull. Soc. Bot.
Genève sér. 2, 5: 221–228 (1913), rev.
Annual herbs. Leaves alternate, flat with entire margins, tomentose on both surfaces. Capitula only
a few together. Involucral bracts papery, hyaline,
stereome undivided. Receptacle flat, paleate. Outer
florets filiform, purple. Central florets perfect, purple. Anthers with flat appendages. Style branches
with much prolonged apex, with hairs dorsally
and apically. Cypselae oblong, with elongated twin
hairs. Pappus absent or a brief crown. About five
species, South America.
Compositae
273
690. Micropus L.
693. Myriocephalus Benth.
Micropus L., Sp. Pl. 927 (1753).
Myriocephalus Benth., Enum. Pl. 61 (1837); Short, Austral.
Syst. Bot 13: 729–738 (2000), rev.
Hyalolepis A. Cunn. ex DC. (1838).
Elachopappus F. Muell. (1862).
Annual herb. Leaves opposite, flat with entire margins, tomentose. Capitula solitary or only a few
together, in leaf axils. Involucral bracts herbaceous
to cartilaginous or apically with a minute papery
portion, hyaline, stereome undivided. Receptacle
flat, epaleate. Outer florets filiform, purple. Central
florets functionally male, purple. Anthers with flat
appendages. Style branches with a much prolonged
sterile apex, with hairs dorsally. Cypselae aborted
or reduced. Outer cypselae laterally compressed,
glabrous. Pappus absent. n = 14. One species, M.
supinus L., Europe, Africa.
Annual herbs. Leaves alternate, flat with entire
margins, tomentose on both surfaces. Capitula
many, clustered into dense secondary heads.
Involucral bracts papery, hyaline, stereome undivided. Receptacle conical, epaleate. All florets
perfect, yellow. Anthers with concave appendages.
Style branches truncate, with hairs apically.
Cypselae obovoid, with elongated twin hairs.
Pappus bristles, when present, with flattened axis,
barbellate, free. n = 8, 10, 12, 14. Ten species,
Australia.
691. Millotia Cass.
Millotia Cass., Ann. Sci. Nat. (Paris) 17: 416 (1829);
Schodde, Trans. Roy. Soc. S. Australia 87: 209–241 (1963),
rev.; Short, Austral. Syst. Bot. 8: 1–47 (1995), rev.; Short &
Anderberg, Austral. Syst. Bot. 8: 49–55 (1995), phylog.
Annual herbs. Leaves alternate or opposite at
least basally, flat with entire margins, tomentose
on both surfaces. Capitula solitary. Involucral
bracts cartilaginous, hyaline, stereome undivided.
Receptacle flat, epaleate. All florets perfect, yellow.
Anthers with concave appendages. Style branches
acute to conical, with hairs dorsally and apically.
Cypselae ellipsoid to turbinate, with elongated
twin hairs. Pappus bristles capillary, barbellate,
connate in a ring. n = 8, 10, 11, 13. Six species,
Australia.
692. Mniodes (A. Gray) Benth.
Fig. 61G
Mniodes (A. Gray) Benth. in Benth. & Hook. f., Gen. Pl. 2:
301 (1873); Cuatrecasas, Folia Biol. Andina 1: 1–7 (1954),
rev.; Dillon & Sagástegui-Alva, Fieldiana, Bot. n.s. 26: 1–70
(1991), rev.
Dioecious, perennial, alpine cushion plants. Leaves
alternate, concave to involute with entire margins,
tomentose distally with an adaxial hair tuft. Capitula solitary. Involucral bracts papery, brownish,
stereome undivided. Receptacle flat, epaleate. Female florets filiform, purple. Male florets purple.
Anthers with flat appendages. Style branches obtuse, with hairs dorsally and apically. Cypselae ellipsoid, with globose twin hairs. Pappus bristles
capillary, barbellate, connate in a ring. Four species,
South America.
694. Neotysonia Dalla Torre & Harms
Neotysonia Dalla Torre & Harms, Gen. Siphon. 540 (1905);
Anderberg, Opera Bot. 104: 1–195 (1991), rev.; Wilson,
Short & Orchard, Muelleria 7: 519–524 (1992), nomencl.
Tysonia F. Muell. (1896), nom. illegit.
Annual herb. Leaves alternate, flat with entire
margins, tomentose on both surfaces. Capitula
solitary. Involucral bracts herbaceous, brownish
to hyaline, stereome undivided. Receptacle flat,
paleate. All florets perfect, yellow. Anthers with
concave appendages. Style branches truncate,
with hairs apically. Cypselae pandurate, glabrous.
Pappus of scales connate into a papery cup. One
species, N. phyllostegia (F. Muell.) Paul G. Wilson
ex J.W. Green, Australia.
695. Nestlera Spreng.
Nestlera Spreng., Anleit. Kenntn. Gew. 2: 568 (1818); Anderberg & Bremer, Ann. Missouri Bot. Gard. 78: 1061–1072
(1991), rev., phylog.
Columellea N.J. Jacquin (1798).
Stephanopappus Less. (1831).
Perennial herb. Leaves alternate or opposite,
flat with entire margins, tomentose on both
surfaces. Capitula solitary or only a few together in
dense corymbs. Involucral bracts papery, brown,
stereome undivided. Receptacle flat, often paleate.
Outer florets radiate, yellow with purple bands.
Central florets perfect, yellow. Anthers with flat
appendages. Style branches truncate, with hairs
apically. Cypselae cylindrical, with few elongated
twin hairs. Pappus of scales, basally connate. n = 5.
274
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
One species, N. biennis (Jacq.) Spreng., southern
Africa.
696. Odixia Orchard
Fig. 66C–D
Odixia Orchard, Brunonia 4: 193 (1981); Orchard, Brunonia
4: 185–197 (1981), rev.
Shrubs. Leaves alternate, revolute with entire margins, tomentose on both surfaces. Capitula many in
corymbs. Involucral bracts papery, white, stereome
undivided. Receptacle conical, paleate or epaleate.
All florets perfect, yellow. Anthers with flat appendages. Style branches truncate, with hairs apically. Cypselae ellipsoid, with short clavate twin
hairs. Pappus absent. Two species, Australia.
697. Oedera L.
Fig. 66A–B
Oedera L., Mant. Pl.: 291 (1771); Harvey, Fl. Cap. 3 (1865),
reg. rev.; Mansfeld, Kew Bull. 6–9: 422–455 (1935), nomencl.;
Anderberg & Källersjö, Bot. J. Linn. Soc. 96: 323–332 (1988),
phylog.; Anderberg & Bremer, Ann. Missouri Bot. Gard. 78:
1061–1072 (1991), rev., phylog.
Eroeda Levyns (1948), nom. illegit.
Shrubs. Leaves alternate or opposite, flat with entire margins, densely tomentose on both surfaces.
Capitula many in dense corymbs, surrounded by
a general involucre of leaves. Involucral bracts papery, brown, stereome undivided. Receptacle flat,
paleate. Outer florets radiate, yellow with purple
bands. Central florets perfect, yellow. Anthers with
flat appendages. Style branches truncate, with hairs
apically. Cypselae cylindrical, glabrous or with few
elongated twin hairs. Pappus of short scales, basally
connate. n = 7. Eighteen species, southern Africa.
698. Oreoleysera K. Bremer
Oreoleysera K. Bremer, Bot. Notiser 131: 450 (1978); Bremer,
Bot. Notiser 131: 449–453 (1978), rev.; Anderberg & Bremer,
Ann. Missouri Bot. Gard. 78: 1061–1072 (1991), rev., phylog.
Fig. 66. Compositae-Gnaphalieae. A, B Oedera genistifolia.
A Flowering branch. B Head. C, D Odixia achleana.
C Flowering branch. D Head. E–H Phaenocoma prolifera.
E Pistillate (outer) floret. F Staminate (central) floret.
G Flowering branch with old and new heads. H Stem with
leafy short shoots. (Drawings by William Murray)
Shrub. Leaves alternate, flat with entire margins,
tomentose on both surfaces. Capitula solitary. Involucral bracts papery, brown, stereome undivided.
Receptacle flat, epaleate. Outer florets radiate, yellow with purple bands. Central florets perfect, yellow. Anthers with flat appendages. Style branches
truncate, with hairs apically. Cypselae cylindrical,
with elongated twin hairs. Pappus of scales and
capillary, barbellate, free bristles. One species, O.
montana (Bolus) K. Bremer, Africa.
Compositae
699. Oxylaena Benth. ex Anderb.
Oxylaena Benth. ex Anderb., Opera Bot. 104: 53 (1991);
Bentham, Icon. Pl. ser. 3, 2: 9, pl. 1109 (1873), rev. sub
Anaglypha; Anderberg, Opera Bot. 104: 1–195 (1991), rev.
Shrub. Leaves alternate, almost flat with revolute
entire margins, tomentose on both surfaces.
Capitula solitary. Involucral bracts cartilaginous,
stereome undivided. Receptacle flat, epaleate.
Outer florets radiate, yellow. Central florets
functionally male, yellow. Anthers with concave
appendages. Style branches obtuse, with hairs
dorsally. Cypselae ellipsoid, glabrous. Pappus
absent. One species, O. acicularis (Benth.) Anderb.,
southern Africa.
700. Ozothamnus R. Br. Figs. 62A, D, F, 63D, AA
Ozothamnus R. Br., Trans. Linn. Soc. London 12: 125 (1817);
Burbidge, Austral. J. Bot. 6: 229–284 (1958), rev.; Breitwieser
& Ward, N. Z. J. Bot. 35: 125–128 (1997), reg. rev.; Ward &
Breitwieser, N. Z. J. Bot. 36: 165–171 (1998), rev.; Breitwieser
et al., N. Z. J. Bot. 37: 399–412 (1999), phylog.
Helichrysum sect. Ozothamnus (R. Br.) Benth. (1867).
Helichrysum subg. Ozothamnus (R. Br.) N.T. Burb. sect.
Ozothamnus (R. Br.) Benth. sensu N.T. Burb. (1958).
Shrubs. Leaves alternate, flat, revolute or concave
with entire margins, tomentose on both surfaces,
abaxially only, or adaxially only. Capitula solitary or
many in corymbs. Involucral bracts papery, brownish or hyaline or coloured, stereome undivided. Receptacle flat, usually epaleate. Outer tubular or filiform florets, when present, yellow or white. Central florets perfect, yellow or white. Anthers with
flat appendages. Style branches truncate, with hairs
apically. Cypselae oblong, ellipsoid, ovoid, obovoid
or turbinate, with elongated or short clavate twin
hairs, or glabrous. Pappus bristles capillary, scabrid
to barbellate, free or connate in a ring or in groups.
n = 14. About 50 species, Australia, New Zealand,
New Caledonia.
701. Parantennaria Beauverd
Parantennaria Beauverd, Bull. Soc. Bot. Genève sér. 2: 255
(1911); Beauverd, Bull. Soc. Bot. Genève sér. 2, 3: 253–260
(1911), rev.
Dioecious perennial herb. Leaves alternate, flat
with entire margins, glabrous. Capitula solitary.
Involucral bracts papery, coloured. Stereome
undivided. Receptacle flat, epaleate. Female florets
filiform, purple. Male florets purple. Style branches
truncate, with hairs dorsally and apically. Cypselae
275
glabrous. Pappus bristles capillary, barbellate,
free. One species, P. uniceps (F. Muell.) Beauverd,
Australia.
702. Pentatrichia Klatt
Pentatrichia Klatt, Bull. Herb. Boissier 3: 436 (1895).
Shrubs. Leaves alternate, flat with dentate margins,
sparsely tomentose on both surfaces. Capitula solitary or only a few together in lax corymbs. Involucral bracts cartilaginous, stereome undivided. Receptacle flat, epaleate. Outer radiate florets, when
present, yellow or white. Central florets perfect, yellow. Anthers with flat appendages. Style branches
obtuse, with hairs dorsally. Cypselae ellipsoid, with
few elongated twin hairs. Pappus bristles capillary,
few, barbellate, free. Four species, Africa.
703. Petalacte D. Don
Petalacte D. Don, Mem. Wern. Nat. Hist. Soc. 5: 552 (1826);
Lundgren, Bot. Notiser 127: 119–124 (1974), rev.; Hilliard &
Burtt, Taxon 29: 507 (1980), nomencl.; Nordenstam, Ann.
Wiener Mus. Naturgesch. 98, B: 403–418 (1996), rev.
Shrub. Leaves alternate, revolute with entire margins, tomentose on both surfaces. Capitula many
in dense corymbs. Involucral bracts papery, white,
stereome undivided. Receptacle flat, paleate. Outer
florets filiform, purple. Central florets functionally
male, purple. Anthers with flat appendages. Style
branches truncate, with hairs apically. Cypselae
ellipsoid, glabrous. Pappus bristles capillary, barbellate, free. One species, P. coronata (L.) D. Don,
southern Africa.
704. Phaenocoma D. Don
Fig. 66E–H
Phaenocoma D. Don, Mem. Wern. Nat. Hist. Soc. 5: 554
(1826); Dyer, The genera of southern African flowering
plants (1975), gen. consp.; Hind, Curtis’s Bot. Mag. 13:
56–62 (1996), rev.
Shrub. Leaves alternate, minute, flat with entire
margins, sparsely tomentose adaxially. Capitula
solitary. Involucral bracts papery, pink to rosy
purple, stereome undivided. Receptacle flat,
epaleate. Outer florets filiform, purple. Central
florets functionally male, purple. Anthers with
flat appendages. Style branches truncate, with
hairs apically. Cypselae ellipsoid, with elongated
twin hairs. Pappus bristles capillary, barbellate,
basally connate. n = 8. One species, P. prolifera (L.)
D. Don, southern Africa.
276
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
705. Phagnalon Cass.
708. Planea P.O. Karis
Phagnalon Cass., Bull. Sci. Soc. Philom. Paris 1819: 174
(1819); Qaiser & Lack, Willdenowia 15: 3–22 (1985), reg.
rev.; Qaiser & Lack, Willdenowia 15: 437–450 (1986), reg.
rev.; Lack & Qaiser, Pl. Syst. Evol. 155: 45–48 (1987), reg. rev.
Gnaphalon Lowe (1868).
Planea P.O. Karis, Bot. J. Linn. Soc. 102: 32 (1990); Karis,
Opera Bot. 99: 1–150 (1989), rev. sub Metalasia; Karis, Bot.
J. Linn. Soc. 102: 23–36 (1990), rev.
Perennial herbs or sometimes shrublets. Leaves alternate, revolute with entire margins, tomentose
on both surfaces. Capitula solitary or only a few
together. Involucral bracts cartilaginous or papery,
brownish or hyaline, stereome undivided. Receptacle flat, epaleate. Outer florets filiform, yellow.
Central florets perfect, yellow. Anthers with flat appendages. Style branches obtuse, with hairs dorsally and apically. Cypselae ellipsoid to turbinate,
with elongated twin hairs. Pappus bristles capillary, barbellate, connate in a ring. n = 9. Fortythree species, western Asia, Europe and northern
Africa.
706. Philyrophyllum O. Hoffm.
Philyrophyllum O. Hoffm. in Engler & Prantl., Nat. Pflanzenfam. 4: 206, 208 (1890); Herman, Bothalia 29: 107 (1999), reg.
rev.
Perennial herb. Leaves alternate, flat with dentate
margins, tomentose on both surfaces. Capitula only
a few together, in lax corymbs. Involucral bracts
cartilaginous, stereome undivided. Receptacle flat,
paleate. Outer florets radiate, yellow. Central florets perfect, yellow. Anthers with flat appendages.
Style branches obtuse, with hairs dorsally. Cypselae ellipsoid, with few elongated twin hairs. Pappus bristles capillary, barbellate, free. Two species,
Africa.
707. Pithocarpa Lindl.
Pithocarpa Lindl., Bot. Reg., Appendix to vols. 1–23: 23
(1839); Lewis & Summerhayes, Kew Bull. 5: 435–440 (1951),
rev.; Lepschi, Nuytsia 13: 61–74 (1999), rev.
Perennial herbs. Leaves alternate, flat with entire
margins, tomentose on both surfaces. Capitula solitary or only a few together. Involucral bracts papery, white, stereome undivided. Receptacle flat,
epaleate. All florets perfect, yellow. Anthers with
concave appendages. Style branches truncate, with
hairs apically. Cypselae obovoid, with short clavate
twin hairs. Pappus absent. n = 13. Two species,
Australia.
Shrub. Leaves alternate, twisted, revolute with entire margins, sparsely tomentose adaxially. Capitula many in dense corymbs. Involucral bracts papery, white, stereome undivided. Receptacle flat,
epaleate. All florets perfect, purple. Anthers with
concave appendages. Style branches truncate, with
hairs apically. Cypselae ellipsoid, glabrous. Pappus
bristles capillary, barbellate, basally connate. One
species, P. schlechteri (L. Bolus) P.O. Karis, Africa.
709. Plecostachys Hilliard & B.L. Burtt
Plecostachys Hilliard & B.L. Burtt, Bot. J. Linn. Soc. 82: 206
(1981).
Perennial herbs. Leaves alternate, flat with entire
margins, sparsely tomentose abaxially. Capitula many in dense corymbs. Involucral bracts
papery, white, stereome undivided. Receptacle flat, epaleate. Outer florets filiform, yellow.
Central florets perfect, purple. Anthers with flat
appendages. Style branches truncate, with hairs
apically. Cypselae ellipsoid, with few elongated
twin hairs. Pappus bristles capillary, barbellate,
free. Two species, South Africa.
710. Pleuropappus F. Muell.
Pleuropappus F. Muell., Trans. Proc. Victorian Inst.
Advancem. Sci. 1: 37 (1855); Short, Muelleria 5: 143–183
(1983).
Annual herb. Leaves opposite, at least basally, flat
with entire margins, tomentose on both surfaces.
Capitula many, forming elongated secondary
heads. Involucral bracts papery, coloured, stereome undivided. Receptacle peg-like, epaleate.
All florets perfect, yellow. Anthers with concave
appendages. Style branches truncate, with hairs
apically. Cypselae obovoid, with globose hairs
without a basal cell. Pappus of jagged-edged
scales. One species, P. phyllocalymmeus F. Muell.,
Australia.
711. Podolepis Labill.
Podolepis Labill., Nov. Holl. Pl. 2: 56, t. 208 (1806), nom.
cons.; Davis, Proc. Linn. Soc. New South Wales 81: 245–286
(1957), rev.; Short, Taxon 35: 610–613 (1986), cytol.
Stylolepis Lehm. (1828).
Panaetia Cass. (1829), pro parte et quoad typum.
Compositae
Annual herbs. Leaves alternate, flat, tomentose on
both surfaces. Capitula solitary or only few together. Involucral bracts papery, brownish, stereome forming two ribs. Receptacle flat, epaleate.
Outer florets radiate or shortly radiate, yellow. Central florets perfect, yellow. Anthers with concave appendages. Style branches truncate, with hairs apically. Cypselae ellipsoid, with globose hairs without
a basal cell, and one cell overtopping the other. Pappus bristles capillary, barbellate, free. n = 3, 7, 8, 9,
10, 11, 12, 20. Twenty species, Australia.
712. Podotheca Cass.
Podotheca Cass., Dict. Sci. Nat. 23: 561 (1822), nom. cons.;
Short, Muelleria 7: 39–56 (1989), reg. rev.
Podosperma Labill. (1806), nom. rej.
Annual herbs. Leaves opposite, at least basally,
flat with entire margins, tomentose on both
surfaces. Capitula solitary. Involucral bracts green,
herbaceous, stereome undivided. Receptacle flat,
epaleate. All florets perfect, yellow. Anthers with
concave appendages. Style branches acute to
conical, with hairs apically. Cypselae ellipsoid to
turbinate, obovoid, with elongated twin hairs.
Pappus bristles with flattened axis, distinctly
plumose, connate in a ring. n = 13, 14, 26. Six
species, Australia.
277
dense terminal clusters. Involucral bracts papery,
hyaline, stereome undivided. Receptacle conical,
epaleate. All florets perfect, yellow. Anthers with
concave appendages. Style branches truncate, with
hairs apically. Cypselae obovoid, with elongated
twin hairs. Pappus bristles with flattened axis, subplumose, barbellate, free. n = 14. Three species,
Australia.
715. Pseudognaphalium Kirp.
Fig. 62B, 63T
Pseudognaphalium Kirp. in Kirp. & L.A. Kuprijanova,
Trudy Bot. Inst. Akad. Nauk S.S.S.R. ser. 1, Fl. Sist. Vyssh.
Rast. 9: 33 (1950); Standley, Field Mus. Nat. Hist. 18:
1419–1538 (1938), reg. rev.; D’Arcy, Ann. Missouri Bot.
Gard. 62: 1033–1053 (1975), reg. rev.; Nash, Fieldiana, Bot.
24: 164–181 (1976), reg. rev.; Hilliard & Burtt, Bot. J. Linn.
Soc. 82: 181–232 (1981), gen. consp.
Perennial, biennial or annual herbs. Leaves alternate, flat with entire margins, tomentose on both
surfaces. Capitula many in corymbs. Involucral
bracts papery, coloured, stereome divided. Receptacle flat, epaleate. Outer florets filiform, yellow.
Central florets perfect, yellow. Anthers with flat
appendages. Style branches truncate, with hairs
apically. Cypselae oblong, with short clavate twin
hairs. Pappus bristles capillary, barbellate, free.
n = 8, 9, 10, 14, 20. About 90 species, Africa, Asia,
Central, North and South America, New Zealand.
713. Pogonolepis Steetz
Pogonolepis Steetz in Lehm., Pl. Preiss. 1: 440 (1845); Short,
Muelleria 5: 185–214 (1983), rev.; Short, Muelleria 6: 237–
253 (1986), rev.
Skirrophorus sect. Pogonolepis (Steetz) A. Gray (1851).
716. Psilocarphus Nutt.
Psilocarphus Nutt., Trans. Amer. Philos. Soc. ser. 2, 7: 340
(1841); Morefield, Madroño 39: 155–157 (1992), reg. rev.
Annual herbs. Leaves alternate or opposite, at least
basally, flat with entire margins, tomentose on both
surfaces. Capitula forming dense secondary heads.
Involucral bracts papery, hyaline, stereome undivided. Receptacle flat, epaleate. All florets perfect,
yellow. Anthers with concave appendages. Style
branches truncate, with hairs apically. Cypselae
obovoid, with globose twin hairs without a basal
cell. Pappus absent. n = 4, 5, 12. Two species, Australia.
Annual herbs. Leaves opposite, flat with entire
margins, tomentose on both surfaces. Capitula
only a few together. Involucral bracts cartilaginous,
hyaline, stereome undivided. Receptacle peg-like,
epaleate. Outer florets filiform, purple. Central
florets functionally male, purple. Anthers with
concave appendages. Style branches with much
prolonged apex, with hairs dorsally. Cypselae
oblong, with short clavate twin hairs. Pappus
bristles mostly absent. n = 14. Eight species, North
and South America.
714. Polycalymma F. Muell. & Sond.
717. Pterochaeta Steetz
Polycalymma F. Muell. & Sond. in Sond., Linnaea 25: 494
(1853); Short, Wilson & Nailon, Muelleria 7: 57–79 (1989),
fruit morph.
Pterochaeta Steetz in Lehm., Pl. Preiss. 1: 456 (1845);
Wilson, Nuytsia 8: 379–438 (1992), rev.
Annual herb. Leaves alternate, flat with entire margins, tomentose on both surfaces. Capitula many in
Annual herb. Leaves alternate, flat with entire
margins, tomentose on both surfaces. Capitula
many in loose racemes. Involucral bracts papery,
278
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
coloured, stereome forming two ribs. Receptacle
flat, epaleate. Outer florets filiform, yellow. Central
florets perfect, yellow. Style branches truncate,
with hairs apically. Cypselae ellipsoid, glabrous.
Pappus bristles capillary, distinctly plumose, free.
n = 12. One species, P. paniculata Steetz, Australia.
718. Pterothrix DC.
Pterothrix DC., Prodr. 6: 279 (1838); Brusse, Bothalia 20:
67–70 (1990), reg. rev.; Koekemoer, Bothalia 29: 65–75
(1999), rev.
Shrubs. Leaves alternate, flat to twisted with entire
margins, sparsely tomentose adaxially. Capitula
solitary or only a few together in dense corymbs.
Involucral bracts papery, brown, stereome undivided. Receptacle flat, epaleate. All florets perfect,
purple. Anthers with flat appendages. Style
branches truncate, with hairs apically. Cypselae
ellipsoid, glabrous. Pappus bristles capillary,
plumose, basally connate. Six species, Africa.
719. Pterygopappus Hook. f.
Pterygopappus Hook. f., London J. Bot. 6: 120 (1847); Breitwieser, Bot. J. Linn. Soc. 111: 183–209 (1993), leaf anat.
Perennial, alpine cushion plant. Leaves alternate,
concave with entire margins, with an adaxial tuft
of stiff hairs. Capitula solitary, terminal. Involucral
bracts papery, pale brown, stereome undivided. Receptacle flat, epaleate. Outer florets tubular, purple.
Central florets male, purple. Anthers with flat appendages. Style branches truncate, undivided, with
hairs apically. Cypselae vestigial. Outer cypselae
obovate, with elongated twin hairs. Pappus bristles capillary, plumose, free. n = 14. One species,
P. lawrencei Hook. f., Australia.
720. Pycnosorus Benth.
Pycnosorus Benth., Enum. Pl. 63 1837; Everett & Doust, Telopea 5: 39–43 (1992), rev.
Perennial or annual herbs. Leaves alternate, flat
with entire margins, tomentose on both surfaces.
Capitula many, forming very dense, globose, secondary heads. Involucral bracts papery, coloured,
stereome undivided. Receptacle peg-like, epaleate.
All florets perfect, yellow. Anthers with concave appendages. Style apices truncate, with hairs apically.
Cypselae obovoid, with elongated twin hairs. Pappus bristles with flattened axis, plumose, connate
in a ring. n = 6, 10. About six species, Australia.
721. Quinetia Cass.
Quinetia Cass., Dict. Sci. Nat. 60: 590 (1830); Anderberg,
Opera Bot. 104: 1–195 (1991), rev.
Annual herb. Leaves alternate, flat with entire
margins, tomentose on both surfaces. Capitula
solitary. Involucral bracts cartilaginous, stereome
undivided. Receptacle flat, epaleate. All florets
perfect, purple. Anthers with flat appendages.
Style branches acute to conical, with hairs apically.
Cypselae turbinate, with elongated twin hairs.
Pappus of scale-like bristles, free. n = 12. One
species, Q. urvillei Cass., Australia.
722. Quinqueremulus Paul G. Wilson
Quinqueremulus Paul G. Wilson, Nuytsia 6: 1 (1987);
Wilson, Nuytsia 6: 1–5 (1987), rev.
Annual herb. Leaves alternate, flat with entire margins, tomentose on both surfaces. Capitula many,
forming loose secondary heads. Involucral bracts
papery, hyaline, stereome undivided. Receptacle
flat, epaleate. All florets perfect, yellow. Anthers
with concave appendages. Style branches truncate,
with hairs apically. Cypselae ellipsoid with five
large ligulate outgrowths at the apex, and with
elongated twin hairs. Pappus, in the conventional
sense, absent. One species, Q. linearis Paul G.
Wilson, Australia.
723. Rachelia J.M. Ward & Breitw.
Rachelia J.M. Ward & Breitw., N. Z. J. Bot. 35: 145 (1997);
Ward, Breitwieser & Lovis, N. Z. J. Bot. 35: 145–154 (1997),
rev.; Breitwieser et al., N. Z. J. Bot. 37: 399–412 (1999), phylog.
Perennial herb. Leaves alternate, flat with entire
margins, tomentose on both surfaces. Capitula solitary in axils of uppermost leaves. Involucral bracts
papery, brownish, stereome undivided. Receptacle conical, epaleate. Outer florets filiform, purple.
Central florets perfect, purple. Anthers with flat appendages. Style branches truncate, with hairs apically. Cypselae fusiform, with projecting papillae,
twin hairs absent. Pappus bristles capillary, scabrid,
connate in groups. x = 14. One species, R. glaria
J.M. Ward & Breitw., New Zealand.
724. Raoulia Hook. f.
Fig. 61A, B
Raoulia Hook. f. in Raoul, Choix Pl. Nouv.-Zél. 20 (1846);
Beauverd, Bull. Soc. Bot. Genève sér. 2, 4: 12–55 (1912),
rev.; Ward, Mauri Ora 10: 11–19 (1982), rev.; Ward, J. Canty
Bot. Soc. 16: 33–41 (1982), rev.; Ward, N. Z. J. Bot. 31:
Compositae
279
21–28 (1993), morph.; Ward, N. Z. J. Bot. 31: 29–42 (1993),
morph.; Breitwieser & Ward, Bot. J. Linn. Soc. 126: 217–235
(1998), leaf anat.; Ward, Etienne Raoul and Canterbury
botany 1840–1996: (1998), history; Ward & Breitwieser,
N. Z. J. Bot. 36: 165–171 (1998), rev.; Breitwieser et al.,
N. Z. J. Bot. 37: 399–412 (1999), phylog.
Psychrophyton Beauverd (1910).
bands. Central florets perfect, yellow. Anthers with
flat appendages. Style branches truncate, with hairs
apically. Cypselae cylindrical, glabrous or with few
elongated twin hairs. Pappus a crown of free scales.
n = 5, 7. Thirteen species, southern Africa.
Perennial mat-forming herbs or alpine cushion
plants. Leaves alternate, flat or concave with
entire margins, generally tomentose on both
surfaces. Capitula solitary, sessile, terminal.
Involucral bracts papery, brownish, hyaline or
coloured, stereome undivided. Receptacle usually
flat, epaleate. Outer florets filiform or tubular,
yellow, white or purple. Central florets generally
perfect, yellow, white or purple. Anthers with flat
appendages. Style branches truncate, with hairs
apically. Cypselae generally oblong to obovoid,
with elongated or short clavate or globose twin
hairs, or rarely glabrous. Pappus bristles capillary,
scabrid or barbellate, connate in groups or free.
n = 14, 28, 42, 56. Twenty-three species, New
Zealand.
Rhodanthe Lindl., Bot. Reg. 1703 (1834); Wilson, Nuytsia 8:
379–438 (1992), rev.
Pteropogon A. Cunn. ex DC. (1838).
Roccardia Neck. ex Voss (1896), nom. illegit., non Raf.
(1838).
Xyridanthe Lindl. (1839).
Acroclinium A. Gray (1852).
Helichrysum sect. Rhodanthe (Lindl.) Baill. (1886).
727. Rhodanthe Lindl.
Fig. 63A, F, R
725. Raouliopsis S.F. Blake
Annual herbs, one species a perennial. Leaves alternate, flat with entire margins, tomentose on both
surfaces. Capitula solitary or only a few together.
Involucral bracts papery, coloured, stereome undivided. Receptacle conical, epaleate. All florets
perfect, yellow. Anthers with concave appendages.
Style branches truncate, with hairs apically. Cypselae obovoid, with elongated twin hairs. Pappus bristles with flattened axis, plumose, connate in a ring.
n = 5, 7, 8, 10, 11, 14. Forty-three species, Australia.
Raouliopsis S.F. Blake, J. Wash. Acad. Sci. 28: 173 (1938);
Blake, J. Wash. Acad. Sci. 28: 172–177 (1938), rev.
728. Rhynchopsidium DC.
Perennial, alpine cushion plants. Leaves alternate,
concave to involute with entire margins, tomentose
distally with an adaxial hair tuft. Capitula solitary,
sessile. Involucral bracts papery, brownish, stereome undivided. Receptacle flat, epaleate. Outer florets filiform, purple. Central florets perfect, yellow. Anthers with flat appendages. Style branches
obtuse, with hairs dorsally. Cypselae ellipsoid, glabrous. Pappus bristles capillary, barbellate, connate
in a ring. Two species, South America.
726. Relhania L’Herit.
Fig. 63G
Relhania L’Herit., Sert. Angl. 1: 24 (1789), nom. cons.; Bremer, Opera Bot. 40: 1–86 (1976), rev.; Bremer, Taxon 25: 207–
208 (1976), nomencl.; Bremer, Taxon 28: 411–412 (1979),
nomencl.; Anderberg & Bremer, Ann. Missouri Bot. Gard.
78: 1061–1072 (1991), rev., phylog.
Shrubs. Leaves alternate or opposite, flat with entire margins, glabrous or sparsely tomentose on
both surfaces. Capitula solitary or many in dense
corymbs. Involucral bracts papery, brown, stereome undivided. Receptacle flat to conical, often
paleate. Outer florets radiate, yellow with purple
Rhynchopsidium DC., Mém. Soc. Phys. Genève 7: 283 (1836);
Anderberg & Bremer, Ann. Missouri Bot. Gard. 78: 1061–
1072 (1991), rev., phylog.
Rhynchocarpus Less. (1832), nom. illegit.
Annual herbs. Leaves alternate or opposite, flat
with entire margins, glabrous or sparsely tomentose on both surfaces. Capitula solitary. Involucral
bracts papery, brown, stereome undivided. Receptacle flat, paleate. Outer florets radiate, yellow with
purple bands. Central florets perfect, yellow. Anthers with flat appendages. Style branches truncate,
with hairs apically. Cypselae ellipsoid, with elongated twin hairs. Pappus of scales, basally connate
or free. n = 5. Two species, Africa.
729. Rosenia Thunb.
Rosenia Thunb., Nova Gen. Pl. 161 (1800); Bremer, Bot. Notiser 129: 97–111 (1976), rev.; Anderberg & Bremer, Ann.
Missouri Bot. Gard. 78: 1061–1072 (1991), rev., phylog.
Polychaetia Less. (1832).
Shrubs. Leaves alternate or opposite, recurved with
entire margins, tomentose on both surfaces. Capitula solitary or only a few together in dense corymbs.
280
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Involucral bracts papery, brown, stereome undivided. Receptacle flat, often paleate. Outer florets
radiate, yellow with purple bands dorsally. Central florets perfect, yellow. Anthers with flat appendages. Style branches truncate, with hairs apically. Cypselae cylindrical, with few elongated twin
hairs. Pappus of scales and a few, free, barbellate,
capillary bristles. n = 7, 14, 28. Four species, southern Africa.
730. Rutidosis DC.
Fig. 63H
Rutidosis DC., Prodr. 6: 158 (1838); Holland, Austrobaileya
4: 199–203 (1994), reg. rev.; Holland, Austrobaileya 5: 565–
572 (1999), rev.
Perennial or annual herbs. Leaves alternate, flat
with entire margins, tomentose on both surfaces.
Capitula solitary. Involucral bracts papery, brownish, stereome divided. Receptacle conical, epaleate.
All florets perfect, yellow. Anthers with concave
appendages. Style branches with much prolonged
apex, with hairs dorsally. Cypselae ellipsoid, with
globose hairs without a basal cell. Pappus of scalelike bristles or broad scales, free. n = 11, 12, 13, 19,
21, 22, 23, 26, 33, 36. Nine species, Australia.
731. Schoenia Steetz
Schoenia Steetz in Lehm., Pl. Preiss. 1: 480 (1845); Wilson,
Nuytsia 8: 361–377 (1992), rev.
Xanthochrysum Turcz. (1851).
Annual herbs. Leaves opposite at least basally, flat
with entire margins, tomentose on both surfaces.
Capitula solitary or only a few together. Involucral bracts papery, white or pink, stereome undivided. Receptacle flat, epaleate. Florets perfect,
innermost functionally male, yellow. Anthers with
concave appendages. Style branches acute to conical, with hairs dorsally. Cypselae obovoid, with
elongated twin hairs. Pappus bristles capillary, barbellate, free. n = 12. Three species, Australia.
732. Scyphocoronis A. Gray
Scyphocoronis A. Gray, Icon. Pl. t. 854 (1851); J. Bot. Kew
Gard. Misc. 4: 225 (1852); Cooke, J. Adelaide Bot. Gard. 7:
273–287 (1985), reg. rev.; Short, Austral. Syst. Bot. 8: 1–47
(1995), rev.; Short & Anderberg, Austral. Syst. Bot. 8: 49–55
(1995), phylog.
Annual herbs. Leaves alternate or opposite at least
basally, flat with entire margins, tomentose on both
surfaces. Capitula solitary. Involucral bracts cartilaginous, greenish, stereome undivided. Recepta-
cle flat, epaleate. All florets perfect, yellow. Anthers
with concave appendages. Style branches conical,
with hairs dorsally. Cypselae turbinate, with elongated twin hairs. Pappus rudimentary, a small cartilaginous crown. Two species, Australia.
733. Siloxerus Labill.
Siloxerus Labill., Nov. Holl. pl. 2: 57 (1806); Short, Muelleria
5: 185–214 (1983b), rev.; Short, Austral. Syst. Bot. Soc.
Newslett. 78: 6–7 (1994), rev.
Styloncerus Spreng. (1818), nom. illegit.
Gyrostephium Turcz. (1851).
Annual herbs. Leaves opposite at least basally,
filiform with entire margins, tomentose on both
surfaces. Capitula many, forming dense secondary
heads. Involucral bracts papery, hyaline, stereome
undivided. Receptacle conical, paleate. All florets
perfect, yellow. Anthers with concave appendages.
Style branches truncate, with hairs apically.
Cypselae obovoid, with globose hairs without
a basal cell. Pappus bristles scale-like, free. Three
species, Australia.
734. Sinoleontopodium Y.L. Chen
Sinoleontopodium Y.L. Chen, Acta Phytotax. Sin. 23: 457
(1985); Chen, Acta Phytotax. Sin. 23: 457–459 (1985), rev.
Dioecious, perennial, alpine cushion plant. Leaves
alternate, concave to involute with entire margins,
tomentose distally with an adaxial hair tuft.
Capitula solitary, sessile. Involucral bracts papery,
brownish, stereome undivided. Receptacle flat,
epaleate. Female florets filiform, yellow. Male
florets yellow. Anthers with flat appendages.
Style branches obtuse, with hairs dorsally and
apically. Cypselae turbinate, glabrous. Pappus
bristles capillary, barbellate, free. One species,
S. lingianum Y.L. Chen, eastern Asia.
735. Sondottia P.S. Short
Sondottia P.S. Short, Muelleria 7: 113 (1989); Short, Muelleria 7: 103–116 (1989), rev.
Annual herbs. Leaves opposite, flat with entire margins, tomentose on both surfaces or glabrous. Capitula many in dense secondary heads. Involucral
bracts cartilaginous, brownish-green or hyaline,
stereome undivided. Receptacle flat, epaleate. Floret solitary, perfect, yellow. Style branches truncate,
with hairs apically. Cypselae obovoid, glabrous except for long apical hairs. Pappus scale-like, connate
in a laciniate cup. n = 3. Two species, Australia.
Compositae
736. Stenocline DC.
Stenocline DC., Prodr. 6:218 (1838).
Shrublets. Leaves alternate, flat or revolute with
entire margins, tomentose. Capitula many in
corymbs. Involucral bracts papery, white or
yellowish, stereome undivided. Receptacle flat,
epaleate. All florets perfect, yellow. Anthers with
flat appendages. Style branches truncate, with
hairs apically. Cypselae oblong, glabrous. Pappus
bristles capillary, barbellate, free. Three species,
Madagascar and (?)Mauritius.
737. Stenophalium Anderb.
Stenophalium Anderb., Opera Bot. 104: 141 (1991); Robinson, Phytologia 55: 121–126 (1984), rev. sub Stenocline; Anderberg, Opera Bot. 104: 1–195 (1991), rev.
Perennial herbs. Leaves alternate, flat with entire
margins, tomentose on both surfaces. Capitula only
a few together, in terminal corymbs. Involucral
bracts papery, white, stereome divided. Receptacle flat, epaleate. All florets perfect, yellow. Anthers
with flat appendages. Style branches truncate, with
hairs apically. Cypselae oblong, glabrous. Pappus
bristles capillary, barbellate, cohering by patent
cilia. Three species, South America.
738. Stoebe L.
Stoebe L., Sp. Pl. 831 (1753); Levyns, J. S. African Bot. 3: 1–35
(1937), rev.; Dyer, The genera of southern African flowering plants (1975), gen. consp.; Nogueira, Bol. Soc. Brot. 51:
127–133 (1977), reg. rev.
Seriphium L. (1753).
Shrubs. Leaves alternate, twisted, revolute, with entire margins, sparsely tomentose adaxially. Capitula solitary or only a few together, in dense corymbs
or spikes. Involucral bracts cartilaginous, stereome undivided. Receptacle flat, epaleate. All florets perfect, purple. Anthers with flat appendages.
Style branches truncate, with hairs apically. Cypselae ellipsoid, glabrous. Pappus bristles capillary,
plumose, basally connate, or rarely absent. n =
8. Thirty-four species, Africa, Madagascar and La
Réunion.
739. Stuartina Sond.
Stuartina Sond., Linnaea 25: 521 (1853); Aston & Cooke,
Muelleria 6: 255–257 (1986), rev.
Annual herbs. Leaves alternate, spathulate, flat with
entire margins, tomentose on both surfaces. Ca-
281
pitula only a few together. Involucral bracts cartilaginous, brownish, stereome undivided. Receptacle conical, epaleate. Outer florets filiform, purple. Central florets perfect, purple. Anthers with
concave appendages. Style branches truncate, with
hairs apically. Cypselae oblong, glabrous. Pappus
absent. Two species, Australia.
740. Stuckertiella Beauverd
Stuckertiella Beauverd, Bull. Soc. Bot. Genève sér. 2, 5: 205
(1913); Beauverd, Bull. Soc. Bot. Genève sér. 2, 5: 205–209
(1913), rev.; Dillon & Sagástegui-Alva, Fieldiana, Bot. n.s.
26: 1–70 (1991), reg. rev.
Annual herbs. Leaves alternate, flat with entire
margins, tomentose on both surfaces. Capitula
only a few together, in terminal clusters. Involucral
bracts papery, brownish, stereome undivided.
Receptacle flat, epaleate. Outer florets filiform,
purple. Central florets functionally male, purple.
Anthers with flat appendages. Style branches
truncate, with hairs apically. Cypselae oblong,
with globose twin hairs. Pappus bristles capillary,
barbellate, connate in a ring. Two species, South
America.
741. Stylocline Nutt.
Stylocline Nutt., Trans. Amer. Philos. Soc. ser. 2, 7: 338
(1840); Wagenitz, Oesterr. Bot. Z. 119: 399–403 (1971), rev.;
Morefield, Madroño 39: 114–130 (1992), rev.
Annual herbs. Leaves alternate, flat with entire
margins, tomentose on both surfaces. Capitula
only a few together. Involucral bracts cartilaginous,
hyaline, stereome undivided. Receptacle peg-like,
epaleate. Outer florets filiform, purple. Central
florets functionally male, purple. Anthers with
concave appendages. Style branches with much
prolonged apex, with hairs dorsally. Cypselae oblong, glabrous. Pappus bristles capillary, scabrid,
free. n = 14. Seven species, North America.
742. Syncarpha DC.
Syncarpha DC., Ann. Mus. Natl Hist. Nat. 16: 205 (1810);
Nordenstam, Comp. Newslett. 17: 2–6 (1989), rev.
Anaxeton Schrank (1824), nom. illegit. non Gaertn. (1791).
Helipterum DC., pro parte (typ. non design.) (1838), nom.
illegit.
Roccardia Neck. ex Voss (1894).
Perennial herbs. Leaves alternate, flat with entire
margins, densely tomentose on both surfaces.
Capitula solitary or only a few together, in lax
282
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
corymbs. Involucral bracts papery, yellow, pink or
brown, stereome undivided or divided. Receptacle
flat, epaleate. All florets perfect, yellow. Anthers
with concave appendages. Style branches truncate,
with hairs apically. Cypselae ellipsoid, with globose
twin hairs. Pappus bristles capillary, plumose or
barbellate, basally connate. n = 7, 11. Twenty-five
species, southern Africa.
743. Syncephalum DC.
Syncephalum DC., Prodr. 6: 282 (1838).
Shrubs. Leaves alternate, flat with entire margins, tomentose abaxially or on both surfaces.
Capitula many in corymbs. Involucral bracts
papery, hyaline, stereome undivided. Receptacle
flat, epaleate. All florets perfect, yellow. Anthers
with flat appendages. Style branches truncate,
with hairs apically. Cypselae oblong, with short
clavate twin hairs. Pappus absent. Five species,
Madagascar.
744. Taplinia Lander
Taplinia Lander, Nuytsia 7: 37 (1989); Anderberg, Opera
Bot. 104: 1–195 (1991), rev.; Lander, Nuytsia 7: 37–42
(1989), rev.
Annual herb. Leaves alternate, flat with entire
margins, tomentose on both surfaces. Capitula
many in terminal panicles composed of many
small corymbs. Involucral bracts cartilaginous,
hyaline, stereome undivided. Receptacle flat,
epaleate. All florets perfect, yellow below, purple
above. Style branches truncate, with hairs apically.
Cypselae ellipsoid, with appressed twin hairs.
Pappus bristles capillary, barbellate, free. One
species, T. saxatilis Lander, Australia.
745. Tenrhynea Hilliard & B.L. Burtt
Tenrhynea Hilliard & B.L. Burtt, Bot. J. Linn. Soc. 82: 232
(1981).
Rhynea DC. (1838), nom. illegit., non Scopoli (1777).
Perennial herb. Leaves alternate, flat with entire
margins, sparsely tomentose abaxially. Capitula
many in dense corymbs. Involucral bracts papery,
white or pink, stereome undivided. Receptacle flat, paleate. Outer florets filiform, purple.
Central florets perfect, purple. Anthers with flat
appendages. Style branches truncate, with hairs
apically. Cypselae ellipsoid, with few elongated
twin hairs. Pappus bristles capillary, barbellate,
free. One species, T. phylicifolia (DC.) Hilliard &
B.L. Burtt, Africa.
746. Thiseltonia Hemsl.
Thiseltonia Hemsl., Icon. Pl. t. 2781 (1905); Wilson, Nuytsia
8: 479–483 (1992), rev.
Annual herb. Leaves alternate, filiform with entire margins, tomentose on both surfaces. Capitula
solitary. Involucral bracts papery, white or yellow,
stereome undivided. Receptacle conical, epaleate.
All florets perfect, yellow. Anthers with concave appendages. Style branches truncate, with hairs apically. Cypselae obovoid, glabrous. Pappus absent.
One species, T. gracillima (F. Muell. & Tate) Paul G.
Wilson, Australia.
747. Tietkensia P.S. Short
Tietkensia P.S. Short, Muelleria 7: 248 (1990); Short,
Muelleria 7: 239–252 (1990), rev.
Rosulate annual herb. Leaves basal, flat with entire
margins, tomentose on both surfaces. Capitula
many in dense secondary heads surrounded by
leaves. Involucral bracts papery, brownish-green
or hyaline, stereome undivided. Receptacle flat,
paleate. Outer florets filiform, yellow. Central
florets perfect, yellow or purplish. Style branches
truncate. Cypselae obovoid, with minute hairs.
Pappus absent. One species, T. corrickiae P.S.
Short, Australia.
748. Toxanthes Turcz.
Toxanthes Turcz., Bull. Soc. Imp. Naturalistes Moscou 24:
176 (1851); Short, Austral. Syst. Bot. 8: 1–47 (1995), rev.;
Short & Anderberg, Austral. Syst. Bot. 8: 49–55 (1995), phylog.
Anthocerastes A. Gray (1852).
Annual herbs. Leaves alternate throughout or opposite at least basally, flat with entire margins, tomentose on both surfaces. Capitula solitary. Involucral bracts cartilaginous, greenish-brown, stereome undivided. Receptacle flat, epaleate. All florets
perfect, yellow. Anthers with concave appendages.
Style branches conical, with hairs dorsally and apically. Cypselae narrowly turbinate, with elongated
twin hairs. Pappus absent. Three species, Australia.
749. Trichanthodium Sond. & F. Muell.
Trichanthodium Sond. & F. Muell. in Sond., Linnaea 25: 489
(1853); Short, Muelleria 7: 213–224 (1990), rev.
Compositae
283
Annual herbs. Leaves alternate, flat with entire margins, tomentose on both surfaces. Capitula many,
forming secondary heads on terminal branches. Involucral bracts cartilaginous, hyaline, stereome undivided. Receptacle flat, epaleate. All florets perfect,
yellow. Style branches truncate. Cypselae obovoid,
glabrous. Pappus scale-like, connate in a ring. n =
3, 4, 7. Four species, Australia.
truncate, with hairs apically. Cypselae oblong with
few short clavate twin hairs. Pappus bristles capillary, barbellate, free. Six species, southern Africa.
750. Trichogyne Less.
Annual herbs. Leaves alternate, flat with entire margins, tomentose on both surfaces. Capitula only
a few together. Involucral bracts papery, white,
stereome divided. Receptacle flat, epaleate. Outer
florets filiform, purple. Central florets perfect, purple. Anthers with flat appendages. Style branches
truncate, with hairs apically. Cypselae oblong, with
short clavate twin hairs. Pappus bristles capillary,
barbellate, free. Seven species, southern Africa.
Trichogyne Less., Linnaea 6: 231 (1831); Hilliard, Bot. J. Linn.
Soc. 82: 293–312 (1981), rev.; Beyers, Bothalia 25: 107–109
(1995), reg. rev.
Perennial herbs. Leaves alternate, almost flat with
involute margins, sparsely tomentose adaxially. Capitula only a few together, in spikes. Involucral
bracts papery, brownish, stereome divided. Receptacle flat, paleate. Outer florets filiform, purple.
Central florets functionally male, yellow. Anthers
with flat appendages. Style branches truncate, with
hairs apically. Cypselae oblong, with short clavate
twin hairs. Pappus bristles capillary, plumose, free.
Eight species, South Africa.
751. Triptilodiscus Turcz.
Fig. 63J
Triptilodiscus Turcz., Bull. Soc. Imp. Naturalistes Moscou
24: 66 (1851); Wilson, Nuytsia 8: 379–438 (1992), rev.
Dimorpholepis A. Gray (1851).
Annual herb. Leaves alternate, flat with entire margins, tomentose on both surfaces. Capitula solitary.
Involucral bracts papery, brownish, inner bracts
cartilaginous, stereome divided. Receptacle conical, epaleate. Outer florets filiform, yellow. Central
florets perfect, yellow. Anthers with concave appendages. Style branches truncate, with hairs apically. Cypselae ellipsoid, with globose hairs without
a basal cell. Pappus bristles plumose, with flattened
axis, free. n = 10. One species, T. pygmaeus Turcz.,
Australia.
752. Troglophyton Hilliard & B.L. Burtt
Troglophyton Hilliard & B.L. Burtt, Bot. J. Linn. Soc. 82: 208
(1981).
Annual or perennial herbs. Leaves alternate, flat
with entire margins, sparsely tomentose on both
surfaces. Capitula solitary or only a few together,
in dense corymbs. Involucral bracts papery, white,
stereome divided. Receptacle flat, epaleate. Outer
florets filiform, purple. Central florets perfect, purple. Anthers with flat appendages. Style branches
753. Vellereophyton Hilliard & B.L. Burtt
Vellereophyton Hilliard & B.L. Burtt, Bot. J. Linn. Soc. 82:
210 (1981).
754. Waitzia J.C. Wendl.
Waitzia J.C. Wendl., Coll. Pl. 2: 13 (1808); Wilson, Nuytsia
8: 461–477 (1992), rev.
Annual herbs. Leaves alternate, flat with entire margins, tomentose on both surfaces. Capitula solitary
or only a few together. Involucral bracts papery,
coloured, stereome forming two ribs. Receptacle
flat, epaleate. All florets perfect, yellow. Anthers
with concave appendages. Style branches conical,
with hairs apically. Cypselae ellipsoid to turbinate,
with globose hairs without a basal cell, and one
cell overtopping the other. Pappus bristles capillary, barbellate, free. n = 10, 12. Seven species,
Australia.
755. Xerochrysum Tzvelev
Figs. 61E, 63I
Xerochrysum Tzvelev, Novosti Sist. Vyssh. Rast. 27: 151
(1990); Bayer, Kew Bull. 56: 1013–1015 (2001), nomencl.
Bracteantha Anderb. & Haegi in Anderb. (1991), nom.
superfl.
Perennial herbs. Leaves alternate, flat with entire
margins, tomentose on both surfaces. Capitula
solitary or only a few together. Involucral bracts
papery, coloured, stereome undivided. Receptacle
flat, epaleate. Outer florets filiform, yellow. Central
florets perfect, yellow. Anthers with concave
appendages. Style branches acute, with hairs
dorsally. Cypselae quadrangular, glabrous. Pappus
bristles capillary, barbellate, free. n = 11, 12, 13,
14, 15. Six species, Australia.
284
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Recently Described Genera not Included
in the Key and Descriptions
Argyrotegium J.M. Ward & Breitw., N. Z. J. Bot. 41:
603–611 (2003).
Segregate of Euchiton. Four species, New Zealand,
Australia.
Castroviejoa Galbany, L. Sáez & C. Benedí, Austral.
Syst. Bot. 17: 581–591 (2004).
Segregate of Helichrysum. Two species, Corsica,
Sardinia.
Leiocarpa P.G. Wilson, Nutysia 13: 595–605 (2001).
Formerly species of Chrysocephalum, Ixiolaena,
Leptorhynchos. Ten species, Australia.
XV. Tribe Astereae Cass. (1819).
G. Nesom and H. Robinson
Annual or perennial herbs, subshrubs, shrubs,
rarely small trees or vines, rarely partially or
totally dioecious. Leaves usually alternate, opposite in a few African and Australian genera,
blades simple to pinnately divided. Heads solitary
or in corymbose, thyrsoid, spicate, racemose or
cyme-shaped cymes, sometimes with modified
secund branches; involucral bracts in (2–)3–5(–9)
series, gradate to nearly equal in length, narrow,
apices acute or rounded, often herbaceous, usually
persistent and reflexed, sometimes caducous;
receptacles flat or convex to conic or cyathiform,
usually alveolate or foveolate, usually glabrous,
sometimes fimbriate or with small to large pales.
Ray florets in 1–2 (to many) series; corolla of
ray florets usually 3–5-veined, apically acute to
truncate or dentate, with yellow or white to blue
limb, sometimes reduced to tube or ray florets
absent; style branches supinate. Disc florets perfect
and fertile or functionally male; corollas narrowly
tubular or funnelform or with limb abruptly
ampliate, rarely asymmetric, lobes usually 5(–4),
short and erect to spreading and sharply recurved
or long and recurved-coiling; thecae bases truncate or slightly auricular, rarely tailed or slightly
calcarate; apical appendage flat, lanceolate, usually
eglandular, rarely lacking. Pollen c. 25-µm diam.
in fluid; spherical; tricolporate and echinate,
with thin perforated tectum continuous between
colpi. Style arms pronate (erect or convergent) at
maturity, spreading radially, with separate lateral
stigmatic lines and lanceolate to deltoid apical
appendages, inner surface between stigmatic
lines and on appendages glabrous, outer surface
with sweeping hairs which are short-papillose to
longer with rounded or less often pointed apices.
Achenes laterally compressed and 2-nerved or
angular to nearly terete and multinerved, rarely
obcompressed (dorsiventrally compressed), surfaces usually with setulae, often with glands, tips
of setulae often spreading or anchor-shaped, cells
of achene wall usually with raphids, without phytomelanin; carpopodium usually annular; pappus
usually of 1–3 or 4 series of barbellate or rarely
plumose bristles, persistent or basally caducous,
outer or all segments sometimes reduced to scales
or awns, or pappus totally lacking.
Base chromosome number x = 9, reduced to
x = 8, 7, 6, 5, 3 and 2. Variation between x = 4, 5 and
9 often by dysploid reduction. Limited data by Herz
(1977) indicate the presence of various flavonoids,
benzofurans and coumarins, unidentified alkaloids, acetylenes with methyltriyne endgroups and
some matricaria esters. No pentaynene acetylenes
have been found, and sesquiterpene lactones are
absent or rare, which is correlated with the rarity
of large non-glutinous glandular punctuations.
Diterpenoids are most common in Machaerantherinae and Solidagininae, clerodanes and
labdanes in many genera with glutinous or viscid
leaf surfaces. Clerodanes are also common in roots
of Solidago, triterpenoids and sterols in many
members of the tribe, distinctive Baccharis-oxide
is characteristic of some Baccharis, and shionone
is found in some Asterinae.
The tribe Astereae occurs widely in temperate
and tropical parts of the World. It comprises c. 3,080
species in 205 genera (in the present account); the
difference in species number from a previous estimate of c. 3,100 (Nesom 1994c) primarily reflects
an increase in the estimated number of species
in Olearia (Lander 1994) and a lower, more accurate estimate for Baccharis (fide D. Giuliano, pers.
comm.).
Astereae are part of the large subfamily Asteroideae sensu stricto. Their echinate, tricolporate, caveate pollen is characteristic of the nonAnthemidean members of that group. The closest
relationship seems to be to Gnaphalieae and possibly Anthemideae. Characteristic for the tribe are
the essentially ecaudate non-calcarate bases of the
anthers, the totally separated stigmatic lines of the
style, and the short to elongate, basically triangular
Compositae
style appendages which are glabrous on the inside
and have sweeping hairs or papillae on the outside. Astereae completely lack phytomelanins in
the achene wall, which occur in most of the helianthoid genera. Astereae was one of the tribes
originally recognized by Cassini (1819), and it has
been recognized in essentially all treatments since
then (Bentham 1873a; Hoffmann 1894; Cronquist
1955; Bremer 1994). There have been few problems of delimitation, but the tribe now includes
Olivaea (De Jong and Beaman 1963), once in Helenieae, Geissolepis (Robinson and Brettell 1972),
Rigiopappus (Robinson and Brettell 1973b) and
Sheareria (Robinson 1981), once in Heliantheae,
Plagiocheilus (Robinson and Brettell 1973d), once
in Anthemideae, and Novenia (Nesom 1994c), once
in Lucilia of Inuleae-Gnaphalieae. A major DNA
study by Noyes and Rieseberg (1999) is the general
basis for the order of subtribes followed here, with
the last nine subtribes forming a North American
clade having a single origin. Revisions in Nesom
(1994c, 2000a) are the primary basis for the subtribal limits followed here, although the DNA data
suggest that some of the basal subtribes, such as
Asterinae, Brachyscominae, Hinterhuberinae and
Podocominae, are somewhat artificial (Noyes and
Rieseberg 1999; Lowrey and Urbatsch 2003). Particularly notable revisions of generic limits are those
of Aster (Nesom 1994d) and Haplopappus (Lane
and Hartman 1996). More complete descriptions
of the genera in North America, Central America,
the Antilles, and Hawaii can be found in Nesom
(2000b). The ETS and ITS study by Roberts and
Urbatsch (2004) shows where further generic revisions may be needed in Solidagininae.
Key to the Subtribes
and Unplaced Genera
1. Heads of two types usually on different plants, dioecious or rarely monoecious
2
– Heads of one type
3
2. Achenes compressed 2. Hinterhuberinae (Aztecaster)
– Achenes prismatic
7. Baccharidinae (p. 310)
3. Anther bases spurred; style base bulbiform
17. Vernoniopsis
– Anther bases not spurred, shortly rounded, or rarely
tailed; style base not or rarely broadened
4
4. Limbs of ray florets, when obvious, usually yellow or
white; achenes often terete to fusiform (compressed in
most Ericameria)
5
– Limbs of ray florets elongate and bluish to purplish or
white, or limbs commonly reduced or lacking; achenes
usually somewhat compressed
17
285
5. Style appendages of fertile disc florets hairy; involucral
bracts usually thickened or keeled (Machaerantherinae and Chrysopsidinae) or not keeled (Ericameria)
6
– Style appendages of fertile disc florets papillose, rarely
hairy; involucral bracts flattened, not keeled
8
6. Teeth and apices of leaves and involucral bracts often
spinulose-tipped; pappus series often gradate; x = 6
and reduced
13. Machaerantherinae p.p. (p. 328)
– Teeth and apices of leaves and involucral bracts not
spinulose-tipped; inner pappus distinctly longer than
outer, or pappus of equal length or nearly so; x = 9
and reduced
7
7. Shrubs; pappus of equal length or nearly so; x = 9
758. Ericameria
– Herbs; inner pappus distinctly longer than outer; x = 9
and reduced
17. Chrysopsidinae (p. 337)
8. Pappus (1–)2(–3)-seriate; leaves often tomentose below
9
– Pappus 1-seriate; leaves with little or no tomentum 11
9. Plants in most parts of Southern Hemisphere to Hawaii
and Mexico
2. Hinterhuberinae (p. 294)
– Shrubs or trees of St. Helena
10
10. Plants with pale hairs
756. Commidendron
– Plants with hairs often blackish 763. Melanodendron
11. Herbs to subshrubs; leaves usually sessile; mostly
American
12
– Shrubs or subshrubs; leaves usually petiolate; Asian or
African
14
12. Involucral bracts usually gradate, outer not or rarely
leaf-like
10. Solidagininae (p. 320)
– Involucral bracts subequal, at least outer leaf-like 13
13. Florets trimorphic, outer pistillate radiate, median pistillate; disc florets functionally male 765. Nannoglottis
– Florets dimorphic, radiate and disc; disc florets perfect
771. Tonestus
14. Disc florets with lobes unequal, outer three lobes
longer
761. Heteroplexis
– Disc florets with lobes equal
15
15. Plants not glutinous; leaf venation strongly ascending
or sublongitudinal; disc florets perfect
764. Microglossa
– Plants glutinous; leaves with veins single or pinnate;
disc florets functionally male
16
16. Leaf venation pinnate; pappus bristles shortly united
at base; ray florets in several series
768. Psiadia
– Leaves 1-nervate; pappus of scales; ray florets 1 or 2
769. Psiadiella
17. Appendages of disc floret styles narrowly oblong to
oblanceolate, longer than the stigmatic lines
11. Pentachaetinae (p. 326)
– Appendages of disc styles deltate to lanceolate, shorter
than the stigmatic lines
18
18. Involucral bracts with orange-resinous ribs or veins
19
– Involucral bracts without prominent resinous veins 22
19. Lower leaves reduced to scales; limb of disc corolla
ampliate; rays few, limbs not coiling 757. Doellingeria
– Lower leaves usually larger than upper; limb of disc
corolla variously shaped
20
20. Achenes terete or compressed with wings; involucral
bracts with tips rounded or obtuse; rays coiling
12. Boltoniinae (p. 327)
286
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
– Achenes compressed, not winged; involucral bracts
with tips mostly acute; rays coiling or straight
21
21. Receptacle without obvious pales, sometimes with
small scales; pistillate florets without pappus and
with rigid, narrowly urceolate corollas
18. Conyzinae (p. 339)
– Receptacle with pales peripherally grading into small
scales on disc; pistillate florets with pappus; corollas
not rigid nor narrowly urceolate 770. Sarcanthemum
22. Appendages of fertile disc styles hairy; involucral
bracts mostly herbaceous or with distinct apical green
herbaceous patch; pappus usually of capillary bristles
23
– Appendages of fertile disc styles papillose; involucral
bracts with herbaceous parts not necessarily in distinct apical patch; pappus of bristles or reduced or
lacking
26
23. Involucral bracts mostly herbaceous
767. Oreostemma
– Involucral bracts with herbaceous part in distinct apical green patch
24
24. Leaves entire or toothed to lobed or pinnatifid, teeth
and apices commonly spinulose-tipped; pappus bristles tending to be flattened at least basally; x = 6 or
reduced
13. Machaerantherinae p.p. (p. 328)
– Leaves entire or rarely toothed, without spinulose tips;
pappus bristles capillary; x = 9 or reduced
25
25. Achenes cylindrical to subcylindrical, 8–12(–18)nerved
760. Eurybia
– Achenes more or less compressed, 2(–8)-nerved
14. Symphyotrichinae (p. 334)
26. Ray florets, when present, (1–)2–4-seriate
27
– Ray florets 1-seriate
29
27. Pappus of bristles in (1–)2–3 series; limbs of ray florets
sometimes elongate and coiling
8. Podocominae (p. 312)
– Pappus reduced or lacking, rarely a single series of
bristles; ray florets with limbs short or lacking
28
28. Achenes not beaked; setulae often with anchor-shaped
tips; receptacles often conical or cyathiform
5. Grangeinae (p. 304)
– Achenes often beaked; setulae without anchor-shaped
tips; receptacles usually convex to flat, rarely conical
6. Lagenophorinae (p. 307)
29. Pappus reduced or lacking; achenes strongly compressed
30
– Pappus usually of capillary bristles
31
30. Plants rosulate with heads solitary on individual
scapes; limbs of ray florets not coiling; setulae of
achene simple
4. Bellidinae (p. 303)
– Plants not rosulate; heads not on individual scapes
from rosettes; limbs of ray florets coiling or short;
setulae of achenes with anchor-shaped tips
3. Brachyscominae (p. 302)
31. Leaves increasing in size upwards from stem base to
near middle, lowest scale-like; involucral bracts often keeled, without distinct herbaceous tips, without
or with narrow hyaline margins; appendages of disc
styles lanceolate
32
– Basal leaves not scale-like, often persistent; involucral bracts not keeled, mostly herbaceous or broadly
hyaline-margined
34
32. Pappus bristles 1–2-seriate, apically attenuate; limbs
of rays coiling
762. Ionactis
– Pappus bristles 2–3-seriate, apically dilated or attenuate; limbs of rays not or scarcely coiling
33
33. Bristles of pappus apically dilated; achene surfaces
eglandular
759. Eucephalus
– Bristles of pappus apically attenuate; achene surfaces
glandular
766. Oclemena
34. Limbs of ray florets, when present, not coiling or reflexed; x = 9 or 5; North America and Mexico
16. Astranthiinae (p. 336)
– Limbs of ray florets, when present, strongly or belatedly coiling
35
35. Involucral bracts mostly herbaceous to base; receptacles not paleate; mostly Palearctic 9. Asterinae (p. 316)
– Involucral bracts usually with broad hyaline margins;
receptacles often paleate
36
36. Achenes terete to compressed; leaves alternate; x = 8;
North America & Mexico 15. Chaetopappinae (p. 335)
– Achenes compressed; leaves alternate or opposite; x =
9 and reduced; Africa and southeast to central Asia
1. Homochrominae (p. 290)
Unplaced Genera
756. Commidendrum DC.
Commidendrum DC., Arch. Bot. (Paris) 2: 334 (1833).
Shrubs to small trees, hairs pale. Leaves clustered
at branch tips, shortly petiolate, subcoriaceous,
obovate to spathulate, gland-dotted, glutinous,
tomentose to sparsely pilose abaxially, venation
pinnate. Inflorescences 1-headed or corymbose,
erect or in pendent clusters; involucres campanulate to hemispheric; bracts 3–5-seriate, gradate,
linear-lanceolate, without resin duct, glabrous
to tomentellous, margins narrowly scarious;
receptacles plano-convex, epaleate. Rays 8–50,
pistillate, white, short-reflexed to long-spreading.
Disc florets 8–50, perfect, yellow, tube long, throat
cylindrical, lobes elongate, reflexed to coiled.
Achenes oblong, terete to subcompressed, 10nerved, glabrous or sparsely setulose, eglandular;
pappus 1-seriate, persistent, bristles smooth below,
barbellate distally. Four species, St. Helena.
757. Doellingeria Nees
Doellingeria Nees, Gen. Sp. Aster. 177 (1832) [1833]; Nesom,
Phytologia 75: 452–462 (1993), emend., key.
Erect perennial herbs; fibrous-rooted; glabrous to
sparsely pubescent, eglandular. Leaves all cauline,
lowest leaves reduced to bracts, upper petiolate,
blades truncate- to cordate-based. Inflorescences
corymbose; involucres cylindric-turbinate; bracts
2–4-seriate, strongly gradate, oblong to oblonglanceolate, apically rounded to acute, with raised
midrib, without herbaceous apical patch; recep-
Compositae
tacles epaleate. Rays 2–11(–16), pistillate, white,
limbs not coiling. Disc florets perfect, yellow, limb
expanded, lobes lanceolate, reflexing to coiling;
style appendages deltate. Achenes plump to compressed, glabrous to sparsely strigose, 5–9-nerved,
nerves with orange duct; pappus 2–3-seriate, outer
series short, longer bristles widened apically. x = 9.
Eleven species, eastern USA, eastern Asia.
Nesom (1993) included eight Asian species in
the genus, but molecular data (Brouillet et al. 2001)
indicate that the North American species are primitive members of the North American clade, whereas
the Asian species are more closely related to typical
Aster.
758. Ericameria Nutt.
Ericameria Nutt., Trans. Amer. Philos. Soc. 2, 7: 518 (1840);
Anderson, General Technical Report INT-200, USDA,
Forest Service, Ogden, Utah: 29–45 (1986), key.
Evergreen ericoid shrubs, often intricately
branched; often glutinous, sometimes tomentose.
Leaves linear to spathulate, sessile to shortpetiolate, usually gland-dotted. Heads solitary,
or in thyrsoid, racemose or corymbose inflorescences; involucres subcampanulate to turbinate;
bracts 3–5-seriate, subequal to strongly gradate,
midvein orange-resinous; receptacles epaleate.
Rays 0–10, pistillate, yellow, limbs coiling. Disc
florets perfect, lobes triangular, usually equal,
erect to spreading-recurved; style appendages
lanceolate to linear-lanceolate, hairy. Achenes narrowly oblong, compressed or less commonly terete,
6–12-ribbed, usually strigose to sericeous; pappus
1–3-seriate, bristles persistent or caducous. x = 9.
Thirty-one species, western USA, north-western
Mexico.
759. Eucephalus Nutt.
Eucephalus Nutt., Trans. Amer. Philos. Soc. 2, 7: 298 (1840).
Perennials, usually rhizomatous, rarely taprooted;
stems simple or distally few-branched, glabrous
or pilose to tomentose, with minute glands below
heads. Leaves all cauline, lowest scale-like, most
large, sessile, entire. Inflorescences (1–)6–20headed, corymbose; involucres campanulate to
hemispheric; bracts 4–6-seriate, gradate, ovate
to lanceolate, often convex, keeled, stramineousindurate, tips often purplish or green, inner
bract margins hyaline; receptacles plano-convex,
epaleate. Rays 0(–5) or 6–20, pistillate, white
to bluish, slightly coiling. Disc florets perfect,
287
yellow, lobes deltate and erect to triangular and
reflexing; style appendages linear-lanceolate.
Achenes obovate, compressed, marginal nerves 2
and sometimes 1–2 nerves on each face; pappus
bristles 2–3-seriate, usually broadened apically,
equally long or outer shorter. x = 9. Eleven species,
north-western USA (incl. California), western
Canada.
760. Eurybia (Cass.) S.F. Gray
Eurybia (Cass.) S.F. Gray, Nat. Arr. Brit. Pl. 2: 464 (1821);
Nesom, Phytologia 77: 256–262 (1994), comb.
Aster subg. Eurybia Cass. (1818).
Herrickia Wooton & Standl. (1913).
Weberaster Å. Löve & D. Löve (1982).
Perennial herbs, rhizomatous; stems and leaves
usually glabrate, few species with stipitate glands.
Leaves sessile, linear to obovate or cordate,
entire to spinulose, usually 3–5-parallel-veined.
Inflorescences usually corymbose, rarely 1- to
few-headed; involucres turbinate to campanulate;
bracts 5–7-seriate, gradate, bases indurate, usually
low-keeled, apically with distinct green patch,
margins usually minutely fringed; receptacles
epaleate. Rays pistillate, blue and coiling or white
and scarcely coiling. Disc florets perfect, becoming
purplish, narrowly tubular or with long tube and
expanded limb; style appendages linear-triangular.
Achenes cylindric, 8–12(–18)-nerved; pappus bristles (1–)2-seriate, often stiff and flattened, often
apically broadened. x = 9. Twenty-eight species,
USA, Canada to Alaska, and north-eastern Asia.
761. Heteroplexis C.C. Chang
Heteroplexis C.C. Chang, Sunyatsenia 3: 266 (1937); Chen,
Guihaia 5: 337–343 (1985), rev.
Scandent or trailing perennial herbs. Leaves shortpetiolate, blades lanceolate; pinnately veined, margins entire to sparsely serrulate. Inflorescences with
glomerules of small heads; involucres campanulate;
bracts 3–4-seriate, gradate, ovate to oblong, obtuse; receptacles convex, epaleate. Pistillate florets
4–7, 1-seriate; corollas tubular, eradiate. Disc florets
4–6, perfect, yellow, funnelform, lobes lanceolate,
unequal, 3 outer lobes longer; style appendages
triangular. Achenes oblong-ovate, almost terete to
slightly compressed, 6-nerved, sparsely setuliferous; pappus of subequal barbellate bristles. Three
species, China.
288
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
762. Ionactis Greene
Ionactis Greene, Pittonia 3: 245 (1897); Nesom & Leary,
Brittonia 44: 247–252 (1992), emend.
Perennial herbs from thick taproots or branched
woody caudex, rarely rhizomatous; puberulous
or glabrous. Leaves cauline, congested, basal
leaves sometimes marcescent, others stiff, narrow,
1-nerved, margins sometimes scarious. Inflorescences 1–3-headed or loosely corymbose;
involucres broadly turbinate to campanulate;
bracts 3–6-seriate, gradate, linear-lanceolate,
strongly keeled, without herbaceous tips; receptacles plano-convex, epaleate. Rays 8–21, pistillate,
limbs violet to bluish or white, coiling. Disc florets
perfect (one species functionally male), narrowly
tubular; style appendages lanceolate. Achenes
narrowly obovate, in rays (2–)3-nerved, in disc
florets 2(–4)-nerved, compressed, strigose to
sericeous (1 species glandular); pappus bristles
1–2-seriate, attenuate, outer series of short bristles
or scales. x = 9. Five species, USA (4 western,
1 eastern).
763. Melanodendron DC.
Melanodendron DC., Prodr. 5: 279 (1836).
Trees; hairs pale to blackish; young stems tomentose. Leaves clustered on branch tips, base broad,
narrowing to pseudopetiole, blade leathery, oblong
to obovate, margins entire, revolute, venation
pinnate, adaxially glabrous, abaxially with sparse
tomentum, obscurely gland-dotted. Inflorescences
corymbose; heads short-pedunculate; involucres
broadly campanulate; bracts 3–4-seriate, gradate,
linear-lanceolate, margins narrowly scarious,
midveins distinct, not resinous; receptacles planoconvex, epaleate. Rays 20–25, pistillate, white,
elongate. Disc florets c. 60–75, perfect, yellow,
tubes elongate, limb narrowly funnelform, lobes
deltate; style appendages deltate. Achenes prismatic to scarcely compressed, 5-nerved, glabrous,
eglandular; pappus bristles 1–2-seriate, persistent,
smooth below, barbellate distally. One species,
M. integrifolium DC., St. Helena.
of dense corymbose to subglobose cymes; involucres campanulate to turbinate; bracts 3–4-seriate,
gradate, ovate to linear-lanceolate to oblonglanceolate; receptacles plano-convex, epaleate.
Rays 3–4-seriate, pistillate, white. Disc florets
perfect, whitish, limbs cylindric-campanulate
to funnelform, lobes deltate, recurved; style appendages lanceolate or conic and papillose at base.
Achenes narrowly turbinate to obovoid, weakly
4-costate, somewhat compressed, pilose-setulose;
pappus bristles 1-seriate, persistent, numerous,
subequal, minutely barbellate. Circa 19 species,
Africa, Madagascar, tropical Asia.
765. Nannoglottis Maxim.
Nannoglottis Maxim., Bull. Acad. Imp. Sci. SaintPétersbourg 27: 480 (1881); Liu et al., Mol. Phylog. Evol.
23: 307–325 (2002), phylog.; Gao & Chen, Comp. Newslett.
40: 34 (2003), gen. class.; Gao et al., Acta Bot. Yunn. 26:
189–190 (2004), gen. class.
Stereosanthus Franch. (1896).
Vierhapperia Hand.-Mazz. (1937).
Perennial herbs; stems, leaves and involucres
stipitate-glandular, usually with greyish-white
tomentum, especially on leaf undersides. Basal
leaves large, petiolate, elliptic-lanceolate, toothed,
pinnately veined, cauline leaves reduced, sessile,
decurrent to clasping. Inflorescences loosely
corymbose; heads large; involucres broadly
campanulate; bracts 2–3-seriate, subequal, herbaceous, oblong-lanceolate. Florets trimorphic; rays
pistillate, yellow. Inner pistillate florets in several
series, tubular, eradiate. Disc florets functionally
male, yellow, limbs broadened, tubular, lobes
oblong-lanceolate, erect to spreading. Fertile
achenes oblong to fusiform, subterete, 8–10ribbed; pappus 1-seriate, caducous, slender above
base in disc achenes. Nine species, south-central
China.
Nannoglottis was placed in Solidagininae by
Nesom (1994c) but its evolutionary origin has since
been shown to be basal or near-basal in Astereae
(Liu et al. 2002).
766. Oclemena Greene
764. Microglossa DC.
Microglossa DC., Prodr. 5: 320 (1836).
Erect to scandent shrubs, not viscid nor glutinous.
Leaves petiolate, membranaceous to coriaceous,
oblong-lanceolate to ovate, entire or serrate, with
strongly ascending secondary veins. Inflorescences
Oclemena Greene, Leafl. Bot. Observ. Crit. 1: 4 (1903).
Perennial herbs; puberulous with coloured hairs;
unbranched below inflorescence. Leaves all cauline,
lowest scale-like, larger towards middle, sessile
to short-petiolate, elliptic to oblanceolate, entire
or serrate, gland-dotted. Inflorescences loosely
Compositae
corymbose, with arching peduncles; involucres
broadly turbinate; bracts 3–4-seriate, gradate,
linear-lanceolate, thin-herbaceous, slightly keeled,
without green patch; receptacles plano-convex,
epaleate. Rays numerous, pistillate, white to
pinkish-purple, long, not coiling. Disc florets
perfect, pink or reddish, lobes deltate, erect; style
appendages lanceolate, distally papillose. Achenes
fusiform to narrowly obovate or oblong, densely
gland-dotted, 4–8-nerved, often compressed with
thicker marginal nerves; pappus bristles 2–3seriate, subequal, outer series sometimes short.
x = 9. Three species, eastern North America.
767. Oreostemma Greene
Oreostemma Greene, Pittonia 4, 23: 224 (1900); Nesom,
Phytologia 74: 305–316 (1993), rev.
Oreastrum Greene (1896), nom. illegit., non Oriastrum
Poepp. (1838).
Perennial, taprooted herbs, sometimes rhizomatous or with branching caudex; eglandular or
stipitate-glandular; upper stems and involucres
loosely tomentose. Leaves in basal rosette linear to oblanceolate, entire, 3-nerved, cauline
leaves reduced. Inflorescence essentially scapose,
1-headed; involucres broadly turbinate; bracts
3–4-seriate, subequal, linear to linear-elliptic,
mostly leaf-like, often with low keel, basal margins
sometimes indurate; receptacles plano-convex,
with subspiniform ridge-junctions, epaleate. Rays
numerous, pistillate, white to purplish. Disc florets
perfect, yellow to reddish, tubular, lobes deltate,
erect; style appendages linear-lanceolate. Achenes
narrowly cylindric, with 5–10 raised nerves,
glabrous or sparsely strigose; pappus bristles
1-seriate, sometimes some in outer series short.
x = 9. Three species, western North America.
768. Psiadia Jacq.
Psiadia Jacq., Pl. Rar. Hort. Schoenbr. 2: 13, t. 152 (1800).
Glabrous or glabrescent to pubescent shrubs;
stems and leaves glabrous and glandular-viscid
to densely tomentose. Leaves petiolate, oblongelliptic to lanceolate, mucronate to acuminate,
entire to serrate, venation pinnate. Inflorescences
corymbose; peduncles to 2 cm long; involucres
hemispheric; bracts 3–5-seriate, gradate, oblong
to lanceolate, obtuse, margins membranaceous
to hyaline; receptacles plano-convex, minutely
paleate, fimbriate or glabrous. Pistillate florets
several-seriate, yellow, radiate or tubular with 2–5
289
lobules. Disc florets functionally male, yellow, narrowly funnelform, lobes deltate, erect-spreading.
Fertile achenes narrowly turbinate, not or scarcely
compressed, 3–6-costate; disc achenes minute;
pappus bristles 1-seriate, persistent, united at
base. Circa 60 species, tropical Africa, Madagascar,
Mascarenes.
769. Psiadiella Humbert
Psiadiella Humbert, Mém. Soc. Linn. Normandie 25: 39, 282
(1923).
Glutinous, ericoid subshrubs with sessile glands,
often hirsute with pale hairs. Leaves oblanceolate to
linear, entire, 1-nerved. Inflorescences corymbose,
heads 2–5, disciform, densely clustered; involucres
campanulate; bracts few, c. 2-seriate, unequal, glandular, obovate-oblong, apices obtuse to rounded,
margins scarious; receptacles flat, bare. Pistillate
florets 1 or 2, yellow, tubular, truncate apically.
Disc florets 3–6, functionally male, yellow, funnelform, lobes deltate, erect; style branches lanceolate, papillose. Achenes oblong-cylindric, truncate
above, weakly 4–5-costate, weakly setulose; pappus
1-seriate, of 2–8 unequal, linear to lanceolate scales.
One species, P. humilis Humbert, Madagascar.
770. Sarcanthemum Cass.
Sarcanthemum Cass., Bull. Sci. Soc. Philom. Paris 1818: 74
(1818).
Shrubs; glabrous, viscous; stems erect, moderately
branched. Leaves petiolate, blades oblongoblanceolate, margins entire in basal half, deeply
oblong-dentate distally. Inflorescences corymbose; involucres hemispheric; bracts 2–3-seriate,
subequal, appressed, ovate-oblong, coriaceous,
margins membranaceous; receptacles plane, with
small scales on disc. Pistillate florets severalseriate, tubular, narrowed above, with small
limb. Disc florets numerous, functionally male,
yellow, thickened and fleshy at base of limb, lobes
5, equal. Fertile achenes obovoid, compressed,
glabrous, striate; disc achenes minute; pappus
of disc achenes of long, flexuous-tipped, basally
connate scales, rudimentary on fertile achenes.
One species, S. coronopus Cass., Rodrigues Island.
771. Tonestus A. Nelson
Tonestus A. Nelson, Bot. Gaz. (Crawfordsville) 37: 262
(1904); Nesom & Morgan, Phytologia 68: 174–180 (1990),
emend; Brouillet et al., Sida 21: 889–900 (2004), phylog.
290
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Perennial herbs from branching caudices or
slender rhizomes; stems, leaves, and involucres
stipitate-glandular or seldom eglandular. Leaves
subclasping, often persistent in rosette, blades
obovate, 3-veined from base, margins spinulosetoothed or entire, upper cauline leaves reduced.
Inflorescences 1- to few-headed, often loosely
corymbose; involucres turbinate to hemispheric
or campanulate; bracts 3–4-seriate, subequal, at
least outer leaf-like; receptacles plano-convex,
epaleate. Rays absent or 1–2-seriate, pistillate,
yellow or white, limbs long, not coiling. Disc florets
perfect, yellow, narrowly tubular; style appendages
lanceolate, papillose. Achenes narrowly oblong
to subfusiform, terete to rather compressed, 5-,
5–8-, or 8–12-nerved, glabrous or strigose; pappus
bristles 1(–2)-seriate, attenuate, sometimes with
short outer series. x = 9. Eight species, western
USA, south-western Canada.
772. Vernoniopsis Humbert
Vernoniopsis Humbert, Mém. Inst. Sci. Madagascar sér. B,
Biol. Vég. 6: 154 (1955).
Shrubs or small trees, moderately branched;
stems with dense, evanescent, bristly hairs. Leaves
oblanceolate, glabrous, entire, basally cuneate and
petioliform, apically rounded, venation pinnate.
Inflorescences terminal, densely corymbose;
involucres narrowly cylindric; bracts 4–5-seriate,
strongly gradate, ovate to linear-lanceolate,
short-acute, smooth outside; receptacles bare.
Rays lacking. Disc florets 1–4, perfect, white,
funnelform with rather abruptly widened limb,
lobes linear-lanceolate, spreading to reflexed;
thecae bases calcarate; style base bulbiform;
style appendages deltate. Achenes cylindrical,
10-ribbed, glabrous; pappus persistent, bristles
coherent in basal ring, barbellate, inner bristles
longer, outer progressively shorter. One species,
V. caudata (Drake) Humbert, Madagascar.
XV.1. Subtribe Homochrominae Benth.
in Benth. & Hook. f. (1873).
Subtribe Feliciinae G.L. Nesom (1994).
Annual to perennial herbs, subshrubs, or shrubs,
strigose to hirsute or villous, lanate in Lachnophyllum. Leaves mostly alternate, opposite in species of
Amellus, Felicia, Engleria, Poecilolepis and Jeffreya,
entire, rarely toothed or lobed. Heads solitary or
few, glomerate in Nolletia and Chrysocoma; involu-
cral bracts not keeled, often with broad hyaline
margins; receptacles plane to plano-convex, with or
without pales. Ray florets 1-seriate, corollas white
to blue, less commonly yellow, coiling. Disc florets
usually bisexual, functionally male in some; style
branch appendages usually triangular-deltate, papillose. Achenes compressed, 2-nerved, eglandular
(glandular in Nolletia), setulae with anchor-shaped
tips only in Amellus; pappus 1–2-seriate (of bristles
and scales), rarely absent (Jeffreya). x = 9, reduced
to 8, 6, 5.
Key to the Genera
1. Receptacle with acuminate pales
773. Amellus
– Receptacle epaleate or with only small deciduous
scale-like pales
2
2. Achenes epappose or with only small apical annulus
779. Jeffreya
– Achenes with few to many pappus bristles
3
3. Pappus bristles basally connate, falling as a unit; annuals or short-lived perennial herbs; Asia
4
– Pappus bristles not or scarcely basally connate; annual
herbs to shrubs; mostly Africa (to Arabia in some
Felicia, to Spain in Nolletia)
5
4. Inflorescence with heads on many branches; involucral
bracts c. 3-seriate
774. Chamaegeron
– Heads solitary; involucral bracts 5–7-seriate
780. Lachnophyllum
5. Heads mostly homogamous, without ray florets
6
– Heads heterogamous, usually with ray florets
7
6. Leaves alternate; inflorescence corymbose; corollas
yellow, without resin glands in lobes
775. Chrysocoma
– Leaves opposite; heads solitary; corollas purplish, with
pairs of elongate, marginal resin glands by sinuses
784. Roodebergia
7. Achenes with wings containing septate resin ducts
776. Engleria
– Achenes not winged
8
8. Leaves opposite or alternate; heads solitary on erect
peduncles; disc florets perfect or sometimes functionally male; pappus never plumose
9
– Leaves alternate; heads solitary on erect peduncles or
grouped; disc florets often functionally male, less often
perfect; pappus sometimes plumose
10
9. Disc floret style appendages relatively evenly
low-papillose; anthers truncate at the base 777. Felicia
– Disc floret style appendages with a ring of markedly
longer collecting hairs at the base of each; anthers
slightly auriculate at the base
782. Poecilolepis
10. Achenes surfaces ‘pocked’ and appearing glandular;
disc florets perfect or functionally male 781. Nolletia
– Achene surfaces smooth or minutely pebbled; disc florets functionally male
11
11. Achenes with tan, smooth faces, with an apical bony
collar; pappus bristles scabrid; limbs of ray florets with
a purple abaxial midstripe
783. Polyarrhena
– Achenes with black, minutely pebbled faces, without
apical bony collar; pappus bristles plumose to scabrid;
limbs of ray florets without an abaxial midstripe 12
Compositae
12. Achenes densely white-papillose from short setulae
with a rounded clavate apex, glabrous in several
species; pappus of disc florets of 2 or 3 bristles, of ray
florets of 1 or 2 bristles or absent
778. Gymnostephium
– Achenes strigose-sericeous with long, filiform setulae;
pappus of disc and ray florets of 8–12 bristles
785. Zyrphelis
291
Genera of Homochrominae
773. Amellus L.
Fig. 67
Amellus L., Syst. Nat., ed. 10, 2: 1225, 1377 (1759), nom.
cons.; Rommel, Mitt. Bot. Staatssamml. München 13:
579–728 (1977), rev.
Erect to spreading annual or perennial herbs or
subshrubs, stems, leaves and involucres variously
densely puberulous to hirsute with stiff hairs.
Leaves alternate to opposite, bases sessile to
nearly sessile, linear or elliptic to oblanceolate.
Inflorescences 1-headed to loosely corymbose;
involucres campanulate to hemispheric; bracts
3–4-seriate, subequal, linear-lanceolate, naviculate; pales linear-lanceolate. Rays 0 to 1–2-seriate,
pistillate, white to purplish, coiling. Disc florets
yellow, throat cylindric to funnelform, often
with resin pockets along veins, lobes deltate,
erect to recurved; style appendages lanceolate to
linear. Achenes obovate, compressed to trigonous,
2–5(–7)-ribbed; pappus usually of c. 5, caducous,
hispid bristles alternating with persistent scales.
Twelve species, southern Africa.
774. Chamaegeron Schrenk
Chamaegeron Schrenk, Bull. Cl. Phys.-Math. Acad. Imp. Sci.
Saint-Pétersbourg 3: 107 (1845); Botschantzev, Fl. U.R.S.S.
25: 118–121 (1959), Engl. transl. 111–113 (1999), reg. rev.
Fig. 67. Compositae-Astereae. Amellus asteroides. A Habit.
B Capitulum with ray and disc florets. C Receptacular palea
with reddish resin duct along keel. D Ray floret showing
style, fertile achene with minute setulae (zwillingshaare),
and coiling of older corolla limb. E Disc floret with fertile
achene showing reddish resin ducts distally along veins of
corolla throat. F Disc floret in longitudinal section showing
anthers with spurred or tailed bases and style with paired
stigmatic lines and triangular, externally papillose style appendages (Drawing by A.R. Tangerini)
Annual or biennial, sparingly branched herbs from
weak taproot, with stipitate glands, sometimes
with longer erect hairs. Leaves cauline, sessile,
narrow to oblanceolate, entire or dentate, with
short, fragile cartilaginous apex. Branches of
inflorescence with terminal, narrowly pedunculate
heads; involucres campanulate; bracts c. 3-seriate,
gradate, herbaceous, inner oblanceolate, apically
fimbriate, margins membranaceous; receptacles
flat, glabrous. Rays 1–2-seriate, pistillate, lilac,
limbs coiling. Disc corollas pale yellow, narrowly
funnelform, with hairs on lower half, outer lobe
separated from other triangular lobes by deeper
sinuses; style appendages acuminate. Achenes
elliptic-oblong to oblanceolate, glabrous or setulose; pappus bristles white, 1-seriate, connate in
basal ring, caducous with ring. x = 9. Four species,
central Asia, Iran, Afghanistan, Pakistan.
775. Chrysocoma L.
Chrysocoma L., Sp. Pl. 2: 840 (1753).
Perennial ericoid herbs or shrubs; glabrous, viscid.
Leaves linear to oblanceolate, usually entire.
292
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Inflorescences corymbose or with heads solitary
on leafy branches, peduncles short to long; involucres hemispheric to broadly campanulate; bracts
few-seriate, imbricate, margins membranaceous,
subserrulate or subentire, midrib darker. Rays
absent. Disc corollas yellow, base cylindric, limb
narrowly funnelform, lobes triangular, erect.
Achenes obovoid, somewhat compressed, pilose;
pappus bristles 1-seriate, caducous, white, as long
as corolla. Twenty species, southern Africa.
lobes deltate to lanceolate, spreading to recurved.
Achenes elliptic-obovate; pappus white, 1-seriate,
bristles sometimes caducous, rarely lacking in rays.
x = 9, reduced to 8, 6, 5. Circa 85 species, Africa,
Arabia.
776. Engleria O. Hoffm.
Branching subshrubs; hairs stout, usually spreading. Leaves almost all cauline, linear, entire.
Inflorescences usually corymbose with numerous heads; involucres campanulate to obconic;
bracts 3–4-seriate, gradate, lanceolate to linearlanceolate, with broad scarious margins. Rays
1-seriate, pistillate, white to blue, narrow. Disc
florets functionally male, yellow, narrowly
funnelform, lobes triangular. Fertile achenes
oblong-obovate, black at maturity, faces pebbled,
eglandular, glabrous or with dense, white, papilliform setulae; pappus 1-seriate, caducous, lacking
in ray florets, or with 1 or 2 plumose or barbellate
bristles, disc florets with more bristles. Eight
species, southern Africa.
Engleria O. Hoffm., Bot. Jahrb. Syst. 10: 273 (1888).
Much-branched perennial herbs or shrubs;
branches terete, striate. Leaves opposite, some
above subopposite, narrowly petiolate, blades
membranaceous, elliptic, acute, dentate. Inflorescence diffuse, with single long-pedunculate
heads terminal or appearing axillary; involucres
broadly ovoid to campanulate; bracts persistent,
4–5-seriate, gradate, linear-lanceolate, outer
squarrose, inner erect-spreading, margins scarious; receptacles convex, epaleate. Ray florets
0 or 1-seriate, pistillate; corollas yellow. Disc
florets 20–30, perfect; corollas yellow, lobes 5,
broadly triangular, recurved; style appendages
linear, obtuse. Achenes obovate, rounded distally
to narrow apical callus, compressed, setulose on
margins and middle of faces, 1-nervate on faces,
margins with narrow wings containing septate
resin ducts; pappus sub-2-seriate, bristles unequal,
barbellate, with shorter narrow scales outside. Two
species, Angola, Namibia.
777. Felicia Cass.
Felicia Cass., Bull. Sci. Soc. Philom. Paris 1818: 165 (1818),
nom. cons.; Grau, Mitt. Bot. Staatssamml. München 9:
195–705 (1973), rev.
Detris Adans. (1763).
Annual or perennial often ericoid herbs or shrubs;
glabrous to strigose, hirtellous or pilose. Leaves
alternate or opposite, sessile or bases winged,
herbaceous to coriaceous, filiform to oblong,
margins usually entire. Inflorescences 1-headed;
peduncles long; involucres hemispheric; bracts c.
2-seriate, subequal to gradate, linear-lanceolate,
dark along midline, sometimes with resin pocket
distally, margins scarious. Rays 1-seriate, rarely
absent, pistillate, blue, violet or rarely white or
yellow. Disc florets sometimes functionally male,
yellow, tubes narrow, limb narrowly funnelform,
778. Gymnostephium Less.
Gymnostephium Less., Syn. Gen. Comp. 185 (1832); Nesom,
Phytologia 76: 85–95 (1994), rev.
Heteractis DC. (1838).
779. Jeffreya Wild
Jeffreya Wild, Kirkia 9: 295 (1974).
Annual herbs; mostly glabrous, glandular-pilose
below heads, stems simple below, sparsely
branched above, terete, 8–10-ribbed. Lower leaves
opposite, others alternate, sessile, bases subclasping, blades linear, entire or remotely serrulate,
narrowly revolute, obtusely callus-tipped. Heads
solitary on long peduncles; involucres broadly
campanulate to subpatelliform; bracts 1–2-seriate,
subequal, margins serrulate-ciliate, not hyaline.
Rays 1-seriate, pistillate, elongate, white to blue or
violet. Disc florets deep yellow to orange-yellow,
tube cylindrical, limb campanulate, lobes deltate,
erect; style appendages linear, minutely puberulous. Achenes ellipsoid, somewhat compressed,
margins slightly costate, smooth, glabrous; pappus
absent or a small annulus. One species, J. palustris
(O. Hoffm.) Wild, Tanzania.
780. Lachnophyllum Bunge
Lachnophyllum Bunge, Beitr. Fl. Russl. 151 (1852);
Botschantzev, Fl. U.R.S.S. 25: 288–289 (1959), Engl. transl.
270–271 (1999), reg. rev.
Compositae
Erect, aromatic, annual or biennial herbs; with
long hairs and stipitate glands, often lanate. Leaves
entire, lowest petiolate, obtuse, middle leaves
sessile, ovate, acuminate. Inflorescence thyrsoid
to racemose; heads pedunculate; involucres
broadly campanulate; bracts 5–7-seriate, gradate,
recurved, outer lanceolate, herbaceous, inner
linear-lanceolate, with midvein green, margins
membranaceous. Rays 2-seriate, pistillate, blue
to pink or violet. Disc florets yellow, with yellow
ducts, tubes slender, limbs narrowly funnelform,
lobes 5, lanceolate, recurving; style appendages
lanceolate, hairy. Achenes oblanceolate, sericeousvillous, abruptly narrowed distally; pappus yellow,
1-seriate, bristles, basally connate. x = 9. Two
species, central Asia.
781. Nolletia Cass.
Nolletia Cass., Dict. Sci. Nat. 37: 479 (1825).
Erect perennial herbs or shrubs; strigose or hispid
to glabrous, rarely with stipitate glands; stems
often clustered. Leaves sessile, linear to subulate.
Heads solitary on branches or in subcorymbose
inflorescences; peduncles long; involucres hemispheric; bracts few-seriate, gradate, linear to
oblanceolate-linear, acute or acuminate, margins
hyaline. Rays 1-seriate, pistillate, yellow, narrowly cylindric, apex fimbriate or 2–3(–5)-lobed.
Disc florets perfect or functionally male, yellow,
cylindric below, limb narrowly funnelform, lobes
triangular, erect-spreading; style appendages
linear. Achenes ellipsoid, surfaces puberulous to
strigillose; pappus 1-seriate, of slender, caducous
setae. Ten species, Africa, one to Spain.
782. Poecilolepis J. Grau
Poecilolepis J. Grau, Mitt. Bot. Staatssamml. München 13:
244 (1977).
Small, decumbent, annual or perennial herbs;
glabrescent. Leaves alternate or opposite, fleshy,
linear, entire, bases subamplexicaul. Inflorescences
of axillary, aphyllous, 1-headed peduncles; involucres broadly campanulate; bracts c. 3-seriate,
weakly to strongly gradate, fleshy, ovoid to oblong,
margins narrowly scarious; receptacles with or
without deciduous scales. Rays to 15, 1-seriate,
pistillate, white, broad. Disc florets to c. 15,
yellow, narrowly funnelform from short, broad
tube, lobes 5, deltate, erect; anther bases slightly
auriculate; style appendages short-deltate, with
a basal ring of longer collecting hairs. Achenes
293
obovate, slightly compressed, smooth, setulose,
2–3-costate; pappus setae few to 8, short, papillose
to scabrid, caducous. Two species, South Africa,
Cape Province, maritime.
783. Polyarrhena Cass.
Polyarrhena Cass., Dict. Sci. Nat. 56: 172 (1828); Grau, Mitt.
Bot. Staatssamml. München 7:347–368 (1970), rev.
Ericoid shrubs; hirsute with non-glandular and/or
stipitate-glandular hairs. Leaves sessile, coriaceous,
ascending to recurved, linear to oblong, margins
dentate to ciliate. Heads solitary on erect peduncles; involucres broadly campanulate; bracts 2–3seriate, mostly subequal, oblong-lanceolate to linear, obtuse to acute, indurate and yellowish at base,
margins narrowly scarious, narrow green patch
distally, with median, yellowish, resin duct. Rays
1–2-seriate, pistillate, white, often purple adaxially. Disc florets perfect or functionally male, yellow, funnelform, lobes triangular, spreading or recurved; style sterile or with deltate appendages.
Achenes ellipsoid-oblong, with pale apical collar;
pappus bristles 1-seriate, caducous. x = 9. Four
species, southern Africa.
784. Roodebergia B. Nord.
Roodebergia B. Nord., Acta Phytotax. Geobot. 53: 101
(2002).
Perennial herbs; primary stems repent, branches
ascending, hirsute-hispid, glanduliferous; leaves
opposite, sessile, narrowly elliptic, entire. Heads
solitary, long-pedunculate; involucral bracts c. 10,
c. 2-seriate, subequal, oblong-lanceolate, glanduliferous and sparsely setiferous. Rays lacking.
Disc florets 15–25, purplish, tubular, wider above,
mostly glabrous, lobes 5, triangular-ovate, each
with elongate resin gland on each margin by sinus.
Achenes elliptic-oblong, subglabrous, with minute
setulae especially proximally; pappus 2-seriate,
bristles numerous, scarcely connate at base, not
falling as unit. One species, R. kitamurana B. Nord.,
southern Africa.
785. Zyrphelis Cass.
Zyrphelis Cass., Ann. Sci. Nat. (Paris) 17: 420 (1829);
Nesom, Phytologia 76: 85–95 (1994), emend., comb.
Homochroma DC. (1836).
Perennial herbs or subshrubs; hairs stout, usually
spreading; plants acaulescent or stems branching.
Leaves almost all cauline, sessile, linear. Inflo-
294
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
rescences with numerous heads or less often
1-headed; peduncles long; involucres broadly
campanulate to obconic; bracts c. 3-seriate,
gradate, ovate to lanceolate, margins broadly
scarious. Rays 1-seriate, pistillate, white to blue or
yellow, narrow, tightly coiling. Disc florets usually
functionally male, funnelform. Fertile achenes
black at maturity, oblong-obovate, faces pebbled,
eglandular, surfaces densely strigose-sericeous,
setulae filiform with evenly connate tips; pappus
plumose, 1-seriate, bristles 8–12, caducous. Ten
species, southern Africa.
XV.2. Subtribe Hinterhuberinae Cuatrec.
(1969).
Perennial herbs, subshrubs, or shrubs. Leaves
alternate, opposite in Pteronia and Olearia, mostly
entire, resinous-glandular, punctuate-glandular
or eglandular. Heads mostly in a corymboid
inflorescence; involucral bracts usually not keeled;
receptacles plano-convex, with or without pales.
Ray florets 1(–3)-seriate, corollas white or yellow,
conspicuous or variously modified or reduced.
Disc florets bisexual or sometimes functionally
male; style branch appendages usually narrowly
lanceolate, papillose. Achenes multinerved, mostly
terete, less commonly compressed and 2-nerved,
commonly glandular, setulae without anchorshaped tips. Pappus (1–)2(–3)-seriate, usually of
even-length bristles. x = 9.
Key to the Genera
1.
–
2.
–
3.
–
4.
–
5.
–
6.
–
Receptacles with pales
2
Receptacles without pales
8
Acaulescent herbs
786. Achnophora
Shrubs
3
Rays present and white; heads borne on abrupt leafless
and ebracteose peduncles
793. Chiliotrichum
Rays absent or yellow; heads sessile or borne on
bracteose peduncles
4
Involucral bracts lacerate-ciliate-margined, scarcely
indurate, persistent; ray florets often present
5
Involucral bracts smooth-margined, indurate, easily
deciduous; ray florets absent
6
Ray florets present; pappus of barbellate bristles
equal to disc corollas in length; achenes not evidently
glandular-viscid
791. Chiliophyllum
Ray florets present or absent; pappus of laceratefimbriate scales c. half as long as disc corollas; achenes
sometimes strongly glandular-viscid
792. Chiliotrichiopsis
Pappus of barbellate bristles equal to disc corolla in
length
805. Nardophyllum
Pappus of broad pales or scales equal or less than equal
to disc corolla length
7
7. Pappus of scales 1–2 mm high; achenes strongly flattened and 2-nerved, strigose-sericeous on margins,
faces glabrous
787. Aylacophora
– Pappus of pales equal to disc corolla in length; achenes
turbinate and multinerved, evenly hirsute-strigose
811. Paleaepappus
8. Dioecious shrubs
788. Aztecaster
– Monoecious herbs, shrubs or trees
9
9. Densely branched, cupressiform, persistently
aromatic shrub
10
– Herbs or shrubs not cupressiform, not notably aromatic
11
10. Pappus of narrowly subulate, caducous scales
801. Lepidophyllum
– Pappus of many long inner bristles and some short
outer scales or bristles
812. Parastrephia
11. Heads homogamous, without pistillate florets
12
– Heads heterogamous, with peripheral pistillate florets
13
12. Heads in corymbose inflorescences; receptacles
columnar, with florets borne on narrow convex tip
802. Llerasia
– Heads usually solitary at ends of branches; receptacles
flat, convex, or concave
815. Pteronia
13. Pistillate florets tubular or subbilabiate, often 5-lobed;
disc florets functionally male
14
– Pistillate florets usually with long or short limbs; disc
florets perfect or functionally male
18
14. Pistillate florets edentate or scarcely dentate at tip 15
– Pistillate corollas with 4 or 5 distinct narrow lobes 16
15. Small bromeliad-like rosettiform plants with thickened, glabrous leaves; pistillate corollas not broadened
distally
806. Novenia
– Small shrubby or subshrubby plants with pubescent,
sometimes stipitate-glandular leaves; pistillate corollas ampliate above middle and slightly constricted at
tip
818. Westoniella
16. Shrubs, often ericoid
799. Hinterhubera
– Erect or creeping herbs
17
17. Pappus present
797. Flosmutisia
– Pappus lacking
800. Laestadia
18. Ray achenes linear or fusiform with distinct distal
rostrum
789. Blakiella
– Ray achenes without rostrum
19
19. Herbs
20
– Shrubs or trees
25
20. Pappus bristles plumose
21
– Pappus bristles not plumose
22
21. Disc corollas narrowly funnelform; Auckland and
Campbell Islands
794. Damnamenia
– Disc corollas goblet-formed, with narrow tube and
abruptly cyathiform limb; S Africa
804. Mairia
22. Disc florets functionally male
23
– Disc florets perfect
24
23. Stems branching, prostrate; leaves cauline; achenes
glandular; pappus bristles almost smooth
796. Floscaldasia
– Stems simple, erect; leaves in a basal rosette; achenes
glandular; pappus bristles scabrid to barbellate
808. Oritrophium
24. Rosulate herbs with narrow mostly 1-nerved leaves;
inflorescences of long-pedunculate, solitary heads
790. Celmisia
Compositae
– Robust herbs, leaf blades broad with many longitudinal veins; inflorescences racemose
813. Pleurophyllum
25. Disc florets functionally male
26
– Disc florets perfect
28
26. Leaves clustered at tips of branches, with weak longitudinal venation; inflorescence a sessile terminal cluster
of heads; ray achenes only slightly compressed, with
9–11 ribs
810. Pacifigeron
– Leaves usually persistent on younger stems, with pinnate venation; inflorescences usually branched with
pedunculate heads; ray achenes distinctly compressed,
with 3–5 ribs
27
27. Pappus of numerous bristles almost as long as disc
corollas; disc florets of one type, all functionally male
795. Diplostephium
– Pappus of 3–9(–14) unequal to subequal, persistent,
somewhat flattened bristles; outer disc florets often
pistillate
816. Remya
28. Anther thecae with long basal tails
814. Printzia
– Anther bases without tails
29
29. Ray corollas yellow
30
– Ray corollas white to bluish or purplish
31
30. Heads 1–3(–12), with obvious rays; involucre broadly
campanulate
817. Rochonia
– Heads c. 15–40 in racemiform or thysiform inflorescence, with short pistillate florets; involucre cylindrical
798. Guynesomia
31. Disc corolla limbs narrowly campanulate, with slightly
spreading lobes; achenes (4–)5–8-ribbed
803. Madagaster
– Disc corolla limbs narrowly funnelform with spreading or recurved lobes; achenes 4–5-ribbed or grooved
32
32. Involucral bracts 2–6-seriate, subglabrous to weakly
tomentose, inner bracts less than 14 mm long; pappus
bristles unequal in length; achenes glabrous to setulose; capitula solitary or in compound inflorescence
807. Olearia
– Involucral bracts 8–9-seriate, densely tomentose, inner bracts more than 14 mm long; pappus bristles
equal in length; achenes densely sericeous; capitula
solitary
809. Pachystegia
Genera of Hinterhuberinae
786. Achnophora F. Muell.
Achnophora F. Muell., Trans. Proc. Roy. Soc. S. Australia 6:
32 (1883).
Acaulescent perennial herbs; rhizome thick, vertical; glabrous. Leaves in basal rosette, linear, basally
sheathing. Inflorescence scapiform, 1-headed; involucre nearly hemispheric; bracts c. 3-seriate, gradate, ovate to obovate, subacute, scarcely scarious distally; receptacles conical, with deciduous
pales. Rays 1-seriate, pistillate, white, long. Disc florets functionally male, yellow, tubular below, narrowly funnelform above, 5(–4) lobes deltoid, erectspreading. Achenes obovate, cuneate at base, some-
295
what 3–4-angled; pappus persistent, segments 8–
12, sublanceolate-subulate, flat, slightly ciliate. One
species, A. tatei F. Muell., Australia, Kangaroo Island.
787. Aylacophora Cabrera
Aylacophora Cabrera, Bol. Soc. Argent. Bot. 4: 261 (1953);
Bonifacino & Sancho, Sida 19: 531–538 (2001), emend.
Intricately branching, subaphyllous shrubs;
branches glabrous with tomentose grooves. Leaves
sparse, linear, entire, acute, with evanescent
tomentum. Heads terminal on branchlets, discoid;
involucres broadly campanulate; bracts 5–6seriate, gradate, ovate to oblong-lanceolate, easily
deciduous, tomentose, margins smooth; receptacular pales oblong-lanceolate, conduplicate, distally
ciliate, acuminate. Florets 20–25, perfect, yellow,
funnelform, lobes lanceolate, erect-spreading.
Achenes obovate, compressed, 2-nerved, margins
and apex strigose-sericeous, faces glabrous;
pappus 1-seriate, of c. 20–25 lanceolate scales c.
1.2 mm long. One species, A. deserticola Cabrera,
central Argentina.
788. Aztecaster G.L. Nesom
Aztecaster G.L. Nesom, Phytologia 75: 64 (1993).
Dioecious shrubs; stems silvery-white tomentose, glabrate. Leaves densely spiralled, often in
axillary fascicles, linear-oblong, entire, flocculent, glutinous, margins revolute. Inflorescences
spicate; heads sessile, subtended by clustered
leaves; involucres turbinate; bracts 3–4-seriate,
gradate, narrowly ovate to lanceolate, yellowish
to greenish-yellow, resinous, narrowly keeled,
margins scarious; receptacles epaleate. Female
heads with 5–10 florets; corollas narrowly tubular, 5-lobed; male heads with 8–9 functionally
male florets; corollas narrowly tubular, lobes
short-triangular, spreading. Fertile achenes
oblong-ovate, compressed with 2 marginal nerves
and 1 nerve on 1 or both faces, strigose; pappus
bristles 1-seriate, numerous. Two species, Mexico.
789. Blakiella Cuatrec.
Blakiella Cuatrec., Webbia 24: 37 (1969).
Perennial subshrubs, hirsutulous and densely
stipitate-glandular. Leaves sessile, base subauriculate, margins crenulate, revolute. Inflorescences
of few clustered long-pedunculate heads; involucres cupulate-campanulate; receptacles epaleate.
296
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Peripheral florets multiseriate, 200–270, pistillate,
yellow, tube subcapillary, distally broadened,
limbs short, slightly constricted at mouth. Disc
florets 55–85, functionally male, yellow, tube
narrow, limb campanulate, lobes triangular. Ray
achenes obovate-fusiform, c. 4-nervate, base
attenuate, distally short-rostrate; disc achenes
narrowly cylindrical, setulose and glandular;
pappus 1-seriate, bristles broadened apically in
disc florets. One species, B. bartsiifolia (S.F. Blake)
Cuatrec., Colombia, Venezuela.
subequal or strongly unequal. Three species,
Argentina, Chile.
792. Chiliotrichiopsis Cabrera
Fig. 68
Chiliotrichiopsis Cabrera, Notas Mus. La Plata, Bot. 2: 172
(1937); Nesom et al., Brittonia 53: 430–434 (2001), emend.
Perennial, glutinous, ericoid shrubs; stems
white-tomentose, glabrescent, brownish-lined,
blackening. Leaves sessile, linear to oblanceolate,
margins entire, revolute, glabrous above, tomen-
790. Celmisia Cass.
Celmisia Cass., Dict. Sci. Nat. 37: 259 (1825), nom. cons.;
Given, N. Z. J. Bot. 7: 400–418 (1969), emend.
Elcisimia B.L. Rob. (1913), nom. rej.
Perennial, mostly rosulate herbs from woody
caudex; stems scapiform, bracteolate. Leaves
mostly in rosettes, subcoriaceous, linear or filiform to elliptic-oblanceolate, with sheathing
bases, abaxially often with white tomentum.
Heads solitary; involucres broadly campanulate to
hemispheric; bracts 4–5-seriate, gradate, linear;
receptacles epaleate. Rays 1-seriate, pistillate,
white or partly violet to purple, not or weakly
coiling. Disc florets perfect, yellow to purplish,
narrowly funnelform, lobes 5, broadly triangular
to lanceolate, spreading; anther bases acute; style
appendages linear, branches papillose to near
base. Achenes elliptic to obovoid, compressed,
ribs marginal and 2 or 3 on each face, glabrous
to sericeous; pappus bristles numerous, unequal.
x = 9. Circa 60 species, New Zealand, Australia.
791. Chiliophyllum Phil.
Chiliophyllum Phil., Linnaea 33: 132 (1836), nom. cons.
Small ericoid shrubs; stems, leaf undersides and
involucres tomentose or glutinous. Leaves sessile,
linear-oblong to obovate, margins entire to revolute. Heads solitary on branchlets, short-pedunculate; involucres broadly campanulate; bracts
c. 15–25, 2–5-seriate, subindurate, persistent,
broadly lanceolate to linear-lanceolate, acute,
margins lacerate-ciliate; receptacular pales rather
conduplicate. Rays c. 8 or 9, 1-seriate, pistillate,
yellow, limbs broad. Disc florets perfect, yellow,
narrowly tubular-funnelform, lobes lanceolate,
spreading or recurved. Achenes oblanceolate,
scarcely compressed, ribbed, setulose, not evidently glandular; pappus bristles 1-seriate,
Fig. 68. Compositae-Astereae. Chiliotrichopsis peruviana.
A Flowering branch. B Capitulum. C Disc floret with palea.
D Receptacular palea. E Corolla and partly exserted anthers
and style of disc floret. F Disc floret, longitudinal section.
G Style of disc floret. H Achene with pappus of long scales.
(Nesom et al. 2001)
Compositae
tose beneath except on midvein. Heads solitary
on short branches, sessile; involucre campanulate;
bracts 4–5-seriate, gradate, persistent, lanceolate to oblong-lanceolate, bases pale, thickened,
median nerve green, margins broadly scarious,
lacerate-ciliate; receptacular pales conduplicate.
Rays pistillate, yellow or absent. Disc florets perfect, yellow, narrowly funnelform, lobes triangular,
recurved; style with stigmatic lines and distal
papillose outer surface overlapping. Achenes
oblanceolate, compressed, strigose-sericeous,
with intermixed resiniferous glands, margins
thickened; pappus of 10–16 lanceolate fimbriate
scales half as long as corollas. Three species,
Argentina, Peru.
793. Chiliotrichum Cass.
Chiliotrichum Cass., Bull. Sci. Soc. Philom. Paris 1817: 69
(1817).
Ericoid shrubs; stems, leaf undersides and
outer involucral bracts pale tomentose. Leaves
subcoriaceous, sessile, linear to oblanceolate,
entire, glabrous adaxially. Heads on clustered
ebracteate peduncles; involucres broadly campanulate to hemispheric; bracts 3–4-seriate, gradate,
caducous, mostly indurate, oblong-lanceolate,
subentire, narrowly scarious, green patches on
outer bracts; receptacular pales weakly conduplicate. Rays 1-seriate, pistillate, white or violet,
coiling. Disc florets perfect, yellow, tube narrow,
limb narrowly campanulate, lobes lanceolate,
recurved; style appendages linear, branches papillose almost to base. Achenes slender, 4–5-nerved,
with long hairs and glandular dots; pappus bristles
multiseriate, about as long as corollas. Two species,
Argentina, Chile.
794. Damnamenia D.R. Given
Damnamenia D.R. Given, N. Z. J. Bot. 11: 786 (1973).
Stoloniferous herbs, with branched caudex; stems
short and erect or scapiform. Primary leaves spirally inserted with imbricated bases, blades linear,
glossy, glabrous, venation simple, lateral veins of
base ending below blade. Inflorescences 1-headed,
on long leafy scapes; involucres campanulate;
bracts 2–3-seriate, subequal, linear-oblanceolate,
with non-glandular hairs; receptacles obconic.
Rays 1-seriate, pistillate, white to pale reddish
distally. Disc florets numerous, perfect, purple or
yellow, with slender tube and abruptly cyathiform
limb, lobes broadly triangular, erect-spreading;
297
anthers with slight tails; style appendages broadly
triangular, with long hairs. Achenes obconic,
setulose; pappus bristles c. 2-seriate, unequal,
plumose. x = 9. One species, D. vernicosa (Hook.
f.) D.R. Given, New Zealand.
795. Diplostephium Kunth
Diplostephium Kunth in H.B.K., Nov. Gen. Sp., ed. folio 4:
75 (1818); Cuatrecasas, Webbia 24: 90–194 (1969), reg. rev.
Shrubs or small trees; stems, leaf undersides and
involucres puberulous to lanate or gland-dotted.
Leaves usually petiolate, coriaceous, margins
plane or revolute. Inflorescences 1-headed to
corymbose or thyrsoid; involucres cylindric to
hemispheric; bracts 4–7-seriate, gradate, ovate to
oblong or linear; receptacles epaleate. Pistillate
florets 1–3-seriate, white to bluish or purplish,
radiate or eradiate. Disc florets functionally male
or rarely perfect, yellow, green or violaceous,
tubular-campanulate, lobes triangular. Achenes
glabrous, setuliferous or gland-dotted; fertile achenes obovoid, 2–3-nerved, somewhat compressed;
disc achenes oblong or linear, 5-nerved; pappus
bristles of disc achenes dilated at tips, outer series
of short setae or squamae. x = 9. Over 70 species,
Costa Rica and Andes to Bolivia.
796. Floscaldasia Cuatrec.
Floscaldasia Cuatrec., Webbia 24: 194 (1969); Sklená &
Robinson, Novon 10: 146–147 (2000), emend.
Minute herbaceous subshrubs; stems branching,
repent, rooting, glabrous; densely foliose branches
decumbent or erect. Leaves sessile, oblong or
trilobed, base vaginate, margins ciliate. Heads
solitary on erect hirsutulous scapes; bracteoles
few; involucres campanulate; bracts 2–3-seriate,
subequal, oblong-lanceolate to oblong, herbaceous; receptacles epaleate. Rays 26–45, pistillate,
reddish to purplish, limbs short. Disc florets 10–
12, functionally male, purplish, tube broad, limb
short-campanulate, lobes deltate; style branches
lanceolate, densely hairy. Fertile achenes obovoid,
scarcely compressed, 2-nervate, glandular along
outer margin; pappus white to brownish, subbiseriate, bristles c. 25, nearly smooth. Two species,
Colombia, Ecuador.
797. Flosmutisia Cuatrec.
Flosmutisia Cuatrec., Anales Jard. Bot. Madrid 42: 415
(1986).
298
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Rhizomatous, rosulate perennial herbs, pilose
and minutely stipitate-glandular. Leaves sessile, elliptic-oblong to obovate. Scapes with
narrow bracteoles; heads solitary; involucres
broadly campanulate; bracts 4–5-seriate, subequal,
oblong-lanceolate to oblanceolate, outer bracts
ciliate on margins and midvein, inner bracts
mostly glabrous; receptacles epaleate. Pistillate
florets 2–3-seriate, white, subbilabiate, 5-lobed,
outer lobes linear, the two inner lobes filiform.
Disc florets functionally male, white, funnelform
with expanded limb, lobes triangular, erectspreading, pilosulous. Fertile achenes ellipsoid,
compressed, densely setuliferous and glandular,
obscurely 5-nerved; disc achenes slender; pappus
whitish, bristles 1-seriate, barbellate. One species,
F. paramicola Cuatrec., Colombian paramo.
798. Guynesomia Bonifacino & Sancho
Guynesomia Bonifacino & Sancho, Taxon 53: 675 (2004).
Shrubs. Stems glandular. Leaves sparse, sessile, linear, clasping, entire, glandular on both surfaces.
Inflorescence racemiform or thyrsiform. Heads c.
15–40, pedunculate, disciform; involucres cylindric; bracts 4–5-seriate, coriaceous, ovate to linear, margins broadly membranaceous; receptacles
epaleate. Pistillate florets 10–11, 1-seriate, yellow,
limb 2–3 mm long, tridentate, lateral lobes reduced.
Disc florets 13–16, perfect, yellow, tubular, lobes
triangular, spreading. Achenes narrowly elliptic,
scarcely compressed, 3–4-ribbed, glandular; pappus bristles 2-seriate, subequal. One species, G. scoparia (Phil.) Bonifacino & Sancho, Chile.
799. Hinterhubera Sch. Bip. ex Wedd.
Hinterhubera Sch. Bip. ex Wedd., Chlor. And. 1: 185 (1857);
Cuatrecasas & Aristeguieta, Bol. Soc. Venez. Ci. Nat. 17:
98–104 (1956), rev.
Small ericoid shrubs, with sessile or stipitate
glands and non-glandular hairs; stems defoliated
below, densely covered with leaf bases or scars.
Leaves linear or oblong, margins revolute. Heads
disciform, sessile or pedunculate on branch
tips; involucres campanulate; bracts 4–5-seriate,
gradate, lanceolate to linear; receptacles epaleate.
Peripheral florets 90–170, pistillate, pluriseriate,
white or yellow, tubular, 4–5 lobules equal or
3 longer. Disc florets 9–60, functionally male,
white or yellow, tube narrow, limb deeply divided
into linear-lanceolate lobes; style appendages
linear, with hair-like papillae. Fertile achenes
obovate-oblong, with setulae and short-stipitate
glands, with 1(–3) marginal nerves; disc achenes
linear; pappus whitish to reddish, with few short
outer setae, bristles of disc achenes broadened
distally. Eight species, Colombia, Venezuela.
800. Laestadia Kunth ex Less.
Laestadia Kunth ex Less., Syn. Gen. Comp. 203 (1832).
Matted herbs, mostly procumbent, gland-dotted,
glutinous, sometimes hirtellous. Leaves crowded,
sessile, sometimes subclasping, oblong to spathulate, entire, 1-nerved. Heads on terminal or
axillary peduncles, disciform; involucres hemispheric; bracts 2–4-seriate, subequal, reflexing,
margins narrowly hyaline, outer bracts herbaceous
distally; receptacle epaleate. Peripheral florets
2–4-seriate, pistillate, regular with 4 or 5 deeply
cut, triangular, spreading or recurving lobes. Disc
florets 15–25, functionally male, white to purple,
abruptly expanded from short tube, limb cut to
near base, lobes triangular. Achenes oblong to
obovoid, nearly terete, 6–10-nerved, gland-dotted;
epappose, callus with ring of glands or on short
glandular beak. Six species, Costa Rica, Hispaniola,
Andes to Bolivia, high elevations.
801. Lepidophyllum Cass.
Lepidophyllum Cass., Bull. Sci. Soc. Philom. Paris 1816: 199
(1816); Cabrera, Bol. Soc. Argent. Bot. 1: 48–58 (1945), rev.
Cupressiform, densely branched shrubs; glabrous,
glutinous; stems subfrondiform with many short
spreading branches. Leaves decussate, scale-like,
closely abutting, as wide as long, rhombic, with
slightly rounded keel. Heads solitary on branch
tips, sessile; involucres narrowly campanulate;
bracts c. 3-seriate, lanceolate, middle indurate,
belatedly caducous, margins thinly scarious;
receptacles epaleate. Pistillate florets 3 or 4, yellow,
radiate or bilabiate with 3 outer lobes short,
coiling. Disc florets few, perfect, yellow, narrowly
funnelform, lobes lanceolate, recurved; style
appendages linear. Achenes oblanceolate, slightly
compressed, with c. 5 broad costae, papillose or
glabrous; pappus of many caducous, narrowly
subulate scales. x = 9. One species, L. cupressiforme
(Lam.) Cass., Patagonia.
802. Llerasia Triana
Llerasia Triana, Ann. Sci. Nat. Bot. 4, 9: 37 (1858); Cuatrecasas, Biotropica 2: 39–45 (1970), emend.
Compositae
Shrubs, small trees or vines, often gland-dotted;
stems and leaf undersides pale-tomentose. Leaves
short-petiolate, coriaceous, entire or dentate, pinnately veined. Inflorescences corymbose; heads homogamous; involucre cylindric; bracts 4–6-seriate,
strongly gradate, mostly obtuse, mostly deciduous;
receptacles columnar, epaleate, florets borne on
narrow convex tip. Florets 3–15, perfect, yellow,
tubular, limb slightly broader, lobes deeply cut,
linear-oblong, spreading; style base bulbous, appendages triangular, hairy. Achenes narrowly prismatic, 3–5-angled, 5–9-nerved, papillose, glanddotted or short-setulose; pappus tawny to white,
2–3-seriate. Ten species, Colombian to Bolivian Andes.
803. Madagaster G.L. Nesom
Madagaster G.L. Nesom, Phytologia 75: 97 (1993).
Shrubs or small trees; tomentose on young
branches, leaf undersides and petioles. Leaves
coriaceous, entire or distally toothed, venation
pinnate. Inflorescences 1-headed to corymbose
or thyrsoid; heads pedunculate; involucres campanulate to hemispheric; bracts 3–4-seriate,
gradate, ovate to oblong or linear, herbaceous at
tips, margins scarious; receptacles epaleate. Rays
1-seriate, pistillate, white to bluish. Disc florets
numerous, perfect, yellow, limbs narrowly campanulate, lobes triangular, erect-spreading; style
appendages short-triangular. Achenes narrowly
oblong-fusiform, (4–)5–8-ribbed, nearly terete to
slightly compressed, strigose, eglandular; pappus
1–2-seriate, bristles dilated apically, outer series
shorter, unequal. Five species, Madagascar.
804. Mairia Nees
Mairia Nees, Gen. Sp. Aster. 247 (1832); Nesom, Phytologia
76: 85–95 (1994), emend.
Annual or perennial herbs; with flexuous hairs;
stems scapiform, 1-headed. Leaves in basal
rosette, thick, broadly obovate. Involucres broadly
campanulate; bracts 3–4-seriate, weakly gradate,
outer rows completely herbaceous, flat; receptacles
epaleate. Rays 1–2-seriate, pistillate, blue or
white, often with staminodia, limbs broad, weakly
coiling. Disc florets perfect, yellow, funnelform,
lobes ovate-deltate, erect to reflexed; style appendages lanceolate. Achenes narrowly oblong
to oblanceolate, compressed, (2–)4–6-nerved,
glandular, densely strigose-sericeous, paired
299
setula tips of uneven lengths, strongly divergent;
pappus 1–2-seriate, persistent, bristles plumose,
sometimes with long scales outside. Three species,
southern Africa.
805. Nardophyllum (Hook. & Arn.) Hook. & Arn.
Nardophyllum (Hook. & Arn.) Hook. & Arn., Companion
Bot. Mag. 2: 44 (1836); Cabrera, Notas Mus. La Plata 17:
55–66 (1954), rev.
Gochnatia subg.(?) Nardophyllum Hook. & Arn. (1835).
Small spreading shrubs; stems, leaf undersides,
and involucres often tomentose; branchlets sometimes spiniform, internodes often angled or with
dark lines. Leaves sessile, subcoriaceous, oblong to
linear, entire. Heads 1–5, terminal or axillary; involucres campanulate; bracts 3–5-seriate, gradate,
caducous, oblong to lanceolate or linear, margins
scarious; receptacles with narrow caducous pales.
Rays lacking. Florets few, perfect, yellow, narrowly
funnelform, lobes triangular to lanceolate, often
recurved; style appendages lanceolate, papillose
outside, branches smooth below. Achenes obovoid,
compressed, 4–5-ribbed, sericeous-villous; pappus bristles as long as corollas. Seven species,
Argentina, Bolivia, Chile.
806. Novenia S.E. Freire
Novenia S.E. Freire, Bol. Soc. Argent. Bot. 24: 296 (1986);
Freire & Hellwig, Taxon 19: 124–125 (1990), comb.
Acaulescent, caespitose, perennial herbs. Leaves
in rosettes, bases involute, densely villous; blades
linear-lanceolate, coriaceous, with 3 grooves adaxially, glabrous. Inflorescences sessile, glomerulate,
1–4-headed; involucres cylindric; bracts c. 3-seriate, subequal, oblong-lanceolate, stramineous,
indurate, glabrous, margins scarious; receptacles
with reduced pales. Pistillate florets 3–9, whitish,
filiform, eradiate. Disc florets functionally male,
whitish, narrowly tubular, lobes oblong-triangular,
erect; style branches linear-lanceolate. Achenes
obovoid to oblong, compressed, with (5–)6 whitish
nerves, with sparse filiform setulae; pappus
bristles 2–3-seriate, persistent. x = 9. One species,
N. acaulis (Wedd. ex Benth. in Benth. & Hook.
f.) S.E. Freire & F. Hellw., Andes from northern
Argentina to southern Peru.
807. Olearia Moench
Olearia Moench, Supp. Meth. 254 (1802), nom. cons.; Cross
et al., Pl. Syst. Evol. 235: 99–120 (2002); Comp. Newslett. 40:
300
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
11 (2004), phylog.
Steetzia Sonder (1853).
Shrubs; glabrous or glutinous to variously
pubescent, hairs sometimes stellate. Leaves
cauline, alternate or opposite, broadly ovate to
linear, often tomentose abaxially. Heads terminal
or axillary, solitary or in corymbose to thyrsoid
inflorescences, sessile to pedunculate; involucres
cylindric to hemispheric; bracts persistent, 2–6seriate, herbaceous, linear to lanceolate, margins
scarious, sometimes resin duct along midvein;
receptacles epaleate. Rays 1-seriate, rarely lacking,
white to blue or purple, coiling. Disc florets few to
numerous, perfect, yellow or purplish, narrowly
funnelform, lobes 5, deltate or lanceolate, recurving; anthers shortly calcarate. Achenes terete or
slightly compressed, usually 5-ribbed, glabrous
or setulose; pappus bristles persistent, often in 2
unequal series. x = 9. Circa 180 species, Australia,
New Zealand, New Guinea.
A molecular-phylogenetic study (Cross et al.
2002) indicates that Olearia is polyphyletic as currently defined, with species divided into three or
four primary, phyletically disparate clades, with
several other smaller ones variously dispersed. Formal taxonomic changes await more detailed sampling and analysis.
808. Oritrophium (Kunth) Cuatrec.
Oritrophium (Kunth) Cuatrec., Cié. (Mexico) 21: 21 (1961);
Cuatrecasas, BioLlania ed. espec. 6: 287–303 (1997), rev.;
Nesom, Sida 18: 523–526 (1998), emend.
Aster L. sect. Oritrophium Kunth (1818).
Perennial herbs from thick erect to horizontal
rhizome. Leaves mostly in basal rosettes, bases
usually densely hairy, blades oblanceolate to linear,
glabrous to sericeous. Inflorescences scapose,
1-headed, bracteoles linear-subulate; involucres
subhemispheric to broadly cylindric or turbinate;
bracts 2–3-seriate, gradate to subequal, linear
to oblong-lanceolate, obscurely veined; receptacles epaleate. Rays 1–2-seriate, pistillate, white,
coiling. Disc florets functionally male, narrowly
funnelform, lobes triangular, erect; style branches
linear, long-papillose. Ray achenes fusiform,
5-veined, several-angled or compressed, glabrous
to sericeous, eglandular; disc achenes linear,
4–5-veined; pappus 1-seriate, bristles subequal.
x = 9. Circa 21 species, mostly Andean, Venezuela
to Bolivia, two in Mexico.
809. Pachystegia Cheeseman
Pachystegia Cheeseman, Man. N. Z. Fl., ed. 2, 910 (1925);
Cross et al., Pl. Syst. Evol. 235: 99–120 (2002), phylog.
Robust spreading shrubs; leaf undersides, peduncles and involucres densely tomentose. Leaves
stoutly petiolate with broadly clasping bases,
blades coriaceous, entire, glabrous adaxially.
Heads solitary on long peduncles; involucres
subglobose to urn-shaped, becoming hemispheric;
bracts to 9-seriate, ovate to lanceolate, outer obtuse, inner acute or with needle-like recurved tips;
receptacles epaleate. Rays 2–3-seriate, pistillate,
white, not or weakly coiling. Disc florets numerous, perfect, yellow, tubes slender, limbs narrowly
funnelform-tubular, lobes 5, lanceolate, spreading,
with distinct yellowish ducts; style appendages
linear, obtuse. Achenes oblong-linear, grooved,
densely sericeous; pappus tawny, 1-seriate, bristles
slightly thickened distally. x = 9. One species,
P. insigne Cheeseman, New Zealand.
810. Pacifigeron G.L. Nesom
Pacifigeron G.L. Nesom, Phytologia 76: 160 (1994).
Shrubs, younger parts with arachnoid tomentum,
persisting in axils. Leaves clustered at branch tips,
coriaceous, obovate-spathulate, entire. Inflorescences of clustered subsessile heads imbedded
in tomentum; involucres campanulate; bracts c.
18–24, 2–3(–4)-seriate, gradate, broadly lanceolate,
yellowish with 3(–5) orange veins; receptacles
epaleate. Rays c. 18–26, 1(–2)-seriate, pistillate,
white or cream, orange-veined, limbs spreading.
Disc florets functionally male, orange-veined, funnelform, limb enclosed in long hairs, lobes broadly
lanceolate. Fertile achenes fusiform, slightly compressed, with 9–11 raised orange-resinous nerves
when young, setulose, eglandular; pappus bristles
30–45, outer series short. One species, P. rapensis
(F. Br.) G.L. Nesom, Polynesia, Rapa Island.
811. Paleaepappus Cabrera
Paleaepappus Cabrera, Bol. Soc. Argent. Bot. 11: 273 (1969);
Bonifacino & Sancho, Sida 19: 531–538 (2001), emend.
Small, divaricately branched shrubs, young
branchlets thorn-like; young branches, young
leaves and involucres tomentose. Leaves small,
subcoriaceous, sessile, oblong to spathulate,
entire, caducous. Heads solitary on bracteose
branches, discoid; involucres campanulate; bracts
Compositae
4–5-seriate, gradate, broadly lanceolate, shortacute, entire, indurate, caducous; receptacles with
conduplicate pales. Rays absent. Florets numerous,
perfect, yellow, tubes narrow, limbs narrowly
campanulate, lobes triangular; style appendages
lanceolate, papillose, branches smooth below.
Achenes oblong-obovoid, sericeous-villous; pappus 2-seriate, of linear-lanceolate scales, as long
as corollas. One species, P. patagonicus Cabrera,
Patagonia.
812. Parastrephia Nutt.
Parastrephia Nutt., Trans. Amer. Philos. Soc. ser. 2, 7: 449
(1841); Nesom, Phytologia 75: 347–357 (1993), rev.
Ericoid to cupressiform shrubs, resinous to partly
tomentose. Leaves alternate, sessile, coriaceous,
scale-like to linear, midvein depressed and tomentose abaxially, remainder of leaf glabrous. Heads
solitary or in small clusters, disciform; involucres
narrowly campanulate; bracts 3–4-seriate, gradate,
oblong to oblong-lanceolate, obtuse to acute,
persistent, mostly indurate, margins thin and
scarious; receptacles epaleate. Pistillate florets
1-seriate, yellow, tubular or with small limb. Disc
florets perfect, yellow, narrowly funnelform, lobes
lanceolate, recurved; style appendages lanceolate,
papillose to base of branches. Achenes obovoid,
compressed, c. 4-ribbed, 2 marginal, sericeous;
pappus bristles 2-seriate, some outer scales or
bristles short. Three species, Argentina, Chile,
Bolivia and Peru.
813. Pleurophyllum Hook. f.
Pleurophyllum Hook. f., Fl. Antarctica 1: 30 (1844); Cross
et al., Pl. Syst. Evol. 235: 99–120 (2002), phylog.
Robust, perennial, rosulate herbs; silvery-sericeous
to white-tomentose. Leaves sessile, basal leaves
large, broadly elliptic to obovate, subacute, entire
or denticulate, coriaceous to subcarnose, with
30–40 parallel veins; cauline leaves narrowly elliptic, with 5–9 veins. Scapes racemose; involucres
depressed-hemispheric; bracts 2–3-seriate, subequal, linear-lanceolate, pilose to lanate; receptacles
epaleate. Rays 15–30(?), 1–3-seriate, pistillate,
whitish to purplish, limbs short or elongate,
3-lobed or deeply tripartite. Disc florets 60 or
more, perfect, purple, tube cylindric, limb campanulate, lobes 4 or 5, long-ovate, recurving; style
appendages deltate. Achenes elongate, compressed
in ray florets, subquadrate in disc florets, ribbed,
densely setulose; pappus bristles 2–3-seriate,
301
sordid, scabrous to subplumose, unequal. Three
species, New Zealand, sub-Antarctic islands south
of New Zealand.
814. Printzia Cass.
Printzia Cass., Dict. Sci. Nat. 37: 488 (1825); Anderberg,
Opera Bot. 104: 1–195 (1991), emend.
Perennial herbs or small shrubs; stems and leaf
undersides pale-tomentose; leaves subsessile, narrowly elliptic to ovate. Inflorescences thyrsoid or
with heads solitary; peduncles short; involucral
bracts 2–5-seriate, gradate to subequal, lanceolate
to ovate, dark in middle, margins pale; receptacle
epaleate. Rays pistillate, 1-seriate, violet to white,
yellowish or greenish, often coiled, rarely filiform,
staminodes sometimes present. Disc florets numerous, perfect, yellow, narrowly funnelform; lobes 5,
triangular, recurving; anther thecae with long basal
tails; style appendages lanceolate, weakly papillate.
Achenes mostly terete, c. 10-nerved, setuliferous;
pappus bristles 1–2-seriate, tips often subplumose.
Six species, southern Africa.
The genus is re-incorporated in Astereae on the
basis of a DNA study by Bayer and Cross (2002), in
spite of the long-tailed anther bases.
815. Pteronia L.
Pteronia L., Sp. Pl., ed. 2, 2: 1176 (1763), nom. cons.; Gen.
Pl., ed. 6, 414 (1764); Hutchinson & Phillips, Ann. S. African
Mus. 9: 277–329 (1917), rev.
Shrubs or shrublets; grey-tomentellous, scabridulous, gland-dotted or glabrous. Leaves alternate or
opposite, sometimes rather fleshy, sessile to petiolate, linear or ovate-oblong to oblanceolate, entire
to serrulate-ciliate. Heads usually solitary, peduncles short; inflorescences sometimes corymbiform
or (P. fasciculata L. f.) numerous single-flowered
heads aggregated into a second-order head;
involucres ovoid to cylindric; bracts 4–7-seriate,
gradate, broadly ovate to linear-oblong, margins
membranaceous to subscarious; receptacles often
fimbriate, epaleate. Rays absent. Florets 1 to many,
perfect, yellow to orange, narrowly funnelform
or limbs campanulate-cylindric, lobes oblonglinear to deltate, erect. Achenes turbinate, often
contracted at apex, villous to glabrous; pappus
bristles c. 2-seriate, persistent, white, stramineous,
or reddish to purple, often connate into basal
ring. Circa 80 species, southern and south-western
Africa to Zambia.
302
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
816. Remya W. Hillebrand ex Benth. in Benth. &
Hook. f.
Remya W. Hillebrand ex Benth. in Benth. & Hook. f., Gen.
Pl. 2, 1: addend. 536 (1873); Wagner & Herbst, Syst. Bot. 12:
601–608 (1987), rev.
Weakly erect or sprawling shrubs, densely tomentose to glabrous. Leaves narrowly elliptic to broadly
ovate or lanceolate, glabrous adaxially, margins
toothed. Inflorescences loosely corymbose; involucres ovoid to globose or hemispheric; bracts 4–5seriate, gradate, coriaceous to chartaceous, ovate
to linear-oblong, margins narrowly scarious; receptacles epaleate. Rays 12–20, pistillate, creamcoloured, limbs short. Disc florets dimorphic in two
species; outer pistillate, tubular; inner functionally
male, yellow, goblet-shaped. Fertile achenes narrowly obovoid, compressed, 3–4-angled, with 1 or
2 nerves on each face, glabrous to strigose; pappus
of 3–9(–14) unequal to subequal, persistent nonbarbellate awns, sometimes reduced in inner disc.
x = 9. Three species, Hawaii.
817. Rochonia DC.
Rochonia DC., Prodr. 5: 345 (1836); Humbert, Fl. Madag.
189, III, 1: 311–317 (1960), reg. rev.
Subshrubs or shrubs, stipitate-glandular, stems,
leaf undersides and involucres tomentose. Leaves
sessile, mostly subcoriaceous, elliptic or lanceolate
to obovate, entire or dentate, 1- or 3-nerved
from base. Inflorescences terminal, 1-headed to
corymbose; peduncles short to long; involucres
hemispheric; bracts 3–4-seriate, gradate, ovate to
oblong-lanceolate, margins scarious; receptacles
epaleate. Rays 1–2-seriate, 14 to c. 35, pistillate,
yellow, coiling. Disc florets numerous, perfect,
yellow, narrowly funnelform, tube short, lobes triangular, erect; style appendages deltate. Achenes
oblong, slightly fusiform, compressed, 5–10costate, setuliferous; pappus bristles 2-seriate,
persistent, subequal or outer shorter. Four species,
Madagascar.
subequal, glabrous to woolly; receptacles epaleate.
Pistillate florets 2–4-seriate, white to reddish or
purple, filiform-tubular or slightly broadened distally, contracted and minutely 5-toothed apically.
Disc florets functionally male, tubular, purple to
reddish, lobes 5, triangular, erect. Fertile achenes
obovate, compressed, 2-nerved or with weak nerve
on each face, strigose to sericeous, usually with
glands apically; pappus bristles 1-seriate, as long
as corollas. Six species, Costa Rica, Panama.
XV.3. Subtribe Brachyscominae G.L. Nesom
(1994).
Annual to perennial herbs. Leaves alternate, entire to toothed or lobed, eglandular or stipitateglandular. Heads solitary to few; involucral bracts
not keeled, commonly with broad hyaline margins; receptacles plane to conical, epaleate. Rays
1-seriate, pistillate, white to blue, coiling. Disc florets bisexual, functionally male in Ceratogyne and
some species of Calotis, corollas short-tubed; style
appendages deltate, papillose. Achenes obovate, 2nerved, flat, commonly winged in Brachyscome, eglandular, faces and margins usually with setulae
with anchor-shaped tips; pappus absent or highly
reduced, less commonly of bristles, 1-seriate. x = 9,
reduced to 8, 7, 6, 5, 4, 3, 2.
Key to the Genera
1. Achenes distally Y- or T-shaped with widely spreading
arms, with wings involute on one side; pappus lacking
821. Ceratogyne
– Achenes without distal arms or projections; pappus
usually present
2
2. Disc florets mostly perfect; pappus of short or minute
scales or absent
819. Brachyscome
– Disc florets usually functionally male; pappus of subulate to spreading awns, often retrorsely barbed
820. Calotis
Genera of Brachyscominae
819. Brachyscome Cass.
818. Westoniella Cuatrec.
Westoniella Cuatrec., Phytologia 35: 472 (1977).
Erect shrubs or prostrate to pulvinate herbs,
glabrous to white-woolly, often gland-dotted.
Leaves imbricate, sessile, rigid, linear to elliptic,
margins revolute, entire to serrate or digitate,
tomentose beneath. Heads solitary or in glomerulae. Involucres campanulate; bracts 4–5-seriate,
Brachyscome Cass., Bull. Sci. Soc. Philom. Paris 1816:
199 (1816); Davis, Proc. Linn. Soc. New South Wales 73:
142–241 (1948), rev.
Brachycome Cass. corr. Cass. (1825), in wide use in horticulture, and arguably the correct orthography.
Annual or perennial herbs, sometimes rhizomatous; glabrous or tomentose to stipitate-glandular;
stems erect or spreading. Leaves in rosettes or
Compositae
cauline, entire to pinnatisect. Peduncles usually
long; involucres campanulate to hemispheric;
bracts 2–3-seriate, subequal, oblong, obtuse,
herbaceous, margins narrowly hyaline; receptacles
convex-conical. Rays 1–2-seriate, numerous, white,
blue, pink or yellow. Disc florets numerous, mostly
perfect, yellow, limbs narrowly campanulate, lobes
triangular-lanceolate, spreading or recurved;
anther appendage rarely lacking; style appendages
deltate to long-triangular. Achenes terete to
compressed, glabrous to setulose or tuberculate,
sometimes winged; pappus absent or of small
white scales. x = 9, reduced to 8, 7, 6, 5, 4, 3 or
2. Circa 70 species, Australia, New Guinea, New
Zealand, New Caledonia.
820. Calotis R. Br.
Calotis R. Br., Bot. Reg. 6: pl. 504 (1820); Davis, Proc. Linn.
Soc. New South Wales 77: 146–188 (1952), rev.
Annual or perennial herbs, prostrate to erect,
sometimes stoloniferous, glabrate to hispid or
hirsute. Leaves cauline and/or basal, petiolate
or sessile, usually toothed or pinnately lobed.
Inflorescences 1-headed or leafy-cymose; involucres campanulate to subglobose; bracts
2–3(–4)-seriate, subequal, herbaceous; receptacles
flat to conical, sometimes with small scales. Rays
1- to several-seriate, limbs blue or purple to white
or yellow, sometimes vestigial. Disc florets usually
functionally male, yellow, limb campanulate, lobes
deltate-lanceolate, erect to spreading. Achenes
obconical, glabrous or strigose to tuberculate,
sometimes winged; pappus of (1–)2–25 subequal
erect to spreading awns, often retrorsely barbed
or ciliate, sometimes with alternating low scales,
rarely of 12–15 short-plumose bristles or absent.
x = 8, reduced to 7, 5, 4. Circa 28–30 species,
mostly Australia.
821. Ceratogyne Turcz.
Ceratogyne Turcz., Bull. Soc. Imp. Naturalistes Moscou 24:
68 (1851).
Annuals or short-lived perennials; taproot weak;
stems strigose, non-glandular; erect or ascending
from rosette. Leaves sessile, obovate or those
subtending heads smaller. Inflorescences seriatecymose; heads sessile in terminal or overtopped
clusters; involucral bracts bracteole-like, 2–5,
1-seriate; receptacles plano-convex. Rays 3–6,
whitish; limbs short. Disc florets c. 3–6, functionally male, narrowly funnelform, lobes 3 or
303
4, deltate-ovate. Fertile achenes lobed, Y- or
T-shaped, compressed, winged, involute on one
face, anchor-shaped setulae on faces, lobe margins
and base; epappose. One species, C. obionoides
Turcz., south-eastern Australia.
XV.4. Subtribe Bellidinae Willk. (1870).
Tribe Bellideae Cass. ex D. Don (1830).
Tribe Bellieae DC. ex Godr. (1850)
Herbs. Leaves alternate in basal rosette, wide,
entire to serrate-dentate, eglandular. Heads solitary on scapes; involucral bracts flat, completely
herbaceous, 2(–3)-seriate; receptacles conical,
epaleate. Ray florets 1-seriate, limbs long, white or
pink-tinged. Disc florets bisexual, with short tube;
style branches with deltate papillose appendages.
Achenes obovate, compressed, with pair of thick
marginal ribs, eglandular, setulae without anchorshaped tips; pappus absent or a short laciniate
crown, apical callus narrow. x = 9.
Key to the Genera
1. Pappus absent or a ring of very short bristles; involucral bracts 2-seriate
822. Bellis
– Pappus with outer ring of 4–6(–10) scales alternating
with inner ring of bristles; involucral bracts 1-seriate
823. Bellium
822. Bellis L.
Bellis L., Sp. Pl. 886 (1753); Webb, Atlas Fl. Europaea 4: 111–
112 (1976), reg. rev.; Fiz et al., Mol. Phylog. Evol. 25: 157–171
(2002), phylog.
Annual or short-lived perennial herbs, sometimes
rhizomatous, roots fibrous; stems usually scapiform. Leaves in rosette, sometimes some on lower
stem, spathulate to oblanceolate, entire to toothed.
Involucres shallowly cupulate; bracts (1–)2-seriate,
subequal, ovate or oval, evenly herbaceous; receptacles conical or hemispheric to flat. Rays pistillate, limbs white above, often pink or purple below, closing upwards at night. Disc florets perfect, yellow, tubular, tube very short, nearly stipelike, lobes erect. Achenes obovate, short-strigose
or ciliate-margined to glabrous, sometimes with
sessile glands; pappus absent or a short crown of
bristles. x = 9. Circa 8 species, Europe, widely introduced.
823. Bellium L.
Bellium L., Mant. Pl. 2: 157 (1771).
304
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Annual or perennial herbs; acaulescent or rarely
caulescent; pilosulous. Leaves elliptic-obovate to
spathulate, bases long-petioliform, tips rounded to
obtuse, margins entire. Involucres broadly campanulate to hemispheric; bracts crowded 1-seriate,
equal, herbaceous in middle, margins scarious, tips
obtuse, rather fimbriate; receptacles hemispheric to
conical. Rays pistillate, limbs white, often reddish
below, not coiling. Disc florets perfect, yellow, narrowly campanulate from short tube, lobes 4 or 5,
broadly triangular, erect. Achenes oblong-obovoid,
setulose on sides; pappus of 4–6(–10) barbellate
bristles as long as achene, alternating with short
scales. x = 9. Four species, southern Europe.
XV.5. Subtribe Grangeinae Benth. in Benth.
& Hook. f. (1873); Fayed, Mitt. Bot.
Staatssamml. München 15: 425–476 (1979),
rev.
Annual to perennial herbs, usually glandular.
Leaves alternate, pinnatifid to pinnatisect. Heads
solitary or few in a loosely corymbose inflorescence, involucral bracts not keeled; receptacles
often conical, with or without pales. Ray florets
usually 2–4-seriate, corolla limb absent or barely
exceeding the involucre, white greenish-yellow or
purplish. Disc florets perfect or often functionally
male, corollas commonly 4-lobed; style branch appendages deltate, papillose. Achenes compressed,
2-nerved, eglandular or glandular, setulae sometimes with anchor-shaped tips; pappus absent,
of a few short bristles, or a low crown of basally
connate scales. x = 9, apparently reduced to 8 in
Erodiophyllum.
– Pappus usually of 12–50 long bristles; receptacles
plano-convex
838. Nidorella
6. Achenes broadly winged; pappus segments fused at
base
833. Grangeopsis
– Achenes not winged; pappus segments not fused at
base
7
7. Annuals; pappus of c. 10 flattened, persistent bristles
in 1 series
828. Dacryotrichia
– Erect perennial herbs; pappus 2–3-seriate, numerous,
unequal, slender caducous bristles 836. Heteromma
8. Perennial herbs; corollas of pistillate florets bilabiate,
with lobules on inner side
839. Plagiocheilus
– Annual or short-lived perennial herbs; corollas of pistillate florets not bilabiate
9
9. Disc florets functionally male
10
– Disc florets perfect
11
10. Anthers without apical appendage; receptacles flat
826. Colobanthera
– Anthers with distinct apical appendage; receptacles
cup-shaped, with pistillate florets borne on both inner
and outer surfaces of cup
827. Cyathocline
11. Pistillate corollas radiate
12
– Pistillate corollas not radiate, sometimes 4-dentate at
apex
14
12. Achene with a cartilaginous apical crown or cup; pappus lacking; ray florets in 1–4 series
830. Egletes
– Achene without cartilaginous apical crown or cup;
pappus of minute teeth or triangular scales
13
13. Leaves pinnatifid with often coarsely toothed oblong
or lanceolate lobes; plants hispid to hirsute; receptacle
hemispheric to conical
835. Gyrodoma
– Leaves simple, entire; plants densely glanduliferous;
receptacle convex to toroid
873. Mtonia
14. Receptacle flat or scarcely convex
834. Grauanthus
– Receptacle hemispheric to conical or columnar
15
15. Pappus a papillose rim, or with 1 or 2 bristles, or lacking; disc corollas campanulate, lobes not spreading
829. Dichrocephala
– Pappus a partially fused corona with a lacerate rim or
paleaceous teeth; disc corollas with widely spreading
lobes
832. Grangea
Genera of Grangeinae
Key to the Genera
1. Receptacles with pales
2
– Receptacles epaleate
3
2. Florets of 2 types, outer florets tubular, without limbs;
receptacles plane
825. Ceruana
– Florets of 3 types, outer pistillate florets radiate; receptacles highly convex to conical
831. Erodiophyllum
3. Pappus of 3–50 elongate, separate, slender or flattened
bristles
4
– Pappus rudimentary or lacking, rarely with 1 or 2
bristles
8
4. Heads heterogamous, with peripheral pistillate florets
5
– Heads homogamous, without differentiated pistillate
florets
6
5. Pappus of 3–8 short bristles; receptacles conic
824. Akeassia
824. Akeassia J.-P. Lebrun & Stork
Akeassia J.-P. Lebrun & Stork, Candollea 48: 332 (1993).
Much-branched annual herbs, longer branches
prostrate-decumbent; stems and leaves weakly
puberulous. Leaves sessile, obovate, narrowed to
base, venation weakly pinnate. Heads clustered
among distal leaves, 3–6, subsessile, subglobose; involucral bracts subbiseriate, subequal,
oblanceolate; receptacles conical. Pistillate florets
1–3-seriate, yellow, short-cylindrical, 4-toothed.
Disc florets bisexual, yellow, tube short, limb
campanulate, glandular, lobes 4, as wide as long.
Ray and disc achenes alike, subcylindric or
ellipsoid, glandular, setulae with anchor-shaped
tips; pappus of (3–)5–8 short bristles. One species,
Compositae
A. grangeoides J.-P. Lebrun & Stork, western Africa
and Congo.
825. Ceruana Forssk.
Ceruana Forssk., Fl. Aegypt.-Arab. 74: 153 (1775).
Erect, coarse, annual or short-lived perennial
herbs; sparsely hirsute, gland-dotted. Leaf bases
slender with short decurrent margins, blades obovate, entire to serrate-pinnatifid. Inflorescences
cymose with racemose branches; heads on short
peduncles, overtopped by branches; involucre
hemispheric; bracts c. 2-seriate, subequal, narrowly oblong, acute, herbaceous, margins very
narrowly scarious; receptacles plane, with pale,
lanceolate, fimbriate, caducous pales. Florets
yellow. Pistillate florets 1-seriate, filiform, with
minute stipitate glands, with 4 narrow apical lobes.
Disc florets perfect, tube short, limb abruptly
campanulate, lobes 5, broadly triangular, scarcely
spreading, with slender yellowish ducts. Achenes
oblanceolate to oblong-obovoid, setulose; pappus
of short, basally connate, bristle-like scales or
setae. One species, C. pratensis Forssk., northern
and western Africa.
826. Colobanthera Humbert
Colobanthera Humbert, Mém. Soc. Linn. Normandie 25:
35, 281 (1923).
Small, erect to spreading, annual or short-lived
perennial herbs, branching mostly from base;
leaves, peduncles and involucres with subsessile glands. Basal leaves subpetiolate, narrowly
oblanceolate, cauline leaves sessile, subamplexicaul, sublinear, usually entire. Heads terminal
and axillary, solitary on long peduncles; involucres broadly campanulate; bracts subbiseriate,
subequal, obtuse, margins scarious, blackish;
receptacles flat. Rays 2–3-seriate, c. 25, pistillate,
white or yellow, narrow, tubes almost lacking. Disc
florets c. 10, functionally male, campanulate from
slender tube, lobes deltoid, erect-spreading; thecae
without apical appendage; style branches oblonglanceolate. Achenes oblong, slightly fusiform, with
many small setulae; pappus none. One species,
C. waterlotii Humbert, Madagascar.
827. Cyathocline Cass.
Cyathocline Cass., Ann. Sci. Nat. (Paris) 17: 419 (1829).
Erect, annual or perennial herbs; hirsute to glandular. Leaves sessile, usually with basal auricles,
305
pinnatifid with pointed teeth. Inflorescences
loosely cymose with corymbose branches; peduncles short; involucres broadly campanulate to
hemispheric; bracts c. 3-seriate; subequal, linearlanceolate, narrowly acute, 1-nerved, herbaceous
medially, margins scarious, ciliate; receptacles
cup-shaped, bearing multiseriate pistillate florets
on outer and inner surfaces of cup. Florets yellow
to purple. Pistillate florets filiform with small oval
limbs. Disc florets few, only in centre of receptacle
cup, functionally male, limb campanulate from
slender tube, lobes broadly triangular, spreading.
Fertile achenes obovoid to fusiform, glabrous;
pappus lacking. x = 9. Three species, tropical Asia.
828. Dacryotrichia Wild
Dacryotrichia Wild, Garcia de Orta, ser. Bot. 1: 67 (1973).
Annual herbs; pilose, sparsely glandular; stems
erect or spreading. Leaves sessile, often narrowed,
subauriculate, pinnatisect, with 2–6 linear to
spathulate lobes. Inflorescences loosely cymose,
c. 10-headed; peduncles 1–2 cm long; involucres hemispheric; bracts 2–3-seriate, oblong
to obovate, acute to obtuse, margins hyaline;
receptacles plano-convex. Ray florets absent. All
florets perfect, orange-yellow, broadly funnelform,
lobes broadly triangular, erect; style appendages
lanceolate, subobtuse. Achenes narrow with base
attenuate, sparsely glandular, biseriate setulae
bearing single, caducous, narrowly vesicular cell
on lower half of apex; pappus persistent, white,
bristles c. 10, flattened, with some shorter outer
bristles. One species, D. robinsonii Wild, Zambia.
829. Dichrocephala L’Hér. ex DC.
Dichrocephala L’Hér. ex DC. in Guill., Arch. Bot. (Paris) 2:
517 (1833).
Annual herbs; straggling to erect; pilose or
glabrescent. Leaf bases clasping-auriculate,
blades lyrate-pinnatifid, coarsely serrate or
serrate-crenate. Inflorescences paniculate; heads
subglobose; involucral bracts 1–2-seriate, subequal, oblong to elliptic, apices rounded to obtuse,
margins membranaceous; receptacle convex to
columnar-ellipsoid, tuberculate to pitted. Rays
multiseriate, pistillate, greenish, yellow-white, or
purplish, tubular, 2–4-dentate or regularly lobed.
Disc florets yellow or reddish, rotate-campanulate
from short tubes, lobes 4–5, deltate to broadly
triangular, erect to spreading. Achenes obovoid
to turbinate, mostly glabrous with glands basally
306
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
and distally; pappus absent, a narrow rim or of
1 to few caducous, short bristles. Ten species,
Africa, Madagascar, Indonesia, tropical and
south-western Asia.
830. Egletes Cass.
Egletes Cass., Bull. Soc. Philom. Paris 1817: 153 (1817);
Shinners, Lloydia 12: 239–247 (1949), rev.; Lloydia 12:
248–250 (1949), emend.
Erect to procumbent, taprooted, annual herbs;
stems much-branched, viscid with usually stipitate
glands. Leaf bases petiolate to sessile, subclasping,
blades obovate to spathulate, lobed or toothed to
pinnatifid or bipinnatifid. Heads usually solitary
and axillary or few in loosely corymbose inflorescences, short-pedunculate; involucres hemispheric
to cupulate; bracts 2–3-seriate, subequal, thinherbaceous, lanceolate; receptacles conical. Rays
1–4-seriate, pistillate, white, filiform, limbs short.
Disc florets perfect, yellow, tubular-funnelform,
3–5-lobed. Achenes oblong to narrowly obovate,
glabrous or gland-dotted, or with hooked setulae,
apical callus a whitish, sometimes flaring crown or
cup; pappus bristles absent. x = 9. Circa 8 species,
neotropical to south-western USA (southern
Texas).
831. Erodiophyllum F. Muell.
Erodiophyllum F. Muell., Fragm. (Mueller) 9: 119 (1875).
Coarse, erect or ascending perennial herbs; stems,
leaves, and involucres hirsute with white hairs.
Leaves herbaceous, lyrate-pinnatifid. Inflorescences terminal or axillary, 1-headed, pedunculate;
involucres campanulate to hemispheric; bracts
c. 2-seriate, subequal, oblong to oblong-linear,
herbaceous; receptacles convex or conical, outer
4–7 rows among pistillate florets with ovate pales
which become indurate with recurved points.
Rays 1-seriate, blue to purple or white. Tubular
pistillate florets 4–7-seriate. Central florets functionally male, yellow, tubes short, limbs narrowly
funnelform, lobes deltate, spreading or recurved.
Fertile achenes obovoid-obconic, glabrous, apex
indurate; pappus of minute scales fused to crenate
rim. x = 8. Two species, Australia.
832. Grangea Adans.
Grangea Adans., Fam. Pl. 2: 121, 563 (1763).
Annual herbs, suberect to prostrate. Leaves
pinnatifid. Inflorescences opposite to leaves or
terminal and corymbose, 3–6-headed; involucres
hemispheric to subglobose; bracts 1–2-seriate,
subequal, inner with membranaceous margins; receptacles hemispheric or conical, naked. Pistillate
florets 2- to many-seriate, yellow, campanulate or
narrowly funnelform, cylindric below, 2–4-dentate.
Disc florets perfect, yellow, campanulate or rotatecampanulate, cylindric below, 4–5-lobed. Achenes
subterete, ellipsoid, compressed or 4-angled, apex
sometimes cartilaginous, sometimes constricted;
pappus obsolete, a lacerate coroniform rim, or
toothed. Ten species, Africa, Madagascar, tropical
Asia.
833. Grangeopsis Humbert
Grangeopsis Humbert, Mém. Soc. Linn. Normandie 25: 34,
281 (1923).
Small, branching perennial herbs, stoloniferous;
stems, leaves and involucres with thin tomentum.
Leaves larger in basal rosette, basal leaves obovate
to oblanceolate, bases petioliform, cauline leaves
oblong-spathulate. Heads solitary on long peduncles; involucres campanulate to hemispheric;
bracts c. 3-seriate, subequal, herbaceous, oblong to
spathulate, obtuse, margins scarious; receptacles
convex. Rays lacking. Florets yellow, tube short,
limb broadly campanulate, lobes (4–)5, subdeltoid,
spreading. Achenes broadly obovate, broadly
winged, ciliate, with rounded to narrow apex
and lacking nerves on faces; pappus of 10–12
hyaline, lanceolate, unequal scales, 1/2–9/10 as
long as corollas, bases fused into ring. One species,
G. perrieri Humbert, Madagascar.
834. Grauanthus Fayed
Grauanthus Fayed, Mitt. Bot. Staatssamml. München 15:
484 (1979).
Erect or ascending annual or perennial herbs,
leaves and involucres hirsute. Leaves sessile,
dentate to lyrato-pinnatifid, generally lanceolate
to linear. Inflorescences subcorymbose; peduncles
short; involucres cupuliform; bracts 2–3-seriate,
subequal, outer bracts narrow, obtuse, 1-nerved,
inner bracts oblong-obovate, margins hyaline,
ciliate; receptacles plano-convex, tuberculate,
centre subacuminate. Pistillate florets c. 3-seriate,
whitish-yellow, tubular with broadened limb,
3- or 4-dentate. Disc florets yellow, tube short
with broadly funnelform limb, lobes 5, oblongtriangular, erect. Achenes yellow, subcylindric,
partial nerve on face, margins setulose, disc
Compositae
307
achenes somewhat smaller; pappus lacking. Two
species, tropical Africa.
dentate ring. One species, M. glandulifera Beentje,
eastern Africa.
835. Gyrodoma Wild
838. Nidorella Cass.
Gyrodoma Wild, Kirkia 9: 294 (1974).
Nidorella Cass., Dict. Sci. Nat. 37: 459, 469 (1826); Wild,
Bol. Soc. Brot. 43: 209–243 (1969), rev.
Decumbent or ascending annual herbs; branching
from base; stems, leaves and outer involucral bracts
hispid to hirsute. Basal leaves short-petiolate, upper leaves attenuate to clasping base, generally
oblong or oblanceolate, coarsely toothed to
pinnatifid with oblong or lanceolate lobes. Heads
solitary on long peduncles; involucres broadly
campanulate to subpatelliform; bracts c. 2-seriate,
subequal, margins membranaceous; receptacles
hemispheric-conical. Rays 1-seriate, pistillate,
white to pale blue, linear, recurved. Disc florets
yellow, tube short, limb widely funnelform, lobes
cut to basal 1/3 of limb, triangular, spreading; style
appendages obtuse. Achenes narrowly turbinate,
minutely setulose; pappus of minute scales c.
0.1 mm long. One species, G. hispida (Vatke) Wild,
Mozambique.
Erect annual or perennial herbs; nearly glabrous
to hirsute or lanate. Leaves mostly cauline,
sessile to pseudopetiolate or subamplexicaul,
obovate to linear-spathulate, entire to pinnatisect.
Inflorescences corymbose with subglobose or
cymose clusters of heads; involucre hemispheric
or campanulate; bracts 1–3-seriate, subequal,
midrib often glandular to pilose, margins hyaline.
Rays 2–3-seriate, pistillate, yellow, limb short or
3–4(–5)-lobed. Disc florets few to many, usually
perfect, yellow, funnelform, lobes triangular,
erect-spreading; style appendages broad, base
cordate and with larger papillae. Achenes obovoid
or ellipsoid, pubescent, often glandular; pappus
bristles persistent, 5 to c. 30. x = 9. Circa 15 species,
tropical and southern Africa.
836. Heteromma Benth. in Benth. & Hook. f.
839. Plagiocheilus Arn. ex DC.
Heteromma Benth. in Benth. & Hook. f., Gen. Pl. 2: 286
(1873).
Plagiocheilus Arn. ex DC., Prodr. 6: 142 (1838).
Polygyne Phil. (1864).
Erect perennial herbs; pubescent. Leaves entire to
serrate-dentate or lyrate-pinnatifid, bases decurrent on stem. Inflorescences corymbose; involucres hemispheric; bracts 2–3-seriate, gradate, outer
bracts narrow, inner bracts apically membranaceous. Rays lacking. Florets yellow, tube short, limb
campanulate, lobes 5, triangular, erect-spreading;
style appendages short-triangular, hirtellous. Achenes oblong-obovoid, usually 3-ribbed, glandular,
setulose; pappus 2–3-seriate, numerous unequal,
caducous bristles. Three species, southern Africa.
Erect to creeping perennial herbs; pilose to hirsute
on leaf-base margins, leaf surfaces or sometimes on
stems. Petioles winged with broadened bases, leaf
blades pinnately to bipinnately dissected. Heads
solitary on axillary peduncles or short-pedunculate
in small terminal clusters; involucres campanulate
to hemispheric; bracts c. 3-seriate, mostly subequal, ovate to oblong, blunt, margins hyaline. Ray
florets multiseriate, pistillate, white, shortly bilabiate with 1 or 2 smaller lobules adaxially. Disc florets functionally male, greenish-yellow to reddishbrown, tube slender, limb narrowly funnelform,
lobes deltate to oblong-lanceolate, erect to spreading. Achenes oblong, constricted but not beaked,
setula tips anchor-shaped; pappus lacking. x = 9.
Seven species, South America.
837. Mtonia Beentje
Mtonia Beentje, Kew Bull. 54: 97 (1999).
Erect, densely glanduliferous, annual herbs; leaves
sessile, narrowly elliptic to lanceolate, entire,
pilose. Inflorescences corymbose; heads 3–4 mm
high, crowded at tips of branches; peduncles short,
with stipitate glands; involucral bracts 2–3-seriate,
subequal, oblong-lanceolate; receptacles convex to
toroid, spiculiferous. Rays 2–3-seriate, c. 30–40,
yellow, limbs short, erect. Disc florets numerous,
yellow, narrowly funnelform, with sticky glands;
5-lobed. Achenes setuliferous; pappus a minutely
XV.6. Subtribe Lagenophorinae G.L. Nesom
(1994).
Annual to perennial herbs or subshrubs. Leaves
alternate, mostly in a basal rosette. Heads borne
singly on scapose stems or few in a loose panicle;
involucral bracts herbaceous; receptacles plane to
convex, conical in Pytinicarpa, sometimes concave
308
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
in Thespis, epaleate. Ray florets (1–)2–4-seriate,
white, limb reduced, usually not coiling, sometimes
reflexing. Disc florets often functionally male; style
branch appendages deltate to lanceolate, mostly
papillose. Achenes compressed, 2-nerved, usually
with persistent or deciduous glands at the apex or
beak; setulae without anchor-shaped tips; pappus
often absent. x = 9.
Key to the Genera
1. Pappus of 6–20, rather stout or persistent, barbellate
bristles
2
– Pappus none or with distorted caducous bristles
3
2. Perennial herbs from short woody stolons; leaves all
basal; pistillate corollas radiate
844. Pappochroma
– Annual herbs in small spreading clumps; leaves alternate; pistillate corollas eradiate
851. Thespis
3. Heads with 9 or fewer florets, only 1 or 2 disc florets
849. Sheareria
– Florets in heads 15 or more
4
4. Disc corollas goblet-shaped, tube abruptly opening
into a broad limb; pistillate florets radiate
5
– Disc corollas tubular to funnelform, gradually opening
into the limb; pistillate corollas with or without limb 8
5. Annual or perennial herbs, not aromatic; leaves herbaceous, flat-margined, sometimes subclasping but not
basally dilated or sheathing, basal leaves often persistent but cauline leaves continuing unreduced in size
halfway to nearly completely up the stem
6
– Perennial herbs to subshrubs or shrubs, at least some
species aromatic; leaves thickened to coriaceous, usually with revolute or deflexed margins, basally dilated
and sheathing, evenly arranged along stems or in rosulate clusters
7
6. Pistillate florets 2–5(–10)-seriate; corollas greenishwhite to pink or purplish; pappus lacking
843. Myriactis
– Pistillate florets 2-seriate, c. 11, disc florets c. 11; pistillate corollas whitish; pappus lacking or with caducous
scraggly bristles
848. Rhynchospermum
7. Leaves of current year in a rosulate or subrosulate
cluster; heads on long, scapiform peduncles; anthers
with a short apical appendage
840. Keysseria
– Leaves densely and more or less evenly arranged on
stems; heads without evident peduncle; anthers without an apical appendage
845. Piora
8. Heads radiate; limb of ray florets strongly developed;
achenes with glands
9
– Heads disciform; limbs of pistillate florets rudimentary or absent
10
9. Involucral bracts narrowly lanceolate to linear, acute
to acuminate; receptacles convex 841. Lagenophora
– Involucral bracts elliptic-obovate to oblong, rounded
to obtuse; receptacles sharply acute 846. Pytinicarpa
10. Disc florets perfect, more numerous than pistillate florets; involucral bracts c. 6, ciliate; achenes obcompressed(?)
847. Rhamphogyne
– Disc florets functionally male, less numerous than pistillate florets; involucral bracts 9–16, denticulate; achenes compressed
11
11. Achene beak conspicuous; ovaries of disc florets completely absent; leaves entire
842. Lagenocypsela
– Achene beak rudimentary or absent; ovaries of disc
florets present, sterile; leaves toothed
850. Solenogyne
Genera of Lagenophorinae
840. Keysseria Lauterb.
Keysseria Lauterb., Feddes Repert. Spec. Nov. Regni Veg. 13:
241 (1914); Koster, Nova Guinea, Bot. 24: 597–608 (1966),
reg. rev.
Perennial, aromatic, spreading herbs or subshrubs,
branches erect or ascending. Leaves fleshy to coriaceous, linear-terete or elliptic to spathulate,
glabrous to tomentose, sometimes glandular,
margins entire to shallowly dentate. Heads solitary, often nodding, peduncles scapiform with
1–6 leaf-like bracteoles; involucres campanulate
or hemispheric; bracts 2–4-seriate, subequal,
herbaceous, midrib prominent, usually glandular,
margins membranaceous, distally fringed. Rays
2–3-seriate, pistillate, white to purplish, short,
linear, recurving. Disc florets perfect or functionally male, yellow to pinkish, funnelform, limb
abruptly widened, lobes (3–)4(–5), triangular.
Fertile achenes obovate to elliptic, glabrate; pappus
absent. x = 9. Twelve species, Indonesia, Hawaii.
841. Lagenophora Cass.
Lagenophora Cass., Bull. Sci. Soc. Philom. Paris 1816: 199
(1816), nom. et orth. cons.; Cabrera, Blumea 14: 285–307
(1966), rev.
Lagenifera Cass. (1816), orth. var. rej.
Perennial stoloniferous herbs or subshrubs,
often matted; usually puberulous; stems erect or
procumbent. Leaves cauline or mostly in rosette,
sessile to petiolate, linear to spathulate, entire or
dentate. Heads solitary, long-pedunculate; involucres turbinate to subrotate; bracts 2–3-seriate,
subequal, more than 10, narrowly oblong, acute or
obtuse, herbaceous, entire or denticulate, margins
scarious. Rays 1–4-seriate, pistillate, white to
purple, coiling. Disc florets fewer than rays, perfect
or functionally male, yellow to reddish, lobes
4 or 5, triangular, erect to spreading. Achenes
obovate to oblanceolate, subfusiform, with short,
glandular and viscid beak or with glandular apical
rim; pappus lacking. x = 9. Fourteen species,
south-eastern Asia, Australasia, Pacific islands,
Central America, South America.
Compositae
842. Lagenocypsela Swenson & K. Bremer
Lagenocypsela Swenson & K. Bremer, Austral. Syst. Bot. 7:
270 (1994).
Small, perennial, rosettiform herbs; with erect
caudex. Leaves in rosette, linear or petiolate with
lanceolate blades, glabrous, entire. Inflorescences
1-headed, on erect pilose peduncles; involucres
campanulate; bracts 9–16, 2–3-seriate, gradate,
rounded, 1-nerved, margins scarious, denticulate.
Pistillate florets 1–2-seriate, red-purple, filiform,
minutely toothed distally. Disc florets few, functionally male, red-purple, narrowly campanulate,
lobes deltate, erect. Achenes fusiform, shortbeaked, glabrous; pappus lacking. Two species,
New Guinea.
843. Myriactis Less.
Myriactis Less., Linnaea 6: 127 (1831); Cabrera, Blumea 14:
285–307 (1966), rev.; Wang & Peng, Comp. Newslett. 41:
58–59 (2004), typ.
Perennial herbs, often rhizomatous, subglabrous
to hirsute-villous. Leaves cauline, bases rather
clasping, blades oblanceolate to ovate or orbicular, crenate-dentate to pinnatisect. Heads 1 to
many, sometimes in subthyrsoid inflorescences,
pedunculate; involucres cupuliform to hemispheric; bracts 2–4-seriate, subequal to gradate,
herbaceous, oblong-lanceolate, with basally keeled
midrib. Rays c. 10–45 or more, 2–5(–10)-seriate,
fertile or sterile, greenish-white to purplish in
age. Disc florets perfect or functionally male,
yellow or green, goblet-shaped, lobes 4 or 5,
lanceolate, spreading to reflexing. Achenes obovate to oblanceolate, faces shiny, eglandular, apex
short-beaked or in disc achenes erostrate, glanddotted; pappus absent. x = 9. Two species, tropical
America, eastern Asia, Indonesia, Philippines.
844. Pappochroma Raf.
Pappochroma Raf., Fl. Tellur. 2: 48 (1836) [1837]; Nesom,
Phytologia 76: 426 (1994), emend.
Lagenithrix G.L. Nesom (1994).
Lagenopappus G.L. Nesom (1994).
Perennial herbs from short stolons, often in mats;
stems and leaves hispid-pilose, stipitate-glandular
or eglandular. Leaves all basal, thick, oblanceolate
to spathulate, entire to distally crenate, 1–3-nerved.
Heads solitary, sessile or on bracteolate scapes; involucres hemispheric; bracts 2–3-seriate, subequal,
oblong-lanceolate, 1-nerved, herbaceous except for
309
scarious margins, often purple-tinged. Rays 25–
36, 1–4-seriate, white, narrow, straight. Disc florets
perfect or functionally male, yellow, funnelform,
tube linear, lobes 4 or 5, triangular. Fertile achenes
oblanceolate-oblong, 2–4-nerved, glabrous except
for viscid glands distally, slightly constricted or
with neck distally; pappus 1–2-seriate, persistent,
bristles 12–50. Nine species, Australia.
845. Piora J.T. Kost.
Piora J.T. Kost., Nova Guinea, Bot. 24: 596 (1966).
Ericoid shrubs; aromatic, glabrous, stems, leaf
undersides and involucres gland-dotted; stems
erect, virgately branched. Leaves caducous below, sessile, with short, sheathing bases, blades
coriaceous or fleshy, 1-nerved, linear, obtuse,
margins entire, recurved. Heads solitary, on
short, sparsely bracteolate peduncles; involucres
obconic-campanulate; bracts 3–4-seriate, gradate,
mostly linear-lanceolate, acute, 1-nerved. Rays
c. 40, 2–3-seriate, pistillate, bluish-white, not
coiling. Disc florets c. 30, functionally male, yellow,
narrowly funnelform, lobes 4, lanceolate, erect,
with orange ducts; anthers with minute, indistinct
apical appendages. Achenes oblong-obovate,
triangular, truncate apically; pappus lacking. One
species, P. ericoides J.T. Kost., New Guinea.
846. Pytinicarpa G.L. Nesom
Pytinicarpa G.L. Nesom, Phytologia 76: 136 (1994); Nesom,
Sida 19: 513–518 (2001), emend.
Perennial, eglandular, rhizomatous herbs. Leaves
in basal rosettes, oblanceolate to linear, with 3
parallel veins, entire or distally toothed, sparsely
villous. Scapes with few filiform bracts; heads
solitary; involucres campanulate; bracts 2–4seriate, all but outermost subequal, oblong, tips
rounded, with thin orange midvein, margins
scarious, laciniate to ciliate; receptacles sharply
conic. Rays 1-seriate, pistillate, white, tube hairy,
limbs 4-veined, coiling. Disc florets functionally
male, orange-veined, tube short, limb tubularcampanulate, lobes deltate, erect. Fertile achenes
narrowly oblong, base narrowed, neck short,
(1–)2(–3)-ribbed on each surface, glabrous, eglandular; pappus lacking. Two species, New Caledonia
and Fiji.
847. Rhamphogyne S. Moore
Rhamphogyne S. Moore, J. Bot. 52: 146 (1914); Swenson &
Bremer, Austral. Syst. Bot. 7: 265–273 (1994), emend.
310
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Small, perennial, caespitose herbs. Leaves densely
inserted, linear-oblanceolate, entire or pinnatifid.
Heads small, solitary, sessile, disciform; involucres
narrowly campanulate; bracts c. 6, acute, ciliate.
Peripheral florets 1-seriate, pistillate, yellow,
tubular, 3-lobed. Disc florets more numerous
than pistillate florets, perfect, yellow, limb narrowly funnelform, lobes 3 or 4, ovate-triangular,
erect-spreading, anthers inserted near base of
corolla; style branches acute, hairy. Achenes
elliptic-obovate, long- and sometimes sinuously
beaked, glabrous; pappus lacking. Two species,
Rodrigues Island, New Guinea.
848. Rhynchospermum Reinw.
Rhynchospermum Reinw., Syll. Pl. Nov. 2: 7 (1825).
Erect perennial herbs; puberulous. Leaves petiolate, blades herbaceous, elliptic to oblanceolate,
remotely dentate. Inflorescences racemose-spicate
to thyrsoid; peduncles usually widely diverging;
heads subdiscoid; involucres hemispheric; bracts
3–4-seriate, weakly gradate, oblong, with distal
green patch, margins narrowly scarious, apices
obtuse, fimbriate. Rays c. 11, 2-seriate, pistillate,
whitish, limbs short, elliptic. Disc florets c. 11,
perfect, yellowish, tubes short, narrow, limb
campanulate, lobes 4, broadly triangular, erect;
style appendages oblong-ovate. Achenes obovate,
gland-dotted distally, ray achenes with apical
neck; pappus lacking or of up to 10 scraggly,
whitish, caducous bristles. x = 9. One species,
R. verticillatum Reinw., eastern and south-eastern
Asia.
849. Sheareria S. Moore
Sheareria S. Moore, J. Bot. 13: 227, t. 165 (1875).
Small annual herbs; sparsely pilose; stems erect,
costate; branches sparse, divergent. Leaves small,
bract-like, linear, entire. Inflorescences mostly 1headed; involucres campanulate; bracts 6 or 7, c.
2-seriate, gradate, ovate to oblong, concave, without obvious venation. Rays 3 or 4, pistillate, white,
with broad limb, apex weakly 5-crenulate. Disc florets 1 or 2, functionally male, yellow, tube slender, limb narrowly funnelform, lobes deltate, erect;
style undivided. Fertile achenes oblong, trigonous,
with narrowly winged, denticulate ribs, glabrous;
pappus lacking. Two species, China.
850. Solenogyne Cass.
Solenogyne Cass., Dict. Sci. Nat. 50: 493 (1827); Davis, Proc.
Linn. Soc. New South Wales 75: 88–194 (1948), rev.; Adams,
Brunonia 2: 3–65 (1979), rev.
Rosulate perennial herbs; glabrous to pilosulous.
Leaves in rosette, oblanceolate to spathulate with
cuneate or pseudopetiolate base, dentate distally.
Heads solitary on elongate peduncles, disciform;
involucres broadly campanulate to hemispheric;
bracts c. 3-seriate, subequal, herbaceous, oblong,
obtuse or rounded, denticulate, margins not or narrowly scarious. Pistillate florets 3–4-seriate, greenish or red-tinged, filiform, oblique or with minute
limbs. Disc florets fewer than pistillate, functionally male, white or greenish to reddish, tube narrow, limb narrowly campanulate, with (3–)4(–5)
erect, deltate lobes; style branches hairy. Achenes
oblong-obovate, short-beaked or beakless, smooth,
glabrous, mostly non-glandular; pappus lacking.
x = 9. Three species, Australia, Tasmania.
851. Thespis DC.
Thespis DC. in Guill., Arch. Bot. (Paris) 2: 517 (1833).
Spreading annual herbs, puberulous to pilose.
Leaves sessile to petiolate, blades ovate to obovate, entire to serrate-dentate. Heads in small
corymbose cymes, short-pedunculate, discoid; involucres hemispheric; bracts c. 2-seriate, subequal,
glabrous, oblong, concave, with broad green band,
margins scarious. Pistillate florets multiseriate;
corollas minutely tubular or lacking. Disc florets
few, functionally male, pale, shortly funnelform,
lobes 4, oblong-ovate, erect, with orange resin
ducts; anther appendages reduced to small
apiculus. Fertile achenes ellipsoid, multinerved,
eglandular, not or slightly compressed, finely
papillose; pappus 1-seriate, of 6–13 scabrid bristles
shorter than achene, sometimes lacking in disc
florets. Three species, south-eastern Asia.
XV.7. Subtribe Baccharidinae Less. (1830).
Subf. Baccharidoideae Burmeist. (1837).
Tribe Heterothalaminae Endl. (1837).
Trees, woody shrubs, or vines, rarely herbs or subshrubs, often dioecious. Leaves alternate, rarely opposite, entire to serrate or pinnatisect, commonly
gland-dotted. Inflorescences congested corymbose
to thyrsoid; involucral bracts flat, mostly herbaceous; receptacles with or without pales. Female
heads with florets tubular or with short white limbs,
Compositae
central florets sometimes appearing functionally
male; male heads with pistillate florets absent or
1–2-seriate; disc style branches with short, papillose appendages. Achenes small, terete to compressed, multinerved, usually eglandular, glandular in some Baccharis and Archibaccharis, setulae
without anchor-shaped tips; pappus 1-seriate, persistent, bristles often apically dilated. x = 9.
Key to the Genera
1. Female heads with a few central functionally male florets
852. Archibaccharis
– Female heads without male florets
2
2. Male and female florets in totally separate heads; plants
totally dioecious or rarely monoecious
853. Baccharis
– Male heads with 2 outer series of pistillate florets; female heads paleate, male heads epaleate
854. Heterothalamus
Genera of Baccharidinae
852. Archibaccharis Heering
Archibaccharis Heering, Jahrb. Hamburg Wissensch. Anst.
21, Beih. 3: 40 (1904); Jackson, Phytologia 32: 81–194 (1975),
rev.; Nesom, Phytologia 71: 152–159 (1991), emend.
Hemibaccharis S.F. Blake (1924).
Perennial herbs, shrubs or vines, functionally dioecious, with stipitate or sessile glands, not glutinous; stems straight, twining or zigzag. Leaves petiolate or sessile. Inflorescences terminal or axillary, strongly cymose or corymbose, unisexual or
vestigially gynomonoecious. Involucral bracts 3–
5-seriate, gradate. Female heads with outer florets pistillate, central florets appearing bisexual
but anthers and achenes usually sterile; pistillate
florets radiate to tubular, sometimes with staminodes. Male heads with gynoecia usually sterile. Fertile achenes compressed, ovate to oblong, 2–5(–7)nerved, mostly 3-angled; pappus 1-seriate, bristles
not elongating at maturity, usually dilated apically
in staminate florets and sometimes in pistillate florets. x = 9. Thirty-two species, mostly Mexico, Central America, 1 or 2 in South America.
853. Baccharis L.
Baccharis L., Sp. Pl. 860 (1753), nom. cons.; Barroso,
Rodriguésia 40: 3–273 (1976), reg. rev.; Barroso & Bueno,
Fl. Illustr. Catarin., Compostas 5, subtribo Baccharidinae:
769–777, 784–1065 (2002), reg. rev.; Cuatrecasas, Revista
Acad. Colomb. Ci. Exact. 13: 5–102 (1967), reg. rev.; Hellwig,
Mitt. Bot. Staatssamml. München 29: 1–456 (1990), rev.;
311
Mahler & Waterfall, Southw. Naturalist 9: 189–202 (1964),
reg. rev.; Matuda, Anales Inst. Biol. Univ. Nac. Mexico 28:
143–174 (1957), reg. rev.
Molina Ruiz & Pav. (1794), nom. illegit. non Cav. (1790).
Sergilus Gaertn. (1791).
Tursenia Cass. (1825).
Pingraea Cass. (1826).
Baccharidastrum Cabrera (1937).
Baccharidiopsis G.M. Barroso (1976).
Neomolina Hellwig (1993), nom. illegit. non Honda &
Sakis. (1930).
Small trees, shrubs or perennial herbs, dioecious
or rarely monoecious; stems sometimes winged.
Leaves rarely opposite, scale-like or absent, lamina
linear to ovate, obovate or pinnatisect, glabrous to
glandular-dotted and glutinous, rarely tomentose.
Inflorescence corymbose or thyrsoid, spicate,
racemose or 1-headed; heads unisexual, discoid.
Involucres cylindric to hemispheric; bracts 3–8seriate, gradate; receptacles plano-convex, usually
epaleate. Female florets filiform-tubular; staminate heads with florets mostly whitish to purplish,
funnelform, lobes cut nearly to base of limb,
spreading-reflexing; style branches rudimentary.
Achenes compressed or nearly terete, glabrous
or hairy, 5–10(–12)-nervate; pappus 1–3-seriate,
bristles sometimes caducous, in female florets
usually elongating beyond involucre, in male
florets often apically broadened. x = 9. Circa 360
species (fide Daniel Giuliano, pers. comm.), mostly
of tropical America, north into central and coastal
USA.
Hellwig (1990, 1993, 1996) proposed and
formally recognized segregate genera from within
Baccharis and made nomenclatural combinations
for representative species, but he has not completed or extended his conceptual modifications.
In a study of Baccharidinae, Giuliano (2001) has
reviewed and modified the infrageneric taxonomy
and also proposed to recognize several segregate
genera, but with concepts which minimally intersect or overlap with those of Hellwig (1990, 1993,
1996).
854. Heterothalamus Less.
Heterothalamus Less., Linnaea 5: 145 (1830); Barroso
& Bueno, Fl. Illustr. Catarin., Compostas 5, subtribo
Baccharidinae: 777–784 (2002), rev.
Polygamo-dioecious, more or less ericoid shrubs,
densely branched, often slightly glutinous. Leaves
linear to elliptical. Inflorescences of small ter-
312
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
minal clusters of heads, often overtopped by
lateral branches; involucres hemispheric; bracts
3–4-seriate, weakly to strongly gradate, oblong to
linear, pale at base, green along midvein distally,
margins hyaline; receptacles convex to subconical.
Female heads with navicular pales, with only yellowish, filiform, short-rayed, pistillate florets. Male
heads epaleate, with outer, sterile, pistillate florets
2-seriate, limbs short; inner florets functionally
male, lobes deltate, reflexing. Fertile achenes
3- or 5-angled, glabrous; pappus bristles short;
male achenes sterile; pappus bristles broadened
apically. Two species, southern Brazil, Uruguay.
XV.8. Subtribe Podocominae G.L. Nesom
(1994).
Annual to perennial herbs, often gland-dotted.
Leaves alternate, entire to toothed or divided.
Heads mostly solitary, paniculate or corymbose
in some genera; involucral bracts herbaceous to
thickened or slightly keeled; receptacles planoconvex, epaleate. Ray florets 2–3-seriate, corollas
white to bluish (yellow in Kippistia), limbs coiling.
Disc florets bisexual or often functionally male;
style appendages usually lanceolate to linear,
papillose. Achenes 2-nerved, compressed, often
with a neck or beak, distally glandular, setulae
without anchor-shaped tips; pappus bristles
(1–)2–3-seriate, often with a much shorter outer
series. x = 9.
Key to the Genera
1.
–
2.
–
3.
–
4.
–
5.
Plants from America
2
Plants from Australia
7
Achenes with a distinct neck or beak
3
Achenes apically rounded or truncate, without a beak
or neck
4
Vestiture arachnoid; leaves cauline, densely overlapping, not clasping; heads 20–35 mm wide, 1–2 on leafy
stems; disc ovaries sterile
855. Asteropsis
Vestiture glabrate to scabrous or hispid-pilose, never
arachnoid; leaves cauline, basal leaves sometimes persistent, loosely disposed along long internodes, clasping to subclasping; heads (5–)7–15(–17) mm wide,
solitary on scapose stems; disc ovaries fertile
869. Podocoma
Leaves herbaceous, not rigid, mostly oblanceolate, entire to serrate or pinnately dissected
5
Leaves rigid, linear and entire or apically trifurcate, or
pinnately dissected
6
Leaves all basal or basal and lower cauline; heads solitary on scapose stems; disc ovaries sterile; achenes
eglandular
860. Inulopsis
– Leaves primarily cauline, basal leaves persistent in
some species; heads on leafy stems; disc ovaries fertile;
achenes glandular
865. Laennecia
6. Stems and leaves sessile- or punctate-glandular, also
arachnoid; disc ovaries fertile
866. Microgynella
– Stems and leaves densely stipitate-glandular, not
arachnoid; disc ovaries sterile
870. Sommerfeltia
7. Achenes with a short neck or long beak
8
– Achenes apically rounded or truncate, without a neck
or beak
11
8. Achenes with a long, filiform beak; receptacles flat or
nearly so; involucres reflexed after fruiting
863. Ixiochlamys
– Achenes with a short neck
9
9. Achenes with a prominently narrowed, basal extension
below the seed, basal extension densely tufted with
stiff, appressed hairs
872. Vittadinia p.p.
– Achenes without a narrowed, basal extension below
the seed, hairs at base not denser than on surface 10
10. Achenes c. 2 mm long, with short, broad neck; pappus
bristles 2-seriate, the outer distinctly shorter than the
inner; involucral bracts incurved after fruiting
857. Dichromochlamys
– Achenes c. 1 mm long, with a short, narrow neck; pappus bristles 1-seriate; involucral bracts never incurved
864. Iotasperma
11. Achenes usually with 2 to numerous facial nerves between the lateral nerves (facial nerves absent in six
species of Vittadinia)
12
– Achenes with only a pair of lateral nerves, these variable in thickness, sometimes obscure or absent
14
12. Achenes with a prominently narrowed, basal extension
below the seed, basal extension densely tufted with
stiff, appressed hairs
872. Vittadinia p.p.
– Achenes narrowly oblong to obovate or oblanceolate,
without a narrowed, basal extension below the seed,
hairs at achene base not denser than on achene surface
13
13. Disc florets with fertile ovaries; achenes usually with
3 pairs of facial nerves; herbs from a woody rootstock
856. Camptacra
– Disc florets with sterile ovaries; fertile achenes with
one pair of facial nerves, rarely none; woody-based
shrubs or shrublets
871. Tetramolopium
14. Achenes with a prominently narrowed, basal extension
below the seed, basal extension densely tufted with
stiff, appressed hairs
872. Vittadinia p.p.
– Achenes without a narrowed, basal extension below
the seed, hairs at achene base not denser than on achene surface
15
15. Disc florets with fertile ovaries; pappus of ray and disc
achenes similar
16
– Disc florets with sterile ovaries; pappus of ray and disc
achenes different
17
16. Ray florets with yellow corollas, ovaries often sterile;
disc corollas 4-merous; achenes usually apically truncate, with a correspondingly broad pappus insertion
864. Kippistia
– Ray florets with white to bluish corollas, ovaries fertile;
disc corollas 5-merous; achenes usually rounded to
a narrow pappus insertion
868. Peripleura
17. Disc corollas with 5 lobes; pappus of ray achenes of
equal length bristles, pappus of disc achenes of unequal bristles or of bristles and scales 867. Minuria
Compositae
– Disc corollas with 3–4 lobes; pappus of ray achenes of
scales or scales and bristles, pappus of disc (sterile)
achenes of bristles
18
18. Leaves oblanceolate; ray achenes with pappus of scales
and bristles
858. Dimorphocoma
– Leaves linear; ray achenes with pappus of scales 19
19. Erect or decumbent herbs; leaves cauline; heads terminal, solitary; pappus scales of ray achenes lanceolate
859. Elachanthus
– Sessile herbs with clustered leaves; heads clustered
among leaf bases; pappus scales of ray achenes broadly
obovate
862. Isoetopsis
Genera of Podocominae
855. Asteropsis Less.
Asteropsis Less., Syn. Gen. Comp. 188 (1832); Nesom,
Phytologia 76: 101–105 (1994), relat.
Perennial herbs; stems, leaves and involucres
loosely tomentose-villous. Leaves densely spirally
inserted, sessile, non-clasping, 1-nerved, linear,
entire. Inflorescences terminal, 1- to few-headed;
involucres hemispheric, 20–30 mm wide; bracts
4–5-seriate, slightly gradate, linear-lanceolate,
with long-attenuate, reddish tips. Rays multiseriate, to 100 or more, pistillate, white, limbs long.
Disc florets functionally male, yellow, tubular,
lobes narrowly triangular; style branches linear,
papillae hair-like. Fertile achenes broadly obovate,
apex short-beaked, surfaces sericeous, eglandular;
pappus bristles c. 2-seriate, persistent, capillary.
One species, A. macrocephala Less., southern
Brazil, Uruguay.
856. Camptacra N.T. Burb.
Camptacra N.T. Burb., Brunonia 5: 11 (1982); Lander, Nuytsia 6: 61 (1987), comb.
Erect perennial herbs from rootstock; loosely tomentose to subglabrous, glandular. Leaves cauline,
sessile, linear to oblong, entire to trifid at tip. Heads
solitary on long peduncles; involucres broadly campanulate to hemispheric; bracts 3–4-seriate, oblong
to linear-lanceolate, with or without membranaceous margins. Rays 1–2-seriate, pistillate, white
to bluish, limbs narrow, short. Disc florets perfect,
yellow, tube short, limb funnelform, lobes triangular, erect-spreading. Achenes narrowly elliptic
to elliptic-oblong, bases not extended nor more
densely hairy, dark or purplish when mature, glandular, each face with 3 slender, non-vascularized
ribs; pappus bristles 1–2-seriate, persistent. x = 9.
Two species, Australia.
313
857. Dichromochlamys Dunlop
Dichromochlamys Dunlop, J. Adelaide Bot. Gard. 2: 235
(1980).
Decumbent annual herbs; stems and leaves villous
to woolly, rarely glabrous, glandular. Leaves
cauline, spathulate, cuneate at base, apex coarsely
toothed. Inflorescences 1-headed, on long, densely
glandular peduncles; involucres hemispheric to
subglobose; bracts 4–5-seriate, strongly gradate,
lanceolate, apices acuminate, median bracts
with broad, white, scarious margins; receptacles
conical, densely glandular, edge with fine hairs.
Florets 100–150. Rays numerous, multiseriate,
pistillate, white to pink. Disc corollas perfect,
yellow. Achenes obovate, smooth, constricted and
glandular distally, sericeous on basal 3/4; pappus
bristles persistent, multiseriate, uneven, outer
series short. One species, D. dentatifolia (F. Muell.)
Dunlop, central Australia.
858. Dimorphocoma F. Muell. & Tate
Dimorphocoma F. Muell. & Tate, Trans. Proc. Roy. Soc.
S. Australia 6: 107 (1883).
Small, ascending annual herbs, with non-glandular
hairs. Leaves cauline, lower leaf bases attenuate, upper subamplexicaul, oblanceolate, entire, midvein
prominent. Heads solitary, pedunculate; involucres
cup-shaped; bracts c. 8, 2-seriate, subequal, lanceolate, herbaceous with scarious margins. Rays 7
or 8, 1-seriate, pistillate, white, limbs short, narrow. Disc florets 3 or 4, functionally male, yellow,
tubular, lobes 3 or 4. Fertile achenes obovate, without neck, sericeous-villous; pappus 2-seriate, outer
row of many bristles, inner row of c. 6 equally long
linear-lanceolate scales; disc florets with 3 or 4 pappus bristles. Two species, Australia.
859. Elachanthus F. Muell.
Elachanthus F. Muell., Linnaea 25: 410 (1853).
Small, erect or decumbent annual herbs; glabrous
or puberulous, non-glandular. Leaves cauline,
sessile, linear, entire. Heads solitary, pedunculate,
disciform; involucres campanulate; bracts c.
2-seriate, subequal with few outer bracts smaller,
oblong to oblanceolate, obtuse, herbaceous with
scarious margins. Pistillate florets 2–3-seriate,
white, tubular, narrowed and 2- or 3-lobed apically. Disc florets 2–5, functionally male, yellow,
narrowly funnelform from narrow tube, lobes 3–5.
Fertile achenes narrowly obovate to obconical,
314
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
slightly compressed, without neck, minutely
setulose to sericeous-villous; pappus c. 2-seriate,
of lanceolate scales; slender disc achenes with
pappus of 1–3 linear, basally connate scales or 3 or
4 bristles. Two species, southern Australia.
860. Inulopsis (DC.) O. Hoffm.
Inulopsis (DC.) O. Hoffm. in Engler & Prantl, Nat. Pflanzenfam. IV/5: 145, 149 (1890); Nesom, Phytologia 76: 115–124
(1994), rev.
Haplopappus Cass. sect. Inulopsis DC. (1836).
Perennial herbs from rhizomes; leaves and
stems glabrate to hirsute-villous, sometimes
gland-dotted, resinous. Leaves all basal or some
cauline, obovate to linear, with 1 or 2 pairs of
longitudinal veins from base. Heads solitary
on scapes; involucre hemispheric to turbinate;
bracts 2–4-seriate, outer 1/2–3/4 as long as inner,
lanceolate to lanceolate-oblong, herbaceous,
1-nerved. Rays 1-seriate, pistillate, white, tube
with biseriate hairs. Disc florets functionally male,
yellow, orange-veined, narrowly funnelform, lobes
deltate, erect. Achenes oblanceolate to elliptic,
sometimes with extra rib on each face, eglandular,
strigose-sericeous, apical neck vestigial; pappus
bristles (1–)2–3-seriate, subequal, attenuate, often
with short outer setae. x = 9. Four species,
southern Brazil, Paraguay, eastern Bolivia.
861. Iotasperma G.L. Nesom
Iotasperma G.L. Nesom, Phytologia 76: 144 (1994).
Erect, taprooted annual herbs; surfaces hispidpilose, sparsely glandular. Leaves cauline, sessile,
subclasping,
oblanceolate-oblong,
coarsely
toothed distally. Inflorescences loosely corymbose;
involucres shallowly cupulate; bracts c. 2-seriate,
subequal, elliptic-lanceolate, green, 3-nerved,
stipitate-glandular, margins scarious, inner bracts
basally indurate. Rays (1–)2–3-seriate, pistillate,
white to purplish. Disc florets functionally male,
yellow, tube linear, limb subabruptly funnelform,
lobes short-triangular. Achenes obovate to
elliptic-obovate, sclerified, setuliferous, rounded
to narrow, gland-dotted bases and apices, prominent apical callus bearing 6–10(–15), 1-seriate,
caducous, bristles. Two species, Australia.
862. Isoetopsis Turcz.
Isoetopsis Turcz., Bull. Soc. Imp. Naturalistes Moscou 34, 1:
174, t. 3 (1851); Robinson & Brettell, Phytologia 26: 73–75
(1973), relat.
Sessile, rosulate annual herbs, glabrous. Leaves
linear, with pale broad bases. Heads sessile among
leaf bases; involucral bracts c. 2-seriate, subequal,
oblong-ovate, obtuse, pale below, green distally,
margins narrowly pale to scarious. Rays 5–6,
pistillate, apomictic(?), greenish or purplish
distally, tubular, narrowed to lobed tip, lobes
usually 3, triangular with setiform tip; bulbous
style base with numerous fibrous strands entering
achene. Disc florets 2–3, functionally male, pale
yellowish, with slender tube, limb flaring, lobes 4;
stamen appendages small, ovate-oblong. Fertile
achenes obovate, nearly terete, 2-nervate, without
neck, sericeous; pappus of c. 8 persistent, obovate,
obtuse, broadly costate scales. One species, I.
graminifolia Turcz., Australia.
863. Ixiochlamys F. Muell. & Sond. ex Sond.
Ixiochlamys F. Muell. & Sond. ex Sond., Linnaea 25: 466
(1853); Dunlop, J. Adelaide Bot. Gard. 24: 241–252 (1980),
rev.
Annual or perennial herbs to small shrubs; glabrous to pilose or hirsute, often stipitate-glandular.
Leaves cauline, sessile, entire to pinnately lobed.
Heads solitary, pedunculate; involucres broadly
campanulate; bracts 3–4-seriate, gradate, herbaceous, lanceolate to linear, reflexed after anthesis,
glabrous or glandular. Rays several-seriate, pistillate, white to pink, limbs often short. Disc
florets usually fewer than rays, functionally male
or perfect, yellow, funnelform from slender tube,
lobes triangular, erect. Fertile achenes smooth,
with few or no hairs or glands, apically narrowed
into filiform beak; pappus bristles persistent, often
with shorter outer bristles. Four species, western
and central Australia.
864. Kippistia F. Muell.
Kippistia F. Muell., Rep. Pl. Babbage’s Exped. 12 (1858); Lander & Barry, Nuytsia 3: 215–219 (1980), emend.
Small, compact, aromatic shrubs. Leaves small, linear, fleshy, glabrous. Heads solitary on branchlets, short-pedunculate; involucres broadly conical;
bracts 2–3-seriate, subequal, linear-lanceolate, fimbriate apically. Rays c. 2-seriate, pistillate, fertile or
sterile, yellow, limbs rather short. Disc florets perfect, yellow, narrowly funnelform, lobes triangular,
erect; style appendages densely hairy. Ray achenes
oblong-linear, sparsely setulose basally; disc achenes elliptic, compressed, glabrous; pappus bristles
unequal, free or bases fused into cup or tube, tube
Compositae
sometimes surmounted by fewer long bristles. One
species, K. suaedifolia F. Muell., southern Australia,
gypsophilous.
865. Laennecia Cass.
Laennecia Cass., Dict. Sci. Nat. 25: 91 (1822); Nesom,
Phytologia 68: 205–228 (1990), rev.
Annual to short-lived perennial, taprooted herbs,
white-tomentose or coarsely hairy, often glanddotted. Leaves sessile, lanceolate or oblanceolate to
oblong, toothed to pinnately lobed, rarely entire.
Inflorescences spicate or racemose to loosely thyrsoid or corymbose; involucres turbinate; bracts
2–5-seriate, gradate, with greenish midvein. Pistillate florets several-seriate, white, filiform-tubular,
eradiate and apically fimbriate or short-radiate.
Disc florets perfect, cream to yellowish, tubularfunnelform, expanded near lobe bases, lobes
lanceolate. Achenes oblanceolate-elliptic to obovate, faces usually with sessile or subsessile glands;
pappus bristles 1–2-seriate, often caducous, with
or without short outer setae or scales. x = 9.
Seventeen species, south-western USA, Mexico to
northern South America (Andean).
866. Microgynella Grau
Microgynella Grau, Mitt. Bot. Staatssamml. München 12:
185 (1975), nom. nov.; Nesom, Phytologia 76:101–105
(1994), emend.
Microgyne Less. (1832), nom. illegit. non Cass. (1827).
Perennial herbs to subshrubs; caudex woody;
stems, leaves and involucres gland-dotted,
resinous, sparingly hispid and with arachnoid
hairs. Leaves densely spirally inserted, linear,
mostly apically trifid with linear lobes. Heads
solitary, on long peduncles; involucres broadly
campanulate; bracts 2–3-seriate, subequal, narrowly oblong to linear, herbaceous with hyaline
margins, inner bracts white-indurate at base. Rays
2-seriate, pistillate, reddish. Disc florets perfect,
reddish, narrowly funnelform from slender tube,
lobes deltate. Achenes broadly oblanceolateconical, apically truncate, not constricted above,
densely sericeous and gland-dotted; pappus c.
2-seriate, bristles reddish-brown. One species,
M. trifurcata (Less.) Grau, south-eastern Brazil,
Argentina, Uruguay.
867. Minuria DC.
Minuria DC., Prodr. 5: 298 (1836); Lander & Barry, Nuytsia
3: 221–237 (1980), rev.
315
Erect annual herbs to small shrubs; glabrous to
puberulous or tomentose, non-glandular. Leaves
alternate or opposite, sessile, linear to ovate
or oblanceolate, entire to serrulate or dentate.
Heads solitary to rarely clustered, pedunculate;
involucres campanulate; bracts 3–4-seriate, weakly
gradate, linear to lanceolate, herbaceous with
greenish midvein and tip, margins scarious. Rays
2–3-seriate, pistillate, limbs white, blue or violet,
narrow. Disc florets functionally male, yellow,
narrowly funnelform, lobes (4–)5, triangular,
recurving. Fertile achenes linear-lanceolate to
obovate, narrowed distally without neck, setulose,
with free pappus bristles; disc achenes with pappus
of unequal bristles, bristles and scales, or cup of
fused scales with 1–8 bristles, bristles usually more
densely barbellate distally. x = 9. Ten species,
Australia.
868. Peripleura (N.T. Burb.) G.L. Nesom
Peripleura (N.T. Burb.) G.L. Nesom, Phytologia 76: 131
(1994).
Vittadinia A. Rich. subg. Peripleura N.T. Burb. (1982).
Erect annual or perennial herbs; strigose or
puberulous to stipitate-glandular. Leaves cauline,
sessile, usually linear-elliptic, entire or with few
teeth or lobes. Heads solitary on leafy stems,
sometimes pedunculate; involucres broadly campanulate; bracts weakly gradate with few outer
bracts, obtuse to acuminate, margins narrow,
membranaceous. Rays 1–2-seriate, pistillate, white
to purplish, short, narrow. Disc florets perfect,
yellow, tubes longer than limb, lobes deltate, erectspreading. Achenes obovate, apically rounded to
pappus, without narrowed base or tuft of hairs,
faces glandular; pappus bristles 1(–2)-seriate,
mostly 1.5–2.5 times as long as achene, belatedly
caducous in species with basally coherent bristles.
x = 9. Nine species, Australia.
869. Podocoma Cass.
Podocoma Cass., Bull. Sci. Soc. Philom. Paris 1817: 137
(1817); Nesom & Zanowiak, Phytologia 76: 106–114 (1994),
emend.
Podopappus Hook. & Arn. (1836).
Perennial herbs, rhizomatous; usually coarsely
pubescent, eglandular. Basal leaves often persistent, cauline leaves clasping. Inflorescences
1-headed or loosely corymbose; involucres
campanulate, (5–)7–15(–17) mm wide; bracts
3–5-seriate, strongly gradate, stiffly indurate,
316
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
narrowly lanceolate or oblong-lanceolate, greenish
along middle, without orange resin ducts. Rays
2–5-seriate, pistillate, white or purple, rarely
yellow, coiling or filiform and erect. Disc florets
fewer than rays, perfect, cream-coloured, tubes
long, limbs shorter, narrowly funnelform, sometimes orange-veined, lobes deltate, erect. Achenes
strigose, eglandular, apically with broad or filiform
neck; capillary pappus bristles in 2(–3) series or
with the outer series short. x = 9. Circa 9 species,
south-eastern South America.
870. Sommerfeltia Less.
Sommerfeltia Less., Syn. Gen. Comp. 189 (1832); Nesom,
Phytologia 76: 101–105 (1994), emend.
Perennial herbs or subshrubs, from woody caudex;
stems, leaves, and involucres with dense sessile
or short-stipitate glands. Leaves densely spirally
arranged, stiff, entire to pinnately dissected,
tips spinose. Inflorescence 1-headed to loosely
corymbose; peduncles long; involucres broadly
campanulate; bracts 3–4-seriate, gradate, lanceolate to linear, thick and indurate, greenish in
middle, carinate, without orange veins, sides hyaline. Rays 1-seriate, pistillate, white. Disc florets
functionally male, yellow, narrowly funnelform,
lobes triangular, erect. Fertile achenes narrowly
obovate, apically rounded, margins and bases
strigose-sericous, faces gland-dotted and sparsely
hairy; pappus bristles 2-seriate, whitish, elongate,
scarcely barbellate. Two species, southern Brazil,
Uruguay, Argentina.
871. Tetramolopium Nees
Tetramolopium Nees, Gen. Sp. Aster. 202 (1832); Lowrey,
Allertonia 4: 204–262 (1986), rev.; van Royen, Alpine Fl.
New Guinea 4: 3245–3311 (1983), reg. rev.
Luteidiscus St. John (1974).
Monoecious or gynomonoecious; erect to decumbent shrubs or subshrubs; often stipitate-glandular
or gland-dotted; branching sympodial. Leaves
clustered at stem tips, sessile or petiolate, lanceolate to spathulate, entire to dentate or shallowly
lobed. Inflorescences 1-headed or corymbose
with 6–50 heads; peduncles bracteolate; involucres
cylindric to turbinate or hemispheric; bracts 3–
4-seriate, gradate, lanceolate to linear-lanceolate.
Rays 5–250, pistillate, yellow or white. Disc
florets perfect or functionally male, yellow or
maroon, limb campanulate, lobes lanceolate,
erect to spreading. Achenes obovoid to rarely
fusiform-cylindric, faces 0–7-nervate, glabrous,
stipitate-glandular and/or strigose; pappus bristles
1–2-seriate, unequal. x = 9. Thirty-eight species,
New Guinea, north-eastern Australia, Hawaii.
872. Vittadinia A. Rich.
Vittadinia A. Rich., Voy. Astrolabe 250 (1832); Burbidge,
Brunonia 5: 1–72 (1982), rev.
Annual to perennial herbs or small shrubs;
strigose to hispid or hirsute, often glandular.
Leaves cuneate-spathulate to oblanceolate, rarely
elliptic to linear-terete, sometimes lobed. Heads
solitary at tips of branches; peduncles short or
long; involucres broadly campanulate; bracts
3–4-seriate, gradate but weakly overlapping,
mostly linear, margins narrowly scarious. Rays 2to several-seriate, pistillate, white to bluish, short,
narrow. Disc florets perfect, yellow, narrow tube
longer than limb, lobes deltate, erect. Achenes
oblanceolate or cuneate, with narrow base bearing
tuft of setulae, apex with broad short neck, faces
with 3–5(–6) slender, often short, false nerves,
glandular; pappus bristles 2–3-seriate, persistent,
not coherent, c. as long as achene. x = 9. Twenty
species, Australia.
XV.9. Subtribe Asterinae (Cass.) Dumort.
(1827).
Subtribe Heterochrominae Benth. (1873), nom. non rite
public.
Herbs. Leaves alternate, mostly entire, sometimes serrate, with or without glandular dots.
Inflorescence corymbose or heads solitary; involucral bracts flat, mostly herbaceous; receptacles
epaleate. Ray florets usually present, 1-seriate, with
long white or blue limbs, short in Psychrogeton.
Disc florets bisexual; style branches with short, papillose appendages. Achenes obovate, compressed,
2-nerved, often with glands, setulae without
anchor-shaped tips; pappus (1–)2(–3)-seriate,
persistent, of equal lengths or with short outer
series. x = 9.
Key to the Genera
1. Disc florets functionally male; pistillate florets tubular
or with short erect limbs
883. Psychrogeton
– Disc florets perfect; pistillate florets lacking or longradiate
2
2. Caespitose plants with scapose single-headed inflorescences
873. Arctogeron
Compositae
– Plants non-caespitose; with usually branching inflorescences
3
3. Outer involucral bracts leaf-like, strongly differentiated from smaller inner bracts
876. Callistephus
– Outer involucral bracts not strongly differentiated
from inner bracts
4
4. Pappus of all achenes short or lacking
5
– Pappus of at least disc florets over 1 mm long
6
5. Pappus totally lacking
882. Miyamayomena
– Pappus bristles not over 1 mm long 880. Kalimeris
6. Halophytic, subsucculent plants; involucral bracts
glabrous; pappus after anthesis becoming 3–4 times
longer than achene
885. Tripolium
– Plants not halophytic; not subsucculent; pappus not
noticeably elongating after anthesis
7
7. Involucral bracts gradate in 3–5 series
8
– Involucral bracts subequal to weakly gradate, in 2–3
series
10
8. Ray florets present and fertile, with ray corollas shorter
than pappus
884. Rhinactinidia
– Ray florets sterile or lacking
9
9. Ray florets lacking
877. Crinitaria
– Ray florets usually present but sterile 878. Galatella
10. Leaves with recurved margins and cuspidate tips; subshrubs
875. Asterothamnus
– Leaf margins mostly flat; plants mostly strictly herbaceous
11
11. Pappus sordid-grey to brown or reddish, bristles
shorter and sometimes lacking in ray florets
879. Heteropappus
– Pappus usually white or sordid white, alike in ray and
disc achenes
12
12. Pappus of only slender bristles, without short outer
series
874. Aster
– Pappus with inner series of slender bristles and outer
series of short, flat bristles
881. Kemulariella
317
white, multiseriate, bristles as long as disc corollas.
One species, A. gramineum DC., Siberia.
874. Aster L.
Aster L., Sp. Pl. 872 (1753). [Aster Tourn. ex L., Syst. ed. 1.
1735.]; Grierson, Notes Roy. Bot. Gard. Edinburgh 26: 67–
163 (1964), reg. rev.; Lippert, Mitt. Bot. Staatssamml.
München 11: 153–258 (1973), rev.; Tamamschyan,
Fl. U.R.S.S. 25: 77–110 (1959), Engl. transl. 72–104 (1999),
reg. rev.
Bellidiastrum Scop. (1760), nom. et orth. cons. prop.
Perennial herbs or shrubs. Leaves basal or cauline
or both. Inflorescence 1-headed to corymbose
or loosely thyrsoid; involucres hemispheric to
campanulate; bracts several-seriate, subequal
to weakly gradate, linear, mostly herbaceous;
receptacles plano-convex, glabrous to spinulose
or puberulous. Rays pistillate, white, bluish, pink
or purplish, limbs coiling. Disc florets usually
yellow, tubular to narrowly funnelform, lobes
triangular, erect to spreading. Achenes obovate
and compressed to cylindric, setuliferous, often
gland-dotted; pappus 1–2-seriate, bristles barbellate, few or no short bristles outside. x = 9. Circa
180 species, mostly Eurasia and Africa, one species
Arctic and western North America.
875. Asterothamnus Novopokr.
Genera of Asterinae
873. Arctogeron DC.
Arctogeron DC., Prodr. 5: 260 (1836); Tamamschyan, Fl.
U.R.S.S. 25: 135–136 (1959), Engl. transl. 125–127 (1999),
reg. rev.
Caespitose, perennial herbs; with branching woody
caudex; older stem bases covered by marcescent
leaves. Young leaves in apical rosettes, bases
somewhat sheathing, with c. 5 stiff marginal cilia,
blades narrow, almost setaceous. Inflorescences
of many erect, flexuously haired scapes bearing
single heads; involucres campanulate; bracts
3-seriate, not or weakly gradate, green, lanceolate,
acuminate, carinate, densely hairy along keel, margins white-membranaceous; receptacles flat. Rays
pistillate, white or pinkish, limbs oblong-ovate,
coiling. Disc florets yellow, narrowly funnelform,
lobes 5, triangular, recurved. Achenes oblong,
somewhat ribbed, densely sericeous; pappus
Asterothamnus Novopokr., Bot. Mater. Gerb. Bot. Inst. Komarova Akad. Nauk S.S.S.R. 13: 334 (1950); Tamamschyan,
Fl. U.R.S.S. 25: 124–129 (1959), Engl. transl. 25: 117–121
(1999), reg. rev.
Much-branched subshrubs, rhizomatous, stems
finely puberulous to greyish-tomentose. Leaves
sessile, small, oval to linear, margins revolute.
Heads solitary or 3–5 in corymbose groups,
pedunculate; involucres obovoid to hemispheric or
campanulate; bracts 3–4-seriate, gradate, oblonglanceolate to linear, margins membranaceous,
whitish, midrib prominent, brownish or reddish;
receptacles flat, margins of alveolae with some
teeth as long as young achenes. Rays 1-seriate
or lacking, pistillate, blue, violet or whitish-pink,
limbs long. Disc florets yellow, lobes triangular,
spreading-recurved. Achenes ovoid to oblanceolate or subfusiform, 3-angled, 1 convex and 2 flat
sides, extra rib on convex side, appressed-setulose;
pappus white or sordid; bristles as long as disc
corollas. Seven species, Eurasia.
318
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
876. Callistephus Cass.
Aster L. subg. Galatea Cass. (1818).
Callistephus Cass., Dict. Sci. Nat. 37: 491 (1825), nom. cons.
Callistemma Cass. (1817).
Perennial herbs with nodose rhizome; stems erect,
mostly unbranched. Leaves sessile, oblong to
narrowly linear, entire, often gland-dotted, 1- or
3-veined. Inflorescences corymbose, occasionally
1-headed; heads pedunculate; involucres obconic
to subhemispheric; bracts 3–4-seriate, gradate,
herbaceous, lanceolate-ovate to oblong, acute to
obtuse, glabrous or puberulous, with 1–3(–5)
veins, margins partly membranaceous; receptacles
slightly convex, alveole margins dentate. Peripheral
florets, 0 to 1–20, usually sterile, limbs pinkishor bluish-violet. Disc florets 5–60(–100), yellow,
narrowly funnelform, lobes lanceolate; style appendages lanceolate or ovate-triangular. Achenes
oblong, somewhat compressed, without obvious
ribs; pappus bristles whitish to pinkish-violet,
2–3-seriate, connate into basal ring. x = 9. Circa
30 species, south-central Europe, Russia, Iran and
India to western China.
Annual herbs, sparingly branched; stems erect,
pilose-hirsute. Leaves mostly cauline, glabrous to
strigose, eglandular, lower leaves petiolate, blades
coarsely toothed, upper leaves reduced, sessile,
spathulate. Inflorescences 1-headed or corymbose
to thyrsoid with few heads; involucres hemispheric;
bracts 2–3(–5)-seriate, leaf-like, long-ciliate proximally, inner 1–2 series membranous-scarious,
often white, broadest distally; receptacles planoconvex. Rays 1–2-seriate, pistillate, bluish, reddish
or white. Disc florets yellow, tube short, limbs
abruptly expanded, lobes deltate-lanceolate, erect
to spreading; style appendages broadly lanceolate.
Achenes compressed, with thickened marginal
nerves, strigose-sericeous, eglandular; inner pappus bristles caducous, outer series of short bristles
or squamae. x = 9. One species, C. chinensis (L.)
Nees, northern China, also cultivated.
877. Crinitaria Cass.
Crinitaria Cass., Dict. Sci. Nat. 37: 460 (1825); Tzvelev, Fl.
U.R.S.S. 25: 173–183 (1959), Engl. transl. 163–172 (1999),
reg. rev.
Linosyris Cass. (1825), nom. illegit. non Linosyris C.G.
Ludw. (1757).
Pseudolinosyris Novopokr. (1918).
Biennial herbs to subshrubs; glabrous to scabrous
or gland-dotted, often tomentose in inflorescence;
stems branched from base or at tip. Leaves cauline,
sessile, narrow with 1 prominent nerve, margins
often involute. Inflorescences corymbose to
1-headed; involucres cylindric to turbinate; bracts
3–5-seriate, subequal to gradate, herbaceous to
subcoriaceous, oblong to linear-subulate, margins
partly scarious; receptacles convex to elongate.
Rays lacking. Florets 4–40, yellow or partly pink,
tubes slender, limbs narrowly campanulate, lobes
(4–)5, lanceolate; style bases sometimes bulbous;
style appendages lanceolate. Achenes obovate
to linear, sericeous-villous or gland-dotted, ribs
obscure, often 1 or 2 on sides; pappus bristles
whitish to pinkish, 2(–3)-seriate, sometimes
basally connate. x = 9. Thirteen species, Eurasia.
878. Galatella Cass.
Galatella Cass., Dict. Sci. Nat. 37: 463, 488 (1825); Tzvelev,
Fl. U.R.S.S. 25: 138–172 (1959), Engl. transl. 128–161 (1999),
reg. rev.
879. Heteropappus Less.
Heteropappus Less., Syn. Gen. Comp. 189 (1832).
Short-lived herbs; puberulous to hispid; stems
branching at base or in inflorescence. Leaves
herbaceous, sessile to pseudopetiolate, larger and
non-persistent in rosette, cauline leaves linear to
oblong or obovate, 1- or weakly 3-nerved, margins
entire or remotely serrate. Inflorescences 1-headed
or corymbose; involucres obconic to rotate; bracts
2–3-seriate, subequal, herbaceous, lanceolate
to linear-lanceolate, margins scarcely scarious;
receptacles plano-convex. Rays 1-seriate, pistillate,
white, blue or purplish, coiled. Disc florets yellow,
narrowly funnelform, lobes triangular, erect,
often zygomorphic; style appendages triangular.
Achenes obovate, compressed, 2-ribbed, sericeous;
pappus bristles sordid to reddish, often with outer
scales, pappus sometimes reduced in rays. x = 9.
Circa 20 species, central and eastern Asia.
880. Kalimeris (Cass.) Cass.
Kalimeris (Cass.) Cass., Dict. Sci. Nat. 37: 464 (1825); Gu &
Hoch, Ann. Missouri Bot. Gard. 84: 762–814 (1997).
Aster L. subg. Kalimeris Cass. (1822).
Perennial taprooted herbs; rhizomatous; stems
branched above. Basal leaves petiolate, nonpersistent, upper leaves reduced, lanceolate to
oblanceolate or linear, entire, coarsely toothed or
lobed, glabrous to strigose, sometimes glandular.
Compositae
Inflorescences 1-headed on leafy peduncles or
corymbose; involucres hemispheric to campanulate; bracts 2–3-seriate, gradate, lanceolate to
obovate, pale-indurate at base, distally greenherbaceous; receptacles convex to subconical. Rays
10–30, pistillate, white to pinkish or bluish, limbs
short, not coiling. Disc florets yellow, tubes short,
limbs campanulate, lobes lanceolate. Achenes
obovoid, compressed, with 2(–4) nerves, glandular
and often strigose; pappus bristles short, 1-seriate,
sometimes reddish. x = 9. Eight species, eastern
Asia, also cultivated.
881. Kemulariella Tamamschyan
Kemulariella Tamamschyan in Komarov, Fl. U.R.S.S. 25: 580
(1959); Tamamschyan, Fl. U.R.S.S. 25: 110–118 (1959), Engl.
transl. 25: 104–111 (1999), reg. rev.
Perennial herbs or subshrubs; stems 1 or many
from nodular rhizomes, glabrous below or with
articulate to hooked hairs. Leaves sessile, ovate to
linear-lanceolate, entire or toothed. Heads usually
solitary, pedunculate; involucres broadly campanulate to hemispheric; bracts 2–3-seriate, subequal to
weakly gradate, herbaceous, oblong-lanceolate to
linear; receptacles convex, conical or hemispheric,
alveole margins narrowly membranaceous. Rays 1–
2-seriate, pistillate, limbs purple or pinkish. Disc
florets yellow, narrowly funnelform, lobes triangular. Achenes oblong, not or slightly compressed,
angular with 3 narrow ribs, appressed-setulose; inner pappus bristles 1-seriate, scales or short flat
bristles outside. x = 9. Six species, Caucasus.
882. Miyamayomena Kitam.
Miyamayomena Kitam., Acta Phytotax. Geobot. 33: 409
(1982).
Gymnaster Kitam. (1937), nom. illegit. non Gymnaster
F. Schütt (1891).
Erect perennial herbs; rhizomes short; stems
branched at top. Leaves glabrous or with short,
coarse hairs, basal leaves with winged petioles, oblong to ovate-oblong, margins coarsely
incised-toothed, cauline leaves sparse, smaller.
Inflorescences loosely corymbose; heads longpedunculate; involucres subglobose to campanulate; bracts 2–3-seriate, subequal, lanceolate;
receptacles conical. Rays 1–2-seriate, pistillate,
limbs blue or white, coiling. Disc florets numerous, yellow, lobes 5; style appendages deltate to
lanceolate, hairy. Achenes oblong, compressed,
319
glabrous, margins ribbed; pappus lacking. x = 9.
Five species, Japan.
883. Psychrogeton Boiss.
Psychrogeton Boiss., Fl. Orient. 3: 156 (1875); Grierson,
Notes Roy. Bot. Gard. Edinburgh 27: 101–147 (1967), rev.
Perennial to biennial herbs; stems usually
clustered, from caudex or rhizomes; hispid to
tomentose. Leaves in basal rosettes or cauline,
herbaceous, lanceolate or oblanceolate to rounded,
entire to pinnatifid. Inflorescences corymbose
or with erect peduncles; involucres obconical to
rotate; bracts c. 3-seriate, subequal, lanceolate to
linear-lanceolate, margins rather broadly scarious,
green patch narrow; receptacles plano-convex.
Pistillate florets 1–2-seriate, white to pink or purple, tubular or with short erect limb. Disc florets
functionally male, rarely perfect, yellow, narrowly
funnelform, lobes somewhat unequal, erect.
Achenes obovate or oblanceolate, compressed,
2-nervate, sericeous; pappus bristles whitish.
x = 9. Twenty species, central Asia, western China.
884. Rhinactinidia Novopokr.
Rhinactinidia Novopokr., Trudy Bot. Inst. Akad. Nauk
S.S.S.R. ser. 1, Fl. Sist. Vyssh. Rast. 7: 114, 134 (1948);
Tamamschyan, Fl. U.R.S.S. 25: 129–135 (1959), Engl. transl.
121–125 (1999), reg. rev.
Rhinactina Less. (1832), non Rhinactina Willd. (1807).
Krylovia Schischk. (1949).
Borkonstia Ignatov (1983).
Erect perennial herbs, from branching caudex; puberulous to pilose. Leaves mostly in basal rosette,
herbaceous, base cuneate or petiolate, blade
obovate-oblanceolate, entire to remotely toothed.
Inflorescences 1-headed to loosely corymbose;
involucres campanulate to hemispheric; bracts
3–4-seriate, gradate, ovate to oblong-lanceolate,
mostly indurate, green distally, tips short-acute,
short-fringed, margins narrowly scarious; receptacles plano-convex, smooth. Rays 1-seriate,
pistillate, pale purple, belatedly coiling. Disc florets
yellow or purplish, narrowly funnelform, lobes 5,
oblong-triangular, somewhat zygomorphic, erect.
Achenes ellipsoid-oblong, compressed, 2-nerved,
sericeous, gland-dotted; pappus bristles whitish,
inner apically broadened, outer shorter. Four
species, central Asia.
885. Tripolium Nees
Tripolium Nees, Gen. Sp. Aster. 152 (1832).
320
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Biennials to short-lived perennials, from fibrousrooted rhizome or caudex, glabrous. Leaves
subsucculent, basal leaves oblanceolate-spathulate,
usually 3-nerved, petiole as long as blade;
cauline leaves sessile. Inflorescences loosely
corymbose, usually 5–20-headed; involucres
campanulate; bracts (1–)2–3(–4)-seriate, gradate, lanceolate to oblong, obtuse to rounded,
evenly thin-herbaceous, 3–5-nerved; receptacles
plano-convex. Rays 10–30, sometimes lacking,
pistillate, blue to purple or white. Disc florets
tubular-funnelform, lobes lanceolate. Achene
faces strigose, sometimes gland-dotted; pappus
1–2-seriate, bristles subequal, accrescent after
anthesis to 11–12 mm, sometimes caducous. x = 9.
One species, T. pannonicum (Jacq.) Dobrocz.,
Eurasia, halophilic, widely introduced.
XV.10. Subtribe Solidagininae O. Hoffm.
(1890).
Herbs or shrubs. Leaves alternate, mostly entire,
less often serrate, with glandular dots, stipitate
glands, or eglandular. Inflorescences corymbose
to thyrsoid, sometimes with secund branches, or
heads solitary; involucral bracts flat, usually basally
indurate with herbaceous apical patch; receptacles
with or without pales. Ray florets 1-seriate or lacking; limbs yellow, sometimes white, or lacking. Disc
florets bisexual, functionally male in Amphiachyris,
Amphipappus or Petradoria; style branches short
to long, with papillose collecting appendages. Achenes terete, multinerved, eglandular, setulae without anchor-shaped tips; pappus usually 1-seriate,
persistent, greatly reduced in the Gutierrezia lineage. x = 9, reduced to 8, 5, 4.
Key to the Genera
1. Pappus absent or a short corona or of minute scales;
x = 5 or 4 (or 8)
2
– Pappus of barbellate bristles or (in some shrubs) of
scales; x = 9
5
2. Shrubs; ray florets barely exceeding the involucre;
x = 4 or 8
898. Gymnosperma
– Herbs or low subshrubs; ray florets conspicuously exceeding the involucre; x = 5
3
3. Disc florets functionally male; disc pappus of 5–8
basally united, linear-spathulate scales
887. Amphiachyris
– Disc florets bisexual; disc pappus absent, a low corona,
or a ring of lanceolate scales
4
4. Heads terminal; ray florets present, disc florets usually more than 3; pappus 1–2-seriate, of stiff scales,
united or reduced in some species, usually longer in
disc
897. Gutierrezia
– Heads axillary; ray florets absent, disc florets 3, white;
pappus 2-seriate, of apiculate scales c. as long as the
corollas
908. Thurovia
5. Shrubs
6
– Herbs or subshrubs
18
6. Pappus of scales, thick and flattened bristles or of short
bristles of dissimilar length, often different on ray and
disc achenes
7
– Pappus of bristles, similar on ray and disc achenes 10
7. Plants spinescent; heads in a compact corymbose cluster; disc florets functionally male; ray pappus of 15–20
flattened, paleate bristles c. 1 mm long, disc pappus of
tortuous-twisted bristles c. 3 mm long
888. Amphipappus
– Plants not spinescent; heads solitary or few in a loose
corymbose cluster; disc florets bisexual; pappus of
awns or scales
8
8. Leaves linear-lanceolate, 3-nerved, resinous-viscid, arcuate; heads discoid, eradiate; pappus of paleate awns
909. Vanclevea
– Leaves linear to oblanceolate or spathulate, 1-nerved,
not resinous-viscid or arcuate; heads discoid or radiate; pappus various
9
9. Heads radiate; involucral bracts ovate; receptacles
epaleate; pappus 1–2-seriate, of thick, broadly
flattened, barbellate, bristle-like scales
886. Acamptopappus
– Heads discoid; involucral bracts oblanceolate; receptacles paleate; pappus 1-seriate, of 5–8 flattened, linearlanceolate scales
894. Eastwoodia
10. Disc corollas regular to slightly irregular; pappus 1–3seriate, bristles sometimes apically dilated
11
– Disc corollas strongly zygomorphic; pappus 1-seriate,
bristles apically attenuate
14
11. Rays present; involucral bracts not ranked; eastern
USA and Caribbean Islands
12
– Rays often absent, when present yellow; involucral
bracts ranked or not ranked; western USA and northwestern Mexico
13
12. Rays yellow; achenes with 8–10 whitish raised nerves;
pappus 2–3-seriate; areoles of leaves subisodiametric
with recessed resinous borders; eastern USA
891. Chrysoma
– Rays white; achenes 5-nerved; pappus 1-seriate; leaf
surfaces punctate-resinous; Caribbean Islands
896. Gundlachia p.p.
13. Pappus bristles apically dilated or attenuate; heads in
compact corymbose clusters; rays present or absent;
involucral bracts in distinct vertical ranks
892. Chrysothamnus
– Pappus bristles apically dilated; heads sessile to shortpedicellate, 2–4 in glomerules, the aggregation loosely
corymbose; rays lacking; involucral bracts not ranked
899. Hesperodoria
14. Leaves hirtellous, not resinous; stems densely foliate
with acicular leaves and axillary clusters; without ray
florets
890. Chihuahuana
– Leaves not hirtellous, resinous, usually punctate; without many axillary clusters of leaves; with or without
ray florets
15
15. Broom-like shrubs with dense, erect, virgate branches;
branches strongly ridged; leaves widely spaced on new
growth, rather erect, not persisting below
910. Xylovirgata
Compositae
– Shrubs unbranched or with many ascending branches,
not broom-like; branches terete to weakly ridged;
leaves rather persistent, rather evenly and closely
spaced
16
16. Leaves involute but appearing terete; without or with
1–3 ray florets
896. Gundlachia p.p.
– Leaves narrow but distinctly flattened; ray florets 5–15
17
17. Involucral bracts weakly gradate; inflorescences
corymbose; stems minutely papillate-scabrous
900. Medranoa
– Involucral bracts strongly gradate; inflorescences
loosely thysiform; stems glabrous 901. Neonesomia
18. Rays white
19
– Rays yellow or absent
20
19. Ray corollas white, disc corollas yellow; achenes glabrous; involucral bracts not keeled, with 3–7 parallel,
longitudinal veins, without an apical patch
902. Oligoneuron p.p.
– Ray and disc corollas white; achenes densely strigosesericeous; involucral bracts keeled, with a single midvein, with a green, sharply delimited apical patch
905. Sericocarpus
20. Heads solitary on leafless or bracteate stems
907. Stenotus
– Heads in corymbose, racemose (Columbiadoria), thyrsoid or secund (Solidago) clusters
21
21. Plants taprooted
22
– Plants fibrous-rooted from a rhizome or caudex 23
22. Leaves net-veined, oblanceolate; inflorescence
branches racemose or weakly corymbose; disc florets
bisexual; pappus bristles in 1 series
893. Columbiadoria
– Leaves with 3–5 raised, parallel veins; inflorescences
distinctly corymbose; disc florets functionally male;
pappus bristles in 2–3 series
904. Petradoria
23. Leaves resinous or punctate or both, but not stipitateglandular; involucral bracts usually indurate at least
below, commonly with a sharply delimited, green apical patch
903. Oreochrysum
– Leaves stipitate-glandular; involucral bracts primarily
herbaceous and even-textured
24
24. Leaves linear to narrowly oblanceolate, distinctly
punctate; disc corollas not orange-veined
25
– Linear mostly broader, punctate (Oligoneuron) or not
(Solidago); disc corollas with orange-resinous veins 26
25. Basal leaves persistent, cauline leaves reduced; heads
discoid, rays absent; involucral bracts in vertical ranks
889. Bigelowia
– Leaves all cauline; heads radiate; rays 15–25; involucral
bracts not in vertical ranks
895. Euthamia
26. Inflorescence strongly corymbose; disc corolla lobes
erect; pappus bristles in (1–)2 series
902. Oligoneuron p.p.
– Inflorescence secund to thyrsoid, rarely weakly
corymbose; disc corolla lobes reflexing-coiling;
pappus bristles in 1 series
906. Solidago
Genera of Solidagininae
886. Acamptopappus (A. Gray) A. Gray
Acamptopappus (A. Gray) A. Gray, Proc. Amer. Acad. Arts
8: 634 (1873); Lane, Madroño 35: 247–265 (1988), rev.
321
Aplopappus sect. Acamptopappus A. Gray (1849).
Taprooted shrubs; stems decumbent, becoming
spinescent, with bark shredding. Leaves lanceolate to obovate, 1-nerved, entire, non-resinous.
Inflorescences 1-headed or loosely corymbose;
involucres broad; bracts c. 3-seriate, broad with
broad laciniate-hyaline margins, base yellowish,
tips green, reflexed. Rays 0 or 5–14, pistillate,
yellow. Disc florets perfect, yellow, funnelform,
lobes lanceolate, reflexing; style appendages
triangular-lanceolate. Achenes obconical, densely
sericeous, with mixed appressed-tortuous and
longer ascending-spreading setulae; pappus
1–2-seriate, of subequal awned scales, attenuate
in ray achenes, spathulate in disc achenes, rarely
also with few short setae. x = 9. Two species,
south-western USA.
887. Amphiachyris (DC.) Nutt.
Amphiachyris (DC.) Nutt., Trans. Amer. Philos. Soc. 2, 7:
313 (1840); Lane, Syst. Bot. 4: 178–189 (1979), rev.
Brachyris Nutt. sect. Amphiachyris DC. (1836).
Taprooted annual herbs; glabrous. Leaves linear to
lanceolate, 1(–3)-nerved, punctate, abaxially with
glandular hairs. Inflorescences corymbose to thyrsoid; heads pedunculate; involucres narrowly campanulate to turbinate; bracts 1–2(–3)-seriate, gradate, 1-nerved, indurated and whitish-glutinous
basally, nerve not green-margined, margins hyaline; receptacles glabrous or pilosulous. Rays 7–
12, pistillate, yellow or orange-tinged. Disc florets functionally male, yellow, tube short, lobes
deltate, erect. Ray achenes obovate, plump, 4–9ribbed, strigose-sericeous, setulae slightly clavate
at tips; ray pappus a low crown or ring of scales; disc
pappus of 5–8 white, basally united, linear scales
with spathulate tips. x = 4, 5. Two species, southcentral USA.
888. Amphipappus Torr. & A. Gray ex A. Gray
Amphipappus Torr. & A. Gray ex A. Gray, Boston J. Nat.
Hist. 5: 107 (1845); Porter, Amer. J. Bot. 30: 481–483 (1943),
rev.
Shrubs, with branchlets and leaves glabrous, becoming leafless and spinescent; bark pale. Leaves
short-petiolate, obovate to narrowly elliptic, entire.
Inflorescences corymbose; involucres turbinatecylindric; bracts 7–12, c. 3-seriate, gradate, ovate
to elliptic, outer bracts keeled. Rays (0–)1–2,
pistillate, yellow, slightly exceeding involucre. Disc
322
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
florets 3–7, functionally male, yellow, narrowly
funnelform, lobes lanceolate, reflexing. Fertile
achenes oblong-elliptic, somewhat compressed,
villous; pappus bristles 15–20, in 1 series, slender,
flattened, paleaeform, basally fused; disc achenes
shorter; pappus tortuous-twisted. x = 9. One
species, A. fremontii Torr. & A. Gray ex A. Gray,
south-western USA.
889. Bigelowia DC.
Bigelowia DC., Prodr. 5: 326 (1836), nom. cons.; Anderson,
Sida 3: 451–465 (1970), rev.; Syst. Bot. 2: 209–218 (1972),
rev.
Subshrubs from short caudex, with short rhizomes; glabrous, initially viscid; stems nearly
scapiform. Leaves in persistent basal rosette,
sessile, linear to oblanceolate, 3-nerved, entire,
gland-dotted, cauline leaves reduced to linear
bracteoles. Inflorescences densely flat-corymbose;
involucres turbinate-cylindric; bracts 3–4-seriate,
unequal, more or less ranked, narrowly lanceolate
to nearly linear, weakly keeled, green-tipped.
Rays absent. Florets 2–6, perfect, yellow, tubular,
lobes lanceolate, recurving; style appendages
lanceolate. Achenes turbinate to subcylindric,
slightly compressed to subquadrate, with 4 or 5
thin resinous nerves, sparsely strigose; pappus
1(–2)-seriate, with few bristles of uneven lengths.
x = 9. Two species, south-eastern USA.
890. Chihuahuana Urbatsch & R.P. Roberts
Chihuahuana Urbatsch & R.P. Roberts, Sida 21, 1: 245
(2004).
Densely branched shrubs. Leaves closely persistent
and in axillary clusters, erect, 3–10 mm long,
sessile, acicular, hirtellous with uniseriate conic
hairs, bases rather decurrent, apices sharply acute,
in section narrowly triangular with thickened
midrib composed mostly of bundle of fibers. Heads
solitary; involucre campanulate; bracts gradate, to
5–6 mm long. Rays lacking. Florets 8–10; corollas
yellow, hairy, lobes ovate-lanceolate, unequal.
Achenes sericeous, pappus of bristles. x = 9. One
species, C. purpusii (Brandegee) Urbatsch & R.P.
Roberts, northern Mexico.
891. Chrysoma Nutt.
Chrysoma Nutt., J. Acad. Nat. Sci. Philadelphia 7: 67 (1834);
Cronquist, Vasc. Fl. SE U.S., Asteraceae 1: 134 (1980), reg.
rev.
Evergreen shrubs; glabrous, glutinous. Leaves
sessile to shortly petiolate, oblanceolate to
narrowly elliptic, entire, 1-nerved, veinlets in
recessed reticulum. Inflorescences cymose; heads
sessile to subsessile; involucres cylindric; bracts
3–4(–5)-seriate, gradate, ranked, lanceolate,
stramineous, with orange-resinous midvein
from base to apex. Rays (0–)1–2(–3), pistillate,
yellow. Disc florets (2–)3–4(–5), perfect, yellow,
elongate-tubular, lobes lanceolate, spreading to
recurving; style appendages linear-triangular.
Achenes turbinate-oblong, with 8–10 raised pale
nerves, densely strigose-sericeous; pappus bristles
2–3-seriate, of uneven lengths. x = 9. One species,
C. pauciflosculosa (Michx) Greene, south-eastern
USA.
892. Chrysothamnus Nutt.
Chrysothamnus Nutt., Trans. Amer. Philos. Soc. 2, 7: 323
(1840), nom. cons. prop.; Nesom & Baird, Phytologia 75:
74–93 (1993), emend.
Suffrutescent herbs or rounded shrubs, muchbranched; stems often resinous. Leaves linear to
spathulate-oblanceolate, 3-nerved, gland-dotted,
often varnished, glabrous to variously pubescent.
Inflorescences corymbose; involucres turbinatecylindric; bracts 4–5-seriate, gradate, ranked,
ovate-lanceolate to triangular-lanceolate, with
golden-resinous midvein, usually with apical
green patch. Rays few or none, pistillate, yellow. Disc florets 4 or 5, perfect, yellow, tubular,
gradually or abruptly broadened into limb; style
appendages linear. Achenes narrowly ellipsoid, 5–
10(–15)-nerved, subterete, glabrous to sericeous,
sometimes glandular near apex; pappus bristles
2–3-seriate, of unequal lengths, tips sometimes
broadened. x = 9. Circa 13 species, south-western
Canada, western USA, north-central Mexico.
893. Columbiadoria G.L. Nesom
Columbiadoria G.L. Nesom, Phytologia 71: 249 (1991).
Perennial taprooted subshrubs; stems strict.
Leaves sessile, eglandular or obscurely punctuateglandular, not resinous, sparsely scabroushispidulous, oblanceolate, entire, net-veined,
reduced upwards, basal leaves usually not persistent. Inflorescence racemose to loosely corymbose,
with 5–15+ heads; involucres cylindric-turbinate,
bracts 5–6-seriate, gradate, lanceolate, basal 2/3
white-indurate, distal half not or slightly keeled,
apex green. Rays c. 5 or 8, pistillate, yellow, not or
Compositae
scarcely coiling. Disc florets perfect, yellow, lobes
deltate, erect; style appendages linear-lanceolate,
much longer than stigmatic lines. Achenes narrowly oblong, slightly compressed, 8-nerved,
moderately strigose; pappus bristles 1-seriate,
apices attenuate. One species, C. hallii (A. Gray)
G.L. Nesom, north-western USA.
894. Eastwoodia Brandegee
Eastwoodia Brandegee, Zoe 4: 397 (1894); Lane, Madroño
35: 247–265 (1988), rev.
Glabrous, usually glutinous shrubs; older stems
with white shedding bark. Leaves not reduced
above, sometimes in axillary fascicles, yellowgreen, linear to oblanceolate, 1-nerved, margins
entire, sparsely ciliate. Inflorescences 1-headed
to corymbose; involucres hemispheric to campanulate, impressed at base; bracts 3–5-seriate,
gradate, stiffly erect, resiniferous, oblanceolate,
indurate below, distal half green, margins narrowly
scarious; receptacles with oblanceolate, caducous
scales. Rays absent. Florets 30–40, perfect, yellow,
cylindric to funnel-shaped, lobes lanceolate,
spreading or reflexing; style appendages lanceolate. Achenes narrowly obconical, 3–4-angled,
strigose-sericeous especially on angles; pappus
of 5–8 linear-lanceolate scales as long as corollas.
x = 9. One species, E. elegans Brandegee, USA
(California).
323
896. Gundlachia A. Gray
Gundlachia A. Gray, Proc. Amer. Acad. Arts 16: 100 (1880);
Lane, Brittonia 48: 532–541 (1996), rev.
Xylothamia G.L. Nesom, Y.B. Suh, D.R. Morgan & B.B.
Simpson (1990).
Evergreen, resinous, punctate shrubs. Stems
non- to many-branched. Leaves usually evenly
spaced, spreading to appressed, sessile to shortpetiolate, linear to obovate or spathulate, flat to
involute-terete. Inflorescences terminal, racemose
to corymbose or thyrsoid; heads 1–5 in clusters or
hidden by leaves; involucres cylindric to narrowly
obconic; bracts 2–5-seriate, linear-lanceolate to
ovate, with apical resin pocket, bases whiteindurate, margins translucent. Rays 3–8 or lacking,
pistillate, corollas white to yellow, sometimes
drying purplish, 1–6.5 mm long. Disc florets 3–50,
white to yellow, lobes slightly irregular, laxly
recurved; style appendages linear-lanceolate to
ovate. Achenes turbinate to cylindric, sparsely
pilose to sericeous; pappus bristles c. 40, subequal.
x = 9. Six species, Caribbean, Mexico, USA (Texas).
Revisions (Urbatsch and Roberts 2004) based
on DNA studies (Urbatsch et al. 2003; Roberts and
Urbatsch 2004) greatly expand Gundlachia to include the type of Xylothamnia Nesom et al., but
exclude most of the latter genus (see Chihuahuana
Urbatsch & R.P. Roberts, Medranoa Urbatsch & R.P.
Roberts, Neonesomia Urbatsch & R.P. Roberts and
Xylovirgata Urbatsch & R.P. Roberts).
895. Euthamia Nutt. ex Cass.
897. Gutierrezia Lag.
Euthamia Nutt. ex Cass., Dict. Sci. Nat. 37: 471 (1825);
Sieren, Rhodora 83: 551–579 (1981), rev.
Gutierrezia Lag., Gen. Sp. Pl. [Elench. Hort. Matriti] 30
(1816); Solbrig, Contr. Gray Herb. 197: 3–42 (1966), rev.;
Lane, Syst. Bot. 10: 7–28 (1985), rev.
Greenella A. Gray (1880).
Perennial herbs, rhizomatous; glabrous to
scabridulous. Leaves cauline, sessile, linear to
linear-lanceolate,
1–3(–5)-veined,
punctate,
rather glutinous. Inflorescence flat- or roundedcorymbose, with glomerate subclusters of heads;
involucres cylindric to turbinate or hemispheric;
bracts 3–5-seriate, gradate, ovate to oblong,
obtuse, mostly chartaceous, glutinous, somewhat
fimbrillate; receptacles sometimes pilosulous.
Rays 15–25, pistillate, yellow. Disc florets (2–)7–
15(–20), perfect, yellow, expanded-tubular, lobes
lanceolate, spreading; style appendages lanceolate.
Achenes oblong to narrowly turbinate, subterete,
faintly 2–4-nerved, strigose; pappus bristles
1-seriate. x = 9. Eight species, mostly central and
south-eastern USA, one south-western Canada to
Baja California Norte.
Annual or perennial, taprooted herbs or small
shrubs. Leaves decurrent, linear to lanceolate
or spathulate, punctate and glutinous, glabrous,
entire. Heads solitary or 3–6 in clusters; involucres
cylindric to campanulate; bracts 2–4-seriate,
gradate, stramineous, 1- or 2-nerved, bases
white-indurate; receptacles flat to columnar, with
uncinate hairs. Rays (0–)1–30, pistillate, yellow
or white, coiling. Disc florets perfect, yellow
or white, lobes deltate, erect; style appendages
linear-lanceolate. Achenes tan to purplish-black,
clavate to cylindric, multinerved, setulose between
ribs, tips of setulae divergent or clavate to bulbous;
pappus 1–2-seriate, of stiff scales, fused or reduced
in some species. x = 4. Sixteen species, mostly
324
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Mexico, also south-western USA, south-western
South America.
898. Gymnosperma Less.
Gymnosperma Less., Syn. Gen. Comp. 194 (1832), nom.
nov.
Selloa Spreng. (1819), nom. illegit. non Selloa Kunth (1818).
Glabrous, glutinous shrubs. Leaves sessile, decurrent, linear to narrowly lanceolate, oblanceolate
or elliptic, uniform up to inflorescence, entire, 1or 3-nerved, punctate. Inflorescences corymbose,
containing compact glomerules of sessile to
subsessile heads; involucres turbinate-cylindric
to elliptic-obovoid; bracts 2–4-seriate, gradate,
mostly white-indurate, little or no green at apices.
Rays 4–9, pistillate, yellow. Disc florets 4–6, perfect
or functionally male, orange-yellow, tubularfunnelform, lobes lanceolate, reflexing-coiling;
style appendages lanceolate. Achenes columnar
or fusiform to compressed, strigillose, without
evident ribs; pappus near-absent or a minute
corona. x = 4 or 8. One species, G. glutinosa
(Spreng.) Less., south-western USA, south to
Guatemala.
899. Hesperodoria Greene
Hesperodoria Greene, Leafl. Bot. Observ. Crit. 1: 173 (1906);
Hall, Publ. Carnegie Inst. Wash. 389: 35, 218–222 (1928),
rev.
Perennial herbs or shrubs; stems glabrous to
hirtellous, white-barked with age. Leaves thick,
linear-lanceolate to oblanceolate, with 3 parallel
nerves, entire, surfaces glabrous to scabrous, one
species punctate and resinous. Inflorescences
loosely corymbose, with 2–4-headed glomerules
or cymose clusters; involucres turbinate to
cylindric-turbinate; bracts 5–6-seriate, subequal
to gradate, not ranked, oblong-lanceolate, tips
greenish, with or without glandular midline. Ray
florets absent. Florets c. 10–20, perfect, yellow,
tubular-funnelform, lobes deeply cut, triangular; style appendages linear-lanceolate. Achenes
subterete, 5–8-nerved, sparsely sericeous; pappus
1–2-seriate, bristles persistent, apically broadened.
x = 9. Two species, USA (Utah, northern Arizona).
900. Medranoa Urbatsch & R.P. Roberts
Medranoa Urbatsch & R.P. Roberts, Sida 21:254 (2004).
Shrubs to 2 m, resinous; branches numerous,
ascending, mostly terete, bark smooth; internodes
1–4 mm long, green, scabridulous. Leaves evergreen, sessile, punctate, narrowly elliptic to ovate,
flat-canaliculate, base decurrent. Inflorescences
terminal, 1-headed or corymbose; involucral bracts
2–3-seriate, weakly gradate, mostly chartaceous.
Rays 5–11; ligules elliptic, 3–7 mm long. Disc
florets 15–22; corollas pale yellow, 4–5 mm long,
lobes spreading or recurved, unequal, 0.8–2.2 mm
long, style branches thickened with doubled veins,
appendages narrowly linear. Achenes turbinate,
pilose; pappus bristles c. 80, setose, subequal.
x = 9. One species, M. parrasana (S.F. Blake)
Urbatsch & R.P. Roberts, Mexico (Coahuila and
Zacatecas).
901. Neonesomia Urbatsch & R.P. Roberts
Neonesomia Urbatsch & R.P. Roberts, Sida 21: 252 (2004).
Shrubs to 3 m, much-branched, aromatic; twigs
ridged, internodes 2–10 mm long. Leaves linear,
elliptic to narrowly oblanceolate, flat with midrib
raised abaxially, with flagelliform hairs, margins
entire to minutely ciliate. Heads terminal, solitary
or loosely paniculate; involucre usually turbinate;
bracts gradate, linear, blunt to subacute, thick,
resinous, stramineous with apical glandular patch.
Rays 5–15; corollas white to yellow, 2–5 mm long.
Disc florets 8–20; corollas 3.5–5 mm long, tube
poorly differentiated from asymmetric 5-lobed
limb, lobes 0.9–3.3 mm long. Achenes strigose to
sericeous; pappus of setose bristles. x = 9. Two
species, Mexico, USA (Texas).
902. Oligoneuron Small
Oligoneuron Small, Fl. SE U.S., ed. 1, 1188 (1903); Nesom,
Phytologia 75: 1–44 (1993), emend.
Unamia Greene (1903).
Erect perennial herbs from short rhizomes; stems
usually unbranched below inflorescence. Leaves
narrowly lanceolate or broadly elliptic, parallelor net-veined, entire or serrulate, often glanddotted, basal leaves often persistent. Inflorescences
corymbose, flat-topped; involucres turbinate to
campanulate or hemispheric; bracts 2–6-seriate,
gradate, oblong to obovate, with 3–7+ translucent
veins. Rays pistillate, yellow to white. Disc florets
perfect, yellow, limbs narrowly campanulate, lobes
narrowly triangular, spreading; style appendages
lanceolate to linear. Achenes oblong-fusiform,
terete to slightly compressed, glabrous, with
5–7(–10–20) thin longitudinal subsurface nerves;
pappus (1–)2-seriate, bristles equal to subequal,
Compositae
apically attenuate or clavate. x = 9. Six species,
eastern North America.
Significant features of Oligoneuron, particularly the flat-topped inflorescence and gland-dotted
leaves, are shared with genera of Solidagininae
other than Solidago L. (Nesom 1993), and species of
Oligoneuron do not hybridize with those of typical
Solidago, within which hybrids in many combinations are easily formed. Recent molecular evidence
(Beck et al. 2004), however, indicates that these six
species probably arose evolutionarily from within
typical Solidago, where they may be better treated.
903. Oreochrysum Rydb.
Oreochrysum Rydb., Bull. Torrey Bot. Club 33: 152 (1906).
Viscid perennial herbs; rhizomes slender, with
scale leaves; stems puberulous or hirtellous and
with stipitate glands. Basal and lower cauline leaves
petiolate, persistent, spathulate to oblanceolate,
upper cauline leaves slightly reduced, usually subclasping, broadly ovate-lanceolate to oblanceolate.
Inflorescences densely corymbose, flat-topped;
involucres campanulate to hemispheric; bracts
3–4-seriate, subequal, herbaceous, outer ovate,
inner broadly lanceolate to oblong, with 1 (or 3)
non-resinous veins. Rays 12–20, pistillate, yellow.
Disc florets perfect, narrowly tubular-funnelform,
lobes triangular, spreading; style appendages linear. Achenes fusiform, compressed, glabrous, with
12–16 raised whitish nerves; pappus 2(–3)-seriate,
bristles equal. x = 9. One species, O. parryi (A.
Gray) Rydb., western USA, north-western Mexico.
904. Petradoria Greene
Petradoria Greene, Erythea 3: 13 (1895); Anderson, Trans.
Kansas Acad. Sci. 66: 632–684 (1963), rev.
Suffrutescent herbs from stout taproot or branching caudex; gland-dotted, resinous. Leaves
persistent in rosette, little reduced upwards,
coriaceous, linear to lanceolate or oblanceolate,
margins entire to scabrous, with 3–5 parallel veins,
venation prominulous. Inflorescences densely
paniculate-corymbose; involucres cylindric to
cylindric-turbinate; bracts 3–6-seriate, gradate,
usually in ranks, ovate to oblong, apices truncate
to attenuate, sometimes weakly keeled. Rays
1–3, pistillate, yellow. Disc florets (1–)2–4(–5),
functionally male, yellow, tubular-funnelform,
lobes lanceolate; style branches linear-lanceolate.
Achenes cylindric to slightly compressed, 6–9nerved, glabrous; pappus 2-seriate, of attenuate
325
bristles. x = 9. One species, P. pumila (Nutt.)
Greene, south-western USA.
905. Sericocarpus Nees
Sericocarpus Nees, Gen. Sp. Aster. 148 (1832) [1833];
Nesom, Phytologia 75: 45–54 (1993), key.
Oligactis Raf. (1836 ) [1837], nom. illegit. non Oligactis
(Kunth) Cass. (1825).
Perennial herbs from rhizomes; puberulous to glabrous, gland-dotted. Leaves basal and cauline, sessile, linear to oblanceolate or elliptic-oblanceolate.
Inflorescences corymbose, flat-topped, with glomerate clusters of heads; involucres cylindric to
turbinate; bracts 2–6-seriate, gradate, coriaceous,
keeled, gibbous and whitish-indurate below, with
green patch distally. Rays 3–8, pistillate, white,
not coiling. Disc florets perfect, white or creamcoloured, funnelform; style appendages lanceolate
to linear-lanceolate. Achenes narrowly obconical
to nearly cylindric, with 4–8 partially sunken
nerves, strigose-sericeous; pappus 2–3-seriate,
of equal, apically broadened bristles. x = 9. Five
species, USA.
906. Solidago L.
Solidago L., Sp. Pl. 878 (1753); Cronquist, Vasc. Fl. SE U.S.,
Asteraceae 1: 116–133 (1980), reg. rev.; Tamamschyan, Fl.
U.R.S.S. 25: 31–50 (1959), Engl. transl. 25: 31–50 (1999),
reg. rev.; Taylor & Taylor, Sida 10: 223–251 (1984), reg. rev.
Brachychaeta Torr. & A. Gray (1842).
Brintonia Greene (1895).
Perennial herbs from rhizome or caudex, rarely
thickly taprooted. Leaves linear or linear-lanceolate
to ovate, some 3-nerved, entire to toothed, some
gland-dotted. Inflorescence cylindric to pyramidal
or racemose with secund or arcuate branches; involucres cylindric to turbinate; bracts 3–4-seriate,
gradate, with an orange-glandular midrib, often
with green patch apically; receptacles sometimes
with caducous pales. Rays pistillate, yellow to
white, usually coiling. Disc florets perfect, yellow, tubular-funnelform, lobes often deeply cut,
lanceolate; style appendages lanceolate. Achenes fusiform-cylindric, 5–8-nerved, glabrous to
strigose; pappus bristles 1-seriate, whitish. x = 9.
Circa 100 species, mostly North America, eight
Mexico, one South America, 10–20 Eurasia.
907. Stenotus Nutt.
Stenotus Nutt., Trans. Amer. Philos. Soc. 2, 7: 334 (1840).
326
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Perennial herbs from woody, branched caudices;
stems crowded, short, glabrous to tomentose,
not resinous nor gland-dotted, sometimes with
stipitate glands. Leaves mostly in rosette, linear
to linear-oblanceolate or spathulate, entire, parallel nerves usually 3. Heads solitary on mostly
leafless peduncles; involucres campanulate to
hemispheric; bracts 2–3(–4)-seriate, strongly to
weakly gradate, herbaceous with hyaline margins,
3(–5)-nerved. Rays usually present, pistillate,
yellow, broad, slightly coiling. Disc florets perfect,
yellow, gradually expanded; style appendages
lanceolate to linear-lanceolate. Achenes subterete to compressed, strigose-sericeous; pappus
bristles 1–2-seriate, subequal, caducous. x = 9.
Six or seven species, Canada, western USA,
north-western Mexico.
908. Thurovia Rose
Thurovia Rose, Contrib. U.S. Natl Herb. 3: 321 (1895).
Annual or short-lived perennial herbs; taprooted
or roots laterally perennating; stems branched
in upper half. Leaves cauline, linear, ascendingappressed, reduced upwards. Inflorescence spicate;
heads solitary in leaf axils, usually 10–20 per stem,
sessile or subsessile; involucres narrowly turbinate;
bracts 5–9-seriate, subequal, outer shorter, herbaceous, inner mostly whitish-indurate with small
herbaceous tips. Rays absent. Florets 3, perfect,
white or pale yellow, limbs expanded, lobes lanceolate, spreading-recurved; style appendages linear.
Achenes obconical, subterete, white-sericeous
with thick setulae; pappus 2-seriate, of c. 10 erose,
subulate scales as long as corollas. x = 5. One
species, T. triflora Rose, south-eastern Texas.
909. Vanclevea Greene
Vanclevea Greene, Pittonia 4: 50 (1899); Anderson &
Weberg., Great Basin Naturalist 34: 151–160 (1974), rev.
Shrub; glabrous, glutinous; stems with shedding
epidermis or whitish bark. Leaves rigid, spreading
to falcate-recurved, linear-lanceolate, entire.
Heads solitary or 2–5 in an open cyme; involucres
broadly turbinate; bracts 3–5-seriate, strongly gradate, narrowly elliptic-lanceolate, attenuate, inner
sometimes slightly squarrose at tips. Rays absent.
Florets c. 30, perfect, yellow, funnelform from
short tube, lobes deltate, erect; style appendages
narrowly triangular-lanceolate. Achenes narrowly
cylindric, subterete, c. 5–8-nerved, glutinous,
sparsely strigose; pappus of (12–)15–18 flattened
narrow segments. x = 9. One species, V. stylosa
(Eastw.) Greene, Utah, Arizona.
910. Xylovirgata Urbatsch & R.P. Roberts
Xylovirgata Urbatsch & R.P. Roberts, Sida 21: 255 (2004).
Virgately branched shrubs to 1 m; stems slender;
branches strongly ridged and angled. Leaves
present mostly on new growth, not persistent,
widely spaced, rather erect, 2–15 mm long, c. 1 mm
wide, resinous, not obviously punctate. Heads
terminal, solitary or in loose racemes; involucres
campanulate to turbinate; bracts strongly gradate,
with thickened apical glandular structure. Rays
3–6, limbs 2- or 3-toothed, 2.3–3.0 mm long, c.
1 mm wide. Disc florets 7–14; corollas yellow,
often drying purplish, 4.0–4.5 mm long, lobes 5,
unequal, 1.3–2.3 mm long. Achenes sericeous;
pappus bristles setose, c. 30, subequal. x = 9. One
species, X. pseudobaccharis (S.F. Blake) Urbatsch
& R.P. Roberts, Mexico (Coahuila).
XV.11. Subtribe Pentachaetinae G.L. Nesom
(2000).
Annual herbs, eglandular. Leaves alternate, entire,
filiform or linear to narrowly oblanceolate. Heads
solitary and long-pedunculate; involucral bracts
in 2–4 gradate series, with hyaline margins, not
keeled; receptacles plano-convex, epaleate. Ray
florets with corollas yellow, white in some Pentachaeta, strongly coiling. Disc florets with style
branch appendages linear-lanceolate, much longer
than the stigmatic lines. Achenes terete to slightly
compressed, narrowly oblong to oblanceolate in
outline, beaked in Tracyina, eglandular; setulae
without anchor-shaped tips; pappus 1(–2)-seriate,
of persistent bristles frequently in multiples of 5,
sometimes laterally dilated at the base and partially
connate, or sometimes completely lacking, or of
long scales (Rigiopappus). x = 9.
Key to the Genera
1. Achenes beaked; pappus of bristles; involucral bracts
distinctly unequal, gradate in 3–4 series
913. Tracyina
– Achenes not beaked; pappus of bristles or scales; involucral bracts in 2–3 equal or subequal series
2
2. Pappus of bristles; receptacles epaleate
911. Pentachaeta
– Pappus of long scales; receptacles paleate between ray
and disc florets
912. Rigiopappus
Compositae
Genera of Pentachaetinae
911. Pentachaeta Nutt.
Pentachaeta Nutt., Trans. Amer. Philos. Soc. 2, 7: 336
(1840); Van Horn, Univ. Calif. Publ. Bot. 65: 1–41 (1973),
rev.
Annual, taprooted herbs, obscurely haired. Leaves
narrowly oblanceolate to linear or filiform, entire,
ciliate. Heads solitary, pedunculate; involucres
campanulate; bracts in 2 equal or 3 subequal
series, herbaceous; receptacles epaleate. Pistillate
florets white, yellow or reddish, non-radiate
or radiate, coiling. Disc florets perfect, yellow,
tubular-funnelform, lobes 3 or 5, shallow, with
fringe of hairs. Achenes oblanceolate, terete to
slightly compressed, strigose; pappus 1-seriate,
3–5, c. 10 or c. 20 barbellate bristles, sometimes
broadened and partially connate at base or completely lacking. Six species, USA (California), Baja
California Norte.
912. Rigiopappus A. Gray
Rigiopappus A. Gray, Proc. Amer. Acad. Arts 6: 548 (1965);
Robinson & Brettell, Phytologia 26: 69–70 (1973), relat.;
Ornduff & Bohm, Madroño 23: 53–55 (1975), relat.
Annual, taprooted, hirsutulous herbs. Leaves
ascending to erect, linear, entire. Heads solitary
on long, leafy or bracteolate peduncles; involucres
broadly turbinate; bracts 2-seriate, subequal,
linear-oblong to lanceolate, acute, hirsutulous,
partially clasping outer achenes; receptacles
paleate between ray and disc florets. Rays 3, 5
or 8(–15), pistillate, rarely absent, yellow, often
purple-tinged. Disc florets perfect, yellow, narrowly cylindrical, lobes 2–4, deltate, with fringe
of hairs. Achenes narrowly cylindric to slightly
fusiform or compressed, strigose with apically
clavate setulae, transversely rugose; pappus of
(0–)3–5 stiff, white, subulate scales. One species,
R. leptocladus A. Gray, north-western USA.
913. Tracyina S.F. Blake
Tracyina S.F. Blake, Madroño 4: 74 (1937); Ornduff &
Bohm, Madroño 23: 53–55 (1975), relat.
Annual, taprooted herbs; glabrous or loosely
pilose below heads. Leaves cauline, ascending to
appressed, linear-oblanceolate, entire, margins
ciliate. Heads solitary on peduncles; involucres
cylindric to broadly ovoid; bracts 3–4-seriate,
distinctly unequal, linear-lanceolate, caducous;
327
receptacles epaleate. Rays 12–20, pistillate, paleor greenish-yellow, red-tinged, filiform, erect.
Disc florets 15–25, greenish-yellow, red-tinged,
filiform-tubular, lobes triangular-ovate, erect,
fringed with papillae. Achenes beaked, narrowly
cylindrical to linear-fusiform, 5-nerved, narrowed
to slender strigose beak; pappus 1–2-seriate, of
numerous, fragile, bristles. One species, T. rostrata
S.F. Blake, California.
XV.12. Subtribe Boltoniinae G.L. Nesom
(2000).
Perennial, herbs or shrubs (Chloracantha), rhizomatous, with persistently green stems and
leaves, thorny in Chloracantha, glabrous. Leaves
alternate, essentially all cauline, entire or fewtoothed. Inflorescences with heads solitary or very
loosely corymbose to thyrsoid; involucral bracts
primarily herbaceous, apically rounded to obtuse,
with three orange-resinous nerves; receptacles
convex to conical, epaleate. Rays 1-seriate, white
to slightly bluish, coiling. Disc florets bisexual,
corollas orange-veined; style branch appendages
deltate, papillose. Achenes terete and multinerved
or flattened, 2-nerved and winged (Boltonia),
setulae without anchor-shaped tips; pappus of
bristles or awns. x = 9.
Key to the Genera
1. Achenes flattened and winged; pappus of 2 lateral awns
915. Boltonia
– Achenes terete, without wings; pappus of bristles 2
2. Herbs; stems 7–20 cm tall, thornless; leaves persistent
914. Batopilasia
– Subshrubs; stems 50–250 cm tall, usually with green
thorns; leaves quickly deciduous 914. Chloracantha
Genera of Boltoniinae
914. Batopilasia G.L. Nesom & Noyes
Batopilasia G.L. Nesom & Noyes, Sida 19: 81 (2000).
Perennial, caespitose, glabrous herbs from thin
woody rhizomes and caudices; stems usually
1–2-branched. Rosette leaves persistent, sessile,
elliptic-oblanceolate, entire, 1-nerved to faintly
3-nerved; cauline leaves few, reduced to linear
bracts. Heads solitary, erect on near-naked peduncles. Involucral bracts 3–4-seriate, gradate,
thin-herbaceous with scarious margins and 1–3
filiform nerves, elliptic-oblanceolate, obtuse to
acute; receptacles plano-convex. Rays 9–18, fertile,
328
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
white, sometimes purplish when dry, coiling at
tips. Disc florets 24–29, perfect, not inflated nor indurated, lobes triangular, erect. Achenes strigose,
terete to slightly compressed, with (2–3–)4(–5)
orange nerves; pappus bristles 1-seriate, with few
outer setae. x = 9. One species, B. byei (S.D. Sundb.
& G.L. Nesom) G.L. Nesom & Noyes, north-western
Mexico.
915. Boltonia L’Hér.
Boltonia L’Hér., Sert. Angl. 27 (1788) [1789]; Cronquist,
Vasc. Fl. SE U.S., Asteraceae 1: 168–169 (1980), reg. rev.
Glabrous perennial herbs, with rhizomes or short
caudices; stems ribbed. Leaves mostly cauline,
entire, oblanceolate to lanceolate, sessile, sometimes decurrent. Inflorescence leafy, corymbose
to thyrsoid; heads few to many, pedunculate.
Involucres shallowly cupulate; bracts 2–4(–5)seriate, gradate to subequal, narrowly oblong
to lanceolate, with orangish midrib; receptacles
convex to conical. Rays 20–40, pistillate, limbs
white to bluish, rarely pink. Disc florets yellow,
cylindrical, tubes short, throat with orange resin
ducts. Achenes compressed, elliptical to ellipticobovate, often winged, often with orangish ducts,
faces sometimes pubescent; pappus absent, a short
corona, or of some small bristles and 2(–4) longer
thick bristles. x = 9. Five species, eastern North
America.
916. Chloracantha G.L. Nesom et al.
Chloracantha G.L. Nesom, Y.B. Suh, D.R. Morgan, S.D.
Sundb. & B.B. Simpson, Phytologia 70: 378 (1991).
Glabrate glaucous perennial subshrubs; stems
erect, branches sharply ascending and sometimes
modified as thorns. Leaves cauline, 1-nerved,
oblanceolate, usually entire, early deciduous.
Inflorescences loosely corymbose. Involucres
broad; bracts 4–5-seriate, gradate, oblong-elliptic
to lanceolate, each with (1–)3(–5) orange-resinous
veins, apices rounded to lanceolate, margins
hyaline; receptacles plano-convex. Rays 10–33,
pistillate, white. Disc florets yellow with orangeresinous veins, narrowly funnelform, lobes deltate.
Achenes compressed, 5(–6)-nerved, glabrous;
pappus bristles 1–2-seriate, usually few shorter
setae outside. x = 9. One species, C. spinosa
(Benth.) G.L. Nesom, south-western USA (east to
Louisiana), south through Mexico to Panama.
XV.13. Subtribe Machaerantherinae G.L.
Nesom (1994).
Annual or perennial herbs to subshrubs (Xylorhiza) or small shrubs (Hazardia), stipitateglandular or resinous-glandular, sometimes
eglandular. Leaves alternate, entire to toothed,
uncommonly lobed to pinnatifid, teeth commonly
spinulose-tipped. Heads solitary to few, rarely
in a loosely corymbose inflorescence; involucral
bracts usually in 4–8 strongly gradate series,
commonly with an apical herbaceous patch;
receptacles convex, with or without pales. Ray
florets 1(–2)-seriate, corollas yellow, white, or blue.
Disc florets bisexual, rarely functionally male;
style branch appendages narrowly triangular,
hairy. Achenes multinerved, commonly obconic,
eglandular; setulae without anchor-shaped tips;
pappus 2–3(–4)-seriate, of bristles graduated in
length and with a tendency to be flattened, at least
basally. x = 6, reduced to 5, 4, 3, 2.
Key to the Genera
1. Style branch appendages truncate or subulate to
deltate or broadly ovate
2
– Style branch appendages narrowly triangular to
linear-lanceolate
4
2. Ray florets absent, outermost disc (tubular) florets zygomorphic with ray-like extensions
925. Lessingia
– Ray florets present
3
3. Ray florets fertile, corollas yellow; disc florets functionally male; stems and leaves without tomentose
vestiture
918. Benitoa
– Ray florets sterile, corollas white to blue or pink; disc
florets fertile (stamens and ovaries); stems and leaves
usually densely to loosely tomentose
919. Corethrogyne
4. Rays white or blue
5
– Rays yellow or absent
9
5. Perennial, spring-flowering subshrubs; heads mostly
13–19 mm high, solitary; rays with lamina mostly 15–
30 mm long
934. Xylorhiza
– Annual or perennial, summer- and fall-flowering
herbs or weak subshrubs; heads mostly 3–12 mm
high, solitary or loosely clustered; rays with lamina
mostly 7–15 mm long
6
6. Leaves deeply pinnatifid to bipinnatifid throughout,
at least many of the teeth and lobes sharply acute
with bristle tips, bristles 0.2–1 mm long; plants annual;
pappus of subulate bristles, dorsiventrally flattened
near base, bases overlapping 926. Machaeranthera
– Leaves entire to toothed or lobed, if pinnatifid to bipinnatifid throughout, then lobes often rounded, with
or without an apiculum but not bristle-tipped; plants
annual to strongly perennial; pappus of filiform to
subulate bristles, terete to flattened near base, bases
overlapping or not overlapping
7
Compositae
7. Plants strongly perennial, usually with a branched
caudex; receptacles covered with scales 0.3–3.0 mm
long, often forming an alveolate reticulum; pappus
bristles subulate, flattened near the base, bases
strongly overlapping
932. Xanthisma p.p.
– Plants taprooted annuals or short-lived perennials; receptacles naked (or scales rarely to 0.5 mm long); pappus bristles filiform, not flattened near base (or slightly
so), bases not (or slightly) overlapping
8
8. Ray florets with prominent pappus; leaves entire to
toothed and plants variously pubescent with glandular
and/or non-glandular hairs
920. Dieteria
– Ray florets with pappus absent or present; if pappus
present, then leaves pinnatifid to bipinnatifid throughout or (if leaves entire to toothed) plants glabrous except for bristles or apiculate or bristle-tipped teeth on
leaf margins
917. Arida
9. Disc corollas with limb campanulate, abruptly broadened from tube into limb
10
– Disc corollas narrowly funnelform, gradually broadened into limb
15
10. Pappus of numerous antrorsely barbellate bristles;
achenes strigose
11
– Pappus a corona or of short scales or a few, slender,
caducous awns or awn-like bristles; achenes glabrous
or glabrate
12
11. Ray florets absent; plants perennial
924. Isocoma
– Ray florets present; plants annual (or biennial?)
930. Rayjacksonia
12. Heads without a basal concavity; pappus absent, a low
crown of scales, or a lacerate corona
13
– Heads with distinct concavity at the peduncle insertion; pappus of 2–5(–12), white, flattened, basally caducous awns
14
13. Rays inconspicuous, barely longer than involucre; achenes strongly flattened; pappus a hyaline corona; heads
sessile to short-pedunculate (0.5–1 mm) in a distinctly
corymbose arrangement
931. Stephanodoria
– Rays conspicuous, much longer than the involucre;
achenes plump, nearly terete to 4–6-sided; pappus
absent, a low crown of awn-like scales, or a lacerate
corona; heads solitary or long-pedunculate (1–5 cm)
in a loosely corymbose arrangement
933. Xanthocephalum
14. Taprooted annuals or perennials with solid stems
from drier habitats; leaves punctuate-resinous;
achenes plump, not winged; pappus of smooth or
minutely antrorsely barbellate awns 921. Grindelia
– Fibrous-rooted annuals with hollow stems from wet
habitats; leaves not evidently punctate or resinous;
achenes flattened, winged; pappus of retrorsely barbellate bristle-like awns
927. Olivaea
15. Heads solitary; receptacles paleate; pappus a ring of
awn-like scales 3–6 mm long
932. Xanthisma p.p.
– Heads solitary or in a corymbose arrangement; receptacles epaleate; pappus of numerous barbellate bristles
16
16. Plants from South America; leaves usually coriaceous,
usually toothed or lobed; heads often solitary on long
nearly leafless peduncles, seldom aggregated; outer
involucral bracts not leaf-like
922. Haplopappus
– Plants from North America; leaves herbaceous to coriaceous, entire to lobed; heads often numerous on
stems; outer involucral bracts sometimes leaf-like 17
329
17. Shrubs
923. Hazardia
– Herbs or weak subshrubs
18
18. Basal leaves persistent, strongly reduced upwards, foliar teeth or lobes, if present, usually not bristle-tipped;
heads usually racemose to spicate, less commonly solitary or loosely corymbose
929. Pyrrocoma
– Basal leaves usually not persistent, reduced upwards
or not, foliar teeth, if present, usually bristle-tipped;
heads solitary or commonly loosely corymbose
19
19. Leaves usually spinulose-toothed or lobed, rarely entire, cauline leaves usually reduced in size upwards,
not subtending the heads; outer involucral bracts not
leaf-like
932. Xanthisma p.p.
– Leaves entire, cauline leaves even-sized, continuing up
the stems and subtending the heads; outer involucral
bracts leaf-like
928. Oonopsis
Genera of Machaerantherinae
917. Arida (R.L. Hartm.) D.R. Morgan & R.L.
Hartm.
Arida (R.L. Hartm.) D.R. Morgan & R.L. Hartm., Sida 20, 4:
1410 (2003).
Machaeranthera Nees sect. Arida R.L. Hartm. (1990).
Annual or rarely short-lived perennial herbs, usually taprooted (rhizomatous in A. blepharophylla
(A. Gray) D.R. Morgan & R.L. Hartm.). Stems erect
to ascending, often much-branched, 10–40 cm, glabrous to variously pubescent with glandular and
non-glandular hairs. Leaves entire, dentate, lacerate to deeply pinnatifid or bipinnatifid. Heads
solitary to loosely thyrsoid-corymbose; involucres
turbinate to depressed hemispheric; bracts 4–8seriate, gradate, appressed, spreading or reflexed,
densely stipitate-glandular to glabrous, bases glabrous, apices with glandular and non-glandular
hairs, not bristle-tipped; receptacles indistinctly
alveolate, naked or nearly so, convex. Ray florets
pistillate, fertile, laminas light to dark blue (absent
in A. carnosa (A. Gray) D.R. Morgan & R.L. Hartm.).
Disc florets perfect, corollas yellow, tubular, lobes
triangular, erect. Achenes narrowly oblong, slightly
compressed laterally, with 5–11 filiform nerves per
face, moderately to densely pubescent; pappus of
white filiform bristles in 2–3 poorly defined series.
x = 5. Nine species, western USA, northern Mexico.
918. Benitoa D.D. Keck
Benitoa D.D. Keck, Leafl. W. Bot. 8: 26 (1956).
Erect, annual, taprooted herbs, scented, viscid,
with dark, tack-shaped glands throughout. Leaves
narrowed to clasping base, linear-lanceolate to
oblanceolate, entire, upper leaves bract-like, apic-
330
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
ulate. Inflorescences loosely cymose-paniculate;
involucres cylindric-turbinate; bracts c. 6-seriate,
gradate, subpersistent, linear-lanceolate, green
along middle, with spreading, slender tips bearing
glandular globule. Rays 5–8(–14), pistillate, yellow,
coiling. Disc florets 9–12, functionally male, yellow, tubes long, limbs abruptly broadened, lobes
deeply cut, linear-lanceolate, erect. Fertile achenes
maculate, triangular and 3-nerved, narrowed
distally; pappus of 3–8 caducous bristles. x = 5.
One species, B. occidentalis (H.M. Hall) D.D. Keck,
California.
919. Corethrogyne DC.
Corethrogyne DC., Prodr. 5: 215 (1836); Saroyan et al.,
Madroño 47: 89–96 (2000), rev.
Perennial herbs, white-tomentose, commonly
glabrescent, sometimes with stipitate glands.
Leaves petiolate to sessile, linear to obovate, entire
to toothed. Inflorescences simple to spicate or
loosely corymbose; heads 1–5(–20); involucres
hemispheric to turbinate; bracts 3–8-seriate,
gradate, often with green apical patch, tips often
spreading to deflexed. Rays 10–40, 1-seriate, pistillate but sterile, white to blue or pink, limbs straight
to slightly coiling. Disc florets perfect, yellow, tubes
long, limbs narrowly cylindrical, lobes narrowly
triangular; style appendages deltate to ovate.
Fertile achenes terete or compressed, 5–8-ribbed,
strigose-sericeous; pappus bristles 1–2-seriate,
tawny, without short outer series. x = 5. One
species, C. californica DC., north-western USA
(Oregon) to Baja California Norte.
920. Dieteria Nutt.
Dieteria Nutt., Trans. Amer. Philos. Soc. ser. 2, 7: 300
(1840).
Machaeranthera Nees subg. Dieteria (Nutt.) Greene (1896).
Taprooted herbs, erect to ascending, muchbranched, 10–100 cm tall, glabrous or with
glandular and/or non-glandular trichomes. Leaves
entire to irregularly serrate or dentate, usually
with spine-tipped teeth. Heads solitary or in
loose panicles; involucres turbinate, campanulate,
or hemispheric; bracts 3–12-seriate, gradate,
appressed, spreading or reflexed, bases indurate,
apices herbaceous. Rays pistillate, white, blue or
purple. Disc florets perfect, yellow, funnelform,
glabrate, lobes triangular, erect. Achenes linear to
obovate, compressed, faces smooth or 4–6-ribbed,
glabrous to strigose; pappus bristles 40–50,
1–3-seriate, white to tawny. x = 4. Three species,
western USA, Mexico.
Ray flowers are sterile or absent in Dieteria
canescens (Pursh) Nutt. var. shastensis (A. Gray)
D.R. Morgan & R.L. Hartm.
921. Grindelia Willd.
Grindelia Willd., Gesell. Naturf. Freunde Berlin Mag.
[Neuesten Entdeck. Gesamten Naturk.] 1: 259 (1807);
Steyermark, Ann. Missouri Bot. Gard. 21: 433–608 (1934),
rev.; Bartoli & Tortosa, Kurtziana 27: 327–359 (1999), rev.;
Lane, Jepson Man.: 271–274 (1993), reg. rev.
Prionopsis Nutt. (1840).
Annual or biennial herbs to subshrubs, taprooted to stoloniferous; with punctate or stipitate
glands or eglandular, often glutinous. Leaves
mostly oblong-lanceolate, entire to spinulose,
glandular-toothed or pinnatifid. Inflorescences
usually 1-headed; involucres campanulate; bracts
4–8-seriate, subequal or gradate, bases usually
sclerified, tips erect to spreading or reflexed. Rays
present or absent, pistillate, yellow to orange.
Disc florets perfect or functionally male, or
outer perfect and central male, tubular, throat
usually abruptly expanded, lobes triangular; style
appendages obtuse-rounded. Achenes sometimes
dimorphic, usually thick-walled, smooth or sculptured, compressed to subquadrate, disc achenes
sometimes compressed with many thin nerves;
pappus bristles white, usually caducous, 2–4,
slender to broad. x = 6. Circa 70 species, USA,
Mexico, southern Andes.
The achenes of Grindelia may be uniform or
dimorphic. They are winged in some rhizomatous
South American species. The pappus usually has
few bristles but some South American species have
up to 18, and the North American G. ciliata (Nutt.)
Spreng. and South American G. prunelloides (Less.)
Bartoli & Tortosa have 2–4 series of 39–50 bristles
which may be basally connate. The inflorescences
are loosely corymbose in some Mexican species.
922. Haplopappus Cass.
Haplopappus Cass., Dict. Sci. Nat. 56: 168 (1828), nom.
cons.; Tortosa & Bartoli, Bol. Soc. Argent. Bot. 37: 115–133
(2002), rev.; Urbatsch et al., Comp. Newslett. 40: 35 (2003),
phylog.
Perennial herbs, subshrubs or shrubs; usually
glabrous and glutinous to stipitate-glandular;
Compositae
stems nearly acaulescent to caulescent and branching. Leaves coriaceous, sessile, entire or dentate
to pinnately lobed. Inflorescences on scapiform
peduncles with scale-like bracteoles and 1 head,
or corymbose to thyrsoid; involucres campanulate to hemispheric; bracts 3–6-seriate, gradate,
coriaceous, tips acute to spiniform, appressed to
recurved; receptacles plano-convex, epaleate. Rays
1-seriate, pistillate or sterile, yellow to purple,
limbs not or weakly coiling. Disc florets numerous,
perfect, yellow, narrowly funnelform, lobes 5,
broadly triangular to lanceolate, erect. Achenes
prismatic or turbinate, angular to terete or slightly
compressed; pappus 1–3-seriate, white, yellowish
or reddish, bristles unequal, scabrid. x = 6, 5. Circa
70 species, southern South America.
The South American Haplopappus sect.
Diplostephioides (Benth. & Hook. f.) Blake has
been treated as the genus Llerasia Triana (Cuatrecasas 1970). All North American and Central
American species identified as Haplopappus have
been transferred to other genera (summary by
Lane and Hartman 1996). Of the 4 sections of
South American Haplopappus, sects. Haplopappus
and Gymnocoma Nutt. are closely related to each
other; sects. Polyphylla Hall and Xylolepis Hall
may be congeneric with Hazardia Greene, based
on morphological and chemical evidence (Clark
1979; Brown and Clark 1981, 1982). Only a single
South American species (Haplopappus glutinosus
Cass., the generitype) has so far been included in
molecular-based phylogenetic analyses.
923. Hazardia Greene
Hazardia Greene, Pittonia 1: 28 (1887); Clark, Madroño 26:
105–127 (1979), rev.
Suffrutescent herbs or shrubs. Leaves usually
clasping or subclasping, deciduous, coriaceous,
glutinous, entire or toothed, glabrous to tomentose, sometimes gland-dotted. Inflorescences
spicate, racemose, narrowly thyrsoid or cymose,
rarely 1-headed; involucres cylindric to turbinate
or campanulate; bracts 5–9-seriate, gradate. Rays
0 or 3–25, pistillate, fertile or sterile, yellow or
drying red-purple. Disc florets perfect or functionally male, yellow, tube narrow, limb tubular
to narrowly funnelform, lobes triangular-ovate,
erect. Achenes fusiform to cuneate, subterete to
compressed, 4–5-nerved, glabrous to sericeous;
pappus of many reddish-brown capillary bristles.
x = 4, 5, 6. Thirteen species, western USA, western
Mexico.
331
924. Isocoma Nutt.
Isocoma Nutt., Trans. Amer. Philos. Soc. 2, 7: 320 (1840);
Nesom, Phytologia 70: 9–114 (1991), rev.
Erect, perennial subshrubs; often glutinous, glabrous to villous, tomentose or hispidulous. Leaves
linear to oblanceolate or obovate, 1-nerved, entire
to toothed or pinnatifid, teeth usually spinosetipped. Inflorescence corymbose or 1-headed;
heads sessile to subsessile; involucres turbinate
or campanulate; bracts (3–)4–6-seriate, gradate,
oblong to linear, chartaceous to subcoriaceous
with herbaceous tips. Rays absent. All florets
perfect, yellow with orange veins, limb abruptly
expanded, lobes triangular, erect, marginal florets
curving outwards as tube elongates. Achenes
obpyramidal, terete to subterete, sericeous, 3–11ribbed, ribs sometimes resinous; pappus bristles
2(–3)-seriate, uneven. x = 6. Sixteen species,
Mexico, south-western USA.
925. Lessingia Cham.
Lessingia Cham., Linnaea 4: 203 (1829); Lane, Jepson Man.:
304–306 (1993), reg. rev.; Markos & Baldwin, Syst. Bot. 26:
168–183 (2001), phylog.
Annual, erect to decumbent, taprooted herbs;
glabrous to white-tomentose, gland-dotted, often
with tack-shaped glands. Leaves in basal rosettes,
petiolate, obovate to spathulate, entire to serrate
or pinnately lobed, sometimes spinose-tipped;
cauline leaves smaller, sessile. Inflorescences
glomerulous, spicate, or thyrsoid; involucres
cylindric to turbinate or campanulate; bracts
3–7-seriate, imbricate, green-tipped. Florets 5–30,
all perfect, yellow, lavender, rose or white; outer
florets zygomorphic, all 5 lobes becoming ray-like;
inner florets tubular-funnelform, lobes linear;
style appendages truncate to subulate. Achenes
turbinate, villous; pappus of many unequal,
reddish-brown bristles, separate or united into
clusters, or of 5 paleaceous awns. x = 5. Seven to 10
species, south-western USA, Baja California Norte.
926. Machaeranthera Nees
Machaeranthera Nees, Gen. Sp. Aster. 224 (1832) [1833];
Turner, Phytologia 62: 207–266 (1987), rev.; Hartman,
Phytologia 68: 439–465 (1990), emend.
Hesperastrum A. Gray (1865).
Annual to perennial herbs, taprooted, erect or
ascending; stems, leaves and involucres glabrous
or with glandular or non-glandular hairs. Leaves
332
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
pinnatifid to bipinnatifid, spinulose. Heads
solitary or in panicles, pedunculate; involucres
turbinate to hemispheric; bracts 3–6-seriate,
gradate, appressed to spreading or reflexed, base
whitish-indurate, distally herbaceous. Rays pistillate, white to blue or purple, coiling. Disc florets
perfect, yellow, funnelform, lobes triangular, erect.
Achenes obovate, strigose, slightly compressed,
rather thick-walled, faces 4–9-ribbed; pappus
white to tawny, 1–3-seriate. x = 4. Two species,
western USA, Mexico.
Molecular data (Morgan 2003) indicate that
Oonopsis (Nutt.) Greene, Pyrrocoma Hook. and
Xanthisma DC. arose evolutionarily from within
the branches of what had recently been considered
the genus Machaeranthera (sensu Hartman 1990).
Accordingly, Morgan and Hartman (2003) have
divided the genus into four smaller monophyletic
genera, including Machaeranthera s. str., Xanthisma, Arida (R.L. Hartman) D.R. Morgan & R.L.
Hartman, and Dieteria Nutt., which more closely
reflect the hypothesized evolutionary patterns.
927. Olivaea Sch. Bip. ex Benth.
Olivaea Sch. Bip. ex Benth. in Hook., Ic. Pl. t. 1103 (1872);
De Jong & Beaman, Brittonia 15: 86–92 (1963), rev.
Oligonema S. Wats. (1891), nom. illegit. non Oligonema
Rostaf. (1875).
Golionema S. Wats. (1891).
Erect, mostly immersed, aquatic annuals from
taproot, simple and glabrous below water surface,
cymosely branched and stipitate-glandular above;
stems hollow. Leaves sessile, linear to lanceolate,
entire, lobed in deciduous submerged leaves.
Heads solitary on leafy peduncles, in groups of few
to 20; involucres hemispheric; bracts c. 3-seriate,
basally fused, chartaceous, herbaceous distally.
Rays 25–50, 1-seriate, pistillate, yellow, coiling.
Disc florets perfect, yellow, tubular, lobes triangular, erect. Achenes quadrate or oblanceolate,
compressed, 2–3-winged or 2-nerved, glabrous,
shiny; pappus of 2–12 caducous, unequal bristles.
x = 6. Two species, south-western Mexico.
928. Oonopsis (Nutt.) Greene
Oonopsis (Nutt.) Greene, Pittonia 3: 45 (1896); Hall, Publ.
Carnegie Inst. Wash. 389: 34, 86–95 (1928), rev.
Stenotus Nutt. sect. Oonopsis Nutt. (1840).
Perennial herbs, with taproots and branched caudices; glabrous or sparsely tomentose. Leaves little reduced upwards, few leaves subtending heads,
linear to narrowly oblanceolate or lanceolate. Inflorescences 1-headed, loosely corymbose with up to
12 heads, or subglomerate; involucres hemispheric
to cylindric-turbinate; bracts 3–6-seriate, gradate,
bases chartaceous, distally green, sharply acuminate. Rays 6–15, pistillate, yellow, not or weakly
coiling. Disc florets perfect; yellow, funnelform,
short-tubed, lobes triangular, erect. Achenes prismatic or narrowly turbinate, strigose or glabrous;
pappus of rather few, brownish, rigid, unequal bristles. x = 5. Four or 5 species, north-central USA.
929. Pyrrocoma Hook.
Pyrrocoma Hook., Fl. Bor.-Am. 1: 306 (1833); Brown,
Jepson Man.: 330–331 (1993), reg. rev.
Homopappus Nutt. (1840).
Erect or decumbent perennial herbs from taproot
or caudex; stems, leaves and involucres glabrous
or tomentose to sericeous, often punctuate- or
stipitate-glandular. Leaves oblanceolate to elliptic
or linear, entire or spinulose-dentate to laciniate;
upper leaves reduced. Inflorescences 1-headed
or spicate to loosely corymbose; involucres
hemispheric to narrowly campanulate; bracts
2–6-seriate, equal to gradate, oblanceolate to
oblong or linear, herbaceous and totally yellowgreen or with indurate whitish base and apical
green patch. Rays 10–80, pistillate, yellow, corollas
sometimes reduced. Disc florets perfect, yellowish,
tubular-funnelform, lobes deltate, erect. Achenes
compressed or 3–4-angled with several nerves
between angles, glabrous to strigose or sericeous;
pappus of few, brownish, rigid, unequal bristles.
x = 6. Circa 14 species, western USA, western
Canada.
930. Rayjacksonia R.L. Hartm. & M.A. Lane
Rayjacksonia R.L. Hartm. & M.A. Lane, Amer. J. Bot. 83:
368 (1996).
Annual to short-lived perennial, taprooted herbs;
stems, bracts and leaves with sessile or stipitate
glands. Leaves linear to oblanceolate, marginal
teeth and apices bristle-tipped. Inflorescence with
1–70 heads per plant, loosely thyrsoid-cymose.
Involucres hemispheric; bracts 4–5-seriate, subequal, linear to lanceolate, 1-nerved, apices erect to
spreading or recurved. Rays pistillate, 18–46, yellow. Disc florets perfect, throat abruptly expanded,
lobes erect-spreading. Achenes strigose-sericeous,
dimorphic, in ray florets broadly ellipsoid to
obovoid, thick-walled, 3-angled with 0–4 ribs
Compositae
on sides, in disc florets broadly ellipsoid-clavate,
compressed, walls thinner, faces 5–9-ribbed;
pappus bristles 3–4-seriate, unequal, thick and
flattened, persistent. x = 6. Three species, central
and south-western USA, north-eastern Mexico.
931. Stephanodoria Greene
Stephanodoria Greene, Erythea 3: 12 (1895); Nesom,
Phytologia 82: 107–113 (1997), rev.
Perennial herbs; stems, leaves and involucres
stipitate-glandular and hirtellous-pilosulous.
Basal leaves persistent, narrowed to petiole, blades
obovate to oblong-elliptic, entire, upper leaves
reduced, sessile. Inflorescence loosely corymbose,
with clusters of 2–6 sessile to subsessile heads;
involucres turbinate; bracts 5–7-seriate, gradate,
tips spreading to reflexed. Rays 20–30, pistillate,
golden-yellow. Disc florets perfect, yellow, outer
bending outwards from elongating tubes. Achenes
oblanceolate to oblong-elliptic, compressed, 2–
3-nerved, stramineous, mostly glabrous, strigose
at apex; pappus a hyaline, erose corona, sometimes awn-like to one side. x = 6. One species,
S. tomentella (B.L. Rob.) Greene, central Mexico,
gypsophilous.
932. Xanthisma DC.
Xanthisma DC., Prodr. 5: 94 (1836); Semple, Rhodora 76:
1–19 (1974), rev.
Eriocarpum Nutt. (1840).
Sideranthus Nutt. ex Nees (1840).
Machaeranthera Nees subg. Sideranthus (Nutt. ex Nees)
R.L. Hartman (1990).
Taprooted herbs, erect or spreading or sprawling,
often much-branched, 3–100 cm tall, glabrous or
with glandular or non-glandular hairs. Leaves
entire or serrate to bipinnatifid, teeth usually
bristle-tipped. Heads solitary or in loose panicles;
involucres turbinate to hemispheric or campanulate; bracts 2–8-seriate, gradate, appressed,
spreading or reflexed, bases chartaceous, apices
herbaceous; receptacles with laciniate scales. Rays
pistillate, white, pink, purple or yellow, sometimes
absent, coiling. Disc florets perfect, yellow, tubular,
lobes triangular, erect. Achenes often dimorphic,
elliptic to obscurely cordate, ray achenes 3-sided,
disc achenes compressed laterally, with faces 3–9nerved, sparsely to densely pubescent; pappus of
white to tawny bristles, often flattened proximally,
in 2–4 series, or of awn-like scales. x = 5, reduced
333
to 4, 3, and 2. Seventeen species, western USA,
northern Mexico.
Xanthisma in its expanded sense comprises
the unispecific genus Xanthisma and the former
Machaeranthera Nees sections Blepharodon (DC.)
R.L. Hartm., Sideranthus (Nutt. ex Nees) R.L.
Hartm., Havardii (R.C. Jacks.) R.L. Hartm., and
Stenoloba R.L. Hartm. (sensu Hartman 1990).
The genus is distinguished by its short, turbinate,
thick-walled, densely pubescent fruits, leaves with
marginal spines, and chromosome numbers based
on x = 4. Rays are either cyanic (blue, purple, pink
or white) or yellow.
933. Xanthocephalum Willd.
Xanthocephalum Willd., Gesell. Naturf. Freunde Berlin
Mag. [Neuesten Entdeck. Gesamten Naturk.] 1: 140 (1807);
Lane, Syst. Bot. 8: 305–316 (1983), rev.; Nesom, Phytologia
66: 482–487 (1989), emend.
Annual or perennial herbs, taprooted or rhizomatous; glandular, often glutinous; stems erect or
prostrate. Leaves sessile, sometimes decurrent,
oblanceolate, entire or spinulose-toothed. Inflorescences 1-headed or loosely corymbose; involucres
hemispheric to campanulate; bracts 2–4-seriate,
gradate, white-indurate basally, green distally. Rays
(0–)14–62, pistillate, yellow to orange-yellow. Disc
florets perfect, yellow, tube narrow, limb abruptly
expanded. Achenes compressed, subterete or
4–6-sided, thick-walled without prominent ribs,
glabrous to strigillose; pappus absent, a lacerate
crown, with awn-like scales or rarely with short
bristles, usually shorter in rays. x = 6. Six species,
western USA, Mexico.
934. Xylorhiza Nutt.
Xylorhiza Nutt., Trans. Amer. Philos. Soc. 2, 7: 297 (1840);
Watson, Brittonia 29: 199–216 (1977), rev.
Perennial herbs to small shrubs; glabrous to
villous or tomentose, sometimes with stipitate
glands; stems often white. Leaves sessile, linear
to lanceolate, oblanceolate or oblong, mostly
1-nerved, margins entire to spinulose-dentate.
Heads solitary on naked peduncles; involucres
campanulate to hemispheric; bracts 3–6-seriate,
gradate, narrowly lanceolate, keeled, bases whiteindurate, tips green. Rays 1–2-seriate, (4–)12–60,
pistillate, white to blue or purple, coiling. Disc florets perfect, yellow, tubes slender, limbs narrowly
funnelform, lobes ovate-deltate, erect-spreading,
resin ducts pale to yellow. Achenes fusiform to
334
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
linear or ovate, somewhat compressed, 4-nerved,
sericeous; pappus bristles 2–3-seriate, stout, often
flattened, unequal. x = 6. Eight species, western
USA, Mexico.
XV.14. Subtribe Symphyotrichinae
G.L. Nesom (1994).
Annual or usually perennial herbs, rarely shrubs
(Ampelaster), eglandular to glandular. Leaves alternate, entire or rarely toothed, not lobed. Inflorescence with heads solitary to few or numerous,
loosely corymbose, thyrsoid, or with branches secund; involucral bracts flat, usually with a herbaceous apical patch; receptacles flat, epaleate. Ray
florets 1(–2–4)-seriate, corollas white to blue, coiling. Disc florets bisexual; style appendages lanceolate, hairy. Achenes obovoid to cylindric, terete
or usually slightly to strongly compressed, multinerved, eglandular, setulae without anchor-shaped
tips; pappus bristles 1-seriate (2-seriate in Symphyotrichum sect. Conyzopsis, often 2-seriate in
Canadanthus). x = 9, reduced to 8, 7, 6, 5, 4.
Key to the Genera
1. Plants annual to short-lived perennial, usually arising
from a slender taproot
2
– Plants perennial, usually arising from long or short
rhizomes and fibrous roots
4
2. Heads and upper stems stipitate-glandular; ray achenes epappose
938. Psilactis p.p.
– Plants completely eglandular; ray achenes pappose 3
3. Involucral bracts evenly herbaceous and of equal
length; pistillate florets in 2–4 series in a broad outer
zone, lamina absent or rudimentary to filiform and
short; disc florets fewer than ray florets; pappus
bristles in 2 series, all of equal length
939. Symphyotrichum sect. Conyzopsis
– Involucral bracts with a green, rhombic apical patch,
basally indurate, gradate in length; pistillate florets in
1(–2) series, lamina prominent or strongly reduced;
disc florets more numerous than ray florets; pappus
bristles of equal length and in a single series
939. Symphyotrichum sect. Oxytripolium p.p.
4. Ray achenes epappose
938. Psilactis p.p.
– Ray achenes pappose
5
5. Involucral bracts with a sharply delimited green apical patch or zone, lower portion of involucral bract
indurate
6
– Involucral bracts evenly herbaceous, without a distinctive green apical patch
7
6. Plants trailing or climbing (not twining) vines; stems
perennial; achenes narrowly cylindric to slightly
fusiform, with 9–12 ribs
936. Ampelaster
– Plants mostly erect, sometimes leaning but never trailing nor even subscandent; stems annual; achenes nar-
rowly obovate, flattened to varying degrees, with (2–
)3–10(–11) ribs
939. Symphyotrichum p.p.
7. Leaves linear, parallel-veined, not clasping; rays white;
achenes terete
935. Almutaster
– Leaves lanceolate to elliptic-lanceolate, net-veined,
subclasping; rays purple; achenes strongly flattened
937. Canadanthus
Genera of Symphyotrichinae
935. Almutaster Å. Löve & D. Löve
Almutaster Å. Löve & D. Löve, Taxon 31: 356 (1982).
Perennial rhizomatous herbs, lower parts glabrous,
stipitate-glandular above. Leaves sessile, thick, narrow, longitudinally 3–5-nerved. Inflorescence with
1 or few heads or loosely corymbose or thrysoid
with 5–10(–30) heads; involucre broad; bracts in
3–4(–5) subequal series, herbaceous without indurated bases. Rays 15–30(–45) in 1 series, pistillate, white to lilac-tinged. Disc florets perfect, tubular, lobes deltate, erect. Achenes fusiform, terete,
7–10-nerved, sparsely strigose; pappus 1-seriate,
of many attenuate, subequal bristles. x = 9. One
species, A. pauciflorus (Nutt.) Å. & D. Löve, central
Canada to Mexico.
936. Ampelaster G.L. Nesom
Ampelaster G.L. Nesom, Phytologia 77: 250 (1994).
Weak, diffusely branched shrubs; stems scrambling, hirsutulous or pilosulous. Leaves sometimes
evergreen, oblanceolate, narrowed to auriculate base, 1-nerved. Heads 1 or 2–8 in loose,
short-pedunculate clusters; involucres broad;
bracts 4–6-seriate, gradate, thick, low-keeled,
linear-oblong to narrowly spathulate, bases white,
apices herbaceous, spreading-reflexed. Rays 40–70,
pistillate, pink or lavender. Disc florets perfect,
tubular. Achenes cylindrical to slightly fusiform,
glabrous, with 9–12 usually whitish ribs; pappus
1-seriate, bristles attenuate. x = 9. One species,
A. carolinianus (Walter) G.L. Nesom, eastern USA
coastal plain.
937. Canadanthus G.L. Nesom
Canadanthus G.L. Nesom, Phytologia 77: 250 (1994).
Perennial thin-rhizomatous herbs; stems single, simple, with long-stipitate glands. Leaves
cauline, sessile, clasping, lanceolate, entire to
serrate, eglandular, lowest leaves much reduced.
Inflorescences usually loosely corymbose, usually
with 3–20 heads on leafy peduncles, sometimes
Compositae
heads solitary; involucre turbinate to hemispheric;
bracts 2–3-seriate, subequal, linear-lanceolate,
thin-herbaceous, stipitate-glandular, without
indurated base or apical green patch, inner bracts
often slightly keeled. Rays 25–40, pistillate, purple,
coiling. Disc florets perfect, tubular, lobes short,
erect. Achenes oblanceolate, strongly compressed,
4–8-nerved; base narrow, stipitate; pappus bristles
1(–2)-seriate, attenuate. x = 9. One species,
C. modestus (Lindl.) G.L. Nesom, USA (Alaska
south to Montana, Michigan), east to south-eastern
Canada (New Brunswick).
335
rowly obovate, compressed, (2–)3–5(–6)-nerved,
sparsely strigose to glabrous; pappus bristles
1(–2)-seriate. x = 8, 7, 6, 5, 4. Circa 92 species,
mostly North America, some Mexico and South
America, few widely naturalized.
Symphyotrichum subg. Symphyotrichum (incl.
Conyzanthus Tamamsch., Brachyactis Ledeb.) includes species with x = 8, 7 and 6, and 5, while
subg. Virgulus (Raf.) Nesom (incl. Lasallea Greene,
Virgulaster Semple) primarily those with independently derived x = 5. Hybrids and backcrosses between the two (x = 13) are common and stabilized
in some areas (Nesom 1994d).
938. Psilactis A. Gray
Psilactis A. Gray, Mem. Amer. Acad. Arts 2, 4: 71 (1849);
Morgan, Syst. Bot. 18: 290–308 (1993), rev.
XV.15. Subtribe Chaetopappinae G.L. Nesom
(2000).
Annual or perennial herbs, usually taprooted;
stems, involucral bracts and often leaves stipitateglandular, sometimes tomentose. Lower leaves
obovate to linear-lanceolate, early deciduous,
upper leaves entire, subclasping. Inflorescence
loosely thyrsoid to corymbose or with branches
1-headed; involucre broad, bracts 2–4-seriate,
subequal to gradate, outer bracts indurate along
basal margins. Rays pistillate, white to bluish,
coiling. Disc florets perfect, tubular. Ray achenes
fusiform to obovate, subterete, 5–14(–18)-nerved,
short-strigose, epappose; disc achenes slightly
longer, with pappus bristles 1-seriate. x = 9, 4, 3.
Six species, mostly USA (Texas) and Mexico, one
to Andes of northern South America.
Annual or short-lived perennial herbs. Leaves
alternate, oblong to oblanceolate-spathulate,
entire. Heads mostly solitary; involucral bracts flat
to convex, not keeled, with broad, hyaline margins;
receptacles usually without pales. Rays 1-seriate,
pistillate, blue to white, strongly coiling. Disc florets
sometimes functionally male; style appendages
obtuse or truncate to triangular. Achenes eglandular or glandular, terete and multinerved (most
Chaetopappa) or obovate, flattened, and 2-nerved
(Monoptilon, some Chaetopappa); setulae without
anchor-shaped tips; pappus of persistent bristles
or scales, or of bristles and scales, commonly
in multiples of 5 (in Chaetopappa), or absent.
x = 8.
939. Symphyotrichum Nees
Key to the Genera
Symphyotrichum Nees, Gen. Sp. Aster. 135 (1832) [1833];
Nesom, Phytologia 77: 267–296 (1994), comb.
Tripolium Nees sect. Oxytripolium DC. (1836).
Virgulus Raf. (1836) [1837].
Tripolium Nees subg. Astropolium Nutt. (1840).
Brachyactis Ledeb. (1845).
Virgulaster Semple (1985).
Mostly perennial herbs, with rhizomes or short
caudex, fibrous-rooted. Leaves petiolate to sessile
and clasping, linear to cordate, entire or toothed.
Inflorescence cylindrical to diffusely thyrsoid
or corymbose; involucres narrowly campanulate
to hemispheric; bracts 3–7-seriate, gradate to
subequal, mostly with pale indurated bases and
herbaceous tips. Rays pistillate; usually long, white
to blue or purplish, coiling. Disc florets perfect, yellow, usually purplish at tips, gradually expanded,
lobes deltate, erect to spreading. Achenes nar-
1. Stems erect to ascending; capitula on distinct peduncles; achenes 2–5-, 8-, or 10-nerved, flattened to terete
940. Chaetopappa
– Stems prostrate to decumbent; capitula closely subtended by foliar bracts; achenes 2-nerved, flattened
941. Monoptilon
940. Chaetopappa DC.
Chaetopappa DC., Prodr. 5: 301 (1836); Shinners, Wrightia
1: 63–89 (1946), rev.; Nesom, Phytologia 64: 448–456 (1988),
key.
Distasis DC. (1836).
Annual or perennial herbs; stems erect or decumbent. Heads terminal, involucres broadly conical
to hemispheric; bracts 2–6-seriate, gradate, elliptic
to linear-lanceolate; receptacles flat to slightly convex. Rays 6–20, white or drying bluish. Disc florets
perfect or central florets functionally male, yellow,
336
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
tubular; style appendages shallowly to narrowly
triangular. Achenes terete to compressed, 2–5- or
8–10-nerved, sparsely strigose, sometimes glanddotted; pappus a hyaline crown, hyaline scales,
awn-like bristles, alternating scales and bristles or
nearly lacking. x = 8. Eleven species, south-central
USA, north-eastern Mexico.
941. Monoptilon Torr. & A. Gray ex A. Gray
Monoptilon Torr. & A. Gray ex A. Gray, Boston J. Nat. Hist.
5: 106 (1845); Keil, Jepson Man.: 319 (1993), reg. rev.
Eremiastrum A. Gray (1854).
Annual taprooted herbs, radiating branches prostrate to decumbent; stems, leaves and involucres
minutely glandular, hispid-hirsute, hairs long,
white. Leaves small. Heads sessile; involucres
campanulate; bracts 1(–2)-seriate, equal, linearlanceolate, purple-tipped. Rays 12–21, white or
rose to purple-tinged. Disc florets mostly perfect,
yellow with orange veins, tubular-funnelform;
style appendages obtuse to truncate. Achenes
obovate, compressed, 2-nerved, sparsely strigose;
pappus a short, toothed cup and single apically
plumose bristle or 1–12 barbellate bristles alternating with scales. x = 8. Two species, south-western
USA, north-western Mexico.
XV.16. Subtribe Astranthiinae G.L. Nesom
(2000).
Annual, biennial or perennial herbs. Leaves alternate, spathulate to linear, entire or few-toothed,
eglandular. Heads solitary; involucral bracts not
keeled, with broad, hyaline margins; receptacles
convex to conical, usually epaleate, paleate in
Geissolepis. Ray florets 1-seriate, corollas white
to bluish or pinkish above (rarely yellow in
Townsendia), usually with a lavender to blue or
pink abaxial midstripe, not reflexing or coiling.
Disc florets with short-tubed corollas; style branch
appendages triangular-lanceolate, papillose.
Achenes oblanceolate to obovate, compressed,
2(–3)-ribbed, commonly with setulae with anchorshaped or uncinate tips (achenes winged and
fringed with setulae with anchor-shaped tips in
Dichaetophora); pappus 1-seriate, of barbellate
bristles or a low crown of setae/bristles and
scales (2-awned in Dichaetophora). x = 9; x = 5
in Astranthium, reduced to 4 and 3; x = 3 in
Dichaetophora.
Key to the Genera
1. Pappus of numerous bristles, often short; x = 9
945. Townsendia
– Pappus of scales, awns, or essentially absent
2
2. Plants succulent, prostrate; pappus of 6–8 narrowly
triangular scales, scale margins with uncinate cilia;
x = 9 or 8; receptacles paleate
944. Geissolepis
– Plants herbaceous, erect to decumbent; pappus of 2
awns or essentially absent; x = 5 or reduced; receptacles epaleate
3
3. Achenes without wings or fringed margins; pappus
absent or a barely perceptible corona
942. Astranthium
– Achenes winged, hairs of marginal fringe with anchorshaped tips; pappus of 2 thin, barbellate awns, usually
with additional minute awns
943. Dichaetophora
Genera of Astranthiinae
942. Astranthium Nutt.
Astranthium Nutt, Trans. Amer. Philos. Soc. 2, 7: 312
(1840); De Jong, Publ. Mus. Michigan State Univ., Biol. Ser.
2: 429–528 (1965), rev.
Annual, biennial or perennial taprooted herbs;
stems erect to creeping, sparsely to densely hairy.
Some leaf bases clasping, lower leaves spathulateobovate, upper leaves narrower. Heads solitary, on
usually long peduncles; involucres campanulate
to hemispheric; bracts 2(–3)-seriate, subequal,
broadly oblanceolate to linear-lanceolate, thinherbaceous; receptacles conical. Rays 10–65(–85),
pistillate, limbs white above. Disc florets perfect,
yellow, tube short, limb rather abruptly expanded,
lobes deltate, erect; style appendages lanceolateacute. Achenes 2-ribbed, faces smooth to papillose,
glabrous or with anchor-shaped setulae; pappus
absent or crown of short setae or scales. x = 3, 4, 5.
Eleven species, Mexico to south-central USA.
943. Dichaetophora A. Gray
Dichaetophora A. Gray, Mem. Amer. Acad. Arts 2, 4: 73
(1849); Shinners, Wrightia 1: 90–94 (1946), rev.
Annuals from thin taproot; stems erect or decumbent, sparsely strigose. Leaves oblanceolate, entire.
Heads terminal, solitary; peduncles 2–9 cm long;
involucres hemispheric; bracts 2-seriate, subequal,
long-ciliate; receptacles convex to conical. Rays 15–
25, pistillate, limbs white above. Disc florets perfect, yellow, tubes short, abruptly expanded into
limb. Achenes obovate to broadly elliptic, faces with
thickened, elliptic central area surrounded by paler,
glabrous wing-like margins, setulae on central area
and edges of wings anchor-shaped; pappus of 2 thin
Compositae
awns, usually minute additional awns along callus
rim. x = 3. One species, D. campestris A. Gray, USA
(south-central Texas, north-eastern Mexico).
944. Geissolepis B.L. Rob.
Geissolepis B.L. Rob., Proc. Amer. Acad. Arts 2, 27: 177
(1892).
Perennial prostrate herbs, with fibrous-rooted
stolons; stems and peduncles sparsely pilose,
glabrescent. Leaves linear-oblanceolate, succulent, strigose adaxially along midvein. Heads on
thick, axillary, bracteate peduncles; involucre
broadly turbinate; bracts 4–5-seriate, strongly
gradate, thick; receptacles steeply conic; paleae
obovate, midvein resinous. Rays 8–12, pistillate,
orange-yellow with orange resin ducts. Disc florets
orange-yellow with orange veins, funnelform,
gradually expanded; style appendages triangular.
Achenes obovate-triangular, slightly compressed,
outer achenes 3–4-angled, with c. 8 pale ribs,
with orange resin pockets distally, strigose with
anchor-shaped setulae; pappus of 6–8 narrow
pales, margins ciliate. x = 8 or 9. One species,
G. suaedifolia B.L. Rob., central Mexico.
945. Townsendia Hook.
Townsendia Hook., Fl. Bor.-Am. 2: 16 (1834); Beaman,
Contr. Gray Herb. 183: 1–151 (1957), rev.; Reveal, Great
Basin Naturalist 30: 23–52 (1970), rev.
Annual, biennial or perennial herbs, taprooted
or rarely stoloniferous; stems and leaves mostly
strigose. Leaves basal or also cauline, spathulate
to linear. Heads terminal, solitary; involucres
hemispheric to campanulate; bracts 2–7-seriate,
gradate, usually broadly hyaline-ciliate-margined;
receptacles convex, rarely conical. Rays white or
bluish to pinkish, one species yellow. Disc florets
perfect, yellow, lobes deltate, erect or incurved.
Achenes oblanceolate to obovate, compressed,
2(–3)-ribbed, smooth or papillose, glabrate or
with anchor-shaped setulae; pappus 1-seriate, of
barbellate bristles or short bristles and squamellae,
ray pappus sometimes shorter than in disc. x = 9.
Twenty-six species, western North America, incl.
two in Mexico.
XV.17. Subtribe Chrysopsidinae G.L. Nesom
(1994).
Annual to perennial herbs. Leaves alternate, mostly
entire, commonly with sessile or stipitate glands.
Heads solitary or in loosely thyrsoid inflorescences;
337
involucral bracts keeled; receptacles epaleate. Ray
florets 1-seriate, yellow, rarely white, coiling
or not coiling. Disc florets bisexual, with large
rectangular-prismatic crystals in the throat; style
appendages narrowly lanceolate, hairy. Achenes
multinervate, usually eglandular; setulae without
anchor-shaped tips; pappus 2–3(–4)-seriate, the
inner 1–2 series of flattened bristles, outer of much
shorter setae, bristles, or scales. x = 9, reduced to
7, 6, 5, 4, 3.
Key to the Genera
1. Rays white; South America
949. Noticastrum
– Rays yellow; North America
2
2. Leaves densely and closely white-woolly-tomentose;
achenes narrowly oblong and strongly flattened, with
7–9 thin, closely adjacent, superficial white nerves on
each side; stems with one head 952. Tomentaurum
– Leaves variously pubescent but not white-woollytomentose; achenes not as above; stems with
1–numerous heads
3
3. Leaves linear, parallel-nerved or with a strong tendency for parallel veins, veins with associated massive
sclerenchyma, prominently superficial and forming
raised ridges
4
– Leaves obovate to oblanceolate, net-veined, veins not
strongly sclerenchymatous, not raised above lamina 5
4. Taprooted annuals to weak perennials; pappus
1-seriate; leaves with minute but evident lacunae on
947. Croptilon
the abaxial surface; x = 6
– Rhizomatous perennials; pappus 2-seriate, of bristles
and a short outer series of setae; leaves without lacunae
on the abaxial surface; x = 9
951. Pityopsis
5. Pubescence of coarse, stiff, osteolate hairs; x = 9
948. Heterotheca
– Pubescence of fine, flexible, flagelliform hairs; x = 5
6
6. Achenes obovate to linear-oblong in outline, nearly
terete to slightly flattened, with 5–10 ribs, ribs often with orange-resinous oil ducts; pappus 2-seriate;
stems mostly with 2–numerous heads
946. Chrysopsis
– Achenes fusiform-cylindric, with 8–16 whitish, superficial ribs; pappus 1-seriate or 2-seriate; stems mostly
with 1 head
950. Osbertia
Genera of Chrysopsidinae
946. Chrysopsis (Nutt.) Elliott
Chrysopsis (Nutt.) Elliott, Sketch Bot. S. Carolina 2: 333
(1824), nom. cons.; Semple, Rhodora 83: 323–384 (1981),
rev.; Nesom, Phytologia 71: 109–121 (1991), emend.
Diplopappus Cass. (1817).
Inula L. sect. Chrysopsis Nutt. (1818).
Annual or rhizomatous perennial herbs, pubescence woolly or sericeous, sometimes arach-
338
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
noid; stems erect to decumbent. Leaves sessile,
basal and lower cauline spathulate to oblanceolate.
Inflorescence terminal, corymbose. Involucres
hemispheric to turbinate; bracts (2–)3–5-seriate,
gradate, linear, not herbaceous; receptacles flat.
Rays 13–34, pistillate, yellow. Disc corolla lobes
deltate, erect. Achenes obovate to linear-oblong,
nearly terete, strigose, ribs 5–10, sometimes with
orange ducts; pappus 2-seriate, inner bristles long,
outer bristles short. x = 5, 3. Circa 11 species,
coastal south-eastern USA.
ing. x = 9. Circa 25–30 species, USA, Canada,
Mexico.
Three sections include sect. Heterotheca with
ray achenes 3-angled, epappose and usually glabrous, and disc achenes compressed; sect. Phyllotheca (Nutt.) Harms with all achenes cylindric
to slightly compressed, pappose and setulose; and
sect. Ammodia (Nutt.) Harms without any short
outer pappus series.
947. Croptilon Raf.
Noticastrum DC., Prodr. 5: 279 (1836); Zardini, Revista
Mus. La Plata, Secc. Bot. 13: 313–424 (1985), rev.
Aplopappus (Haplopappus) sect.(?) Leucopsis DC. (1836).
Leucopsis (DC.) Baker (1882).
Croptilon Raf., Fl. Tellur. 2: 47 (1836) [1837]; Smith,
Rhodora 67: 217–238 (1965), rev.; Smith, Sida 9: 59–63
(1981), emend.
Isopappus Torr. & A. Gray (1842).
Annual to weak perennial taprooted herbs, usually sparsely hispid, usually stipitate-glandular.
Leaves sessile, linear to narrowly oblanceolate
or lanceolate, often falcate, entire to serrate,
long-ciliate basally, parallel veins 3, arachnoid
hairs in abaxial areoles. Involucres narrowly
turbinate to subcylindric; bracts 3–5-seriate,
gradate, lanceolate to linear-lanceolate. Rays 5–10,
pistillate, yellow to orangish, coiling. Disc corollas
yellow, lobes triangular, erect. Achenes elongate,
terete to few-angled, with 6–14 superficial pale
nerves, strigose to sericeous, resin ducts present
in 1 species; pappus 1-seriate, bristles persistent,
rigid, reddish-brown. x = 6, 5, 4. Three species,
south-eastern USA to north-eastern Mexico.
949. Noticastrum DC.
Perennial herbs or subshrubs, scapiform with
basal rosettes or with stolons, decumbent to erect;
pubescence non-glandular, stipitate-glandular
and/or woolly. Leaves often copiously silky near
base. Inflorescence 1- to few-headed. Involucre
campanulate to turbinate; bracts 3–6-seriate, gradate, linear-lanceolate, often reddish-margined.
Rays pistillate, white to purplish, rarely yellow. Disc
corollas yellow, throat tubular, lobes short, erect.
Achenes subfusiform, somewhat compressed,
setuliferous, with 16–26 costae, usually 5–10
of these prominent, carpopodium asymmetric;
pappus to 4-seriate, bristles usually tawny, some
short outer bristles present. x = 9. Circa 19 species,
Andean, southern South America.
950. Osbertia Greene
948. Heterotheca Cass.
Heterotheca Cass., Bull. Sci. Soc. Philom. Paris 1817: 137
(1817); Semple, Syst. Bot. 13: 547–558 (1988), rev.; Nesom,
Phytologia 69: 282–294 (1990), rev.; Semple, Univ. Waterloo
Biol. Ser. 37: i–iv, 1–164 (1996), rev.
Annual or perennial taprooted or rhizomatous
herbs, often hispid-pilose, often aromatic or viscid,
glands sessile or short-stipitate. Leaves entire or
shallowly toothed, basal often petiolate, upper
sessile. Inflorescences 1-headed or corymbose
with few heads. Involucres campanulate; bracts
4–8-seriate, gradate, lanceolate, rigid; receptacles
plano-convex. Rays absent or pistillate, yellow to
orangish, coiling. Disc corollas slightly expanded
above. Achenes cylindrical and 8–14-nerved or
compressed with 2 ribs, glabrous to short-strigose
or sericeous; pappus bristles tawny to whitish,
sometimes a crown of short bristles or lack-
Osbertia Greene, Erythea 3: 14 (1895); Turner & Sundberg,
Pl. Syst. Evol. 151: 229–239 (1986), rev.; Nesom, Phytologia
71: 132–135 (1991), emend.
Erect, perennial, stoloniferous herbs, pilose and
eglandular to puberulous and glandular with
septate glandular hairs; stems usually single.
Leaves of basal rosette linear-oblanceolate to
elliptical; cauline leaves reduced, clasping. Heads
solitary; peduncles long. Involucres hemispheric
to turbo-campanulate; bracts 4–6-seriate, gradate,
linear-lanceolate; receptacles convex, alveolae
deep. Rays pistillate, yellow, often purplish beneath, not coiling. Disc florets perfect, yellow,
tube and limb weakly delimited. Achenes teretefusiform, appressed-villous, with 8–16 superficial
nerves, carpopodium asymmetric; pappus bristles
1-seriate, also a short outer series in 1 species.
x = 5. Three species, Mexico, Guatemala.
Compositae
951. Pityopsis Nutt.
Pityopsis Nutt., Trans. Amer. Philos. Soc. 2, 7: 317 (1840);
Semple & Bowers, Univ. Waterloo Biol. Ser. 29: 1–34 (1985),
rev.
Perennial herbs from short fibrous-rooted rhizomes, silky-sericeous with mostly appressed
hairs, often stipitate-glandular. Leaves mostly
basal or basal and cauline, reduced on upper stem,
linear to lanceolate or oblanceolate, sometimes
falcate, entire, parallel-veined. Inflorescence
corymbose, with few to many heads. Involucre
mostly cylindric-turbinate; bracts 4–8-seriate,
lanceolate, usually with apical green patch. Rays
8–35, pistillate, yellow, coiling. Disc corollas
gradually expanded, lobes deltate, erect. Achenes
fusiform, terete to slightly compressed, strigose;
pappus 2-seriate, outer bristles shorter. x = 9.
Circa seven species, south-eastern USA, one into
Mexico, Central America.
952. Tomentaurum G.L. Nesom
Tomentaurum G.L. Nesom, Phytologia 71: 129 (1991).
Perennial, scapose herbs from slender rhizomes,
tomentose, thicker uniseriate hairs absent, with
sessile or short-stipitate orange-tipped glands.
Leaves in basal rosette or with clasping bases on
lower part of stem, sessile, obovate to oblanceolate,
entire. Heads solitary, nodding in bud. Involucres
hemispheric; bracts 5–6-seriate, gradate, narrowly
ovate-lanceolate, margins purplish with hyaline
flange, outer bracts herbaceous, inner herbaceous only distally; receptacles plano-convex.
Rays 16–21, pistillate, yellow, coiling. Achenes
linear-oblong, compressed, each side 7–9-nerved,
densely white-sericeous; pappus of 45–60 white
bristles in several series, a few short outer setae.
One species, T. niveum (S. Wats.) G.L. Nesom,
north-western Mexico.
XV.18. Subtribe Conyzinae Horan. (1847).
Tribe Erigeroneae Gren. & Godr. (1850).
Annual to perennial herbs. Leaves alternate,
mostly entire, less commonly toothed to lobed.
Heads solitary to few in a loosely corymboid
inflorescence, densely paniculate in some Conyza;
involucral bracts persistent, not keeled, with
orange-resinous veins; receptacles usually convex,
epaleate. Ray florets 1(–3)-seriate; corollas white
to blue, rarely yellow, coiling, reflexing or straight.
Disc florets bisexual, with orange resin canals
339
along veins; style branch appendages deltate,
papillose. Achenes 2-nerved (rarely multinerved),
compressed, eglandular; setulae without anchorshaped tips; pappus 1–2(–3)-seriate, inner series
of barbellate bristles, outer of scales or setae. x = 9
(x = 5, reduced to 4 and 3 in Aphanostephus).
Key to the Genera
1. Pistillate florets tubular or with short erect limbs 2
– Pistillate florets with long limbs
3
2. Perennial herbs from woody rhizomes or tubers; achenes subcylindrical
954. Apopyros
– Annual or perennial herbs without tubers; achenes
compressed
955. Conyza
3. Shrubs
956. Darwiniothamnus
– Herbs
4
4. Achenes 4-angled to fusiform-cylindrical
5
– Achenes compressed
6
5. Leaves broad to linear-lanceolate to pinnatifid; hairs
of stems and leaves appressed to spreading; achenes
usually 4-angled, thick-walled, with subsurface nerves
953. Aphanostephus
– Leaves filiform to linear-oblanceolate; stems and
leaves long-pilose with ciliate hairs; achenes fusiformcylindrical, with numerous, raised, orange-resinous
nerves
960. Neja
6. Rays yellow; plants taprooted
958. Hysterionica
– Rays white or bluish to pinkish (rarely yellow); plants
taprooted or rhizomatous
7
7. Involucral bracts usually with 1 resinous longitudinal
vein
957. Erigeron
– Involucral bracts with 3 longitudinal veins
959. Leptostelma
Genera of Conyzinae
953. Aphanostephus DC.
Aphanostephus DC., Prodr. 5: 310 (1836); Turner, Phytologia 56: 81–101 (1984), rev.
Annual or perennial taprooted, erect to decumbent herbs; hairs soft to stiff. Leaves sessile or
petiolate, oblanceolate to linear-lanceolate, entire
to pinnatifid. Heads solitary; involucres broadly to
depressed hemispheric; bracts 3–5-seriate, weakly
gradate, ovate-lanceolate, not reflexing with age,
margins broadly scarious; receptacles convex to
conical. Rays pistillate, 1(–2)-seriate, white above,
with purplish median stripe beneath, closing upwards at night, not coiling. Disc corollas yellow,
tubular to funnelform, tube or limb sometimes
swollen and indurate, lobes deltate, erect. Achenes 4-angled or sometimes subterete, with 4–12
grooves or ribs, strigose or glabrous; pappus of
short cilia, or with awns with or without bristle
340
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
tips. x = 5 reduced to 4 or 3. Four species, southcentral USA, Mexico.
Aphanostephus differs in chromosome number
and morphology, achene morphology (including
vestiture), and pappus structure from all other
members of subtribe Conyzinae, but molecular
data (Noyes 2000) indicate that its evolutionary
origin was sister to a group of Caribbean species
of Erigeron L.
954. Apopyros G.L. Nesom
Apopyros G.L. Nesom, Phytologia 76: 177 (1994).
Erect perennial herbs, from rhizomes or tubers,
stiffly pilose-hirsute to nearly glabrous, eglandular. Leaves cauline, stiffly erect, shiny-indurate,
sessile, entire, 3(–5)-nerved, basal leaves scale-like.
Inflorescence 1-headed or thyrsoid-corymbose;
involucres broadly campanulate; bracts 3–4seriate, weakly gradate, with resinous midvein,
margins narrowly hyaline. Peripheral florets
pistillate, tubular, with 3–4 triangular teeth. Disc
florets narrowly tubular, base a wide indurate
disc. Achenes subcylindric, strigose, eglandular,
nerves 5, orange-resinous when young, becoming
paler; pappus bristles (1–)2-seriate, attenuate,
sometimes with short outer series. Two species,
Brazil, Paraguay, Argentina.
955. Conyza Less.
Conyza Less., Syn. Gen. Comp. 203 (1832), nom. cons.;
Cuatrecasas, Webbia 24: 198–228 (1969), reg. rev.; Wild,
Bol. Soc. Brot. 43: 247–276 (1969), reg. rev.; Zardini, Bol.
Soc. Argent. Bot. 17: 31–46 (1976), rev.
Conyzella Fabric. (1759).
Leptilon Raf. (1818).
Annual or perennial herbs, nearly glabrous to
coarsely hispid-pilose. Leaves linear to oblanceolate, entire to pinnatifid. Inflorescences broadly
ellipsoid to columnar or corymbose; involucres
cylindric to hemispheric; bracts 2–4-seriate, subequal to weakly gradate, outer bracts with 3 resinous
nerves; receptacle sometimes a hypanthium-like
cup. Peripheral florets pluriseriate, pistillate,
whitish, eradiate or with rudimentary limbs. Disc
florets 3–35, perfect, whitish, tubular-funnelform,
tube c. 2/3 of length. Achenes glabrous to sparsely
strigose; pappus bristles 1-seriate, sometimes
lengthening after anthesis, rarely with short outer
series of setae. x = 9. Circa 60–100 species, primarily tropical and subtropical, some introduced
pantropically.
Species of Conyza native to Africa apparently
are more similar and perhaps more closely related to genera of subtribe Grangeinae. Molecular
data indicate that American Conyza, as currently
viewed, is polyphyletic, having arisen several times
from within the nexus of Erigeron L. (Noyes 2000).
The taxonomy of subtribe Conyzinae is currently
under review (Nesom, unpubl. data).
956. Darwiniothamnus Harling
Darwiniothamnus Harling, Acta Horti Berg., Bd. 20, 3: 108
(1962).
Shrubs, glabrous to strigose, sometimes minutely
glandular. Leaves clustered at ends of branches,
narrowly linear to oblanceolate or lanceolate,
sessile or short-petiolate, entire to mucronate.
Inflorescences leafy-bracteate corymbose or
nearly 1-headed; buds erect; involucres cupulate to
hemispheric; bracts 4–6-seriate, strongly gradate,
1-nerved. Peripheral florets pistillate, pluriseriate,
whitish, limbs exceeding involucre, coiling. Disc
florets yellow, tubular-funnelform, constricted in
lower 1/2–1/3, limb sometimes strongly inflated.
Achenes oblong, sparsely to densely strigose,
somewhat dimorphic, ray achenes compressed,
2(–4)-nerved, disc achenes shorter and broader,
3–5(–6)-nerved; pappus bristles 1-seriate. x = 9.
Two species, Galapagos Islands.
957. Erigeron L.
Erigeron L., Sp. Pl. 683 (1753); Botschantzev, Fl. U.R.S.S. 25:
191–288 (1959), Engl. transl. 25: 180–269 (1999), reg. rev.;
Cronquist, Brittonia 6: 121–302 (1947), rev.; Cronquist,
Intermount. Fl. 5: 305–351 (1994), reg. rev.; Halliday, Fl.
Eur. 4: 116–120 (1976), reg. rev.; Nesom, Phytologia 66:
415–455 (1989), rev.; 67: 67–93 (1989), emend.; 72: 157–208
(1992), rev.; Solbrig, Contr. Gray Herb. 191: 3–79 (1962),
rev.
Trimorpha Cass. (1817).
Polyactis Less. (1832).
Achaetogeron A. Gray (1849).
Astradelphus Remy (1849).
Wyomingia A. Nelson (1899).
Annual to perennial herbs, rarely shrubs, with
caudex or rhizomes; stems, leaves, involucres
glabrous to variously pubescent, often with stipitate glands. Basal rosettes sometimes persistent,
cauline leaves linear to lanceolate or spathulate,
entire to pinnatifid. Inflorescences 1-headed
or corymbose to thyrsoid; involucres turbinate
Compositae
to hemispheric; bracts 2–4(–7)-seriate, equal
to strongly gradate, with 1(–3) resinous veins;
receptacles flat to conical. Rays 1–4-seriate, rarely
absent, white to blue or pink, rarely yellow, reflexing, coiling, or straight. Disc florets yellow, limb
sometimes inflated; style appendages lanceolate,
acute or obtuse. Achenes compressed with 2–4
ribs, seldom subterete with 6–14 ribs, glabrous
to strigose; pappus bristles 1-seriate, sometimes
caducous or lacking, sometimes with short outer
setae, scales, or a corona. x = 9. Circa 400 species,
North to South America, West Indies, Galapagos,
Eurasia.
In the analysis of molecular data by Noyes
(2000), subtribe Conyzinae comprises Erigeron,
American Conyza Less., Trimorpha Cass., Darwiniothamnus Harling, the four genera of the
Leptostelma group (Leptostelma D. Don, Hysterionica Willd., Neja D. Don, Apopyros Nesom),
and Aphanostephus DC., but the cladistically basal
and terminal taxa of the subtribe are Erigeron.
Formal recognition of any of these genera, for
consistency in cladistic classification, would
require further and much finer generic splitting
of species traditionally considered to be Erigeron.
Nesom (1989, 1990, 1994a) and Nesom and Noyes
(1999) have divided the species of ‘traditional’
North and Central American Erigeron into 21
sections, emphasizing variation in habit, vestiture,
inflorescence structure, bud position, lamina behaviour, cypsela morphology, and other features.
Australian species formerly identified as Erigeron
have been transferred to other genera (Nesom
1994b, 1998 [2000]), as has a Mexican species
(Nesom and Noyes 2000).
958. Hysterionica Willd.
Hysterionica Willd., Gesell. Naturf. Freunde Berlin Mag.
[Neuesten Entdeck. Gesamten Naturk.] 1: 140 (1807).
Annual or perennial taprooted herbs; caudex
simple; stems sometimes branched at base. Leaves
closely inserted, little reduced above, blades
oblanceolate, glandular-pubescent. Heads 1 to
several, peduncles short or long; involucres hemispheric; bracts 1–2-seriate, subequal, lanceolate,
acute, glandular-pubescent and hirsute outside.
Rays 1–3-seriate, pistillate, yellow or white, limbs
broad to filiform. Disc florets yellow, tubular,
lobes deltoid, erect. Achenes compressed, setulose;
pappus 2-seriate, of bristles and an outer series
of short linear scales. x = 9. Seven species,
south-eastern South America.
341
959. Leptostelma D. Don
Leptostelma D. Don in Sweet, Brit. Fl. Gard. ser. 2, 38 (1830).
Coarse perennial herbs, erect from decumbent
bases; stems, leaves and involucres often puberulous to hispid; stems often broad with broad pith.
Leaves cauline, herbaceous, broadly inserted, narrowed to base, rarely pseudopetiolate, blades ovate
to oblanceolate, venation pinnate, margins serrate.
Inflorescences corymbose to rather subumbellate;
heads large; peduncles short to long; involucres
campanulate; bracts 2–3-seriate, subequal, herbaceous to chartaceous, oblong-lanceolate, 3-nerved,
margins scarious. Rays 1–2-seriate, 80 or more,
pistillate, whitish to creamy or yellow, narrow,
not coiling. Disc florets yellow, tube slender, limb
narrowly funnelform, lobes 5, lanceolate, erect.
Achenes obovate-oblong, somewhat compressed,
2–3-nerved, mostly glabrous; pappus white,
1–3-seriate, of 30–40 flexuous bristles. x = 9. Five
species, south-eastern South America.
960. Neja D. Don in Sweet
Neja D. Don in Sweet, Hort. Brit. (Sweet), ed. 2, 299 (1830);
Espinar, Darwiniana 22: 537–549 (1980), reg. rev.
Perennial herbs, usually from taproot or branched
caudex. Leaves mostly basal, filiform to linearoblanceolate, crinkly-pilose with long cilia.
Heads solitary, long-pedunculate or on sparsely
bracteolate stems; involucre broadly turbinate to
hemispheric; bracts 2–4-seriate, gradate, narrowly
triangular. Rays pistillate, 1-seriate, yellow or
white. Disc florets yellow, tubular, lobes deltate,
erect. Achenes fusiform-cylindrical, with 7–10
prominulous, orange-resinous nerves, strigose
between nerves; pappus bristles 1–3-seriate, outer
series scarcely differing or of short setaceous
bristles or scales. Circa six species, south-eastern
South America, Cuba.
Recently Described or Reinstated Genera
not Included in the Key and Descriptions
Allittia P.S. Short, Muelleria 20: 54 (2004).
Segregate of Brachyscome. Two spp., south-eastern
Australia and Tasmania.
Cuniculotinus Urbatsch, R.P. Roberts and Neubig,
Sida 21: 1618 (2005).
Solidagininae. One sp., Cuniculotinus (Chrysothamnus) gramineus (H.M. Hall) Urbatsch, R.P. Ro-
342
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
berts & Neubig (= Petradoria discoidea L.C.
Anderson), western USA.
Herrickia Wooton & Standley [reinstated], Contr.
U.S. Natl Herb. 16: 186 (1913).
Eurybia sect. Herrickia (Wooton & Standley) G.L. Nesom.
Machaerantherinae grade. Four spp., southwestern USA.
Hullsia P.S. Short, Muelleria 20: 58 (2004).
Segregate of Brachyscome. One sp., H. argillicola
P.S. Short, north Australia.
Lorandersonia Urbatsch, R.P. Roberts and Neubig,
Sida 21: 1619 (2005).
Solidagininae. Seven spp., western USA and northwestern Mexico.
Nestotus R.P. Roberts, Urbatsch & Neubig, Sida 21:
1650 (2005).
Solidagininae. Two spp., western USA.
Novaguinea D.J.N. Hind, Kew Bull. 59: 177 (2004).
Lagenophorinae. One sp., N. rudalliae D.J.N. Hind,
New Guinea.
Pembertonia P.S. Short, Muelleria 20: 62 (2004).
Brachyscome sect. Heteropholis F. Muell. One sp.,
P. latisquamata (F. Muell.) P.S. Short, western Australia.
Toiyabea R.P. Roberts, Urbatsch & Neubig, Sida 21:
1652 (2005).
Solidagininae. One sp., T. (Haplopappus) alpina
(L.C. Anderson & S. Goodrich) R.P. Roberts, Urbatsch & Neubig, western USA.
Triniteurybia Brouillet, Urbatsch & R.P. Roberts,
Sida 21: 898 (2004).
Basal grade to Machaerantherinae. One sp., T. aberrans (A. Nelson) Brouillet, Urbatsch & R.P. Roberts,
western USA.
XVI. Tribe Anthemideae Cass. (1819).
Tribe Cotuleae Benth. (1868).
Tribe Ursinieae H. Rob. & Brettell (1973).
Ch. Oberprieler, R. Vogt and L.E. Watson
Annual, biennial or perennial, hapaxanthic or
pollacanthic herbs, subshrubs or shrubs, rarely
spinescent. Indumentum rarely absent, usually
of short biseriate glandular hairs (glands) and
uniseriate basifixed, T-shaped medifixed or stellate
hairs. Leaves usually alternate, rarely opposite or
basally crowded, generally variously dissected,
dentate, serrate, lobed, pinnatifid to pinnatisect,
rarely entire, sometimes vermiform, rarely succulent. Capitula solitary or in lax to dense corymbs,
panicles or racemes, or in glomerules, often
pedunculate, rarely sessile, heterogamous and
radiate or disciform, or homogamous and discoid.
Involucres often hemispherical, sometimes obconical, cylindrical or urceolate. Phyllaries in 2–7 rows,
imbricate, sometimes with one to several resin
canals, almost always with scarious margins and
apex. Receptacles flat, meniscoid, hemispherical,
conical or narrowly conical, glabrous or hairy,
paleate or epaleate. Paleae persistent or caducous,
flat or canaliculate, sometimes with a central resin
canal. Ray florets female and fertile or sterile, or
neuter; limb white, white with a yellow base, yellow,
or rarely blue-violet, pinkish or reddish. Outer
disc florets (in disciform capitula) in one to several
rows, female, usually fertile; corolla tubular and
with 0–5 apical lobes, usually yellow, rarely absent.
Central disc florets hermaphrodite or functionally
male; corolla tubular or funnel-shaped, actinomorphic, rarely slightly zygomorphic, yellow or rarely
whitish or reddish, with 3–6 apical lobes; lobes
rarely with central resin sacs. Stamens equal in
number to and alternating with the corolla lobes;
upper part of the filament with cells with thickened
walls, forming a split cylindrical or balusterform
anther collar (filament collar); anthers usually
with an ovate, triangular or subtriangular to
subulate apical appendage, generally rounded,
rarely shortly tailed at the base; pollen usually
spiny, sometimes rugose or smooth, tricolporate
(hexa-panto-colporate in Adenanthellum); exine
ecaveate (caveate in Ursinia), with a thick foot
layer, large basal columellae (lacking or only
occasionally or vestigially found in Ursinia) with
distal branches (branches complex and interwoven
in Artemisia and Crossostephium), and a double
Compositae
tectum formed by infratectal columellae of uniform length (Vezey et al. 1994). Style with a slender
or bulbous base, usually situated on a cup-shaped
stylopodium (nectary); style branches usually free,
sometimes (in functionally male florets) fused,
usually linear, rarely elliptic or ovate in outline,
apically truncate, penicillate, and usually with two
parallel, sometimes curved, papillate stigmatic
areas on their adaxial surfaces. Achenes various
in form, often obovoid, obconical or cylindrical
in outline and circular, triquetrous or angled in
cross-section, sometimes dorsiventrally flattened
and elliptic or rhombic in cross-section, usually
with more or less prominent ribs, sometimes even
winged, sometimes without ribs; apex ecoronate
and marginally rounded or truncate, or with
an entire to lacerate (sometimes adaxially more
developed) corona, or with a corona (‘pappus’)
formed by individual, sometimes basally fused,
scales or bristle-like scales, or with an (usually
adaxial) auricle; pericarp without a carbonised
layer, from extremely thin and delicate to thick
and sclerified, often with resin ducts or sacs and
specialised, myxogenic epidermal cells. Embryo
sac development monosporic or tetrasporic, rarely
bisporic. Base chromosome numbers generally
x = 9, sometimes x = 10, rarely x = 6, 7, 8, 11, 13
or 17.
Comprising 111 genera with c. 1,800 species,
distributed worldwide (extratropical) but with
main concentrations in central Asia, the Mediterranean region and southern Africa.
Members of the tribe are well known as
aromatic plants, and some are utilised for their
pharmaceutical and/or pesticidal value. Three
main classes of chemical substances are of
considerable systematic significance: acetylenes
(Greger 1977), sesquiterpene lactones (Seaman
1982) and flavonoids (Bohm and Stuessy 2001).
All recent molecular studies based on coding
and/or non-coding sequences from the chloroplast
genome (Kim and Jansen 1995; Bayer and Starr
1998; Panero and Funk 2002) indicate a more
or less well-supported sister-group relationship
of Anthemideae with Astereae. This is also supported by a cladistic study of the family based
on morphological and phytochemical characters
by Karis (1993b). The circumscription of Anthemideae remains relatively unchanged since
early, artificial classification systems (Lessing 1832;
Hoffmann 1894; Bentham 1873a) and more recent
ones (Poljakov 1967; Reitbrecht 1974; Heywood
and Humphries 1977; Bremer and Humphries
343
1993), with Cotula and Ursinia included in the
tribe despite considerable debate (Bentham
1873a; Robinson and Brettell 1973c; Heywood
and Humphries 1977; Jeffrey 1978; Gadek et al.
1989; Bruhl and Quinn 1990, 1991; Bremer and
Humphries 1993; Kim and Jansen 1995). The
tribe was monographed on the basis of a mostly
morphological phylogeny (Bremer and Humphries
1993), but there is little congruence with any of the
previous classifications. To complicate matters further, molecular phylogenies for the tribe as a whole
(Watson et al. 2000) are largely incongruent with
either the morphological phylogeny (Bremer and
Humphries 1993) or the previously proposed
classifications (Poljakov 1967; Reitbrecht 1974;
Heywood and Humphries 1977). This is also true
for molecular phylogenies for the Mediterranean
genera alone (Francisco-Ortega et al. 1997; Oberprieler and Vogt 2000; Oberprieler 2002, 2004a,
b, 2005). There is little support for monophyly of
most of the subtribes of Bremer and Humphries
(1993), and there is substantial disagreement with
previously proposed sister-group relationships
of genera and subtribes. A molecular phylogeny
for the tribe based on the chloroplast gene ndhF
(Watson et al. 2000) revealed a basal grade of
Southern Hemisphere genera which are sister to
a mostly Northern Hemisphere clade. The latter
consists of a basal grade of predominantly Asian
genera, which includes generic assemblages related
to Artemisia and Chrysanthemum (i.e. the former
Dendranthema), sister to a monophyletic group
of Mediterranean and more widespread, northwestern Eurasian genera. This monophyletic clade
was also found to be characterised by a large, 15-bp
deletion in nrDNA ITS (Francisco-Ortega et al.
1997; Oberprieler and Vogt 2000; Oberprieler 2002,
2004a, b, 2005). A strongly supported clade within
this monophyletic group included a large number
of Mediterranean and Macaronesian genera of
the tribe (e.g. genera allied to Argyranthemum,
Chamaemelum and Leucanthemum). In addition
to findings of ndhF (Watson et al. 2000) and
ITS studies (Francisco-Ortega et al. 1997), this
group is also characterised by an apomorphic
5-bp deletion in the trnL-trnF intergenic spacer
of the chloroplast genome (Oberprieler and Vogt
2000). Molecular studies also have helped to
resolve generic delimitation and circumscription
in several groups of the tribe, and have led to
their formal recognition. Oberprieler (2001), using
ITS data, provided evidence that Anthemis subg.
Anthemis is more closely related to Tripleuro-
344
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
spermum than to A. subg. Cota, which resulted in
the resurrection of Cota as an independent genus.
Oberprieler (2002) also clarified the delimitation
of Chamaemelum and Cladanthus which was
formalised with new combinations (Oberprieler
and Vogt 2002). Additional studies by Kornkven
et al. (1998, 1999) and Torrell et al. (1999) and
(more comprehensively) of Watson et al. (2002)
and Vallès et al. (2003) have shown that generic
delimitation in subtribe Artemisiinae sensu
Bremer and Humphries (1993) is problematic.
For this treatment, we followed the results of
the cited papers, insofar as the segregate genus
Seriphidium is merged with Artemisia. In addition,
the generic independence of several other small
genera of this subtribe (i.e. Sphaeromeria, Neopallasia, Crossostephium, Filifolium) is questionable
(Watson et al. 2002). However, we refrained from
merging them into Artemisia until a more complete sampling of molecular and morphological
data is available. Accordingly, due to the lack
of a comprehensive study for Tanacetum, it is
presently treated in the broad sense of Bremer and
Humphries (1993). Further investigations may lead
to considerable rearrangements in and around
this polyphyletic genus. Clearly, the subtribal
classification proposed by Bremer and Humphries
(1993) needs to be revised on the basis of expanded
molecular and more detailed morphological data.
In the present treatment, therefore, genera are
arranged in a linear manner according to the
results of Watson et al. (2000), i.e. in a primarily
geographic representation of the members of
the tribe, beginning with the southern African
representatives, followed by the central and eastern
Asian ones, and ending with the Mediterranean
genera. Within these four major groups (three
grades and the Mediterranean clade), genera are
arranged alphabetically. However, to retain as
much information as possible concerning several
generic assemblages, monophyletic groups of
genera were retained and named after one of their
members whenever possible.
Key to the Genera
1. Receptacle distinctly paleate with paleae subtending
florets (in Anthemis, rarely with the marginal part
of receptacle epaleate; in Schistostephium and Tanacetum, rarely with the central part of receptacle epaleate)
2
– Receptacle completely epaleate (sometimes pilose
or hirsute) or capitula few-flowered and presence of
paleae unclear
51
2. Ray florets present
3
– Ray florets absent
28
3. Limb of ray florets yellow or white with a yellow base,
rarely abaxially reddish
4
– Rays white (without a yellow base) or pink to reddish
on both sides
13
4. Phyllaries in 2 unequal rows, the outer scarious with
very wide margins, the inner connate and densely villous; paleae densely villous
967. Eriocephalus
– Phyllaries imbricate and subequal, not in 2 unequal
rows; paleae glabrous or sparsely villous
5
5. Achenes ± circular or quadrangular in cross-section,
with or without distinct ribs
6
– Achenes dorsiventrally flattened, with prominent lateral ribs or wings
11
6. Apex of achene with a corona of 5–10 large ovate or
circular scales and sometimes 5 additional subulate
scales
989. Ursinia
– Apex of achene with a shallow corona, an auricle, or
rarely a few small scales, or ecoronate
7
7. Achenes of disc florets with 5 distinct ribs
8
– Achenes of disc florets with c. 10 distinct, often tuberculate ribs, or without distinct ribs
9
8. Leaves linear, entire or apically few-lobed (southern
Africa)
989. Ursinia
– Leaves elliptical or obovate-spathulate, serrate (W
Mediterranean)
1060. Lepidophorum
9. Indumentum of stellate hairs
1054. Mecomischus
– Plants glabrous or with an indumentum of basifixed
or medifixed hairs
10
10. Corolla of disc florets basally saccate, adaxially clasping the upper half or more of achene; indumentum of
basifixed hairs
1053. Cladanthus
– Corolla of disc florets basally not saccate, not clasping
the apex of achene; indumentum of medifixed hairs or
lacking
1033. Anthemis
11. Achenes with distinct lateral wings, without additional
ribs
1029. Anacyclus
– Achenes without distinct lateral wings, if winged, then
wings only narrow and achenes with additional ribs
12
12. Achenes with 2 lateral ribs and rarely with an additional adaxial rib; pericarp rather thin
1028. Achillea
– Achenes with 2 lateral ribs or narrow wings and 3–10
additional ribs on each face; pericarp rather thick
1034. Cota
13. Achenes dorsiventrally flattened, with prominent lateral ribs or wings
14
– Achenes ± circular or quadrangular in cross-section,
with or without distinct ribs
18
14. Phyllaries in 2 unequal rows, the outer scarious with
very wide margins, the inner connate and densely villous; paleae densely villous
967. Eriocephalus
– Phyllaries imbricate and subequal, not in 2 unequal
rows; paleae glabrous or sparsely villous
15
15. Achenes with distinct lateral wings, without additional
ribs
16
– Achenes without distinct lateral wings, if winged, then
wings only narrow and achenes with additional ribs
17
16. Leaves vermiform; corolla of disc florets basally deeply
and equally saccate, clasping apex of achene both
adaxially and abaxially
1031. Leucocyclus
Compositae
– Leaves pinnatisect; corolla of disc florets basally only
slightly saccate, clasping apex of achene only adaxially
1029. Anacyclus
17. Achenes with 2 lateral ribs and rarely with an additional adaxial rib; pericarp rather thin 1028. Achillea
– Achenes with 2 lateral ribs or narrow wings and 3–10
additional ribs on each face; pericarp rather thick
1034. Cota
18. Phyllaries in 2 unequal rows, the outer scarious with
very wide margins, the inner connate and densely villous; paleae densely villous
967. Eriocephalus
– Phyllaries imbricate and subequal, not in 2 unequal
rows; paleae glabrous or sparsely villous
19
19. Achenes conspicuously villous 972. Lasiospermum
– Achenes glabrous, sometimes papillose
20
20. Indumentum of stellate hairs
1054. Mecomischus
– Plants glabrous or indumentum of basifixed or medifixed hairs
21
21. Phyllaries in 1–2 rows, readily deciduous at maturity;
paleae subulate, readily deciduous; apex of achenes
broadened into a bowl-shaped corona with radiating
teeth or lobes; achenes persistent at maturity
1033. Anthemis
– Phyllaries imbricate, not readily deciduous at maturity; paleae persistent or readily deciduous; apex of
achenes without a bowl-shaped corona; achenes deciduous or persistent at maturity
22
22. Apex of achenes with 5–10 large, obovate scales and
sometimes 5 additional subulate scales 989. Ursinia
– Apex of achenes with a shallow corona, an auricle, or
sometimes small scales, or ecoronate
23
23. Achenes without distinct ribs, sometimes 3–4-angled
or 3–4-ribbed
24
– Achenes with 5–20 distinct ribs
26
24. Anthers caudate
974. Osmitopsis
– Anthers ecaudate
25
25. Pericarp very thin, formed only by mucilage cells in
longitudinal rows; corolla of disc florets basally saccate, laterally clasping the apex of achene
1052. Chamaemelum
– Pericarp rather thick, with scattered mucilage cells;
corolla of disc florets basally not saccate
1033. Anthemis
26. Achenes with 5 ribs
1035. Gonospermum
– Achenes with (6–)10 or more ribs
27
27. Shrubs with opposite (rarely alternate) leaves; achenes
with 10–18 ribs; pericarp without myxogenic cells;
anthers with polarised endothecial tissue (southern
Africa)
984. Eumorphia
– Herbs or subshrubs with alternate leaves; achenes with
10 smooth or tuberculate ribs; pericarp with myxogenic cells; anthers with unpolarised endothecial tissue (N Hemisphere)
1033. Anthemis
28. Pericarp with resin sacs or ducts
29
– Pericarp without resin sacs or ducts (but often with
glandular hairs)
34
29. Achenes densely villous
972. Lasiospermum
– Achenes glabrous or with glandular hairs only
30
30. Achenes with 2 lateral and 0–1 adaxial ribs; pericarp
with a single resin sac or with 5 longitudinal resin
canals
31
– Achenes with 4–18 ribs (sometimes with 2 more distinct lateral ribs and 3 additional inconspicuous ones);
31.
–
32.
–
33.
–
34.
–
35.
–
36.
–
37.
–
38.
–
39.
–
40.
–
41.
–
42.
–
345
pericarp with longitudinal rows of resin sacs in ribs or
scattered epidermal resin sacs
32
Annual herbs; pericarp with a single resin sac apically
in the adaxial rib
1069. Lonas
Perennial herbs and shrubs; pericarp with 5 longitudinal resin canals along vascular bundles 1028. Achillea
Perennial herbs; pericarp with scattered epidermal
resin sacs; indumentum of basifixed hairs
1030. Heliocauta
Shrubs or shrublets; pericarp with longitudinal rows
of resin sacs in ribs; indumentum of stellate hairs or
absent
33
Achenes with a corona of basally united, fimbriate
scales; anthers with an apical appendage without
a resin sac
963. Hymenolepis
Achenes apically rounded or with a short, thickened
rim (sometimes with a pseudo-pappus of long-stalked
glands); anthers with an apical appendage with or
without a resin sac
962. Athanasia
Phyllaries in 2 unequal rows, the outer scarious with
very wide margins, the inner connate and densely villous; paleae densely villous
967. Eriocephalus
Phyllaries imbricate and subequal, not in 2 unequal
rows; paleae glabrous or sparsely villous
35
Apex of achenes with a corona of 5–10 large, obovate
scales and sometimes 5 additional subulate scales
989. Ursinia
Apex of achenes with a shallow corona, an auricle, or
of small scales, or ecoronate
36
Corolla of disc florets 4-lobed; central part of receptacle epaleate
987. Schistostephium
Corolla of disc florets 5-lobed; receptacle paleate
throughout
37
Achenes dorsiventrally flattened
38
Achenes circular or quadrangular in cross-section 41
Achenes with 2 lateral and 1 adaxial, flimsy ridges;
pericarp consisting only of large myxogenic cells in
longitudinal rows
1055. Rhetinolepis
Achenes with prominent lateral wings or ribs; pericarp
with scattered myxogenic cells
39
Achenes with distinct lateral wings, without additional
ribs
1029. Anacyclus
Achenes without distinct lateral wings, if winged, then
wings only narrow and achenes with additional ribs
40
Achenes with 2 lateral ribs and rarely with an additional adaxial rib; pericarp rather thin 1028. Achillea
Achenes with 2 lateral ribs or narrow wings and 3–10
additional ribs on each face; pericarp rather thick
1034. Cota
Corolla of disc florets basally saccate and clasping apex
or more of achene; achenes ecoronate
42
Corolla of disc florets basally not saccate, not clasping
apex of achene; achene coronate or ecoronate
44
Pericarp formed only of longitudinal rows of myxogenic cells; achenes only with 2 lateral and 1 adaxial,
flimsy ridges; indumentum absent or of basifixed hairs
1052. Chamaemelum
Pericarp without or with scattered myxogenic cells;
achenes 3–5-angled, with more conspicuous ribs; indumentum absent or of basifixed or medifixed hairs
43
346
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
43. Corolla of disc florets clasping only apex of achene;
indumentum absent or of medifixed hairs
1056. Santolina
– Corolla of disc florets clasping upper half of achene
(laterally even more); basal part of floret persistent
on achene; indumentum densely sericeous, greyishwhite, of basifixed hairs
1032. Otanthus
44. Capitula in a long panicle
999. Artemisia
– Capitula solitary or in lax to dense corymbs
45
45. Leaves opposite
985. Gymnopentzia
– Leaves alternate
46
46. Anthers caudate (southern Africa) 971. Inulanthera
– Anthers not caudate (N Hemisphere)
47
47. Indumentum exclusively of basifixed hairs
48
– Indumentum absent or of medifixed hairs
49
48. Stems sparsely branched and leafy; capitula in dense
corymbs
1013. Handelia
– Stems loosely branched and almost leafless; capitula
in lax corymbs
1017. Sclerorhachis
49. Capitula in dense corymbs
1035. Gonospermum
– Capitula solitary or in lax corymbs
50
50. Only marginal florets subtended by paleae; leaves pinnate with linear and pungent segments; plant suffruticose
1037. Tanacetum (T. paleaceum)
– All florets or at least the central ones subtended by
paleae; leaves variously divided, but never with pungent segments; annuals or perennials 1033. Anthemis
51. Ray florets present
52
– Ray florets absent
122
52. Ray florets yellow or white with a yellow base, rarely
abaxially reddish
53
– Rays florets white (without a yellow base) or pink,
reddish, bluish-violet on both sides
74
53. Corolla of disc florets apically 4-lobed
54
– Corolla of disc florets apically 5-lobed
55
54. Achenes tetragonal in cross-section, apex with minute
scales
970. Inezia
– Achenes dorsiventrally flattened, with 2 distinct lateral, wing-like ribs, apex ecoronate (rays not true ray
florets but outer disc florets with one corolla lobe expanded to a limb)
964. Cotula
55. Pericarp with resin sacs or ducts
56
– Pericarp without resin sacs or ducts
62
56. Resin sacs or ducts in ribs of achene
57
– Resin ducts vallecular between ribs of achene
58
57. Shrubs or shrublets; achenes circular in cross-section,
with 10–18 rounded ribs, some of them with resin sacs
986. Phymaspermum
– Annual herb; achenes dorsiventrally flattened, with 2
lateral broad wings, both with resin ducts
975. Adenoglossa
58. Adaxial ribs of achenes basally fused into a ± distinct
callus
59
– Ribs of achenes basally not fused into a callus
60
59. Achenes with an adaxial auricle as long as or even
longer than corolla
1064. Glossopappus
– Achenes with an adaxial auricle much shorter than
corolla
1063. Coleostephus
60. Limb of ray florets white with a yellow base
1066. Mauranthemum
– Limb of ray florets yellow, sometimes becoming red 61
61. Limb of ray florets golden yellow
1062. Chrysanthoglossum
– Limb of ray florets pale yellow
1068. Rhodanthemum (Rh. maresii)
62. Achenes of ray florets triquetrous in cross-section,
winged; achenes of disc florets laterally flattened and
adaxially and abaxially winged or sometimes terete to
prismatic
63
– Achenes of ray florets similar to achenes of disc florets;
all achenes circular (or sometimes triangular) in crosssection, always without wings
66
63. Shrubs or shrublets
1048. Argyranthemum
– Annual herbs
64
64. Indumentum of glandular hairs; wings of achenes with
apical spines
1050. Heteranthemis
– Indumentum absent or of basifixed, eglandular hairs;
wings of achenes (if present) without apical spines 65
65. Corolla of disc florets red; achenes of disc florets laterally flattened and adaxially and abaxially winged
1051. Ismelia
– Corolla of disc florets yellow; achenes of disc florets
prismatic with a narrow adaxial wing or terete and
inconspicuously ribbed
1049. Glebionis
66. Annual herbs
67
– Shrubs, subshrubs, or perennial herbs
70
67. Leaves 3(–5)-lobed
998. Tridactylina
– Leaves variously pinnatisect
68
68. Achenes 3-angled; ray florets fertile; indumentum absent
1058. Endopappus
– Achenes with 5–7 ribs; ray florets sterile; indumentum
of medifixed hairs
69
69. Limb of ray florets yellow; achenes of disc florets ecoronate, with 1 adaxial, 2 lateral and 2 abaxial ribs
1043. Prolongoa
– Limb of ray florets white with a yellow base; achenes of
disc florets with a scarious corona, and with 5–7 ribs
1041. Hymenostemma
70. Achenes densely hairy
1011. Trichanthemis
– Achenes glabrous or with sessile glands
71
71. Achenes without myxogenic cells 1037. Tanacetum
– Achenes with myxogenic cells on ribs
72
72. Apex of achenes with scarious corona
1042. Leucanthemopsis
– Apex of achenes ecoronate, marginally rounded 73
73. Basally strongly woody shrublets with few-lobed
leaves, lobes linear
993. Brachanthemum
– Perennial herbs or subshrubs; leaves with rather broad
lobes, rarely entire or serrate 994. Chrysanthemum
74. Annual herbs
75
– Shrubs, shrublets, or perennial herbs
95
75. Corolla of disc florets (3–)4-lobed
76
– Corolla of disc florets 5-lobed
80
76. Ray florets stalked
964. Cotula
– Ray florets sessile
77
77. Capitula very small (2–5 mm); phyllaries in 1–2 rows;
leaves 3–5-lobed
1036. Nananthea
– Capitula larger; phyllaries in 2–5 rows; leaves variously
pinnatisect
78
78. Achenes of central disc florets without myxogenic cells
981. Oncosiphon
– Achenes of central disc florets with myxogenic cells 79
79. Achenes with 5, mainly adaxial ribs 1044. Matricaria
– Achenes inconspicuously 3-ribbed or without distinct
ribs
977. Foveolina
80. Achenes with resin sacs or ducts
81
Compositae
– Achenes without resin sacs or ducts (but often with
glandular hairs)
86
81. Achenes with 10 ribs and 10 longitudinal, vallecular
resin ducts between ribs
1066. Mauranthemum
– Achenes without or with 3–5 ribs, with apical resin
sacs or resin ducts in ribs
82
82. Achenes with (1–)2(3–5) abaxial-apical resin sacs, and
2 lateral and 1 adaxial distinct ribs
1038. Tripleurospermum
– Achenes with resin canals or resin sacs in ribs
83
83. Achenes somewhat dorsiventrally flattened and with 2
lateral, inconspicuous ribs; corolla lobes with central
resin sacs
1047. Aaronsohnia
– Achenes with 3 or more ± distinct ribs, sometimes
slightly flattened; corolla lobes without or sometimes
with central resin sacs
84
84. Achenes of ray florets laterally winged, wings with apical teeth; apex with an adaxial auricle 1057. Daveaua
– Achenes of ray florets terete or sometimes flattened
but not winged, variously ribbed
85
85. Achenes with 5 adaxial-lateral ribs and a small corona
or auricle; lateral ribs with resin canals
1044. Matricaria
– Achenes with 3 adaxial-lateral, rounded ribs and 2
abaxial weaker ribs; 3–5 resin canals or apical resin
sacs in the ribs; apex of achene with a corona of 7–10
obovate scales
1059. Heteromera
86. Achenes actinomorphic, with 5–10 equidistantly arranged, equal ribs
87
– Achenes without ribs or with unequally arranged, ±
unequal ribs
90
87. Ray florets yellowish-white; leaves 3–5-lobed
998. Tridactylina
– Ray florets white; leaves variously pinnatisect
88
88. Achenes with myxogenic cells scattered over the pericarp; ribs rather inconspicuous; apex with an erosedenticulate, sometimes unilateral corona
1033. Anthemis
– Achenes with myxogenic cells on the ridges of the 10
distinct ribs, ecoronate
89
89. Indumentum of medifixed hairs; phyllaries with
brown scarious margins
1040. Castrilanthemum
– Indumentum absent or of basifixed hairs; phyllaries
with pale to light brown scarious margins
1070. Nivellea
90. Achenes with 1 adaxial and 2 lateral thick ribs and
with 2–3 adaxial weaker ribs; apex with an adaxial
stiff auricle
1071. Otospermum
– Achenes with mainly adaxial-lateral ribs or with inconspicuous ribs; apex with an entire or scaly corona,
or a scarious auricle
91
91. Pericarp and corona white and spongy, abaxially thin
(southern Africa)
977. Foveolina
– Pericarp not white and spongy; corona scarious (N
Hemisphere)
92
92. Achenes laterally pilose; apex of disc achenes with
a fimbriate whitish corona; apex of ray achenes with
auricle
1022. Microcephala
– Achenes glabrous, but often with sessile glands; apex
of achenes various
93
93. Achenes somewhat dorsiventrally flattened and with 2
lateral weak ribs; corolla lobes with central resin sacs
1047. Aaronsohnia
347
– Achenes with 3 or more adaxial and lateral, distinct
ribs; corolla lobes with or without central resin sacs 94
94. Achenes with 5 adaxial-lateral ribs; apex with an auricle or a small corona
1044. Matricaria
– Achenes with 3 adaxial-lateral ribs; apex with a stiff
corona
1058. Endopappus
95. Corolla of disc florets (3–)4-lobed
96
– Corolla of disc florets 5-lobed
100
96. Ray florets stalked
964. Cotula
– Ray florets sessile
97
97. Leaves entire, lanceolate to linear (ericoid)
98
– Leaves lobed or pinnatisect
99
98. Receptacle glabrous; pericarp without myxogenic cells
970. Inezia
– Receptacle hairy; pericarp with myxogenic cells in
rounded, scattered groups
988. Thaminophyllum
99. Receptacle hairy; leaves with few oblong to rounded,
mucronate lobes
973. Lidbeckia
– Receptacle glabrous; leaves 1–2-pinnatisect
968. Hilliardia
100. Apex of achenes with a corona of 4–12 linear to
obovate-oblong scales at least half as long as the
corolla, or with many (25–50) bristle-like scales as
long as or longer than the corolla
101
– Apex of achenes marginally rounded, or with an auricle, an entire corona, or a corona of short scales 103
101. Apex of achenes with 25–50 bristle-like scales equalling or exceeding the corolla in length 1007. Allardia
– Apex of achenes with a corona of 4–12 linear to
obovate-oblong scales half as long as or equalling the
corolla in length
102
102. Pericarp densely hairy
1011. Trichanthemis
– Pericarp glabrous
1010. Richteria
103. Achenes with resin sacs or canals
104
– Achenes without resin sacs or canals (but often with
glandular hairs)
110
104. Achenes with 5 or c. 10 longitudinal, vallecular resin
ducts between ribs
105
– Achenes with resin ducts or sacs in ribs, or with
abaxial-apical resin sacs
107
105. Achenes c. 1 mm long
1066. Mauranthemum
– Achenes more than 1 mm long
106
106. Leaves entire, dentate-serrate, or up to 3-pinnatisect,
sessile; florets not becoming reddish at maturity; achenes with c. 10 rounded ribs
1065. Leucanthemum
– Leaves 1–3-pinnatisect, long-petiolate; florets becoming reddish at maturity; achenes with 5 or c. 10 protruding, often wing-like ribs 1068. Rhodanthemum
107. Achenes circular or somewhat compressed in crosssection, with 10–18 distinct ribs
108
– Achenes triquetrous or flattened in cross-section, with
2 lateral and 1 adaxial ribs
109
108. Pericarp with myxogenic cells; achenes with 10–18
rounded ribs; resin sacs only in some of the ribs; shrubs
or shrublets
986. Phymaspermum
– Pericarp without myxogenic cells; achenes with 10
ribs, all of them with longitudinal resin canals; perennial herb
961. Adenanthellum
109. Achenes dorsiventrally flattened, laterally winged,
with 2 lateral resin canals
978. Leucoptera
– Achenes triquetrous, with 2 lateral and 1 adaxial ribs,
laterally not winged, with (1–)2(–3–5) abaxial-apical
resin sacs
1038. Tripleurospermum
348
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
110. Achenes of ray florets triquetrous, winged; achenes of
disc florets laterally compressed and 2-winged, sometimes terete
1048. Argyranthemum
– Achenes of ray florets and disc florets subequal, not
winged, terete
111
111. Pericarp with myxogenic cells
112
– Pericarp without myxogenic cells
116
112. Apex of achene with an entire or scaly corona
113
– Apex of achene marginally rounded
115
113. Indumentum of medifixed hairs; apex of achene with
an entire, flimsy corona
1042. Leucanthemopsis
– Indumentum of basifixed hairs; apex of achene with
an oblique, entire corona or with a corona of 2–6 scales
114
114. Apex of achene with a corona of 4–6 separate, mainly
abaxial, subulate scales unequal in length (eastern
Asia)
1024. Opisthopappus
– Apex of achene with an oblique, adaxially longer
corona, or of one large adaxial and one smaller abaxial
scale (South Africa)
976. Cymbopappus
115. Basally strongly woody shrublets; leaves few-lobed
with linear lobes; involucres hemispherical to cylindrical, up to 7 mm in diameter; capitula with 1–10 ray
florets; limb of ray florets up to 8 mm long, broadly
elliptical to elliptical
993. Brachanthemum
– Herbs or subshrubs; leaves entire, serrate, or with
rather broad lobes; involucres meniscoid, (6–)8–20(–
40) mm in diameter; capitula with more than 10 ray
florets; limb of ray florets 8–60 mm long, elliptical to
linear
994. Chrysanthemum
116. Leaves opposite
117
– Leaves alternate
118
117. Shrubs with entire to few-lobed leaves (southern
Africa)
984. Eumorphia (E. prostrata)
– Creeping, suffruticose perennial with serratepinnatifid leaves (Iberian peninsula)
1045. Phalacrocarpum
118. Apex of achenes with an entire, toothed, lobed, or
scaly, acute corona; receptacle glabrous
119
– Apex of achenes ecoronate, marginally rounded, or
with a rounded rim; receptacle glabrous or sparsely
hairy
120
119. Glabrous shrub; leaves serrate, crowded at the end of
the woody branches
1023. Nipponanthemum
– Perennial herbs or basally woody subshrubs; indumentum absent or present; leaves variously pinnatisect
1037. Tanacetum
120. Ray florets sterile; receptacle glabrous
1021. Leucanthemella
– Ray florets fertile; receptacle glabrous or sparsely hairy
121
121. Leaves entire, narrowly linear; receptacle sparsely
hairy
996. Hulteniella
– Leaves variously lobed or dissected; receptacle glabrous
992. Arctanthemum
122. Achenes with dark vallecular resin canals between pale
ribs and with myxogenic cells on the ribs
123
– Achenes without vallecular resin canals, with or without ribs and myxogenic cells
125
123. Annual herb
1061. Chlamydophora
– Perennial herbs or subshrubs
124
124. Adaxial ribs of achenes basally fused into a ± distinct
callus
1067. Plagius
– Achene base without an adaxial callus
1065. Leucanthemum
125. Disc florets apically (3–)4-lobed
126
– Disc florets apically 5(–6)-lobed
139
126. Achenes dorsiventrally flattened with 2 lateral ± distinct ribs or wings
127
– Achenes terete or only slightly dorsiventrally flattened,
sometimes triquetrous
130
127. Shrublets or subshrubs; capitula in lax to dense
corymbs (rarely solitary)
987. Schistostephium
– Annual or perennial herbs; capitula solitary
128
128. Capitula sessile in leaf axils; outer disc florets without
corolla and with style persistent and spinescent in fruit
966. Soliva
– Capitula pedunculate; outer disc florets with corolla
and with style not persistent in fruit
129
129. Outer disc florets with corolla inflated and with a hollow space between the outer and the inner epidermis;
central disc florets female-sterile
965. Leptinella
– Outer disc florets with corolla not inflated; central disc
florets fertile
964. Cotula
130. Outer disc florets with corolla inflated and with a hollow space between the outer and the inner epidermis
965. Leptinella
– Outer disc florets with corolla not inflated
131
131. Perennials
132
– Annuals
133
132. Capitula discoid; all florets hermaphrodite; achenes
with 5 distinct ribs and an apical corona of scales;
pollen spiny; indumentum of basifixed hairs
1014. Lepidolopsis
– Capitula disciform, with an outer row of female
florets; achenes without conspicuous ribs, apically
rounded; pollen smooth; indumentum of medifixed
hairs
1001. Filifolium
133. Capitula disciform, with an outer row of female florets
134
– Capitula discoid; all florets hermaphrodite
136
134. Delicate, somewhat succulent herb; capitula smaller
than 5 mm in diameter
1036. Nananthea
– Habit variable; capitula usually larger than 5 mm in
diameter
135
135. Pericarp either white and spongy with 1 adaxial and
2 lateral weak ribs, abaxially thin, or achenes almost
terete and thin-walled all around
97. Foveolina
– Pericarp not white and spongy; achenes with 5 adaxiallateral ribs
1044. Matricaria
136. Achenes without myxogenic cells
981. Oncosiphon
– Achenes with myxogenic cells
137
137. Achenes with an apical corona covered with myxogenic cells
980. Myxopappus
– Achenes ecoronate or with a corona not covered with
myxogenic cells
138
138. Achenes with myxogenic cells on the abaxial surface
(southern Africa)
977. Foveolina
– Achenes with myxogenic cells on all sides (northern
Africa, Near East)
1039. Brocchia
139. Capitula disciform, with an outer row of female florets
140
– Capitula discoid; all florets hermaphrodite
163
140. Pollen spiny (sometimes with short spines only) 141
– Pollen smooth
150
141. Achenes triquetrous with 1 adaxial and 2 lateral, thick
ribs and sometimes with 2 abaxial weaker ribs, abaxi-
Compositae
–
142.
–
143.
–
144.
–
145.
–
146.
–
147.
–
148.
–
149.
–
150.
–
151.
–
152.
–
153.
–
154.
–
155.
–
156.
–
157.
–
158.
–
159.
–
ally and apically with 2 distinct (sometimes 1 or 3–5)
resin sacs
1038. Tripleurospermum
Achenes various, not with abaxial-apical resin sacs 142
Annuals
143
Perennial herbs, subshrubs or shrubs
145
Achenes somewhat dorsiventrally flattened and with
2 lateral weak ribs
1047. Aaronsohnia
Achenes circular in cross-section
144
Pericarp either white and spongy with 1 adaxial and
2 lateral weak ribs, abaxially thin, or achenes almost
terete and thin-walled all around
977. Foveolina
Pericarp not white and spongy; achenes with 5 adaxiallateral ribs
1044. Matricaria
Achenes dorsiventrally flattened, with or without lateral wings
969. Hippia
Achenes terete, without wings
146
Achenes sparsely to densely hairy; receptacle hairy
1006. Artemisiella
Achenes glabrous; receptacle glabrous
147
Style branches brownish; corolla lobes erect; pericarp
without myxogenic cells
997. Phaeostigma
Style branches yellowish; corolla lobes spreading; pericarp with or without myxogenic cells
148
Pericarp with myxogenic cells
990. Ajania
Pericarp without myxogenic cells
149
Achene apex marginally rounded
990. Ajania
Achene apex with a short to well-developed, entire
to dentate, acute rim (or sometimes with an adaxial
auricle)
1037. Tanacetum
Achene apex with a corona or with small scales
1000. Crossostephium
Achene apex marginally rounded, ecoronate
151
Capitula arranged in lax to dense corymbs, generally
erect
152
Capitula arranged in long panicles, rarely in racemes
or spiciform or subglobose synflorescences, sometimes pendent
154
Annual herbs
991. Ajaniopsis
Perennial herbs
153
Central disc florets functionally male, compressed into
a resinous mass; achenes obliquely obovoid, always
without ribs (E Asia)
1001. Filifolium
Central disc florets hermaphrodite; achenes rarely
obovoid, usually obovoid-oblong to cylindrical and
with 3–5 ribs (North America) 1005. Sphaeromeria
Corolla lobes glabrous
155
Corolla lobes hairy
157
Capitula 30–50-flowered
1005. Sphaeromeria
Capitula usually with fewer than 30 florets
156
Central disc florets of two kinds; outer perfect, inner
completely sterile with reduced ovaries; annual or biennial herbs
1003. Neopallasia
Central disc florets all perfect or all female-sterile with
reduced ovaries; annuals, biennials, and perennials
999. Artemisia
Outer female florets without corolla; indumentum of
stellate hairs
1002. Mausolea
Outer female florets with corolla; indumentum absent
or of basifixed, medifixed, or stellate hairs
158
Indumentum of stellate hairs
1019. Kaschgaria
Indumentum absent or of basifixed or medifixed hairs
159
Achenes hairy
160
Achenes glabrous (but with glands)
161
349
160. Shrublet with older branches forming long spines
1004. Picrothamnus
– Plant without spinescent branches
999. Artemisia
161. Capitula 30–50-flowered
1005. Sphaeromeria
– Capitula usually with fewer than 30 florets
162
162. Capitula densely congested into glomerules arranged
in a spike, or rarely capitula solitary in interrupted,
partly congested spikes; marginal female florets in the
axils of canaliculate involucral bracts
1027. Turaniphytum
– Capitula usually forming a long panicle, sometimes
much reduced and racemose, spiciform or subglobose;
involucral bracts not canaliculate
999. Artemisia
163. Achenes triquetrous with 1 adaxial and 2 lateral, thick
ribs and sometimes with 2 abaxial weaker ribs, abaxially and apically with 2 distinct (sometimes 1 or 3–5)
resin sacs
1038. Tripleurospermum
– Achenes various, not with abaxial-apical resin sacs 164
164. Capitula in panicles, in racemes, or in spiciform to
subglobose synflorescences (densely aggregate at the
stems in Marasmodes)
165
– Capitula solitary or in lax to dense corymbs, sometimes in a pseudo-umbel
167
165. Apex of achenes marginally rounded, ecoronate
999. Artemisia
– Apex of achenes with a corona of scales
166
166. Pericarp without myxogenic cells 1014. Lepidolopsis
– Pericarp with myxogenic cells mainly abaxially and on
the ribs
979. Marasmodes
167. Annual herbs
168
– Perennial herbs, subshrubs, or shrubs (in Pseudohandelia, sometimes biennial and monocarpic)
175
168. Pollen smooth
995. Elachanthemum
– Pollen spiny
169
169. Pericarp of all achenes without myxogenic cells
1037. Tanacetum
– Pericarp of at least the central achenes with myxogenic
cells
170
170. Achenes without ribs or with 2 lateral weak ribs 171
– Achenes with 3 or more, ± distinct ribs
172
171. Achenes without ribs, circular in cross-section; apex
truncate to marginally rounded; pericarp with longitudinal rows of myxogenic cells; corolla lobes without
central resin sacs; indumentum of medifixed hairs
1025. Stilpnolepis
– Achenes somewhat dorsiventrally flattened and with 2
lateral weak ribs; apex with an adaxial auricle, sometimes marginally rounded; pericarp with myxogenic
cells abaxially; corolla lobes sometimes with central
resin sacs; indumentum of basifixed hairs
1047. Aaronsohnia
172. Achenes with unbranched hairs adaxially between ribs
1022. Microcephala
– Achenes glabrous
173
173. Indumentum absent or of medifixed hairs
1033. Anthemis
– Indumentum of basifixed hairs
174
174. Achene apex with a corona covered with myxogenic
cells
980. Myxopappus
– Achene apex marginally rounded or with a corona not
covered with myxogenic cells
977. Foveolina
175. Achenes with a corona of 4–20 linear, bristle-like to
obovate-elliptical scales at least as long as the achene
176
350
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
– Achenes ecoronate or with a corona of short scales, an
auricle, or an acute or rounded rim
178
176. Pericarp glabrous or sparsely hairy; corona scales
brownish
1008. Cancrinia
– Pericarp densely hairy; corona scales white
177
177. Capitula on short, nodding peduncles
1012. Ugamia
– Capitula on elongated, straight peduncles
1011. Trichanthemis
178. Achenes with myxogenic cells
179
– Achenes without myxogenic cells
185
179. Achenes with 10–18 ribs, some with resin sacs
986. Phymaspermum
– Achenes with fewer than 10 ribs, always without resin
sacs
180
180. Corolla tube swollen and with thick vascular strands;
shrubs or subshrubs with ± ericoid leaves
181
– Corolla tube not or only slightly swollen and with thin
vascular strands; perennial herbs or subshrubs with
few-lobed, pinnatifid, or pinnatisect leaves
182
181. Achene apex with 7–10 elliptical, adaxially longer
scales
979. Marasmodes
– Achene apex with an adaxial auricle, a corona of 3–
5 adaxially longer scales, or marginally rounded and
ecoronate
982. Pentzia
182. Achenes ecoronate
183
– Achenes with a corona of scales
184
183. Achenes narrowly cylindrical and somewhat arcuate;
pericarp tuberculate; capitula in a dense corymb or
a pseudo-umbel
1016. Pseudohandelia
– Achenes obovoid and ± straight; pericarp smooth; capitula solitary or in lax corymbs 993. Brachanthemum
184. Apex of achenes with an often oblique corona (adaxially larger than abaxially) formed by free or basally
united scales; capitula usually in lax to dense corymbs,
rarely solitary; involucre hemispherical, with phyllaries in 2–3 rows
1026. Tanacetopsis
– Apex of achenes with an adaxial auricle or 3–6 adaxial
scales; capitula usually solitary, rarely in lax corymbs;
involucre usually cylindrical to conical, with phyllaries
in 4–5 rows, rarely hemispherical 1046. Xylanthemum
185. Anthers tailed
186
– Anthers not tailed
187
186. Achene apex with a corona of small, entire to toothed
scales terminating each of the 8–10 ribs
971. Inulanthera
– Achene apex bordered by a ± distinct, rounded rim;
achenes with 5–8 inconspicuous ribs; ribs sometimes
with longitudinal resin canals
1018. Hippolytia
187. Pericarp conspicuously tuberculate or rugose
983. Rennera
– Pericarp with smooth longitudinal ribs
188
188. Achenes with longitudinal rows of resin sacs in ribs
189
– Achenes without any resin sacs or canals
190
189. Achene apex with a corona of basally united, fimbriate
scales
963. Hymenolepis
– Achene apex marginally rounded or sometimes with
a short, thickened rim
962. Athanasia
190. Leaves opposite
985. Gymnopentzia
– Leaves alternate or basally crowded
191
191. Capitula small and numerous, in a dense corymb;
plant basally villous and with 3–4-pinnatisect leaves
1015. Polychrysum
– Capitula various in size, solitary, or in lax to dense
corymbs; plants basally not densely villous; leaves entire, dentate, or 1–4-pinnatisect
192
192. Involucre cylindrical to conical; phyllaries in 4–7 rows
1020. Lepidolopha
– Involucre hemispherical; phyllaries in 1–4 rows 193
193. A compact, basally woody subshrub; indumentum
of basifixed hairs; capitula solitary; phyllaries in 1–2
rows, all of almost the same length; receptacle sparsely
hairy
1009. Cancriniella
– Perennial herbs; indumentum absent or of medifixed
and/or basifixed hairs; capitula solitary or in lax to
dense corymbs; phyllaries in 2–5 rows, of different
lengths; receptacle glabrous or sparsely hairy
194
194. Apex of achenes with an adaxial auricle or with a crenate to dentate rim
1037. Tanacetum
– Apex of achenes with an often oblique (adaxially more
developed) but abaxially developed corona formed by
free or basally united scales
1026. Tanacetopsis
XVI. Group A
[Southern Hemisphere Genera]
961. Adenanthellum B. Nord.
Adenanthellum B. Nord., Bot. Notiser 132: 160 (1979);
Nordenstam, Bot. Notiser 129: 137–165 (1976), rev.
Adenanthemum B. Nord. (1976), non H. Conwentz (1886).
Perennial herb. Indumentum of basifixed hairs.
Leaves alternate, serrate, glandular-punctuate.
Capitula solitary or in lax corymbs, more or
less distinctly pedunculate, radiate. Involucre
broadly campanulate. Phyllaries in 3–4 rows, with
scarious margins. Receptacle convex, glabrous,
epaleate. Ray florets female, fertile; limb white,
confluent with the achene, tube absent. Disc florets
hermaphrodite, fertile; corolla 5-lobed, yellow,
confluent with the achene; tube not spongy;
apical anther appendage ovate-elliptical. Achenes
oblong, with 10 ribs, somewhat compressed; apex
marginally rounded; pericarp without myxogenic
cells, with 10 resin canals in the ribs. One species,
A. osmitoides (Harv.) B. Nord. South Africa,
Swaziland.
XVI.1. Athanasia Group (962–963)
962. Athanasia L.
Fig. 69
Athanasia L., Sp. Pl., ed. 2: 1180 (1763); Källersjö, Opera
Bot. 106: 1–75 (1991), rev.
Morysia Cass. (1824).
Holophyllum Less. (1832).
Stilpnophyton Less. (1832).
Pristocarpha E. Mey. ex DC. (1838).
Saintmorysia Endl. (1838).
Bembycodium Kunze (1842).
Compositae
351
section and with 5–12(–18) sharply angled ribs, or
dorsiventrally flattened and with two lateral and
one adaxial rib; apex marginally rounded or with
a short, thickened rim; pericarp with longitudinal rows of resin sacs in ribs. x = 8 (polyploidy).
Thirty-nine species, South Africa, Namibia.
963. Hymenolepis Cass.
Hymenolepis Cass., Bull. Sci. Soc. Philom. Paris 1817: 138
(1817); Bremer & Källersjö, Nordic J. Bot. 5: 517–520 (1985),
rev.
Oligodora DC. (1838).
Phaeocephalus S. Moore (1900).
Shrubs. Indumentum absent or of stellate hairs.
Leaves alternate, entire, dentate or lobed. Capitula in dense corymbs, discoid, shortly pedunculate.
Involucre narrowly ellipsoid-cylindrical to obconical. Phyllaries in 3–4 rows, without or with only
narrow scarious margins. Receptacle flat, paleate,
rarely epaleate; paleae flat, narrowly elliptical. Florets hermaphrodite, fertile; corolla 5-lobed, yellow.
Achenes obovoid, circular in cross-section, with
5–10 rounded ribs; apex with a corona of basally
united, fimbriate scales; pericarp with longitudinal rows of resin sacs in ribs. Seven species, South
Africa.
XVI.2. Cotula Group (964–966)
964. Cotula L.
Fig. 69. Compositae-Anthemideae. Athanasia dentata.
A Habit. B Capitulum. C Tubular floret. D Achene.
(Drawings by Ch. Oberprieler)
Asaemia (Harv.) Harv. ex Benth. (1873).
Shrubs or shrublets. Indumentum absent or of stellate, rarely basifixed hairs. Leaves alternate, rarely
opposite, entire to pinnatifid, sometimes succulent. Capitula in dense corymbs and pedunculate
or solitary and sessile along branches, discoid. Involucre narrowly ellipsoid-cylindrical, hemispherical or spherical to urceolate. Phyllaries in 2–5
rows, often containing resin sacs. Receptacle flat
to hemispherical, paleate, rarely epaleate; paleae
flat or canaliculate, narrowly elliptical, rarely with
a central resin duct. Florets hermaphrodite, fertile;
corolla 5-lobed, yellow or whitish, tube with longstalked glands, scattered or aggregated into a basal
ring, rarely with short-stalked glands, or glabrous.
Achenes obovoid or cylindrical, circular in cross-
Cotula L., Sp. Pl.: 891 (1753).
Cenia Comm. ex Juss. (1789).
Lancisia Lam. (1797).
Strongylosperma Less. (1832).
Machlis DC. (1837).
Otochlamys DC. (1838).
Pleiogyne Koch (1843).
Sphaeroclinium (DC.) Sch. Bip. (1844).
Gymnogyne Steetz (1845).
Ctenosperma Hook. f. (1847).
Annual or perennial herbs. Indumentum absent
or of basifixed hairs. Leaves alternate, sometimes
opposite or basally crowded, lobed or 1–3pinnatisect, sometimes entire. Capitula solitary,
discoid or disciform, sometimes shortly radiate,
pedunculate; peduncles sometimes inflated. Involucre hemispherical or obconical. Phyllaries in
2–3 rows, with central resin ducts and pale or light
to dark brown scarious margins. Receptacle flat to
hemispherical, epaleate. Ray florets or outer disc
florets female, fertile, stalked; limb (when present)
352
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
white. Inner disc florets hermaphrodite, fertile;
corolla 3–4-lobed, yellow, basally sometimes
adaxially saccate; lobes usually with central resin
canals, one lobe occasionally forming a radiate
limb. Achenes obovoid, dorsiventrally flattened
and elliptical in cross-section, with 2 distinct,
lateral, wing-like ribs; apex marginally rounded,
ecoronate; pericarp rather thin, generally hairy,
sometimes with myxogenic cells and/or 2 lateral
resin canals. x = 8, 9, 10 (polyploidy). Fifty-five
species, southern and eastern Africa, Australia,
South America, Mexico, southern oceanic islands.
Some species widespread and naturalised.
narrowly elliptical in cross-section, with 2 lateral,
wing-like ribs; ribs sometimes projecting into
apical teeth; apex marginally rounded; pericarp
glabrous, without myxogenic cells or resin sacs.
x = 10 (polyploidy). Eight species, South America,
North America, Australia. One species widespread
as weed.
965. Leptinella Cass.
Shrubs. Indumentum of basifixed hairs. Leaves opposite, sometimes alternate, often fascicled on lateral brachyblasts, mostly ericoid, entire or lobed
to pinnatisect. Capitula in dense corymbs, radiate
or disciform. Involucre hemispherical or urceolate.
Phyllaries in 2 unequal rows, the outer scarious,
with very wide brown to reddish scarious margins,
the inner (= paleae of marginal florets) connate
and hairy. Receptacle flat, paleate; paleae elliptical
to linear, canaliculate, villous. Ray florets or outer
disc florets female, fertile; limb (when present)
yellow, white or reddish. Inner disc florets functionally male, with fused style branches; corolla 5lobed, yellow or mauve. Achenes obovoid, circular
or dorsiventrally flattened and elliptical in crosssection, with 5 equally arranged or 2 lateral ribs;
apex marginally rounded; pericarp lanate or pilose,
without myxogenic cells or resin sacs. x = 9 (polyploidy). Thirty-two species, South Africa, Namibia,
Botswana, Lesotho.
Leptinella Cass., Bull. Sci. Soc. Philom. Paris 1822: 127
(1822); Lloyd, N. Z. J. Bot. 10: 277–372 (1972), rev.
Symphyomera Hook. f. (1847).
Perennial or facultative annual herbs. Indumentum of basifixed hairs. Leaves alternate or opposite,
pectinate or 1–2-pinnatisect. Capitula solitary,
discoid or disciform, pedunculate. Involucre
hemispherical, sometimes umbonate. Phyllaries in
2 rows, with rather narrow, pale scarious margins.
Receptacle flat to conical, epaleate. Outer florets
female, fertile, tubular, yellow, apically 4-lobed,
inflated to form a hollow space between the outer
and inner epidermis. Inner florets functionally
male; corolla yellow, apically 4-lobed. Achenes
obovoid to cylindrical, circular or dorsiventrally
flattened and elliptic in cross-section, without
distinct ribs; apex marginally rounded; pericarp
glabrous or hairy, sometimes with myxogenic cells
and/or minute resin canals. x = 13 (polyploidy).
Thirty-three species, New Guinea, Australia, New
Zealand, South America, Falkland Islands.
967. Eriocephalus L.
Eriocephalus L., Sp. Pl.: 926 (1753); Müller, Herman & Kolberg, Flora of Southern Africa 33, 4/1: 1–63 (2001), rev.
Cryptogyne Cass. (1827), non Hook. (1876).
Monochlaena Cass. (1827).
968. Hilliardia B. Nord.
Hilliardia B. Nord., Opera Bot. 92: 147 (1987).
966. Soliva Ruiz & Pav.
Soliva Ruiz & Pav., Fl. Peruv. Prodr.: 113 (1794).
Gymnostyles Juss. (1804).
Annual herbs. Indumentum of basifixed hairs.
Leaves alternate, 2–3-pinnatisect. Capitula solitary, sessile in leaf axils, disciform. Involucre
hemispherical. Phyllaries in 2 rows, with narrow,
pale or light brown scarious margins. Receptacle
hemispherical to conical, epaleate. Outer florets
female, fertile; tube and limb absent; style spinescent at maturity. Inner florets functionally male;
corolla 3–4-lobed, yellow; style branches fused.
Achenes obovoid, dorsiventrally flattened and
Scrambling shrub. Indumentum of long and fine,
basifixed hairs. Leaves alternate, 1–2-pinnatisect.
Capitula in lax corymbs, pedunculate, radiate. Involucre hemispherical. Phyllaries in 2–3 rows, with
broad, pale or light brown to reddish scarious margins. Receptacle conical to elongated, epaleate. Ray
florets female, fertile; limb white, apically deeply
bifid. Disc florets hermaphrodite, fertile; corolla 4lobed, yellow. Achenes obovoid to subglobose, circular in cross-section, with 2–3 thin adaxial ribs;
apex marginally rounded; pericarp very thin and
translucent, glabrous, without myxogenic cells or
resin sacs. x = 10. One species, H. zuurbergensis
(Oliver) B. Nord., South Africa.
Compositae
969. Hippia L.
Hippia L., Mant. Pl.: 158, 291 (1771).
Shrublets or subshrubs. Indumentum of basifixed
hairs. Leaves alternate, entire, dentate to pinnatisect. Capitula in lax to dense corymbs, rarely solitary, disciform. Involucre hemispherical. Phyllaries
in 3 rows, with narrow, pale scarious margins. Receptacle hemispherical to conical, epaleate. Outer
florets female, fertile; limb absent; tube much reduced. Inner florets functionally male, with fused
style branches; corolla 5-lobed, sometimes with 2
larger and 3 smaller lobes, white or yellow. Achenes
obovoid, dorsiventrally flattened, narrowly elliptical in cross-section, with 2 lateral, wing-like ribs;
apex marginally rounded; pericarp glabrous, papillose to sparsely pubescent, without myxogenic
cells or resin sacs. Eight species, South Africa.
970. Inezia E. Phillips
Inezia E. Phillips, Bull. Misc. Inform. 1932: 297 (1932).
Perennial herbs. Indumentum of basifixed hairs.
Leaves alternate, entire. Capitula solitary, pedunculate, radiate. Involucre obconical to hemispherical.
Phyllaries in 3–5 rows, with narrow scarious
margins. Receptacle conical, epaleate. Ray florets
female, fertile or sterile; limb yellow or white,
adaxially and basally laterally pilose, confluent
with the achene, tube short or absent. Disc florets
hermaphrodite, fertile; corolla 4-lobed, yellow.
Achenes obovoid to cylindrical, dorsiventrally
slightly flattened, 4-angled; apex marginally
rounded or with minute scales; pericarp without
myxogenic cells or resin sacs. x = 10. Two species,
South Africa, Swaziland.
971. Inulanthera Källersjö
Inulanthera Källersjö, Nordic J. Bot. 5: 539 (1986).
Shrubs or subshrubs. Indumentum absent or of
basifixed hairs. Leaves alternate, entire to lobed.
Capitula in lax corymbs, discoid. Involucre hemispherical to spherical. Phyllaries in 3 rows, with
narrow, pale to light brown scarious margins. Receptacle flat, paleate, rarely epaleate; paleae linear, flat to shallowly canaliculate, with a central
resin duct. Florets hermaphrodite, fertile; corolla
5-lobed, yellow; anthers basally caudate. Achenes
obovoid, circular in cross-section, with (8–)10 ribs;
apex with a corona of small, entire to toothed scales
terminating each rib; pericarp smooth, sometimes
papillose, without myxogenic cells or resin sacs, but
353
with protruding, light brown glands. Ten species,
South Africa, Swaziland, Lesotho, Angola, Zimbabwe, Madagascar.
972. Lasiospermum Lag.
Lasiospermum Lag., Gen. Sp. Pl.: 31 (1816); Müller, Herman
& Kolberg, Flora of Southern Africa 33, 4/1: 64–73 (2001),
rev.
Annual or perennial herbs. Indumentum absent
or of long basifixed hairs. Leaves alternate, 1–2pinnatisect, rarely entire. Capitula solitary or in lax
corymbs, radiate or discoid, pedunculate. Involucre hemispherical. Phyllaries in 2–4 rows, usually
with several longitudinal resin ducts and light scarious margins. Receptacle flat to conical, paleate;
paleae flat, elliptical to obovate, with a central resin
duct. Ray florets (when present) female, fertile;
limb white or reddish. Disc florets hermaphrodite,
fertile; corolla 5-lobed, yellow or reddish. Achenes
obovoid, circular in cross-section, with 6 ribs; apex
marginally rounded; pericarp densely villous, with
longitudinal rows of resin sacs in ribs. x = 9,
10. Four species, South Africa, Namibia, Lesotho,
Egypt.
973. Lidbeckia P.J. Bergius
Lidbeckia P.J. Bergius, Descr. Pl. Cap.: 306 (1767).
Subshrubs. Indumentum of basifixed hairs. Leaves
alternate, glandular-punctate, pinnately to digitately lobed. Capitula solitary, pedunculate, radiate.
Involucre hemispherical to cylindrical. Phyllaries
in 3–4 rows, without scarious margins. Receptacle
convex, hairy, epaleate. Ray florets female, sterile
or neuter; limb white, sometimes dorsally pilose,
tube sometimes laterally pilose, confluent with the
achene. Disc florets hermaphrodite, fertile; corolla
4-lobed, yellow, sometimes pilose; stylopodium
large and persistent in fruit. Achenes ellipsoid,
laterally pilose, with 3–8 ribs; apex marginally
rounded. Two species, South Africa.
974. Osmitopsis Cass.
Osmitopsis Cass., Bull. Sci. Soc. Philom. Paris 1817: 154
(1817); Bremer, Bot. Notiser 125: 9–48 (1971); Bremer, Bot.
Notiser 129: 21–24 (1976), rev.
Shrubs or subshrubs. Indumentum absent or of
basifixed hairs. Leaves alternate, entire to lobed.
Capitula solitary or in lax corymbs, radiate. Involucre campanulate. Phyllaries in 2–4 rows, often with scarious margins. Receptacle flat to con-
354
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
ical, paleate; paleae narrowly elliptical to obovate,
canaliculate and enclosing the florets. Ray florets
female, fertile, sterile or neuter; limb white, occasionally pilose abaxially, tube occasionally pilose.
Disc florets hermaphrodite and fertile or functionally male; corolla 5-lobed, yellow; anthers basally
caudate; stylopodium sometimes large and persistent in fruit. Achenes obovoid, ellipsoid or obovate,
3–4-angled or -ribbed; apex with a corona of subulate to triangular, basally fused scales, or marginally
rounded. x = 10. Nine species, South Africa.
XVI.3. Pentzia Group (975–983)
975. Adenoglossa B. Nord.
Adenoglossa B. Nord., Bot. Notiser 129: 137 (1976).
Annual herb. Indumentum absent, first stem internode with basifixed hairs. Leaves alternate or opposite, entire, succulent. Capitula solitary, pedunculate, radiate. Involucre hemispherical. Phyllaries
in 2–3 rows, without scarious margins, with central
resin canals. Receptacle conical, epaleate. Ray florets female, fertile; limb short, yellow. Disc florets
hermaphrodite, fertile; corolla 5-lobed, slightly zygomorphic with 2 smaller adaxial and 3 larger abaxial lobes, yellow. Achenes obovate, dorsiventrally
strongly flattened and laterally winged; apex with
5–6 scales; pericarp with myxogenic cells on abaxial
surface and with 2 lateral resin canals. One species,
A. decurrens (Hutch.) B. Nord., South Africa.
976. Cymbopappus B. Nord.
Cymbopappus B. Nord., Bot. Notiser 129: 150 (1976).
Shrubs or subshrubs. Indumentum woolly, of basifixed hairs. Leaves alternate, crowded, linear or
pinnatisect. Capitula solitary, pedunculate, radiate. Involucre hemispherical to cylindrical. Phyllaries in 3–4 rows, with brownish scarious margins.
Receptacle convex to subconical, epaleate. Ray florets female, fertile; limb white or pinkish. Disc florets hermaphrodite, fertile; corolla 5-lobed, yellow,
tube swollen, with thick vascular strands. Achenes
oblong, with 5 ribs; apex with an oblique, adaxially
longer corona, or with a large adaxial and a smaller
abaxial scale; pericarp with myxogenic cells on the
ribs and on the abaxial surface, without resin sacs.
Three species, South Africa.
977. Foveolina Källersjö
Foveolina Källersjö, Bot. J. Linn. Soc. 96: 316 (1988);
Källersjö, Bot. J. Linn. Soc. 96: 299–322 (1988), rev.
Annual herbs. Indumentum of basifixed hairs.
Leaves alternate, 2-pinnatisect. Capitula solitary,
pedunculate, discoid, disciform or radiate. Involucre hemispherical. Phyllaries in 3–5 rows,
with or without central resin canals. Receptacle
flat-conical, epaleate. Ray florets (when present) female, fertile; limb white. Achenes inconspicuously
3-ribbed; apex with an adaxial auricle or a corona;
pericarp abaxially covered with myxogenic cells,
laterally with two rows of short-stalked glands
immersed in small cavities, adaxially white and
spongy with numerous druses, abaxially very thin
and translucent. Outer florets (of disciform capitula) filiform, female. Achenes almost terete; apex
marginally rounded; pericarp without myxogenic
cells, spongy and wrinkled. (Inner) disc florets
hermaphrodite, fertile; corolla 4–5-lobed, yellow,
tube often basally saccate. Achenes as in ray florets
or obovoid, with a marginally rounded apex and
a thin pericarp, abaxially with myxogenic cells.
Five species, South Africa, Namibia.
978. Leucoptera B. Nord.
Leucoptera B. Nord., Bot. Notiser 129: 140 (1976).
Glabrous shrublets. Leaves alternate or opposite,
entire or lobed, somewhat succulent. Capitula solitary, pedunculate, radiate. Involucre meniscoid to
hemispherical. Phyllaries in 2–3 rows, with wide
scarious margins. Receptacle convex, epaleate. Ray
florets female, fertile; limb white, often becoming pink-reddish. Disc florets hermaphrodite, fertile; corolla 5-lobed, yellow. Achenes dorsiventrally
flattened, laterally winged; apex with 3 adaxiallateral scales; pericarp abaxially covered with myxogenic cells, with 2 lateral resin canals. x = 9. Three
species, South Africa.
979. Marasmodes DC.
Marasmodes DC., Prodr. 6: 136 (1838).
Glabrous shrubs. Leaves alternate, entire or occasionally lobed. Capitula small, closely aggregated,
rarely solitary, discoid. Involucre hemispherical
to urceolate. Phyllaries in 4–5 rows, with scarious margins. Receptacle flat, epaleate. Florets
hermaphrodite, fertile; corolla 5-lobed, yellow,
tube swollen and with thick vascular strands.
Achenes circular in cross-section, with 5 ribs; apex
with a corona of 7–10 elliptical, adaxially longer
scales; pericarp with myxogenic cells mainly
abaxially and on the ribs. Four species, South
Africa.
Compositae
980. Myxopappus Källersjö
Myxopappus Källersjö, Bot. J. Linn. Soc. 96: 314 (1988);
Källersjö, Bot. J. Linn. Soc. 96: 299–322 (1988), rev.
Annual herbs. Indumentum of basifixed hairs.
Leaves alternate, variously pinnatisect. Capitula
solitary, pedunculate, discoid. Involucre meniscoid to hemispherical. Phyllaries in 3–5 rows, very
narrow and acute, without scarious margins, often
with central resin canals. Receptacle convex to
conical, epaleate. Florets hermaphrodite, fertile;
corolla 4- or 5-lobed, yellow. Achenes obovoid to
oblong, triquetrous in cross-section, with 1 adaxial
and 2 lateral ribs; apex with a corona covered with
myxogenic cells; pericarp spongy and containing
numerous druses, with myxogenic cells abaxially
and on the ribs. x = 7. Two species, South Africa,
Namibia.
981. Oncosiphon Källersjö
Oncosiphon Källersjö, Bot. J. Linn. Soc. 96: 310 (1988);
Källersjö, Bot. J. Linn. Soc. 96: 299–322 (1988), rev.
Annual herbs. Indumentum absent or of basifixed
hairs. Leaves alternate, variously pinnatisect. Capitula solitary or in corymbs, pedunculate, radiate or discoid. Involucre hemispherical to cylindrical. Phyllaries in 3–4 rows, with scarious margins, with or without central resin canals. Receptacle flat to convex or conical, epaleate. Ray florets
(when present) female, fertile; limb white. Disc florets hermaphrodite, fertile; corolla 4-lobed, yellow,
tube conspicuously swollen and brittle. Achenes
cylindrical, circular in cross-section, with 4 ribs;
apex with a mainly abaxial entire or toothed rim;
pericarp without myxogenic cells. x = 6, 8. Eight
species, South Africa, Lesotho, Namibia.
982. Pentzia Thunb.
Pentzia Thunb., Prodr. Pl. Cap. 2: 145 (1800).
Shrubs or subshrubs. Indumentum absent or
of medifixed hairs. Leaves alternate or rarely
opposite, entire to pinnatisect. Capitula solitary
or in corymbs, pedunculate, discoid. Involucre
hemispherical. Phyllaries in 3–5 rows, with pale or
brown scarious margins, sometimes with central
resin canals. Receptacle convex to conical, epaleate.
Florets hermaphrodite, fertile; corolla 5-lobed,
yellow, tube swollen and with thick vascular
strands. Achenes narrowly obovoid, with 5 ribs;
apex with an adaxial auricle, an adaxially longer
corona of 3–5 scales, or marginally rounded;
355
pericarp with myxogenic cells abaxially and along
the ribs. x = 6, 7, 8, 9. Twenty-seven species, South
Africa, Namibia, Morocco, Algeria, Chad, Somalia,
Yemen.
983. Rennera Merxm.
Rennera Merxm., Mitt. Bot. Staatssamml. München 2: 335
(1957); Herman, Bot. J. Linn. Soc. 129: 367–377 (1999), rev.
Perennial herbs or shrublets. Indumentum absent
or of basifixed hairs. Leaves alternate or opposite at
base, entire to pinnatisect. Capitula solitary, pedunculate, discoid. Involucre hemispherical. Phyllaries
in 3–4 rows, with central resin canals. Receptacle flat to convex, epaleate. Florets hermaphrodite,
fertile; corolla 5-lobed, yellow, tube narrow, limb
wide. Achenes obovoid or ellipsoid, 5-angled; apex
marginally rounded or with a thick, spreading to
revolute rim; pericarp without myxogenic cells,
conspicuously tuberculate to rugose. Four species,
Namibia, South Africa, Botswana.
XVI.4. Phymaspermum Group (984–986)
984. Eumorphia DC.
Eumorphia DC., Prodr. 6: 2 (1838).
Shrubs or shrublets. Indumentum absent or of basifixed hairs. Leaves opposite, rarely alternate, entire
or lobed. Capitula solitary or in lax corymbs, pedunculate, radiate. Involucre hemispherical. Phyllaries in 2–3 rows, usually with a central resin duct,
with light to dark brown scarious margins. Receptacle flat to conical, paleate, rarely epaleate; paleae
linear, canaliculate, with a central resin duct. Ray
florets female, fertile; limb white, sometimes purplish. Disc florets hermaphrodite, fertile; corolla 5lobed, yellow, sometimes purplish. Achenes cylindrical, circular in cross-section, with 10–12(–18)
rounded ribs; apex with an entire or dentate, thickened rim; pericarp often minutely papillose, without myxogenic cells or resin sacs. Six species, South
Africa, Lesotho, Swaziland.
985. Gymnopentzia Benth.
Gymnopentzia Benth. in Benth. & Hook. f., Gen. Pl. 2: 1235
(1876).
Shrub. Indumentum absent or of basifixed hairs.
Leaves opposite, entire or lobed, connate and
sheathing at base. Capitula in dense corymbs,
pedunculate, discoid. Involucre hemispherical.
356
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Phyllaries in few rows, with ciliate scarious
margins. Receptacle flat to hemispherical, epaleate
(occasionally paleate). Florets hermaphrodite,
fertile; corolla 5-lobed, yellow, often purplish.
Achenes ellipsoid, circular in cross-section, with
10 rounded ribs; apex truncate, without a corona
or thickened rim; pericarp with very long papillae,
without myxogenic cells or resin sacs. One species,
G. bifurcata Benth., South Africa, Lesotho.
986. Phymaspermum Less.
Phymaspermum Less., Syn. Gen. Comp.: 253 (1832).
Brachymeris DC. (1837).
Adenachaena DC. ‘Adenochaena’ (1838).
Iocaste E. Mey. ex Harv. (1865).
Shrubs or shrublets. Indumentum absent or of basifixed hairs. Leaves alternate, entire to lobed. Capitula solitary or in lax corymbs and pedunculate,
rarely solitary and sessile along branches, radiate
or discoid. Involucre hemispherical to spherical,
rarely cylindrical to obconical. Phyllaries in 3–4
rows, often with longitudinal resin ducts. Receptacle flat-conical, epaleate. Ray florets female, fertile;
limb white or yellow, sometimes purplish. Disc florets hermaphrodite, fertile; corolla 5-lobed, yellow,
sometimes purplish; tube rarely pubescent with
long hairs. Achenes obovoid or cylindrical, circular in cross-section, with 10–18 rounded ribs; apex
with an entire or dentate rim or corona; pericarp
often minutely papillose, with ovoid myxogenic trichomes and resin sacs in some of the ribs. Nineteen species, South Africa, Swaziland, Zimbabwe,
Namibia.
987. Schistostephium Less.
Schistostephium Less., Syn. Gen. Comp.: 251 (1832).
Peyrousea DC. (1838).
Shrublets or subshrubs. Indumentum of basifixed
hairs. Leaves alternate, entire to pinnatisect.
Capitula in lax to dense corymbs, rarely solitary,
disciform or discoid. Involucre hemispherical,
sometimes obconical or urceolate. Phyllaries in
2–3 rows, with narrow, pale scarious margins.
Receptacle hemispherical to conical, epaleate or
with few marginal paleae. Outer florets female,
fertile; limb absent. Inner florets hermaphrodite,
fertile (innermost sometimes functionally male);
corolla 4-lobed, yellow. Achenes obovoid, dorsiventrally flattened, elliptical in cross-section,
with 2 lateral, wing-like ribs; apex truncate or
marginally rounded; pericarp glabrous or papillose, sometimes with myxogenic cells, always
without resin sacs. Twelve species, South Africa,
Mozambique, Zimbabwe, Swaziland.
988. Thaminophyllum Harv.
Thaminophyllum Harv. in Harvey & Sonder, Fl. Cap. 3: 155
(1865); Bond, J. S. African Bot. 46: 157–166 (1980), rev.
Shrublets. Indumentum of basifixed hairs. Leaves
alternate, crowded, entire, lanceolate to linear, ericoid. Capitula solitary or in lax corymbs, radiate.
Involucre hemispherical. Phyllaries in 3–4 rows,
with narrow, often reddish, scarious margins. Receptacle convex to conical, hairy, epaleate. Ray florets female, sterile; limb white to somewhat purplepink, tube confluent with the achene. Disc florets
hermaphrodite, fertile; corolla 4-lobed, yellow; stylopodium large and persistent in fruit. Achenes
ellipsoid, 3–4-angled; apex marginally rounded;
pericarp with large myxogenic cells in rounded,
scattered groups. x = 10. Three species, South
Africa.
989. Ursinia Gaertn.
Ursinia Gaertn., Fruct. Sem. Pl. 2: 462 (1791), nom. cons.;
Prassler, Mitt. Bot. Staatssamml. München 6: 363–478
(1967), rev.
Sphenogyne R. Br. (1813).
Ursiniopsis E. Phillips (1951).
Annual or perennial herbs or shrublets. Indumentum absent or of basifixed hairs. Leaves alternate,
entire to 2-pinnatisect, sometimes succulent.
Capitula solitary or in lax corymbs, radiate or
discoid, pedunculate. Involucre hemispherical.
Phyllaries in 3–7 rows, with narrow to broad scarious margins. Receptacle hemispherical, paleate;
paleae canaliculate, elliptical or narrowly linear
with an apical limb. Ray florets usually neuter,
sometimes female and fertile; limb yellow, orange,
white or reddish. Disc florets hermaphrodite,
fertile; corolla 5-lobed, yellow, purplish. Achenes
cylindrical or obovoid, straight or curved, circular
in cross-section, with 5 ribs and a basal tuft of
hairs or glabrous; apex with a uniseriate pappus of
5–10 ovate or circular scales, or a biseriate pappus
of 5 outer such scales and 5 inner subulate ones, or
rarely epappose; pericarp rarely with myxogenic
cells. x = 5, 7, 8. Thirty-nine species, South Africa,
Namibia, Botswana, Ethiopia.
Compositae
XVI. Group B [Asian Grade]
XVI.5. Ajania Group (990–998)
990. Ajania Poljakov
Ajania Poljakov, Bot. Mater. Gerb. Bot. Inst. Komarova
Akad. Nauk S.S.S.R. 17: 419 (1955); Shih, Acta Phytotax.
Sin. 32: 365–368 (1994), rev.
357
marginally rounded; pericarp without myxogenic
cells or resin sacs. x = 9 (polyploidy). Three
species, Arctic Eurasia, Siberia, Japan, Arctic
North America.
993. Brachanthemum DC.
Brachanthemum DC., Prodr. 6: 44 (1838); Zhao, Acta Sci.
Nat. Neimonggol 27: 805–807 (1996), rev.
Perennial herbs or subshrubs. Indumentum of
medifixed and basifixed hairs. Leaves alternate,
serrate to pinnatisect, rarely entire. Capitula small,
in corymbs or rarely solitary, disciform. Involucre
hemispherical to cylindrical. Phyllaries in 3–4
rows, with rather wide, light to dark scarious margins. Receptacle convex to conical, epaleate. Outer
row of florets female, fertile; corolla slender, 2–3-,
rarely 4–5-lobed. Central florets hermaphrodite,
fertile; corolla 5-lobed, yellow. Achenes obovoid,
with 4–6 ribs; apex marginally rounded; pericarp
with myxogenic cells in rows, without resin sacs.
x = 9 (polyploidy). Thirty-nine species, central
Asia, China, Japan.
Small shrublets, woody at base. Indumentum of
basifixed, medifixed or stellate hairs. Leaves alternate, few-lobed. Capitula solitary or in lax corymbs,
pedunculate, radiate or rarely discoid. Involucre
hemispherical to cylindrical. Phyllaries in 4–5 rows,
with pale to brownish membranous margins. Receptacle flat to convex or conical, glabrous or hairy,
epaleate. Ray florets female, fertile; limb yellow, yellowish or white. Disc florets hermaphrodite, fertile;
corolla 5-lobed, yellow. Achenes obovoid to oblong,
with 5–7 ribs; apex marginally rounded; pericarp
with myxogenic cells, without resin sacs. x = 9. Ten
species, central Asia, Mongolia, China.
991. Ajaniopsis C. Shih
994. Chrysanthemum L.
Ajaniopsis C. Shih, Acta Phytotax. Sin. 16: 86 (1978).
Chrysanthemum L., Sp. Pl. 887 (1753); Gen. Pl., ed. 5: 379
(1754), nom. et typ. cons.
Dendranthema (DC.) Des Moul. (1860).
Annual herb. Leaves alternate, pinnatisect and fewlobed. Capitula small, in dense corymbs, disciform. Involucre hemispherical. Phyllaries in 2 rows,
without or with narrow dark scarious margins.
Receptacle convex, epaleate. Outer row of florets
female, fertile; corolla 0–2-lobed, tapering above,
apically densely pilose with erect, straight hairs.
Central florets hermaphrodite, fertile; corolla 5lobed, yellow, apically densely pilose with erect,
straight hairs. Achenes obovoid, 3–6-ribbed; apex
marginally rounded; pericarp thin, with 3–6 rows
of myxogenic cells, without resin sacs. One species,
A. peniciliformis C. Shih, China, Tibet.
992. Arctanthemum (Tzvelev) Tzvelev
Arctanthemum (Tzvelev) Tzvelev, Novosti Sist. Vyssh. Rast.
22: 274 (1985).
Perennial herbs. Indumentum absent or of basifixed hairs. Leaves rosulate to alternate, apically
lobed-serrate. Capitula solitary, pedunculate,
radiate. Involucre meniscoid to hemispherical.
Phyllaries in 3–4 rows, with dark brown scarious
margins. Receptacle convex to conical, epaleate.
Ray florets female, fertile; limb white. Disc florets
hermaphrodite, fertile; corolla 5-lobed, yellow.
Achenes ellipsoid to obovoid, with 5–8 ribs; apex
Perennial herbs or subshrubs. Indumentum absent
or of medifixed or basifixed hairs. Leaves alternate,
pinnatisect, lobed, serrate or rarely entire. Capitula in lax corymbs or solitary, pedunculate, radiate.
Involucre meniscoid. Phyllaries in 2–4 rows, generally with brown scarious margins. Receptacle convex to conical, epaleate (paleate in some cultivars).
Ray florets female, fertile; limb white, pink or yellowish. Disc florets hermaphrodite, fertile; corolla
5-lobed, yellow. Achenes obovoid, with 5–8 ribs;
apex marginally rounded; pericarp thin, generally
with myxogenic cells in rows, without resin sacs.
x = 9 (polyploidy). Forty-one species, Asia (Mongolia, Russia, China, Japan, Korea), eastern Europe.
Some species widely cultivated as ornamentals.
995. Elachanthemum Y. Ling & Y.R. Ling
Elachanthemum Y. Ling & Y.R. Ling, Acta Phytotax. Sin. 16:
62 (1978).
Annual herb. Leaves alternate, pinnatisect, fewlobed or entire. Capitula in lax corymbs, pedunculate, discoid. Involucre hemispherical to suborbicular. Phyllaries in 3–4 rows, with broad scarious
margins. Receptacle subconical, epaleate. Florets
358
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
hermaphrodite, fertile (the inner sterile); corolla
5-lobed, yellow, tubular. Achenes obovoid, without
ribs; apex oblique and marginally rounded; pericarp with longitudinal rows of myxogenic cells,
without resin sacs. x = 9. One species, E. intricatum (Franchet) Y. Ling & Y.R. Ling, Mongolia,
China.
fertile; corolla 5-lobed, yellow. Achenes obovoid to
cylindrical, with 5 ribs; apex marginally rounded
or with a rim; pericarp with myxogenic cells, without resin sacs. One species, T. kirilowii (Turcz. ex
DC.) Sch. Bip., eastern Siberia.
XVI.6. Artemisia Group (999–1005)
996. Hulteniella Tzvelev
Hulteniella Tzvelev in B.A. Jurtzev, Arktich. Fl. S.S.S.R. 10:
117 (1987).
Perennial herb. Indumentum of basifixed hairs.
Leaves rosulate to alternate, linear. Capitula solitary, pedunculate, radiate. Involucre meniscoid to
hemispherical. Phyllaries in 3–4 rows, with dark
brown scarious margins. Receptacle convex to conical, sparsely hairy, epaleate. Ray florets female,
fertile; limb white. Disc florets hermaphrodite, fertile; corolla 5-lobed, yellow. Achenes ellipsoid to
obovoid, with 5 ribs; apex truncate or with a rim;
pericarp without myxogenic cells or resin sacs.
x = 9. One species, H. integrifolium (Richardson)
Tzvelev, Arctic Eurasia, Arctic North America.
999. Artemisia L.
Fig. 70
Artemisia L., Sp. Pl.: 845 (1753).
Oligosporus Cass. (1817).
Seriphidium Fourr. (1869).
Artanacetum (Rzazade) Rzazade (1956).
997. Phaeostigma Muldashev
Phaeostigma Muldashev,
Leningrad) 66: 586 (1981).
Bot.
Zhurn.
(Moscow
&
Perennial herbs or subshrubs. Indumentum of
medifixed hairs. Leaves alternate, pinnatifid to
shallowly lobed or entire. Capitula small, in dense
corymbs, disciform. Involucre hemispherical.
Phyllaries in 3–4 rows, with brown scarious
margins. Receptacle convex, epaleate. Outer row of
florets female, fertile; corolla slender, 2–4-lobed.
Central florets hermaphrodite, fertile; corolla
5-lobed, yellow; style branches brownish. Achenes
obovoid, with 4–6 ribs, apically ribs projected into
minute teeth; pericarp without myxogenic cells or
resin sacs. x = 9. Three species, China.
998. Tridactylina (DC.) Sch. Bip.
Tridactylina (DC.) Sch. Bip. in Webb & Berthelot, Hist. Nat.
Iles Canaries 3 (2, 2): 245 (1844).
Annual herb. Indumentum of basifixed and medifixed hairs. Leaves alternate, few-lobed. Capitula
in a lax corymb, pedunculate, radiate. Involucre
meniscoid to hemispherical. Phyllaries in 3 rows,
with dark brown scarious margins. Receptacle flat
to convex, epaleate. Ray florets neuter; limb yellow or yellowish-white. Disc florets hermaphrodite,
Fig. 70. Compositae-Anthemideae. Artemisia absinthium.
A Habit. B Capitulum. C Hermaphrodite and tubular florets.
D Achene. (Drawings by Ch. Oberprieler)
Compositae
Annual or perennial herbs, subshrubs or shrubs.
Indumentum absent or of basifixed, medifixed
or stellate hairs. Leaves alternate, variously lobed
or dissected, rarely entire. Capitula discoid or
disciform, usually arranged in a long panicle but
this sometimes much reduced and racemose, spiciform or subglobose. Involucrum hemispherical
to cylindrical or urceolate. Phyllaries in 2–7 rows,
with more or less scarious margins. Receptacle flat
to conical, sometimes pilose, epaleate or occasionally paleate. Outer female florets usually tapering
above, with 2–4 teeth, or truncate, commonly
oblique at orifice. Central florets hermaphrodite
and fertile or functionally male; corolla 5-lobed,
yellow or sometimes reddish-violet or purple.
Achenes obovoid to elliptical, cylindrical or
flattened, usually glabrous but occasionally hairy;
apex marginally rounded; pericarp with or without
rows of myxogenic cells, without resin sacs. x = 7,
8, 9, 10, 11, 17 (polyploidy). Circa 522 species,
Northern Hemisphere, South America, southern
Africa, Pacific Islands.
359
apex marginally rounded; pericarp consisting of
large myxogenic cells in longitudinal rows, without resin sacs. x = 9. One species, F. sibiricum (L.)
Kitam., Far East (Siberia, Mongolia, China, Korea).
1002. Mausolea Poljakov
Mausolea Poljakov, Trudy Inst. Bot. (Alma-Ata) 11: 170
(1961).
Shrub. Indumentum of medifixed hairs. Leaves
alternate, few-lobed or entire. Capitula small,
subglobose, few, more or less sessile in a reduced
panicle, disciform. Involucre hemispherical. Phyllaries in 2 rows, with narrow scarious margins.
Receptacle convex to hemispherical, epaleate.
Outer florets female, fertile, without corolla; style
branches dilated, lanceolate, flat, acute. Central
florets functionally male; corolla 5-lobed, apically
with medifixed hairs. Achenes obovoid; apex
marginally rounded; pericarp densely pilose,
without myxogenic cells or resin sacs. x = 9
(polyploidy). One species, M. eriocarpa (Bunge)
Poljakov, Iran, Afghanistan, central Asia.
1000. Crossostephium Less.
Crossostephium Less., Linnaea 6: 220 (1831).
Shrub. Indumentum tomentose, of medifixed
hairs. Leaves alternate, narrowly spathulate,
apically few-lobed or entire, succulent. Capitula
rather small and rounded, paniculate, disciform.
Involucre hemispherical. Phyllaries in 3 rows,
outer bracts tomentose, inner bracts scarious.
Receptacle hemispherical, epaleate. Outer female
florets tubular, 2–3-lobed, yellow. Central florets
hermaphrodite, fertile; corolla 5-lobed, yellow.
Achenes obovoid, weakly 5-ribbed; apex with
a lacerate corona or with small scales. x = 9.
One species, C. chinense (L.) Makino, Philippines,
Taiwan, southern Japan, China. Widely cultivated
in eastern Asia.
1001. Filifolium Kitam.
Filifolium Kitam., Acta Phytotax. Geobot. 9: 157 (1940).
Perennial herb, woody at base. Indumentum absent or of medifixed hairs. Leaves alternate, pinnatisect. Capitula in corymbs, disciform. Involucre
hemispherical. Phyllaries in 3 rows, with scarious
margins. Receptacle conical, epaleate. Outer row of
florets female, fertile, tapering above, minutely 2–
4-dentate. Central florets functionally male; corolla
(4–)5-lobed, compressed in a resinous mass, yellowish. Achenes obliquely obovoid, without ribs;
1003. Neopallasia Poljakov
Neopallasia Poljakov, Bot. Mater. Gerb. Bot. Inst. Komarova
Akad. Nauk S.S.S.R. 17: 429 (1955).
Annual or biennial herbs. Indumentum of medifixed hairs. Leaves alternate, pectinate-pinnatisect.
Capitula small and rounded in a narrow spiciform panicle, disciform. Involucre cylindrical to
urceolate. Phyllaries in 2 rows, with broad scarious margins. Receptacle narrowly conical, epaleate.
Outer florets female, fertile; corolla narrowly tubular, without apical lobes. Central florets of two
kinds, outer hermaphrodite and fertile, inner completely sterile with reduced ovaries; corolla 5-lobed,
bluish or pinkish. Achenes in one row at the base
of the receptacle, oblong-obovoid, somewhat compressed or triquetrous; apex marginally rounded;
pericarp with many rows of myxogenic cells, without resin sacs. x = 9. Three species, central Asia,
southern Siberia, Mongolia, China.
1004. Picrothamnus Nutt.
Picrothamnus Nutt., Trans. Am. Philos. Soc., ser. II, 7: 417
(1841).
Shrublet, woody at base, with older branches
forming long spines. Indumentum of medifixed
hairs. Leaves alternate, few-lobed. Capitula small
and subglobose, solitary or few together along
360
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
the branches, almost sessile, disciform. Involucre
hemispherical. Phyllaries in 1–2 rows, with narrow
hyaline scarious margins. Receptacle hemispherical, epaleate. Outer florets female, fertile; corolla
tubular. Central florets functionally male; corolla
5-lobed, with long arachnoid hairs. Achenes
obovoid; apex marginally rounded; pericarp
densely arachnoid-pilose, without myxogenic cells
or resin sacs. x = 9 (polyploidy). One species,
P. desertorum Nutt., North America.
1005. Sphaeromeria Nutt.
Sphaeromeria Nutt., Trans. Am. Philos. Soc., ser. II, 7: 401
(1841); Holmgren, Shultz & Lowrey, Brittonia 28: 255–262
(1976), rev.
Chamartemisia Rydb. (1916).
Vesicarpa Rydb. (1916).
Perennial herbs or subshrubs. Indumentum of
medifixed hairs. Leaves alternate to rosulate,
pinnatisect and few-lobed or entire. Capitula
apparently in corymbs but sometimes somewhat
paniculate or capitate or solitary, disciform.
Involucre hemispherical or obconical. Phyllaries in
2–3 rows, with light to dark scarious margins. Receptacle flat to conical, rarely pubescent, epaleate.
Outer row of florets female, tapering above; corolla
sometimes glandular, rarely with hairs at the apex.
Central florets hermaphrodite, fertile; corolla
5-lobed, pale or bright yellow, rarely with hairs
at the apex. Achenes obovoid-oblong, sometimes
faintly 3–5-ribbed; apex marginally rounded or
with a rim, a corona or with small scales; pericarp
sometimes with myxogenic cells, without resin
sacs. x = 9. Nine species, North America, Mexico.
1006. Artemisiella Ghafoor
Artemisiella Ghafoor, Candollea 47: 636 (1992).
Perennial herb, woody at base. Indumentum of
medifixed hairs. Leaves alternate, basally crowded,
2–3-pinnatisect. Capitula in a simple raceme,
pedunculate, disciform. Involucre hemispherical.
Phyllaries in 3–4 rows, with rather broad, brown
scarious margins. Receptacle hemispherical, pilose, epaleate. Outer florets female, fertile; corolla
tubular, 4-lobed, densely hairy. Inner florets
hermaphrodite, fertile; corolla 5-lobed, yellow,
densely hairy. Achenes obovoid, quadrangular in
cross-section; apex with a more or less distinct,
rounded rim; pericarp sparsely or (in achenes of
outer florets) densely hairy. x = 9. One species,
A. stracheyi (Hook. f. & Thompson ex Clarke)
Ghafoor, Ladakh, Tibet, Nepal, Bhutan, southern
China.
XVI.7. Cancrinia Group (1007–1012)
1007. Allardia Decne
Allardia Decne in Jacquemont, Voy. Inde 4: 87 (1836).
Waldheimia Kar. & Kir. (1842).
Perennial herbs. Indumentum absent or of
basifixed hairs. Leaves alternate, lobed to pinnatisect. Capitula solitary, pedunculate, radiate.
Involucre hemispherical. Phyllaries in 3–4 rows,
with broad, blackish-brown scarious margins.
Receptacle hemispherical, epaleate. Ray florets
female, fertile or sterile, or neuter; limb white,
pink or bluish violet. Disc florets hermaphrodite,
fertile; corolla 5-lobed, yellow or bluish violet.
Achenes obconical, circular in cross-section, with
5–10 protruding ribs; apex with a corona formed
by 25–50 bristle-like, apically brownish scales as
long as or longer than the achene body; pericarp
glabrous to densely hairy, without myxogenic cells
or resin sacs. Eight species, Afghanistan, central
Asia, Mongolia, China.
1008. Cancrinia Kar. & Kir.
Cancrinia Kar. & Kir., Bull. Soc. Imp. Naturalistes Moscou
15: 124 (1842).
Perennial herbs. Indumentum of basifixed hairs.
Leaves alternate, 1–2-pinnatipartite to -pinnatisect.
Capitula solitary, pedunculate, discoid. Involucre hemispherical. Phyllaries in 2–3 rows, with
broad, pale to dark brown or blackish scarious
margins. Receptacle hemispherical, epaleate.
Florets hermaphrodite, fertile; corolla 5-lobed,
yellow, glabrous. Achenes obconical, circular in
cross-section, with 5–6(–10) inconspicuous ribs;
apex with a corona formed by 5–12 linear or
narrowly elliptical, apically brownish scales as
long as or longer than the achene body; pericarp
glabrous to sparsely hairy, without myxogenic cells
or resin sacs. Four species, central Asia, Mongolia,
China.
1009. Cancriniella Tzvelev
Cancriniella Tzvelev in Komarov, Fl. U.R.S.S. 26: 292, 876
(1961).
Subshrub. Indumentum of long basifixed hairs.
Leaves alternate, palmately lobed to pinnatipartite.
Capitula solitary, pedunculate, discoid. Involucre
Compositae
hemispherical. Phyllaries in 1–2 rows, all of nearly
the same length, the outer with narrow, the inner
with broad, pale scarious margins. Receptacle
hemispherical, sparsely hairy, epaleate. Florets
hermaphrodite, fertile; corolla 5-lobed, yellow,
glabrous. Achenes narrowly obconical, circular
in cross-section, with 10 distinct ribs; apex with
a corona formed by 6–10 rather short, apically
rounded scales much shorter than the achene
body; pericarp glabrous, without myxogenic cells
or resin sacs. One species, C. krascheninnikovii
(Rubtzov) Tzvelev, central Asia.
1010. Richteria Kar. & Kir.
Richteria Kar. & Kir., Bull. Soc. Imp. Naturalistes Moscou
15: 126 (1842).
Subshrubs. Indumentum absent or of long basifixed hairs. Leaves alternate, basally crowded,
2–3-pinnatisect. Capitula solitary, rarely 2–3 per
shoot forming a lax corymb, pedunculate, radiate.
Involucre hemispherical. Phyllaries in 2–3 rows,
with broad, dark brown to blackish scarious
margins. Receptacle hemispherical, epaleate. Ray
florets female, fertile; limb white or pinkish. Disc
florets hermaphrodite, fertile; corolla 5-lobed,
yellow, glabrous. Achenes obconical, circular in
cross-section, with 5–10 rather inconspicuous
ribs; apex with a corona formed by 5–10 obovate to
linear-elliptical, apically brownish scales shorter
than the achene body; pericarp glabrous, without
myxogenic cells or resin sacs. x = 9 (polyploidy).
Six species, Iran, Afghanistan, central Asia,
Mongolia, China, Himalaya.
361
by (4–)5–10(–12) ovate to linear-elliptical scales
as long as or adaxially distinctly longer than the
achene body; pericarp densely hairy, sometimes
with myxogenic cells, without resin sacs. Nine
species, central Asia.
1012. Ugamia Pavlov
Ugamia Pavlov, Vestn. Akad. Nauk Kazakhsk. S.S.R. 8: 25
(1950).
Subshrub. Indumentum of basifixed hairs. Leaves
alternate, 1–2-pinnatipartite to -pinnatisect. Capitula solitary, on very short, nodding peduncles,
discoid. Involucre hemispherical. Phyllaries in 4–5
rows, with narrow to broad, light to dark brown
scarious margins. Receptacle flat to hemispherical,
epaleate. Florets hermaphrodite, fertile; corolla 5lobed, yellow. Achenes obconical, circular in crosssection, with 5 distinct and 5–10 inconspicuous
ribs; apex with a corona of 10–20 narrow, linearelliptical scales as long as or longer than the achene body; pericarp densely hairy, without myxogenic cells or resin sacs. One species, U. angrenica
(Krasch.) Tzvelev, central Asia.
XVI.8. Handelia Group (1013–1017)
1013. Handelia Heimerl
Handelia Heimerl, Oesterr. Bot. Z. 71: 215 (1922).
Basally woody, hapaxanthic, biennial to perennial
herb with one to few thick stems filled with soft
pith. Indumentum of basifixed hairs. Leaves alternate, up to 3-pinnatisect. Capitula numerous, in
dense corymbs, discoid. Involucre hemispherical
to spherical. Phyllaries in 3 rows, with broad,
1011. Trichanthemis Regel & Schmalh.
pale scarious margins. Receptacle hemispheriTrichanthemis Regel & Schmalh., Trudy Imp. S.-Peterburgsk.
cal, paleate; paleae narrowly elliptical to linear,
Bot. Sada 5: 617 (1877).
moderately canaliculate. Florets hermaphrodite,
Glossanthis Poljakov (1959).
fertile; corolla 5-lobed, yellow. Achenes cylindrical
Subshrubs or perennial herbs. Indumentum of to narrowly obovoid, circular to elliptical in
long basifixed hairs. Leaves alternate, apically trifid cross-section, with 5 inconspicuous ribs; apex with
to 1–2-pinnatisect. Capitula solitary, pedunculate, a short, abaxially more-developed rim; pericarp
radiate or discoid. Involucre hemispherical or with myxogenic cells, without resin sacs. x = 9.
obconical. Phyllaries in 3–5 rows, with narrow to One species, H. trichophylla (Schrenk) Heimerl,
broad, pale to brown scarious margins. Receptacle Afghanistan, Pakistan, central Asia, China.
flat to hemispherical, glabrous or sparsely to
densely hairy, epaleate. Ray florets (when present) 1014. Lepidolopsis Poljakov
female and fertile; limb white, yellow or pinkish;
Lepidolopsis Poljakov, Bot. Mater. Gerb. Bot. Inst. Komarova
tube usually hairy. Disc florets hermaphrodite,
Akad. Nauk S.S.S.R. 19: 374 (1956).
fertile; corolla 5-lobed, yellow, usually hairy.
Achenes obconical, circular in cross-section, with Basally woody, pollacanthic, perennial herb with
5–8(–10) distinct ribs; apex with a corona formed one to few rather thick stems filled with soft pith. In-
362
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
dumentum of basifixed hairs. Leaves alternate, 2–3pinnatisect. Capitula numerous, in a long panicle,
shortly pedunculate, discoid. Involucre hemispherical. Phyllaries in 3–4 rows, the inner with broad,
pale scarious margins. Receptacle hemispherical
to conical, epaleate. Florets hermaphrodite, fertile;
corolla (4–)5(–6)-lobed, yellow. Achenes narrowly
obovoid, circular in cross-section, with 5 conspicuous ribs; apex with a slanted corona of abaxially prolonged, dentate to laciniate scales; pericarp
without myxogenic cells or resin sacs. x = 9. One
species, L. turkestanica (Regel & Schmalh.) Poljakov, Iran, Afghanistan, central Asia.
1015. Polychrysum (Tzvelev) Kovalevsk.
Polychrysum (Tzvelev) Kovalevsk. in Vvendensky, Fl.
Uzbek. 6: 148 (1962).
Basally woody, monocarpic, perennial herb with
usually one, rarely 2–3 rather thick stems filled
with soft pith. Indumentum of basifixed hairs.
Leaves alternate, 3–4-pinnatisect. Capitula numerous, in a large and dense corymb, discoid.
Involucre hemispherical to spherical. Phyllaries
in 2–3 rows, the inner with broad, pale scarious margins. Receptacle flat to hemispherical,
epaleate. Florets hermaphrodite, fertile; corolla
5-lobed, yellow. Achenes narrowly cylindrical,
circular in cross-section, with 5–7 ribs; apex with
an irregularly dissected corona of many acute
lobes; pericarp without myxogenic cells or resin
sacs. x = 9. One species, P. tadshikorum (Kudr.)
Kovalevsk., Afghanistan, central Asia.
1016. Pseudohandelia Tzvelev
Pseudohandelia Tzvelev in Komarov, Fl. U.R.S.S. 26: 878
(1961).
Basally woody, hapaxanthic, biennial to perennial
herb with usually one, rarely 2–4 thick stems
filled with soft pith. Indumentum of basifixed
hairs. Leaves alternate, 2–3-pinnatisect. Capitula
numerous, in a dense corymb or pseudo-umbel,
discoid. Involucre hemispherical. Phyllaries in
2–3 rows, the outer and middle with broad,
the inner with narrow, pale scarious margins.
Receptacle hemispherical to conical, epaleate.
Florets hermaphrodite, fertile; corolla 5-lobed,
yellow. Achenes narrowly cylindrical, arcuate,
circular in cross-section, with 4–5 inconspicuous
ribs; apex ecoronate; pericarp tuberculate, with
myxogenic cells, without resin sacs. One species,
P. umbellifera (Boiss.) Tzvelev, Iran, Afghanistan,
central Asia, China.
1017. Sclerorhachis (Rech. f.) Rech. f.
Sclerorhachis (Rech. f.) Rech. f., Anz. Österr. Akad. Wissensch., Math. Naturwissensch. Kl. 105: 242 (1969).
Monocarpic or pollacanthic, biennial to perennial
herbs with one to few rather thick stems filled with
soft pith. Indumentum of basifixed hairs. Leaves alternate, but often basally crowded, 3–4-pinnatisect,
with a sclerified, persistent rhachis. Capitula in
lax corymbs, discoid. Involucre hemispherical, umbonate. Phyllaries in 2–4 rows, with pale scarious
margins. Receptacle flat to hemispherical, paleate;
paleae persistent to readily deciduous, subulate.
Florets hermaphrodite, fertile; corolla 5-lobed, yellow. Achenes obovoid, circular in cross-section,
with 4–7 inconspicuous ribs; apex ecoronate; pericarp with myxogenic cells, without resin sacs. x = 9.
Four species, Iran, Afghanistan.
1018. Hippolytia Poljakov
Hippolytia Poljakov, Bot. Mater. Gerb. Bot. Inst. Komarova
Akad. Nauk S.S.S.R. 18: 288 (1957).
Perennial herbs. Indumentum of medifixed hairs.
Leaves alternate, 2–3-pinnatipartite to -pinnatisect.
Capitula rarely solitary, usually numerous (2–15),
in lax to dense corymbs, shortly pedunculate to
subsessile, discoid. Involucre hemispherical. Phyllaries in 2–3 rows, the outer with narrow, the inner
with broad, pale to dark brown or blackish scarious
margins. Receptacle flat to hemispherical, epaleate.
Florets hermaphrodite, fertile; corolla 5-lobed, yellow; anthers basally caudate. Achenes cylindrical
to narrowly obovoid, circular in cross-section, with
5–8 inconspicuous ribs; apex with a more or less
distinct, rounded rim; pericarp glabrous, without
myxogenic cells, sometimes with longitudinal resin
canals. x = 9. Nineteen species, central Asia, Mongolia, China, Himalaya.
1019. Kaschgaria Poljakov
Kaschgaria Poljakov, Bot. Mater. Gerb. Bot. Inst. Komarova
Akad. Nauk S.S.S.R. 18: 282 (1957).
Subshrubs. Indumentum of stellate hairs. Leaves
alternate, entire or few-lobed. Capitula rather
small and few in an elongated panicle, at the summit fasciculate, disciform. Involucre cylindrical.
Phyllaries in 2–4 rows, with scarious margins.
Receptacle conical, epaleate. Outer florets female,
Compositae
tapering above; corolla 2–3-lobed, with stellate
hairs. Central florets hermaphrodite, fertile;
corolla 5-lobed, yellow, apically with stellate
hairs. Achenes obovoid; apex marginally rounded;
pericarp without myxogenic cells or resin sacs.
x = 9. Two species, Mongolia, Kazakhstan, China.
1020. Lepidolopha C. Winkl.
Lepidolopha C. Winkl., Trudy Imp. S.-Peterburgsk. Bot.
Sada 13: 236 (1894).
Shrublets with basally woody, virgate and sometimes leafless stems. Indumentum of medifixed
hairs. Leaves alternate, sometimes crowded basally
or fasciculate on brachyblasts, entire or 3-lobed.
Capitula solitary or numerous in lax to dense
corymbs, on short to long peduncules, discoid.
Involucre cylindrical to obconical. Phyllaries in
4–7 rows, with narrow, pale scarious margins.
Receptacle flat to hemispherical, epaleate. Florets
hermaphrodite, fertile; corolla 5-lobed, yellow.
Achenes obconical, round in cross-section, with
6–10 ribs; apex with a corona formed by 8–10
linear-elliptical scales; pericarp glabrous, without
myxogenic cells or resins sacs. x = 6, 8. Nine
species, central Asia.
1021. Leucanthemella Tzvelev
Leucanthemella Tzvelev in Komarov, Fl. U.R.S.S. 26: 137
(1961).
Decaneurum Sch. Bip. (1844), non DC. (1833).
Perennial herbs. Indumentum absent or of basifixed and medifixed hairs. Leaves alternate, entire
or serrate, glandular-punctate. Capitula solitary
or in lax corymbs, pedunculate, radiate. Involucre
meniscoid to hemispherical. Phyllaries in 2–3
rows, with brown scarious margins. Receptacle
convex, epaleate. Ray florets female, sterile; limb
white. Disc florets hermaphrodite, fertile; corolla
5-lobed, yellow. Achenes narrowly obovoid,
slightly curved or straight, with 8–12 ribs; apex
with a marginal rim; pericarp without myxogenic
cells or resin sacs. x = 9. Two species, eastern
Europe (Slovakia, Ukraine, Bulgaria, Romania,
Yugoslavia, Hungary), Far East (Mongolia, China,
Korea, Japan). Widely cultivated as ornamentals.
1022. Microcephala Pobed.
Microcephala Pobed., Bot. Mater. Gerb. Bot. Inst. Komarova
Akad. Nauk S.S.S.R. 21: 356 (1961); Podlech, Mitt. Bot.
Staatssamml. München 12: 655–682 (1976), rev.
363
Annual herbs. Indumentum of basifixed hairs.
Leaves alternate, pinnatisect. Capitula solitary,
pedunculate, radiate or discoid. Involucre hemispherical. Phyllaries in 2–3 rows, with broad pale
scarious margins. Receptacle conical, hollow,
epaleate. Ray florets female, fertile; limb white.
Disc florets hermaphrodite, fertile; corolla 5-lobed,
yellow or reddish. Achenes circular to slightly
dorsiventrally flattened in cross-section, with
3–5 adaxially arranged ribs; apex with a lacerate
or fimbriate, adaxially somewhat longer corona;
pericarp with myxogenic cells abaxially and on the
ribs, adaxially between the ribs with unbranched
hairs of 3–8 cells with spiral cell wall thickenings.
x = 7. Five species, central Asia, Afghanistan, Iran,
Pakistan, Mongolia, China.
1023. Nipponanthemum Kitam.
Nipponanthemum Kitam., Acta Phytotax. Geobot. 29: 168
(1978).
Glabrous shrub. Leaves alternate, crowded at the
end of the woody branches, serrate. Capitula
solitary, pedunculate, radiate. Involucre meniscoid
to hemispherical. Phyllaries in 4 rows, with pale
brown scarious margins. Receptacle convex,
epaleate. Ray florets female, fertile; limb white.
Disc florets hermaphrodite, fertile; corolla 5-lobed,
yellow. Achenes narrowly obovoid,slightly curved
or straight, with c. 10 ribs; apex with a lacerate
corona or with small scales; pericarp without
myxogenic cells or resin sacs. x = 9. One species,
N. nipponicum (Franchet ex Maxim.) Kitam.,
Japan.
1024. Opisthopappus C. Shih
Opisthopappus C. Shih, Acta Phytotax. Sin. 17: 110 (1979).
Perennial herbs with basally woody stems. Indumentum of basifixed hairs. Leaves alternate,
1–2-pinnatisect. Capitula solitary or 2–3 in lax
corymbs, pedunculate, radiate. Involucre hemispherical. Phyllaries in 3–4 rows, with broad
membranous margins. Receptacle hemispherical
to conical, epaleate. Ray florets female, fertile;
limb white or pinkish. Disc florets hermaphrodite,
fertile; corolla 5-lobed, yellow. Achenes obovoid,
with 3–5 ribs; apex with a corona of 4–6 separate,
mainly abaxial subulate scales of unequal length;
pericarp with myxogenic cells, without resin sacs.
Two species, China.
364
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
1025. Stilpnolepis Krasch.
Stilpnolepis Krasch., Bot. Mater. Gerb. Bot. Inst. Komarova
Akad. Nauk S.S.S.R. 9: 207 (1946).
Annual herb. Indumentum of medifixed hairs.
Leaves opposite or alternate, pinnatisect, few-lobed
or entire. Capitula in lax corymbs, pedunculate,
nodding, discoid. Involucre hemispherical to
urceolate. Phyllaries in 4–5 rows, the outer ovate
and with broad scarious margins, the inner
broadly elliptical to obovate, completely scarious.
Receptacle convex to subconical, epaleate. Florets
hermaphrodite, fertile; corolla 5-lobed, yellow,
with a distinct tube and a broadened, crateriform
limb. Achenes narrowly obovoid, without ribs;
apex truncate to marginally rounded; pericarp
thin, with longitudinal rows of myxogenic cells,
without resin sacs. One species, S. centiflora
(Maxim.) Krasch., Mongolia, China.
1026. Tanacetopsis (Tzvelev) Kovalevsk.
Tanacetopsis (Tzvelev) Kovalevsk. in Schreder, Fl. Uzbek. 6:
138 (1962).
Perennial herbs or subshrubs. Indumentum of
basifixed and medifixed hairs. Leaves alternate,
1–4-pinnatsect. Capitula rarely solitary, usually
numerous (2–40), in lax to dense corymbs, discoid,
pedunculate. Involucre hemispherical. Phyllaries
in 2–3 rows, the outer with narrow, the inner with
broad, pale to dark brown or blackish scarious
margins. Receptacle hemispherical, glabrous or
sparsely hairy, epaleate. Florets hermaphrodite,
fertile; corolla 5-lobed, yellow, usually glabrous,
rarely hairy. Achenes obconical, circular in crosssection, with 5–7 ribs; apex with an often oblique,
adaxially more-developed corona formed by free
or basally united scales of various shapes; pericarp
glabrous, usually with myxogenic cells, without
resin sacs. Twenty-three species, Iran, Afghanistan,
central Asia.
1027. Turaniphytum Poljakov
Turaniphytum Poljakov in Komarov, Fl. U.R.S.S. 26: 880
(1961).
Perennial herbs, somewhat woody at base. Indumentum of basifixed and medifixed hairs.
Leaves alternate or rosulate, pinnatisect. Capitula
disciform; synflorescence a spike of glomerules
with densely congested capitula, or rarely capitula
solitary in interrupted, partly congested spikes.
Involucre cylindrical. Phyllaries in 3 rows, with
broad hyaline scarious margins. Receptacle convex
to hemispherical, epaleate. Outer florets female,
subtended by scaphoid inner involucral bracts;
corolla unequally crenate at the apex. Central
florets functionally male; corolla 5-lobed, apically
with long rigid somewhat reddish hairs. Achenes obliquely oblong-obovoid; apex marginally
rounded; pericarp with rows of myxogenic cells,
without resin sacs. x = 9 (polyploidy). Two species,
Turkmenistan, Iran, Afghanistan, Kazakhstan.
XVI. Group C [Eurasian Grade]
XVI.9. Achillea Group (1028–1032)
1028. Achillea L.
Achillea L., Sp. Pl.: 896 (1753).
Millefolium Mill. (1754).
Ptarmica Mill. (1754).
Arthrolepis Boiss. (1849).
Perennial herbs and subshrubs. Indumentum of
basifixed, sometimes asymmetrically medifixed
hairs. Leaves alternate, rarely entire, usually
dentate to 4-pinnatisect, sometimes vermiform.
Capitula usually in dense corymbs, rarely solitary,
pedunculate to subsessile, radiate or rarely discoid.
Involucre hemispherical to cylindrical. Phyllaries
in 2–3 rows, with pale to black scarious margins.
Receptacle flat, hemispherical or conical, rarely
much elongated, paleate; paleae ± canaliculate,
sometimes with a central resin duct. Ray florets
female, fertile; limb white, yellow or pink; tube
± flattened. Disc florets hermaphrodite, fertile;
corolla 5-lobed, white, yellow or pink, basally
inconspicuously saccate, clasping top of achene.
Achenes obovoid, dorsiventrally flattened, with 2
lateral and rarely with 1 additional adaxial rib; apex
marginally rounded; pericarp with myxogenic
cells on ribs, with or without longitudinal resin
sacs. x = 9 (polyploidy). Circa 115 species, Europe,
Asia, northern Africa. Some species naturalised
in North America, Australia, New Zealand, South
Africa.
1029. Anacyclus L.
Anacyclus L., Sp. Pl.: 892 (1753); Humphries, Bull. Brit.
Mus. (Nat. Hist.), Bot. 7: 83–142 (1979), rev.
Cyrtolepis Less. (1831).
Annual or perennial herbs. Indumentum of
basifixed hairs. Leaves alternate, 1–3-pinnatisect,
flat to terete. Capitula solitary or in lax corymbs,
Compositae
sometimes densely aggregated, discoid or radiate; peduncles sometimes inflated at maturity.
Involucre hemispherical or obconical. Phyllaries
in 3 rows, with narrow to broad, light to dark
brown scarious margins. Receptacle flat to conical,
paleate; paleae obcuneate to obovate, flat or
canaliculate. Ray florets female, fertile; limb white
or yellow, sometimes abaxially red; tube flattened,
persistent on achenes. Disc florets hermaphrodite,
fertile; corolla yellow, actinomorphic to zygomorphic with 2 larger, cucullate lobes, basally flattened
and very slightly saccate. Achenes obconical,
dorsiventrally flattened, with 2 lateral wing-like
ribs; apex sometimes with a narrow marginal
corona or a thin, lacerate appendage; pericarp with
small myxogenic cells, without resin sacs. x = 9.
Twelve species, northern Africa, southern Europe,
Near East.
1030. Heliocauta Humphries
Heliocauta Humphries, Bot. Notiser 130: 155 (1977).
Perennial, creeping herb. Indumentum of basifixed
hairs. Leaves in a basal rosette, 3-pinnatisect. Capitula solitary, pedunculate, discoid. Involucre hemispherical. Phyllaries in 3 rows, with laterally pale to
light brown, apically dark brown scarious margins.
Receptacle conical, paleate; paleae narrowly elliptical to narrowly obovate, moderately canaliculate.
Florets hermaphrodite, fertile; corolla 5-lobed, yellow. Achenes narrowly obovoid, somewhat dorsiventrally flattened, circular to elliptical in crosssection, 4–5-ribbed with 2 distinct lateral ribs; apex
with a minute corona formed by a dentate rim; pericarp without myxogenic cells, with elongated resin
sacs. x = 9. One species, H. atlantica (Litard. &
Maire) Humphries, northern Africa.
1031. Leucocyclus Boiss.
Leucocyclus Boiss., Diagn. Pl. Orient., ser. I, 11: 13 (1849).
Perennial herb. Indumentum of basifixed hairs.
Leaves alternate, 2-pinnatisect, vermiform. Capitula solitary, long-pedunculate, radiate; peduncles
slightly inflated at maturity. Involucre hemispherical. Phyllaries in 3 rows, sometimes with dark
brown tips. Receptacle flat to convex, paleate;
paleae linear-elliptical. Ray florets female, fertile;
limb white, oblong to subrotund; tube flattened,
basally saccate and both adaxially and abaxially
clasping top of achene. Disc florets hermaphrodite,
fertile; corolla yellow, basally clasping top of
achene. Achenes dorsiventrally flattened, with two
365
lateral wing-like ribs; apex marginally rounded;
pericarp thin on faces, thick and sclerenchymatic
in ribs. One species, L. formosus Boiss., Turkey.
1032. Otanthus Hoffmanns. & Link
Otanthus Hoffmanns. & Link, Fl. Portug. 2: 364 (1834).
Diotis Desf. (1799), non Schreber (1791).
Neesia Spreng. (1818), nom. rej., non Blume (1835), nom.
cons.
Densely sericeous, greyish-white, suffruticose
perennial. Indumentum of basifixed hairs. Leaves
alternate, entire to dentate, ± succulent. Capitula
in dense corymbs, pedunculate to subsessile,
discoid. Involucre hemispherical. Phyllaries in 3
rows, almost without scarious margins. Receptacle
convex, paleate; paleae ± canaliculate, with
a central resin duct. Florets hermaphrodite, fertile;
corolla 5-lobed, yellow, basally conspicuously
swollen, laterally clasping upper half of achene or
more; basal part persistent on achenes. Achenes
obovoid, round to tetragonal, with 4–5 weak
ribs; apex marginally rounded; pericarp without
myxogenic cells or resin sacs. x = 9. One species,
O. maritimus (L.) Hoffmanns. & Link, Europe,
northern Africa, south-western Asia.
XVI.10. Anthemis Group (1033–1038)
1033. Anthemis L.
Anthemis L., Sp. Pl.: 893 (1753); Benedí I González, Thesis
Barcelona (1987); Georgiou, Thesis Patras (1990); Ghafoor
& Al Turki, Candollea 52: 457–474 (1997); Oberprieler,
Bocconea 9: 1–328 (1998); Ghafoor & Ali, Comp. Newslett.
38: 1–41 (2002), rev.
Maruta (Cass.) Gray (1821).
Lyonnetia Cass. (1825).
Ammanthus Boiss. & Heldr. ex Boiss. (1849).
Annuals, biennials, or short- to long-lived
perennial herbs and subshrubs. Indumentum of
medifixed hairs. Leaves alternate, dentate to lobed
or 1–3-pinnatisect. Capitula solitary or in lax
corymbs, pedunculate, discoid or radiate; peduncles sometimes inflated or curved at maturity.
Involucre hemispherical or obconical, sometimes
umbonate. Phyllaries in 3–5 rows, with pale to
brown or black scarious margins. Receptacle
hemispherical or conical to narrowly conical,
paleate or epaleate marginally or totally; paleae
subulate or elliptical to obovate, flat. Ray florets
(when present) female and fertile, or neuter; limb
366
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
white, yellow or pink; tube sometimes hairy. Disc
florets hermaphrodite, fertile; corolla 5-lobed,
yellow, rarely pink, sometimes hairy. Achenes
obovoid to obconical, circular or quadrangular
in cross-section, usually with c. 10 smooth or
tuberculate ribs, rarely without distinct ribs; apex
with a shallow, often adaxially more-developed
corona or auricle, or ecoronate; pericarp with
small myxogenic cells; epicarpic cells with crystal
sand. x = 9 (polyploidy). Circa 175 species, Europe,
south-western Asia, northern and eastern Africa.
Naturalised in North America, Australia, New
Zealand, South Africa.
1034. Cota J. Gay
Cota J. Gay in Gussone, Fl. Sicul. Syn. 2: 866 (1845).
Annuals, biennials, or short- to long-lived perennial herbs. Indumentum of medifixed hairs, rarely
of basifixed hairs. Leaves alternate, 1–3-pinnatisect;
segments often pectinately divided. Capitula solitary or laxly corymbose, radiate or discoid, pedunculate; peduncles slender or inflated at maturity. Involucre hemispherical or obconical, sometimes umbonate. Phyllaries in 3–5 rows, with pale
to dark scarious margins, often cartilaginous. Receptacle hemispherical, paleate; paleae elliptical to
obovate, flat, often cartilaginous, persistent. Ray
florets (when present) female, fertile; limb white,
yellow or pinkish. Disc florets hermaphrodite, fertile; corolla 5-lobed, yellow or pinkish. Achenes obconical, dorsiventrally flattened, with prominent
lateral ribs, smooth or with 3–10 ribs on each face;
apex with a rounded rim or a short corona; pericarp
with small myxogenic cells; epicarpic cells without or with single, octahedral or prismatic crystals.
x = 9 (polyploidy). Forty species, Europe, southwestern Asia, northern Africa.
1035. Gonospermum Less.
Gonospermum Less., Syn. Gen. Comp.: 263 (1832).
Lugoa DC. (1838).
Basally woody perennial herbs and shrubs. Indumentum of medifixed hairs. Leaves alternate,
1–2-pinnatipartite to -pinnatisect. Capitula in lax
to dense corymbs, pedunculate, discoid or radiate.
Involucre obconical to hemispherical. Phyllaries in
2–4 rows, with rather narrow, pale to dark brown
scarious margins. Receptacle hemispherical,
paleate; paleae narrowly elliptical, moderately
canaliculate. Ray florets (when present) female,
fertile; limb white. Disc florets hermaphrodite,
fertile; corolla 5-lobed, yellow. Achenes cylindrical
to narrowly obovoid, circular in cross-section,
with 5 conspicuous ribs; apex with a corona of 5
teeth projecting from the ribs; pericarp without
myxogenic cells or resin sacs. x = 9. Five species,
Canary Islands.
1036. Nananthea DC.
Nananthea DC., Prodr. 6: 45 (1838).
Small annual herb; glabrous, sometimes with
some basifixed hairs. Leaves alternate, 3–5-lobed,
with a long petiole, succulent. Capitula very
small, solitary, pedunculate, radiate or disciform.
Involucre hemispherical. Phyllaries in 1–2 rows,
with broad, pale scarious margins. Receptacle
conical, epaleate. Ray florets female, fertile; limb
white or reduced. Disc florets hermaphrodite,
fertile; corolla 4-lobed, yellow. Achenes obovoid,
circular in cross-section, with c. 8 inconspicuous
ribs; apex marginally rounded; pericarp with
large myxogenic cells, without resin sacs. x = 9.
One species, N. perpusilla (Loisel.) DC., southern
Europe (Corsica, Sardinia).
1037. Tanacetum L.
Tanacetum L., Sp. Pl.: 843 (1753).
Balsamita Mill. (1754).
Pyrethrum Zinn (1757).
Gymnocline Cass. (1816).
Omalanthus Less. (1832), non Juss. (1824).
Omalotes DC. (1838).
Psanacetum (Necker ex Less.) Spach (1841).
Hemipappus K. Koch (1851).
Spathipappus Tzvelev (1961).
Poljakovia Grubov & Filat. (2001).
Perennial herbs, rarely annuals or subshrubs. Indumentum absent or of basifixed and/or medifixed, sometimes stellate hairs. Leaves alternate,
rarely entire or dentate, usually 1–3-pinnatipartite
to -pinnatisect. Capitula solitary or in lax to dense
corymbs, pedunculate to subsessile, radiate, disciform or discoid. Involucre hemispherical. Phyllaries in 3–5 rows, with narrow to broad, pale to
brown or blackish scarious margins. Receptacle flat
to hemispherical, epaleate, rarely paleate. Ray florets female or neuter, fertile or sterile; limb yellow, white or pink. Disc florets all hermaphrodite
or (in disciform capitula) the outer female, fertile; corolla 5-lobed, yellow. Achenes obconical or
Compositae
cylindrical, circular in cross-section, with 5–10(–
15) ribs; apex marginally rounded or with a short
to well-developed, entire or dentate corona, rarely
with an adaxial auricle; pericarp without myxogenic cells or resin sacs. x = 9 (polyploidy). Circa
160 species, Europe, Asia, northern Africa, North
America. Some species widely cultivated.
1038. Tripleurospermum Sch. Bip.
Tripleurospermum Sch. Bip., Tanaceteen: 31 (1844).
Gastrosulum Sch. Bip. (1844).
Rhytidospermum Sch. Bip. (1844).
Dibothrospermum Knaf (1846).
Trallesia Zumag. (1849).
Annual or perennial herbs. Indumentum absent or
of basifixed hairs. Leaves alternate, 1–3-pinnatisect.
Capitula solitary or in corymbs, pedunculate, discoid, disciform or radiate. Involucre hemispherical
to cup-shaped. Phyllaries in 2–5 rows, with narrow to broad pale to brown or black scarious margins. Receptacle convex to conical, epaleate. Ray
florets female, fertile; limb white or sometimes pale
pink. Disc florets hermaphrodite, fertile; corolla 5lobed, yellow or greenish, lobes usually with a resin
sac. Achenes triquetrous with 1 adaxial and 2 lateral ribs and sometimes 1–2 abaxial ribs, often
rugose or tuberculate abaxially and between ribs;
apex with a corona, an adaxial auricle, or with few
scales, sometimes marginally rounded; pericarp
sometimes with myxogenic cells, with (1–)2(–3–5)
abaxial-apical resin sacs. x = 9 (polyploidy). Forty
species, Europe, northern Africa, temperate Asia,
North America. One species widespread as a weed.
367
XVI.11. Leucanthemopsis Group
(1040–1043)
1040. Castrilanthemum Vogt & Oberprieler
Castrilanthemum Vogt & Oberprieler, Anales Jard. Bot.
Madrid 54: 342 (1996).
Annual herb. Indumentum of medifixed hairs.
Leaves alternate, 1–2-pinnatisect. Capitula solitary, pedunculate, radiate. Involucre meniscoid to
hemispherical. Phyllaries in 4 rows, with broad
brown scarious margins. Receptacle convex,
epaleate. Ray florets female, fertile; limb white.
Disc florets hermaphrodite, fertile; corolla 5-lobed,
yellow with reddish lobes. Achenes obovoid, with
10 ribs; apex marginally rounded; pericarp with
myxogenic cells along the ribs, without resin sacs.
x = 9. One species, C. debeauxii (Degen, Hervier &
É. Rev.) Vogt & Oberprieler, south-western Europe
(Spain).
1041. Hymenostemma Willk.
Hymenostemma Willk., Bot. Zeitung 22: 253 (1864).
Annual herb. Indumentum of medifixed hairs.
Leaves alternate, pinnatisect. Capitula solitary,
pedunculate, radiate. Involucre meniscoid to
hemispherical. Phyllaries in 4 rows, with light
brown scarious margins. Receptacle convex,
epaleate. Ray florets female, sterile; limb white,
yellow at base. Disc florets hermaphrodite, fertile;
corolla 5-lobed, yellow. Achenes obovoid, with 5–7
ribs; apex with a scarious corona; pericarp with
myxogenic cells along the ribs, without resin sacs.
x = 9. One species, H. pseudanthemis (Kunze)
Willk., south-western Europe (Spain).
1039. Brocchia Vis.
1042. Leucanthemopsis (Giroux) Heywood
Brocchia Vis. in Spongia, Comment. Med. 2: 218 (1836).
Leucanthemopsis (Giroux) Heywood, Anales Inst. Bot.
Cavanilles 32: 181 (1975); Heywood, Anales Inst. Bot.
Cavanilles 12: 313–374 (1954, under Tanacetum L.);
Heywood, Anales Inst. Bot. Cavanilles 32: 175–187 (1975),
rev.
Annual herb. Indumentum of basifixed hairs.
Leaves alternate, pinnatipartite to pinnatisect,
sometimes entire to lobed. Capitula solitary or
in lax corymbs, discoid, pedunculate. Involucre hemispherical. Phyllaries in 2 rows, with
narrow, pale membranous margins. Receptacle
hemispherical to conical, epaleate. Disc florets
hermaphrodite, fertile; corolla yellow, apically 4lobed. Achenes obovoid, circular in cross-section,
with c. 4 inconspicuous lateral and adaxial ribs;
apex slanting, marginally rounded; pericarp with
large, elongated myxogenic cells, without resin
sacs. x = 9. One species, B. cinerea (Del.) Vis.,
northern Africa, Near East.
Perennial herbs, sometimes woody at base. Indumentum of medifixed hairs. Leaves alternate,
serrate-dentate to pinnatisect. Capitula solitary,
pedunculate, radiate. Involucre meniscoid to
hemispherical. Phyllaries in 4 rows, with pale to
dark brown scarious margins. Receptacle convex,
epaleate. Ray florets female, fertile; limb yellow,
white, white with yellow base, or reddish. Disc
florets hermaphrodite, fertile; corolla 5-lobed, yellow, lobes sometimes reddish. Achenes narrowly
368
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
obovoid, with (3–)5–10 ribs; apex with a scarious
corona; pericarp with myxogenic cells along the
ribs, without resin sacs. x = 9 (polyploidy). Nine
species, central, southern and eastern Europe,
north-western Africa (Morocco).
1043. Prolongoa Boiss.
Prolongoa Boiss., Voy. Bot. Espagne: 320 (1840), nom. cons.
Annual herb. Indumentum of medifixed hairs.
Leaves alternate, pinnatisect. Capitula solitary,
pedunculate, radiate. Involucre meniscoid to
hemispherical. Phyllaries in 4 rows, with broad
whitish scarious margins. Receptacle convex,
epaleate. Ray florets neuter, sterile; limb yellow.
Disc florets hermaphrodite, fertile; corolla 5-lobed,
yellow. Achenes obovoid, slightly dorsiventrally
compressed, with 1 adaxial and 2 lateral and 2
abaxial ribs; apex marginally rounded, in sterile
ray achenes with a scarious corona; pericarp with
myxogenic cells along the ribs, without resin sacs.
x = 9. One species, P. hispanica G. López & C.E.
Jarvis, south-western Europe (Spain).
1044. Matricaria L.
Matricaria L., Sp. Pl.: 890 (1753); Gen. Pl., ed. 5.: 380 (1754).
Chamomilla Gray (1821).
Lepidotheca Nutt. (1841).
Lepidanthus Nutt. (1841), non Nees (1830).
Cenocline Koch (1843).
Akylopsis Lehm. (1852).
Cotulina Pomel (1874).
Annual herbs. Indumentum absent or of basifixed
hairs. Leaves alternate, pinnatisect. Capitula
solitary or in lax corymbs, pedunculate, disciform
or radiate. Involucre hemispherical to cup-shaped.
Phyllaries in 2–3 rows, with broad pale or brown
scarious margins. Receptacle conical, hollow,
epaleate. Ray florets female, fertile; limb white,
sometimes reduced. Disc florets hermaphrodite,
fertile; corolla 4–5-lobed, yellow or greenish.
Achenes obovoid-oblong, circular to slightly
dorsiventrally flattened in cross-section, with 3–5
adaxially arranged ribs; apex marginally rounded
or with a small corona, in ray achenes sometimes
with adaxially longer corona; pericarp with myxogenic cells abaxially and on the ribs, lateral ribs
sometimes with resin canals. x = 9 (polyploidy).
Six species, Europe, northern Africa, Macaronesia,
western, south-western and central Asia, western
North America. Some species widespread as
weeds.
1045. Phalacrocarpum (DC.) Willk.
Phalacrocarpum (DC.) Willk., Bot. Zeitung 22: 252 (1864);
Nieto Feliner, Anales Jard. Bot. Madrid 39: 53–60 (1982),
rev.
Perennial herb. Indumentum of medifixed hairs.
Leaves opposite, sheathing at base, serrate-dentate
to bipinnatisect. Capitula solitary, pedunculate, radiate. Involucre meniscoid to hemispherical. Phyllaries in 4 rows, with dark brown scarious margins.
Receptacle convex, sometimes hairy, epaleate. Ray
florets female, fertile, white or reddish. Disc florets
hermaphrodite, fertile or functionally male; corolla
5-lobed, yellow. Achenes narrowly obovoid, slightly
curved or straight, with 7–9 ribs; apex marginally
rounded; pericarp without myxogenic cells or resin
sacs. x = 9. Two species, south-western Europe
(Spain, Portugal).
1046. Xylanthemum Tzvelev
Xylanthemum Tzvelev in Komarov, Fl. U.R.S.S. 26: 284, 877
(1961).
Shrublets with basally woody, virgate and sometimes leafless stems. Indumentum of medifixed and
basifixed hairs. Leaves alternate, 1–2-pinnatipartite
to -pinnatisect. Capitula solitary or numerous in
lax corymbs, on long peduncles, discoid. Involucre
hemispherical or cylindrical to obconical. Phyllaries in 4–5 rows, the outer with narrow, the inner
with broad, dark brown scarious margins. Receptacle flat, epaleate. Florets hermaphrodite, fertile;
corolla 5-lobed, yellow, sometimes hairy. Achenes
obconical, circular in cross-section, with 5–6 distinct ribs; apex with an adaxial auricle or a corona
formed by 3–6 adaxial, elliptical scales shorter than
the achene body; pericarp glabrous, with myxogenic cells, without resin sacs. x = 9 (polyploidy).
Eight species, Iran, Afghanistan, central Asia.
rou
XVI. Group D
[Mediterranean Clade]
1047. Aaronsohnia Warb. & Eig
Aaronsohnia Warb. & Eig, Bull. Agric. Exp. Stat. Tel-Aviv 6:
39 (1927).
Annual herbs. Indumentum of basifixed hairs.
Leaves alternate, pinnatisect. Capitula solitary, pedunculate, discoid, disciform or radiate. Involucre
hemispherical. Phyllaries in 3–4 rows, with pale
scarious margins. Receptacle convex to conical,
Compositae
369
epaleate. Ray florets female, fertile or sterile; limb
white. Disc florets hermaphrodite, fertile; corolla
5-lobed, yellow, lobes sometimes with a central
resin sac. Achenes obovoid, circular to slightly
dorsiventrally flattened in cross-section; apex
with an adaxial, whitish auricle or sometimes
marginally rounded; pericarp with myxogenic
cells abaxially and sometimes with two lateral
resin canals. x = 9. Two species, northern Africa,
Near East.
XVI.12. Argyranthemum Group (1048–1051)
1048. Argyranthemum Webb
Fig. 71
Argyranthemum Webb in Webb & Berthelot, Hist. Nat. Iles
Canaries 3 (2, 2): 258 (1844); Humphries, Bull. Nat. Hist.
Mus., ser. Bot. 5: 147–240 (1976), rev.
Monoptera Sch. Bip. (1844).
Preauxia Sch. Bip. (1844).
Stigmatotheca Sch. Bip. (1844).
Scyphopappus B. Nord. (1976).
Subshrubs. Indumentum absent or of basifixed
hairs. Leaves alternate, serrate-dentate to variously
dissected. Capitula solitary or in lax corymbs,
pedunculate, radiate. Involucre meniscoid to
hemispherical. Phyllaries in 3–4 rows, with broad
pale to brown scarious margins. Receptacle
convex to conical, epaleate. Ray florets female,
fertile; limb white, rarely yellow or reddish. Disc
florets hermaphrodite, fertile; corolla 5-lobed,
yellow, rarely with reddish lobes. Achenes of
ray florets triquetrous and generally strongly
3-winged, sometimes united into groups, wings
sometimes reduced, often apically projected into
a corona; achenes of disc florets generally laterally
compressed and 2-winged, sometimes terete,
sometimes united; apex often with a corona;
pericarp without myxogenic cells or resin sacs.
x = 9. Twenty-four species, Macaronesia (Madeira,
Salvage Islands, Canary Islands). Some species
widely cultivated as ornamentals.
1049. Glebionis Cass.
Glebionis Cass. in Cuvier, Dict. Sci. Nat. 41: 41 (1826).
Chrysanthemum sensu auct., non L. (1753) quoad typ.
cons.
Xantophtalmum Sch. Bip. (1844).
Annual herbs. Indumentum absent or of basifixed hairs. Leaves alternate, serrate-dentate to
2-pinnatisect. Capitula solitary or in lax corymbs,
pedunculate, radiate. Involucre meniscoid to
Fig. 71. Compositae-Anthemideae. Argyranthemum foeniculaceum. A Habit. B Ray floret. C Tubular floret. D Achenes
of ray (above) and disc florets (below). (Drawings by Ch.
Oberprieler)
hemispherical. Phyllaries in 3–4 rows, with broad
pale or light brown scarious margins, with resin
canals. Receptacle convex, epaleate. Ray florets
female, fertile; limb yellow or white with yellow
base. Disc florets hermaphrodite, fertile; corolla
5-lobed, yellow. Achenes of ray florets triquetrous
and 2–3-winged; achenes of disc florets cylindrical and 10-ribbed or adaxially 1-winged; apex
marginally rounded; pericarp without myxogenic
cells or resin sacs. x = 9. Two species, Europe,
northern Africa, Caucasus, south-western Asia,
Macaronesia. Widespread as weeds.
370
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
1050. Heteranthemis Schott
Heteranthemis Schott, Isis (Oken) 1818: 822 (1818).
Pinardia Cass. (1826).
Annual herb. Indumentum of viscid-glandular,
basifixed hairs. Leaves alternate, serrate-dentate to
pinnatisect. Capitula solitary or in lax corymbs,
pedunculate, radiate. Involucre meniscoid to
hemispherical. Phyllaries in 4 rows, with broad
pale scarious margins, with resin canals. Receptacle convex to conical, epaleate. Ray florets female,
fertile; limb yellow. Disc florets hermaphrodite,
fertile; corolla 5-lobed, yellow. Achenes of ray
florets triquetrous and 3-winged; achenes of disc
florets laterally flattened and 1-winged, wings
projected into apical spines; pericarp without
myxogenic cells or resin sacs. x = 9. One species,
H. viscidehirta Schott, south-western Europe,
north-western Africa, introduced in Palestine.
1051. Ismelia Cass.
Ismelia Cass. in Cuvier, Dict. Sci. Nat. 41: 40 (1826).
Annual herb. Indumentum absent or of basifixed
hairs. Leaves alternate, pinnatisect. Capitula solitary or in lax corymbs, pedunculate, radiate. Involucre meniscoid to hemispherical. Phyllaries in
4 rows, with broad whitish scarious margins, with
resin canals. Receptacle convex, epaleate. Ray florets female, fertile; limb yellow or white with yellow
base. Disc florets hermaphrodite, fertile; corolla 5lobed, red or purple. Achenes of ray florets triquetrous and 3-winged; achenes of disc florets laterally flattened and 1–2-winged; apex marginally
rounded or with a short lacerate corona; pericarp
without myxogenic cells or resin sacs. x = 9. One
species, I. carinata (Schousb.) Sch. Bip., northwestern Africa (Morocco). Cultivated as an ornamental.
broad, dark brown scarious margins, sometimes
deflexed at maturity. Receptacle hemispherical to
conical, paleate; paleae flat or slightly canaliculate.
Ray florets female, fertile or sterile; limb white,
sometimes strongly reduced and hidden by phyllaries. Disc florets hermaphrodite, fertile; corolla
5-lobed, yellow, basally saccate, laterally clasping
the top of achene. Achenes obovoid, round in crosssection, with 2 lateral and 1 adaxial, flimsy ridges;
apex marginally rounded; pericarp consisting of
large myxogenic cells in longitudinal rows, without resin sacs. x = 9. Two species, southern and
western Europe, northern Africa.
1053. Cladanthus Cass.
Cladanthus Cass., Bull. Sci. Soc. Philom. Paris 1816: 199
(1816).
Ormenis (Cass.) Cass. (1823).
Perennial or annual herbs or subshrubs. Indumentum of basifixed hairs. Leaves alternate, dentate to
2-pinnatisect. Capitula solitary or in lax corymbs,
sessile (and then plant branched immediately below capitula) or pedunculate, radiate. Involucre
hemispherical. Phyllaries in 3 rows, flat to canaliculate, with broad, light scarious margins. Receptacle hemispherical to narrowly conical, paleate;
paleae canaliculate and geniculate, with a central
resin duct. Ray florets female, fertile or sterile; limb
orange, yellow, or white with a yellow base. Disc
florets hermaphrodite, fertile; corolla 5-lobed, orange or yellow, basally strongly saccate, adaxially
clasping the upper half or more of achene. Achenes obovoid, round in cross-section, with 2 lateral and 1 adaxial, flimsy ridges; apex slanting,
marginally rounded; pericarp consisting of large
myxogenic cells in longitudinal rows, without resin
sacs. x = 9. Five species, southern Europe, northern
Africa, south-western Asia.
1054. Mecomischus Coss. ex Benth. & Hook. f.
XVI.13. Chamaemelum Group (1052–1056)
1052. Chamaemelum Mill.
Chamaemelum Mill., Gard. Dict. Abr., ed. 4: 315 (1754).
Marcelia Cass. (1825).
Perideraea Webb (1838).
Perennial or annual herbs. Indumentum absent or
of basifixed hairs. Leaves alternate, 2–3-pinnatisect.
Capitula solitary or in lax corymbs, pedunculate,
radiate, disciform or discoid. Involucre hemispherical. Phyllaries in 3 rows, flat, with narrow, light or
Mecomischus Coss. ex Benth. & Hook. f., Gen. Pl. 2: 418
(1873).
Fradinia Pomel (1874).
Perennial or annual herbs. Indumentum of stellate hairs. Leaves alternate, sometimes basally opposite, entire to lobed. Capitula solitary, long-pedunculate, radiate. Involucre hemispherical, umbonate. Phyllaries in 3 rows, the outer with narrow, the inner with broad, light brown scarious
margins. Receptacle hemispherical, paleate; paleae
canaliculate, with a central resin duct. Ray flo-
Compositae
rets neuter, sterile; limb white or yellow. Disc florets hermaphrodite, fertile; corolla 5-lobed, yellow,
basally slightly saccate, hardly clasping the top of
achene. Achenes obovoid, slightly dorsiventrally
flattened, with 2 lateral and 1 adaxial, flimsy ridges;
apex marginally rounded; pericarp with large myxogenic cells in longitudinal rows, without resin
sacs. Two species, north-western Africa.
1055. Rhetinolepis Coss.
Rhetinolepis Coss., Bull. Soc. Bot. France 3: 707 (1856).
Annual herb. Indumentum of medifixed hairs.
Leaves alternate, entire to palmately lobed. Capitula small, in few-headed corymbs, rarely solitary,
shortly pedunculate, discoid. Involucre obconical
to urceolate. Phyllaries in 3 rows, flat to canaliculate, with very narrow, light scarious margins; the
inner with a central resin duct. Receptacle conical,
paleate; paleae canaliculate, somewhat geniculate,
with a central resin duct. Florets hermaphrodite,
fertile; corolla 5-lobed, yellow, basally inconspicuously saccate, hardly clasping the top of achene.
Achenes obovoid, slightly dorsiventrally flattened,
with 2 lateral and 1 adaxial, flimsy ridges; apex
marginally rounded; pericarp consisting of large
myxogenic cells in longitudinal rows, without resin
sacs. One species, R. lonadioides Coss., northern
Africa.
1056. Santolina L.
Santolina L., Sp. Pl.: 842 (1753).
Shrublets. Indumentum of medifixed hairs. Leaves
alternate, entire to dentate or pinnatisect, sometimes vermiform. Capitula solitary, pedunculate,
discoid. Involucre hemispherical, sometimes umbonate. Phyllaries in 3–4 rows, flat to carinate, with
light to dark brown or sometimes purple scarious margins. Receptacle hemispherical, paleate;
paleae ± canaliculate, with a central resin duct.
Florets hermaphrodite, fertile; corolla 5-lobed, yellow to whitish, basally saccate, clasping the apex of
achene. Achenes obconical, 3–5-angled, sometimes
slightly dorsiventrally flattened; apex marginally
rounded; pericarp with or without myxogenic cells,
without resin sacs. x = 9 (polyploidy). Circa thirteen species, southern Europe, northern Africa.
1057. Daveaua Willk. ex Mariz
Daveaua [Daveana] Willk. ex Mariz, Bol. Soc. Brot. 9: 206,
220, 243, 258 (1891).
371
Annual, glabrous herbs. Leaves alternate, pinnatisect. Capitula solitary, pedunculate, radiate. Involucre meniscoid to hemispherical. Phyllaries in
3 rows, with dark brown scarious margins. Receptacle conical, epaleate. Ray florets female, fertile
or sterile; limb white. Disc florets hermaphrodite,
fertile; corolla 5-lobed, yellow. Achenes of ray florets dorsiventrally flattened, with 3 adaxial ribs,
laterally winged, wings projected into apical teeth;
apex with an adaxial auricle. Achenes of disc florets
obovoid, circular in cross-section, with 5 ribs; apex
marginally rounded; pericarp with myxogenic cells
and with 3–5 resin sacs apically in the ribs. One
species, D. anthemoides Mariz, south-western Europe, north-western Africa (Morocco).
1058. Endopappus Sch. Bip.
Endopappus Sch. Bip., Bonplandia 8: 369 (1860).
Glabrous annual herb. Leaves alternate, pinnatisect. Capitula solitary, pedunculate, radiate.
Involucre meniscoid to hemispherical. Phyllaries
in 3–4 rows, with pale to brown scarious margins.
Receptacle flat to convex, epaleate. Ray florets
female, fertile; limb white or yellow. Disc florets
hermaphrodite, fertile; corolla 5-lobed, yellow.
Achenes dorsiventrally flattened, 3-angled with
one adaxial and two lateral ribs; apex with
a corona; pericarp with myxogenic cells abaxially
and along the ribs, without resin sacs. x = 9. One
species, E. macrocarpus Sch. Bip., northern Africa
(Morocco, Algeria, Tunisia, Libya).
1059. Heteromera Pomel
Heteromera Pomel, Bull. Soc. Sci. Phys. Algérie 11: 60
(1874).
Annual herbs. Indumentum of basifixed hairs.
Leaves alternate, pinnatisect. Capitula solitary,
pedunculate, radiate. Involucre meniscoid to
hemispherical. Phyllaries in 3–4 rows, with light
brown to brown scarious margins. Receptacle convex to conical, epaleate. Ray florets female, fertile
or sterile; limb white. Disc florets hermaphrodite,
fertile; corolla 5-lobed, yellow or basally reddish.
Achenes cylindrical to obovoid, circular or dorsiventrally flattened in cross-section, with 3–5
ribs; apex with an adaxially longer scarious corona
or auricle, with a short, basally callose corona, or
with 5–9 obovate scales; pericarp with myxogenic
cells along the ribs and on the abaxial surface, with
3–5 resin canals or apical sacs in the ribs. x = 9.
Two species, northern Africa.
372
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
1060. Lepidophorum Neck. ex DC.
Lepidophorum Neck. ex DC., Prodr. 6: 19 (1838).
Annual, glabrous herb. Leaves alternate, serrate.
Capitula solitary, pedunculate, radiate. Involucre
meniscoid to hemispherical. Phyllaries in 3–4 rows,
with narrow brown scarious margins. Receptacle
convex, paleate; paleae flat to canaliculate, narrowly
elliptical to obovate, with a central resin duct. Ray
florets female or neuter, sterile; limb yellow. Disc
florets hermaphrodite, fertile; corolla 5-lobed, yellow. Achenes of ray florets flat; apex with c. 4 free or
basally connate scales. Achenes of disc florets narrowly obovoid, 5-ribbed; apex marginally rounded;
pericarp with myxogenic cells along the ribs, without resin sacs. x = 9. One species, L. repandum (L.)
DC., south-western Europe.
XVI.14. Leucanthemum Group (1061–1068)
1061. Chlamydophora Ehrenb. ex Less.
Chlamydophora Ehrenb. ex Less., Syn. Gen. Comp.: 265,
448 (1832).
Glabrous annual herb. Leaves alternate or basally
opposite, entire or lobed, somewhat succulent.
Capitula solitary, pedunculate, discoid. Involucre
meniscoid to hemispherical. Phyllaries in 3–4 rows,
with light brown, sometimes red-tinged, scarious
margins. Receptacle convex, epaleate. Florets
hermaphrodite, fertile; corolla (4–)5-lobed, yellow
or somewhat reddish. Achenes ellipsoid, with
c. 10 ribs; apex with a large scarious and adaxially
longer corona; pericarp with myxogenic cells along
the ribs and with resin canals between the ribs.
x = 9. One species, C. tridentata (Delile) Ehrenb.
ex Less., northern Africa, eastern Mediterranean
region.
1062. Chrysanthoglossum B.H. Wilcox, K. Bremer
& Humphries
Chrysanthoglossum B.H. Wilcox, K. Bremer & Humphries,
Bull. Brit. Mus. (Nat. Hist.), Bot. 23: 143 (1993).
Annual, biennial or perennial herbs. Indumentum
absent or of basifixed hairs. Leaves alternate,
pinnatisect. Capitula solitary, pedunculate, radiate. Involucre meniscoid to hemispherical.
Phyllaries in 4 rows, with broad whitish to brown,
scarious margins. Receptacle convex, epaleate.
Ray florets female, sterile or sometimes fertile;
limb yellow. Disc florets hermaphrodite, fertile;
corolla 5-lobed, yellow. Achenes of ray florets
dorsiventrally compressed; apex with a scarious
adaxial auricle or marginally rounded. Achenes of
disc florets obovoid, with c. 10 conspicuous ribs;
apex with a corona with myxogenic cells on the
outside; pericarp with myxogenic cells along the
ribs and with resin canals between the ribs. x = 9.
Two species, northern Africa.
1063. Coleostephus Cass.
Coleostephus Cass. in Cuvier, Dict. Sci. Nat. 41: 43 (1826).
Myconia Neck. ex Sch. Bip. (1844).
Kremeria Durieu (1846).
Myconella Sprague (1928).
Annual herbs. Indumentum absent or of basifixed
hairs. Leaves alternate, serrate-dentate. Capitula
solitary or in lax corymbs, pedunculate, radiate.
Involucre meniscoid to hemispherical. Phyllaries
in 4 rows, with narrow to broad, whitish or pale
brown, scarious margins. Receptacle convex to conical, epaleate. Ray florets female, fertile or sterile;
limb yellow, sometimes white with yellow base. Disc
florets hermaphrodite, fertile; corolla 5-lobed, yellow. Achenes cylindrical, slightly curved to arcuate, with c. 10 ribs; base adaxially with a prominent
whitish callus; apex with a scarious adaxial auricle,
a scarious corona, or rarely marginally rounded;
pericarp with myxogenic cells along the ribs and
with resin canals between the ribs. x = 9. Three
species, Mediterranean region, Macaronesia.
1064. Glossopappus Kunze
Glossopappus Kunze, Flora 29: 748 (1846).
Glabrous annual herb. Leaves alternate, entire to
serrate-dentate. Capitula solitary, pedunculate,
radiate. Involucre meniscoid to hemispherical.
Phyllaries in 4 rows, with a fleshy central part and
apically broad whitish to pale brown, scarious
margins. Receptacle conical, epaleate. Ray florets
female, fertile or sterile; limb yellow or rarely white
with yellow base. Disc florets hermaphrodite,
fertile; corolla 5-lobed, actinomorphic or slightly
zygomorphic by two lobes having elongate appendages, yellow, basally abaxially saccate and
clasping top of achene. Achenes cylindrical, with
c. 10 ribs; base adaxially with a prominent, whitish
callus; apex with a large scarious adaxial auricle,
or rarely marginally rounded; pericarp with myxogenic cells along the ribs and with resin canals
between the ribs. x = 9. One species, G. macrotus
(Durieu) Briq. & Cavill., south-western Europe,
northern Africa.
Compositae
1065. Leucanthemum Mill.
Leucanthemum Mill., Gard. Dict. Abr., ed. 4: 769 (1754);
Vogt, Ruizia 10: 1–261 (1991); Villard, Ber. Schweiz. Bot.
Gesell. 80: 96–188 (1970); Horvatic, Acta Bot. Croat. 22:
203–218 (1963), rev.
Phalacrodiscus Less. (1832).
Perennial herbs, rarely subshrubs. Indumentum
absent or of basifixed hairs. Leaves alternate, entire, dentate-serrate, lobed, or up to 3-pinnatisect.
Capitula solitary or in lax corymbs, pedunculate,
radiate or rarely discoid. Involucre meniscoid to
hemispherical. Phyllaries in 4–5 rows, with pale to
blackish scarious margins. Receptacle flat to convex, epaleate. Ray florets female, fertile; limb white.
Disc florets hermaphrodite, fertile; corolla 5-lobed,
yellow. Achenes obovoid to cylindrical, with c. 10
ribs; apex marginally rounded or with a scarious
corona, or an adaxial auricle; pericarp with myxogenic cells along the ribs and with resin canals
between the ribs. x = 9 (polyploidy). Forty-three
species, Europe, Siberia. Some species widely cultivated as ornamentals or widespread as weeds.
1066. Mauranthemum Vogt & Oberprieler
Mauranthemum Vogt & Oberprieler, Taxon 44: 377 (1995).
Leucoglossum B.H. Wilcox, K. Bremer & Humphries (1993),
nom. illegit.
Annual, rarely perennial, glabrous herbs. Leaves
alternate, dentate-serrate to 2-pinnatisect. Capitula solitary, pedunculate, radiate. Involucre hemispherical. Phyllaries in 3–4 rows, with pale to dark
brown or black scarious margins. Receptacle conical, epaleate. Ray florets female or neuter, fertile
or sterile; limb white, rarely yellow at base. Disc
florets hermaphrodite, fertile; corolla 5-lobed, yellow. Achenes ellipsoid to obovoid, with c. 10 ribs;
apex with a scarious corona, an adaxial auricle,
or (in disc achenes) marginally rounded; pericarp
with myxogenic cells along the ribs and with resin
canals between the ribs. x = 9. Four species, southwestern Europe, north-western Africa. One species
widely cultivated as an ornamental.
1067. Plagius L’Hér. ex DC.
Plagius L’Hér. ex DC., Prodr. 6: 135 (1838); Vogt &
Oberprieler, Willdenowia 36(Special Issue):47–68 (2006),
rev.
Herbaceous or suffruticose perennials. Indumentum absent or of basifixed, sometimes glandular
hairs. Leaves alternate, serrate-dentate. Capitula
373
solitary or in lax corymbs, pedunculate, discoid.
Involucre meniscoid to hemispherical. Phyllaries
in 3–4 rows, with pale to brown scarious margins. Receptacle flat to convex, epaleate. Florets
hermaphrodite, fertile; corolla 5-lobed, yellow.
Achenes cylindrical, slightly curved to arcuate,
with c. 10 ribs; base adaxially with a prominent,
whitish callus; apex with an oblique, adaxially
longer corona or marginally rounded; pericarp
with myxogenic cells along the ribs, and with resin
canals between the ribs. x = 9 (polyploidy). Three
species, southern Europe (Corsica, Sardinia),
northern Africa.
1068. Rhodanthemum (Vogt) B.H. Wilcox, K.
Bremer & Humphries
Rhodanthemum (Vogt) B.H. Wilcox, K. Bremer &
Humphries, Bull. Brit. Mus. (Nat. Hist.), Bot. 23: 141 (1993).
Perennial herbs or subshrubs. Indumentum of
basifixed or medifixed hairs. Leaves alternate,
rosulate, 1–3-pinnatisect, long-petiolate. Capitula
solitary, pedunculate, radiate. Involucre hemispherical. Phyllaries in 3–5 rows, with pale to dark
brown scarious margins. Receptacle convex to
hemispherical, epaleate. Ray florets female, fertile;
limb white, red-violet or rarely yellow. Disc florets
hermaphrodite, fertile; corolla 5-lobed, yellow
or reddish. Achenes narrowly obovoid, straight
or slightly curved, with 5–10 often strongly protruding ribs; apex with an often adaxially longer
corona; pericarp with myxogenic cells along
the ribs and with resin canals between the ribs.
x = 9. Fourteen species, south-western Europe,
north-western Africa.
1069. Lonas Adans.
Lonas Adans., Fam. Pl. 2: 118, 572 (1763).
Glabrous annual herb. Leaves alternate, pinnatisect. Capitula in a dense corymb, discoid. Involucre hemispherical to cylindrical. Phyllaries in 3–
4 rows, with narrow pale scarious margins. Receptacle narrowly conical, paleate; paleae narrowly
obovate, flat to slightly canaliculate, with a central
resin canal. Florets hermaphrodite, fertile; corolla
5-lobed, yellow. Achenes narrowly obovoid, round
in cross-section or slightly dorsiventrally flattened,
with 1 adaxial and 2 lateral ribs; apex with a scarious corona; pericarp with myxogenic cells abaxially
and on the ribs and with a resin sac apically in the
adaxial rib. x = 9. One species, L. annua (L.) Vines
& Druce, southern Europe (Italy), northern Africa.
374
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
1070. Nivellea B.H. Wilcox, K. Bremer &
Humphries
Nivellea B.H. Wilcox, K. Bremer & Humphries, Bull. Brit.
Mus. (Nat. Hist.), Bot. 23: 140 (1993); Vogt & Oberprieler,
Bot. J. Linn. Soc. 122: 123–135 (1996), rev.
Annual herb. Indumentum absent or of basifixed
hairs. Leaves alternate, lobed and lacerate. Capitula solitary, pedunculate, radiate. Involucre meniscoid and hemispherical. Phyllaries in 4–5 rows,
with broad pale brown to brown scarious margins.
Receptacle convex, epaleate. Ray florets female, fertile; limb white. Disc florets hermaphrodite, fertile;
corolla 5-lobed, yellow. Achenes narrowly obovoid,
slightly curved or straight, with c. 10 ribs; apex
marginally rounded; pericarp with myxogenic cells
along the ribs, without resin sacs. x = 9. One
species, N. nivellei (Braun-Blanq. & Maire) B.H.
Wilcox, K. Bremer & Humphries, north-western
Africa (Morocco).
1071. Otospermum Willk.
Otospermum Willk., Bot. Zeitung 22: 251 (1864).
Otocarpum Willk. (1892).
Glabrous annual herb. Leaves alternate, pinnatisect. Capitula solitary or in a lax corymb, pedunculate, radiate. Involucre meniscoid to hemispherical. Phyllaries in 3–4 rows, with brown scarious
margins. Receptacle convex to conical, epaleate.
Ray florets female, fertile; limb white. Disc florets hermaphrodite, fertile; corolla 5-lobed, yellow. Achenes basally connate with the receptacle,
narrowly obovoid, circular to dorsiventrally flattened in cross-section, with one abaxial, two lateral,
and 1–3 adaxial ribs; apex with an adaxial auricle;
pericarp with myxogenic cells along the ribs, without resin canals. x = 9 (polyploidy). One species,
O. glabrum (Lag.) Willk., south-western Europe,
northern Africa.
XVII. Tribe Inuleae Cass. (1819).
Tribe Plucheeae (Benth.)
A. Anderb. (1989).
A.A. Anderberg and P. Eldenäs
Trees, shrubs, subshrubs, perennial or annual
herbs. With elongated or tuberous (Jasonia)
rhizome. Stems with (e.g. Doellia, Laggera, Porphyrostemma, Limbarda, Rhanterium, Telekia
and Tessaria) or without resin ducts, generally
without fibres in the phloem. Laticiferous tissue
absent. Leaves alternate or subopposite, sometimes
(Cylindrocline, Monarrhenus) clustered at branch
tips, or confined to a basal rosette (Rhodogeron,
Sachsia), simple or pinnatifid to pinnatisect,
entire, serrate or dentate, sometimes fleshy (e.g.
Iphiona p.p., Limbarda), often glandular and hairy
with septate hairs (unseptate in Pterocaulon);
leaf-base sometimes clasping, or conspicuously
decurrent on the stem (Calostephane, Geigeria,
Laggera, Ondetia, Pterocaulon). Capitula solitary,
in corymbs, panicles, or in more or less pronounced spike-like to spherical secondary heads
(Caesulia, Neojeffreya, Pseudoblepharispermum,
Pterocaulon, Sphaeranthus and Triplocephalum),
heterogamous or homogamous, radiate, disciform
or discoid. Involucral bracts generally in many
rows, herbaceous or cartilaginous; stereome undivided. Receptacle flat, convex or concave, epaleate
or paleate. Female or rarely neuter marginal florets, when present, in one to several rows. Corolla
radiate, minutely radiate, tubular or filiform,
generally three-lobed, yellow, pink, mauve, purple
or white. Cypselas and pappus generally similar
to those of disc florets, sometimes flattened. Disc
florets perfect or functionally male, 4- or 5-lobed,
lobes generally short but sometimes long, yellow,
pink, mauve, purple or sometimes white. Anthers
ecalcarate or exceptionally calcarate; basally with
long or short, branched or unbranched tails, rarely
ecaudate; endothecial tissue radial or polarized.
Cells of filament collar flattened or mammillose.
Pollen caveate; sexine with one baculate layer
between the spines and two baculate layers in the
spine bases (Leins 1971). Style bifid or undivided;
style branches most often with acute sweepinghairs ending above the furcation or with obtuse
sweeping-hairs reaching far below the furcation;
stigmatic surface apically confluent and basally
separated in two distinct lines, rarely covering the
entire inner surface of the style branches. Cypselas
generally with five vascular strands, ellipsoid,
turbinate or triquetrous, often with sclerenchymatic ribs; sometimes with resiniferous ducts or
cavities; glabrous or with glandular hairs and/or
elongated non-myxogenic twin-hairs; twin-hairs
straight, anchor-shaped or uncinate; epidermal
cells with one large elongated oxalate crystal, with
crystal sand, or without crystals. Cypselas often
much reduced in functionally male florets. Pappus
of capillary bristles, capillary bristles and small
scales in combination, large scales only, sometimes
rim-like or absent.
Compositae
Inuleae are a mainly Eurasian and East and
South African tribe, but some genera (e.g. Pluchea)
have a worldwide distribution. About 66 genera and
687 species.
Inuleae (incl. Plucheeae) are a monophyletic
group most closely related to Heliantheae s.lat.,
as evidenced by molecular data from the ndhF
gene of the plastid genome (Kim and Jansen 1995;
Eldenäs et al. 1999; Anderberg et al. 2005). The
support for this is very robust, although data
from morphology (Bremer 1987) and trnL/trnF
(Bayer and Starr 1998) suggest that Inuleae s.lat.
are basal in subfamily Asteroideae. The close
relationship between Inuleae and Heliantheae is
supported also by the fact that genera such as
Blepharispermum, Athroisma and Anisopappus,
which have always been placed in Inuleae, have
been shown to belong to the Heliantheae clade, and
not to the Inuleae-Plucheeae group. As they differ
considerably from other Heliantheae, experts in
that tribe have been reluctant to accept them, and
consequently Panero and Funk (2002) placed them
in the new tribe Athroismeae. The DNA sequence
data from ndhF clearly support the separation
of Inuleae-Gnaphaliinae into the separate tribe
Gnaphalieae (Bremer 1987; Anderberg 1989,
1991a), but the distinction between Inuleae s.str.
and Plucheeae as outlined by Anderberg (1989,
1991a, b, c) is more complex. Anderberg et al.
(2005) found support for two major monophyletic
lineages in the Inuleae-Plucheeae complex. One
lineage corresponded to the core Inuleae s.str.,
with styles having acute sweeping-hairs ending
above the furcation, and cypselas provided with
one large oxalate crystal in each epidermis cell.
The other lineage was Plucheeae, which included
all genera placed there by Anderberg (1991c),
but also five genera earlier included by him
in Inuleae s.str. (Anderberg 1989, 1991b), viz.
Antiphiona, Calostephane (incl. Mollera), Geigeria,
Ondetia and Pegolettia. These five genera have
style branches with acute sweeping-hairs, like
the Inuleae s.str., but lack the characteristic,
large elongated crystal in the cells of the cypsela
epidermis. The monophyly of Inuleae s.str. is
robust, but the support for the plucheoid lineage
is weak. Since Inuleae and Plucheeae have been
found to form a robustly supported clade together,
but their interrelationships indicate that the core
plucheoid group of genera has evolved from inuloid ancestors, there is little sense in treating them
as two separate tribes. Several genera belonging to
Inuleae s.lat. have been shown to be polyphyletic
375
as presently circumscribed (Anderberg et al. 2005)
but, since no formal reclassification has yet been
presented, we have in the following maintained
the traditional generic classification, adding notes
on the generic status when relevant. The tribal
position of some genera has not been confirmed
by analysis of molecular data, but most of them are
likely to be members of Inuleae as circumscribed
here. Only Nanothamnus and Feddea have tentatively been treated here as members of Inuleae and
of uncertain position.
Key to the Genera10
1. Cypsela epidermis cells each with one large calcium
oxalate crystal. Flowers often yellow, rarely white. Capitula usually with long or short ray florets
2
– Cypsela epidermis cells without crystals or with many
small crystals. Flowers often purple, sometimes yellow
or whitish. Capitula with or without ray florets
29
2. Receptacle paleate
3
– Receptacle epaleate
12
3. Involucral bracts concrescent into a cupule. Receptacle
strongly concave
1088. Anvillea
– Involucral bracts not forming a cupule. Receptacle flat
or convex
4
4. Paleae villous. Cypselas villous with coiled hairs
1089. Lifago
– Paleae not villous. Cypselas glabrous or sparsely hairy
5
5. Leaves up to 50 cm long, petiolate with cordate lamina
1097. Telekia
– Leaves much smaller, neither distinctly petiolate nor
with cordate lamina
6
6. Inner involucral bracts enclosing outer florets. Pappus
of very broad flattened bristles or absent
1079. Rhanterium
– Inner involucral bracts not enclosing outer florets 7
7. Leaves with raised reticulate venation. Pappus of flattened bristles
1078. Rhanteriopsis
– Leaves not with raised reticulate venation. Pappus of
scales or absent
8
8. Capitula homogamous or minutely radiate. Cypselas
ellipsoid
1098. Chrysophthalmum
– Capitula with well-developed ray florets. Cypselas
more or less triquetrous
9
9. Cypselas with chambered secretory cavities along the
margins
1091. Asteriscus
– Cypselas without chambered secretory cavities along
the margins
10
10. Shrub. Leaves needle-shaped, or somewhat dentate
1090. Ighermia
– Herbs. Leaves flat, not needle-shaped
11
11. Capitula subtended by a row of leaf-like bracts
1092. Pallenis
– Capitula not subtended by leaf-like bracts
1093. Buphthalmum
10 With the exception of genera 1136. Nanothamnus Thoms.
and 1137. Feddea Urb.
376
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
12. Pappus of bristles, or of capillary bristles and short
scales
13
– Pappus absent
28
13. Pappus of bristles only
14
– Pappus of bristles and short scales
23
14. Pappus bristles of unequal length, in many rows 15
– Pappus bristles more or less equal in length, in one to
several rows
16
15. Leaves trifid, pinnatifid, needle-shaped or terete. Disc
floret corolla epidermis with long needle-shaped crystals; cell walls sinuous
1085. Iphiona p.p.
– Leaves simple, flat and elliptic. Disc floret corolla epidermis cells without long needle-shaped crystals; cell
walls straight
1077. Vieraea
16. Leaves conspicuously fleshy, semiterete, sometimes
apically tridentate. Cypselas with five secretory ducts
1087. Limbarda
– Leaves neither fleshy nor apically tridentate
17
17. Capitula with filiform female florets
1074. Blumea s.str.
– Capitula with only perfect florets, or with radiate female florets
18
18. Anthers with distinctly polarized endothecial tissue
19
– Anthers with radial endothecial tissue
20
19. Cypselas contracted, pappus bristles basally connate
into a cupule
1086. Dittrichia
– Cypselas not contracted, pappus bristles not forming
a cupule
1072. Duhaldea
20. Capitula small, few-flowered. Pappus bristles generally few
21
– Capitula larger, many-flowered. Pappus bristles many
1094. Inula
21. Leaves terete or almost so, somewhat fleshy
1076. Schizogyne
– Leaves flat, not fleshy
22
22. Capitula homogamous, florets perfect. Plants erect
with divaricate branches. Leaves lanceolate to narrowly oblong
1100. Varthemia
– Capitula generally heterogamous, outer female florets
radiate to almost tubular. When all florets perfect, then
plants compact and leaves elliptic 1099. Pentanema
23. Disc floret epidermis cells with long needle-shaped
crystals
24
– Disc floret epidermis cells without long needle-shaped
crystals
25
24. Corolla epidermis cell walls sinuous
1085. Iphiona p.p.
– Corolla epidermis cell walls straight
1084. Perralderia
25. Shrubs with dichotomously branched stem
1083. Allagopappus
– Herbs or normally branched shrubs
26
26. Anthers with radial endothecial tissue 1080. Pulicaria
– Anthers with polarized endothecial tissue
27
27. Capitula radiate. Inner involucral bracts not scarious
nor ciliate at apex
1081. Jasonia
– Capitula discoid. Inner involucral bracts scarious and
ciliate at apex
1082. Chiliadenus
28. Capitula with short ray florets. Cypselas not contracted
into a beak
1095. Amblyocarpum
– Capitula with tubular female florets. Cypselas contracted into a beak
1096. Carpesium
29. Receptacle paleate
30
– Receptacle epaleate
33
30. Capitula radiate, with yellow, radiate female florets
1106. Ondetia
– Capitula discoid, with whitish or pink filiform female
florets
31
31. Capitula arranged in spherical secondary heads 32
– Capitula in dense corymbs but not in spherical secondary heads
1112. Cylindrocline
32. Pappus of capillary bristles
1124. Neojeffreya
– Pappus absent
1127. Pseudoblepharispermum
33. Capitula arranged in spherical or spike- or racemelike secondary heads (many Pterocaulon with dense
raceme-like or spike-like arrangement of capitula) 34
– Capitula solitary or more or less congested but never
arranged in spherical or spike-like secondary heads or
dense raceme-like capitulescences
37
34. Pappus of capillary bristles. Secondary heads spikelike or spherical
1111. Pterocaulon
– Pappus absent, or a short scale-like rim
35
35. All secondary heads sessile in leaf axils 1073. Caesulia
– Secondary heads terminal
36
36. Herbs or subshrubs. Leaves decurrent
1126. Sphaeranthus
– Shrub. Leaves not decurrent 1125. Triplocephalum
37. Involucral bracts straw-coloured with a dark longitudinal midstripe
1109. Iphionopsis
– Involucral bracts not with a dark contrasting longitudinal midstripe
38
38. Delicate perennial herbs with a basal leaf-rosette and
a leafless scape with capitula in a terminal corymb 39
– Stems not scapose and with leaves not in a basal rosette
40
39. Capitula discoid
1108. Sachsia
– Capitula radiate
1107. Rhodogeron
40. Styles with acute sweeping-hairs ending at or above
the furcation
41
– Styles with obtuse sweeping-hairs extending down the
shaft below the furcation
48
41. Capitula with well-developed yellow ray florets
42
– Capitula without ray florets
43
42. Receptacle with numerous bristles
1105. Geigeria
– Receptacle naked, without bristles
1103. Calostephane
43. Capitula heterogamous, with filiform outer florets and
bisexual or functionally male disc florets
44
– Capitula homogamous with only one kind of functional floret
46
44. Leaves entire. Florets purple 1115. Pechuel-loeschea
– Leaves dentate to lobed
45
45. Leaves large and deeply lobed. Disc florets functionally
male
1075. Merrittia
– Leaves dentate or entire, rarely lobed. Disc florets generally perfect
1074. Blumea p.p.
46. Functionally dioecious shrubs
1110. Cratystylis
– Herbs or shrublets with perfect florets
47
47. Leaves pinnatifid or pinnatisect, pappus of barbellate
bristles
1102. Antiphiona
– Leaves entire or dentate, or if leaves pinnatifid, then
pappus plumose
1104. Pegolettia
48. Cypselas with conspicuous red resiniferous ducts or
cavities
49
– Cypselas without red resiniferous ducts or cavities 50
49. Cypselas with longitudinal resiniferous ducts. Pappus
of capillary bristles
1132. Doellia
Compositae
– Cypselas with 2- or 3-celled resiniferous cavities arranged in rows. Pappus absent or a short rim
1133. Porphyrostemma
50. Capitula radiate or disciform with minutely radiate or
almost tubular female florets
51
– Capitula disciform with filiform female florets
52
51. Leaves broad, florets purple
1134. Streptoglossa
– Leaves filiform, florets whitish 1135. Allopterigeron
52. Pappus absent, a short rim, or a few scale-like bristles
53
– Pappus of capillary bristles
57
53. Receptacle deeply concave. Pappus absent
1114. Litogyne
– Receptacle flat or slightly convex. Pappus absent or of
scale-like bristles
54
54. Capitula axillary near the base of the stem. Cypselas
with uncinate hairs
1121. Thespidium
– Capitula not axillary near the base of the stem. Cypselas not with uncinate hairs
55
55. Pappus absent, or exceptionally of free scale-like bristles
1113. Epaltes
– Pappus of scale-like bristles which are connate into
tube at base
56
56. Capitula large. Involucral bracts very broad, obtuse
1122. Coleocoma
– Capitula small. Involucral bracts not very broad, acute
1123. Delamerea
57. Shrubs. Capitula congested into dense corymbs.
Leaves confined to the distal part of the branches
1120. Monarrhenus
– Trees, shrubs or herbs. Capitula not congested into
dense corymbs. Leaves scattered along the stem or
branches
58
58. Tree. Capitula heterogamous, with only one functionally male floret.
1119. Tessaria
– Shrubs or herbs. Male florets, if present, more than
one
59
59. Perennial herbs with decurrent leaves. Anther tails
long and branched. Cypselas more than 3 mm long
1101. Stenachaenium
– Shrubs or herbs. Cypselas less than 3 mm. Anther tails
short or inconspicuous
60
60. Leaves bipinnatifid, anthers ecaudate
1116. Adelostigma
– Leaves not pinnatifid, or anthers caudate
61
61. Disc florets functionally male with reduced cypselas.
Pappus bristles generally with erecto-patent teeth 62
– Disc florets perfect with developing cypselas. Bristles
often with adpressed teeth
63
62. Capitula solitary, terminal and > 1 cm long
1118. Karelinia
– Capitula in corymbs, or seldom solitary but then <
1 cm long
1117. Pluchea
63. Anthers ecaudate
64
– Anthers caudate
1128. Pseudoconyza
and
1074. Blumea p.p.
64. Flowers yellow. Leaves sharply dentate, not decurrent
1129. Blumeopsis
– Flowers purple. Leaves often decurrent
65
65. Pappus with numerous bristles
1130. Laggera
– Pappus with few bristles
1131. Nicolasia
377
Genera of Inuleae
1072. Duhaldea DC.
Duhaldea DC., Prodr. 5: 366 (1836).
Inula L. sect. Cappa DC. (1836).
Shrubs or perennial herbs. Leaves alternate,
simple, hairy. Capitula heterogamous, radiate or
disciform, solitary or in terminal corymbs. Receptacle epaleate, with scale-like ridges. Marginal
florets female. Corolla yellow to white, radiate to
miutely radiate. Disc florets perfect. Corolla yellow
or whitish. Anthers ecalcarate, with branched tails;
apical appendix truncate and almost emarginate;
endothecial tissue polarized. Style branches with
acute sweeping-hairs not reaching the furcation.
Cypselas ellipsoid, hairy; epidermis with elongated
crystals. Pappus of barbellate, capillary bristles in
one row. 2n = 18, 20, 40. Fourteen species, Asia,
possibly one species in Africa.
1073. Caesulia Roxb.
Caesulia Roxb., Pl. Corom. 1: 64. t. 93 (1759).
Annual herb. Leaves alternate, linear-lanceolate,
serrate, glabrous. Capitula in dense axillary
secondary heads, homogamous, discoid, oneflowered. Involucral bracts two, enclosing the
fruit, bracts with resiniferous ducts. Receptacle
epaleate. Florets perfect. Corolla whitish, deeply
lobed. Anthers ecalcarate, with branched tails;
endothecial tissue polarized. Style branches with
semi-acute sweeping-hairs not reaching the furcation. Stigmatic area covering almost the entire
inner surface. Cypselas dorsiventrally flattened;
pericarp one-layered, epidermis crystals missing.
Pappus missing (but with two awns formed by the
enclosing bracts). One species, C. axillaris Roxb.,
India.
1074. Blumea DC.
Blumea DC., Prodr. 5: 447 (1836), nom. cons.; Randeria,
Blumea 10: 176–317 (1960), rev.
Shrubs or herbs. Leaves alternate, simple, dentate,
serrate or lobed, hairy. Capitula heterogamous, disciform, in loose or dense corymbs or panicles,
or more or less solitary. Receptacle epaleate, with
scale-like ridges. Marginal florets female, in several rows. Corolla yellow or purple, filiform. Disc
florets perfect. Corolla yellow, white or purple. Anthers minutely calcarate, tailed; endothecial tissue radial or polarized. Style branches with acute
378
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
sweeping-hairs not reaching the furcation (type)
or obtuse sweeping-hairs reaching below the furcation. Cypselas narrowly oblong, hairy, shorter
than the corolla; epidermis with elongated crystals
(type) or not. Pappus of barbellate, capillary bristles in one row, with patent (type) or adpressed
teeth. 2n = 16, 18, 20, 22, 30, 32, 36, 44, 48, 54.
About 100 species, mainly subtropical and tropical
Asia.
Blumea is a heterogeneous assemblage with
considerable variation in micromorphological
characters. The type of the genus, B. balsamifera
(L.) DC., shares its characteristics with Duhaldea
DC. of Inuleae s.str., and the overall similarity
between Blumea balsamifera and Duhaldea cappa
(Buch.-Ham. ex D. Don) Pruski & Anderb. is also
striking. Molecular data have shown that many
other Blumea species belong to the same clade
as the type and Duhaldea, but also that some
species belong to other clades than the type. Two
former Blumea species (now in Doellia Sch. Bip.)
have obtuse sweeping-hairs reaching below the
furcation, and pappus bristles with adpressed
teeth, and their floral features more resemble
plucheoid genera such as Pseudoconyza Cuatr.
(Blumea aurita (L. f.) DC.). Blumea gariepina DC.
has acute sweeping-hairs and is closely related
to Antiphiona Merxm. The relationships with
Merrittia Merr. are still unclear.
1075. Merrittia Merr.
Merrittia Merr., Philip. J. Sci. Bot. 5: 396 (1910).
Herb. Leaves alternate, deeply and acutely lobed,
sparsely hairy. Capitula heterogamous, disciform,
small, several in loose corymbs. Receptacle
epaleate. Marginal florets female; corolla filiform.
Disc florets functionally male. Anthers tailed;
endothecial tissue radial. Style branches with
acute sweeping-hairs reaching below the furcation.
Cypselas sparsely hairy. Pappus of free, barbellate,
capillary bristles in one row; each bristle with
patent teeth. One species, M. benguetensis Merr.,
Philippines.
perfect. Corolla yellow. Anthers minutely calcarate,
with branched tails; endothecial tissue radial. Style
branches with acute sweeping-hairs not reaching
the furcation. Cypselas ellipsoid; epidermis imbricate, with elongated crystals. Pappus of barbellate,
capillary bristles in one row. 2n = 18. Two species,
Canary Islands.
1077. Vieraea Webb & Berth.
Vieraea Webb & Berth., Hist. Nat. Isles Canar. 3, 2: t. 84
(1839).
Shrub. Leaves alternate, coarsely serrate, glabrous,
glaucous, fleshy. Capitula heterogamous, radiate,
solitary. Receptacle epaleate, smooth. Marginal florets female. Corolla yellow, radiate. Disc florets
perfect. Corolla yellow. Anthers ecalcarate, with
branched tails; endothecial tissue polarized. Style
branches with acute sweeping-hairs not reaching
the furcation. Cypselas ellipsoid; epidermis with
elongated crystals. Pappus of unequal, barbellate,
capillary bristles in more than one row. 2n = 16.
One species, V. laevigata Webb & Berth., Canary
Islands.
1078. Rhanteriopsis S. Rauschert
Rhanteriopsis S. Rauschert, Taxon 31: 557 (1982); Wiklund,
Bot. J. Linn. Soc. 95: 27–42 (1987), rev., phylog.
Postia Boiss. & Blanche (1875), nom. illegit.
Subshrubs. Leaves alternate, elliptic, with alveolate
surface, hairy. Capitula solitary, heterogamous, radiate or homogamous, discoid. Receptacle paleate;
paleae folded. Marginal female florets present or
often absent. Corolla yellow, radiate. Disc florets
perfect. Corolla yellow. Anthers minutely calcarate,
with short tails; endothecial tissue radial. Style
branches with acute sweeping-hairs not reaching
the furcation. Cypselas subcylindrical; epidermis
with elongated crystals. Pappus of few, very broad,
flattened bristles in one row. Four species, Middle
East.
1079. Rhanterium Desf.
1076. Schizogyne Cass.
Schizogyne Cass., Dict. Sci. Nat. 56: 23 (1828).
Rhanterium Desf., Fl. Atl. 2: 291 (1799); Wiklund, Bot.
J. Linn. Soc. 93: 213–246 (1986), rev., phylog.
Shrubs. Leaves alternate, linear, somewhat fleshy,
glabrous or sericeous. Capitula heterogamous,
disciform, in dense terminal corymbs. Receptacle
epaleate. Marginal florets female. Corolla yellow,
minutely radiate to almost tubular. Disc florets
Shrublets. Leaves alternate, lanceolate, hairy.
Capitula solitary, terminal, heterogamous, radiate. Receptacle smooth, paleate; paleae folded.
Involucre hemispherical; inner involucral bracts
enclosing the outer florets. Marginal florets female.
Compositae
379
Corolla yellow, radiate. Disc florets perfect. Corolla
yellow. Anthers minutely calcarate, with short
tails; endothecial tissue polarized. Style branches
with acute sweeping-hairs not reaching the furcation. Cypselas ribbed; epidermis with elongated
crystals. Pappus of very broad, flattened bristles in
one row, or absent. 2n = 24. Three species, North
Africa to Middle East.
Corolla yellow. Anthers minutely calcarate, with
branched tails; endothecial tissue polarized. Style
branches with acute sweeping-hairs not reaching
the furcation. Cypselas ellipsoid, hairy; epidermis
with elongated crystals. Pappus of barbellate,
capillary bristles in one row with an outer row of
small scales. 2n = 18. One species, J. tuberosa DC.,
Spain.
1080. Pulicaria Gaertn.
1082. Chiliadenus Cass.
Pulicaria Gaertn., Fruct. 2: 461 (1791); Gamal-Eldin,
Phanerog. Monogr. 14 (1981), reg. rev.; Wagenitz &
Gamal-Eldin, Bot. Jahrb. Syst. 104: 91–113 (1983), rev.
Francoeuria Cass. (1825).
Platychaete Boiss. (1845).
Sclerostephane Chiov. (1929).
Chiliadenus Cass., Dict. Sci. Nat. 34: 34 (1825); Brullo,
Webbia 34: 289–308 (1979), rev.
Shrubs, shrublets, perennial or annual herbs.
Leaves alternate, simple, entire to dentate, hairy.
Capitula solitary or in more or less dense corymbs,
heterogamous, radiate or disciform or homogamous, discoid. Receptacle epaleate, often with
with scale-like ridges. Marginal female florets
present or absent. Corolla yellow, radiate or
minutely radiate. Disc florets perfect. Corolla
yellow. Anthers minutely calcarate, with branched
tails; endothecial tissue radial. Style branches with
acute sweeping-hairs not reaching the furcation.
Cypselas ellipsoid to cylindrical, in a few species
with a tube-shaped distal outgrowth surrounding
the base of the corolla, often apically contracted
and glandular, hairy; epidermis cells with elongated crystals. Pappus of barbellate, capillary to
more or less flattened bristles in one row with
an outer cup of connate scales (one species with
bristles only). 2n = 12, 14, 16, 18, 20, 36. About 85
species, Europe, Africa, Arabia, Asia.
Pulicaria is heterogeneous and not monophyletic as presently circumscribed. Genera such
as Jasonia Cass. and Dittrichia Greuter have
been shown to have their closest relatives within
Pulicaria (Anderberg et al. 2005). The status of
Francoeuria Cass. also needs to be clarified.
Perennial herbs or shrublets. Leaves alternate,
sessile, ovate to lanceolate, glandular, hairy. Capitula homogamous, discoid, solitary or in dense
raceme-like corymbs. Receptacle with scale-like
ridges, epaleate. Florets perfect. Corolla yellow.
Anthers long, minutely calcarate, with very short
tails; endothecial tissue polarized. Style branches
with acute sweeping-hairs not reaching the furcation. Cypselas ellipsoid, hairy, apically constricted
and glandular; epidermis with elongated crystals.
Pappus of barbellate, capillary bristles in one row
with an outer row of narrow scales. 2n = 16, 18.
Ten species, Mediterranean region.
1083. Allagopappus Cass.
Allagopappus Cass., Dict. Sci. Nat. 56: 21 (1828).
Shrubs, stem subdichotomously branched. Leaves
confined to tips of branches, alternate, glabrous,
dentate, viscid. Capitula small, homogamous, discoid, in umbel-like corymbs. Receptacle smooth,
epaleate. Florets perfect. Corolla yellow. Anthers
minutely calcarate with short tails; endothecial tissue polarized. Style branches with acute sweepinghairs not reaching the furcation. Cypselas elliptic,
hairy and apically glandular; epidermis with elongated crystals. Pappus of barbellate capillary bristles in one row with an outer row of narrow scales.
2n = 20. Two species, Canary Islands.
1081. Jasonia Cass.
1084. Perralderia Coss.
Jasonia Cass., Dict. Sci. Nat. 24: 200 (1822); Brullo, Webbia
34: 289–308 (1979), rev.
Perralderia Coss., Bull. Soc. Bot. France 6: 394 (1859);
Eldenäs, Bot. J. Linn. Soc. 102: 157–173 (1990), rev., phylog.
Fontquera Maire (1931).
Perennial herb with tuberous rhizome. Leaves
alternate, simple, lanceolate to linear-lanceolate,
hairy. Capitula heterogamous, radiate. Receptacle
epaleate, with scale-like ridges. Marginal florets
neuter. Corolla yellow, radiate. Disc florets perfect.
Shrublets or subshrubs. Leaves alternate, somewhat fleshy, pinnatisect to bi- or tripinnatisect;
axils with hair tufts. Capitula heterogamous,
radiate or homogamous, discoid. Receptacle
380
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
epaleate, with scale-like ridges. Neuter marginal
florets present or absent. Corolla yellow, radiate;
epidermal cells papillose. Disc florets perfect.
Corolla yellow or purple-tinged. Anthers minutely
calcarate, with short tails; endothecial tissue polarized. Style branches with acute sweeping-hairs
not reaching the furcation. Cypselas ellipsoid,
hairy; epidermis with elongated crystals. Pappus
of barbellate, capillary bristles in one row with an
outer row of short scales. 2n = 18. Three species,
North Africa.
1085. Iphiona Cass.
Fig. 72
Iphiona Cass., Bull. Sci. Soc. Philom. Paris 1817: 153 (1817),
nom. cons.; Anderberg, Nordic J. Bot. 5: 169–194 (1985),
rev., phylog.
Grantia Boiss. (1845).
Hirschia Baker (1895).
Perralderiopsis S. Rauschert (1982).
Shrubs or perennial herbs. Leaves alternate, sessile, entire, lobed or pinnatisect, often fleshy, sometimes spiny, hairy or glabrous. Capitula solitary or
few, heterogamous, radiate to minutely radiate or
homogamous, discoid. Receptacle epaleate, with
scale-like ridges. Neuter marginal florets present
or absent. Corolla yellow, radiate. Disc florets perfect. Corolla yellow; epidermis with sinuous cell
walls, with needle-like crystals. Anthers minutely
calcarate, with short tails; endothecial tissue polarized. Style branches with acute sweeping-hairs not
reaching the furcation. Cypselas ellipsoid, conspicuously ribbed; epidermis with elongated crystals.
Pappus of unequal, spreading, barbellate, capillary
bristles in several rows, or equally long capillary
bristles in one row with an outer row of small scales.
2n = 18, 20. Twelve species, north-eastern Africa,
Arabia to Pakistan.
1086. Dittrichia Greuter
Dittrichia Greuter, Exsiccat. Genavensium fasc. 4: 71
(1973).
Fig. 72.
Compositae-Inuleae. Iphiona anthemidifolia.
A Habit. B Involucral bracts. C Ray floret. D Disc floret.
E Stamen. F Style. (Anderberg 1985)
Annual or perennial herbs. Leaves alternate,
lanceolate to linear, glandular, viscid, dentate,
hairy. Capitula heterogamous, radiate, many
in terminal racemes. Receptacle epaleate, with
scale-like ridges. Marginal florets female. Corolla
yellow, radiate or minutely radiate. Disc florets
perfect. Corolla yellow. Anthers ecalcarate, with
branched tails; endothecial tissue polarized. Style
branches with acute sweeping-hairs not reaching
the furcation. Cypselas ellipsoid, hairy, apically
Compositae
381
somewhat constricted, glandular; epidermis
with elongated crystals. Pappus of barbellate,
capillary bristles in one row, basally connate and
concrescent into a cupule. 2n = 16, 18, 20. Two
species, Mediterranean, introduced in America
and Australia.
1087. Limbarda Adans.
Limbarda Adans., Fam. 2: 125, 570 (1763).
Shrublet. Leaves alternate, fleshy, glabrous, subterete, sessile, entire or apically three-lobed. Capitula heterogamous, radiate, solitary or in loose
corymbs. Receptacle epaleate. Marginal florets female. Corolla yellow, radiate. Disc florets perfect.
Corolla yellow. Anthers ecalcarate, with branched
tails; endothecial tissue radial. Style branches with
acute sweeping-hairs not reaching the furcation.
Cypselas ellipsoid, with five secretory ducts; epidermis with elongated crystals. Pappus of barbellate, capillary bristles in several rows. 2n = 18.
One species, L. crithmoides (L.) Dumort., Mediterranean region.
1088. Anvillea DC.
Fig. 73
Anvillea DC., Prodr. 5: 487 (1836); Anderberg, Nordic J. Bot.
2: 297–305 (1982), rev.
Anvilleina Maire (1940).
Shrublets. Leaves alternate, lanceolate to cuneate
or spathulate, hairy. Capitula heterogamous, radiate or homogamous, discoid, solitary. Involucre
concrescent into hard spiny cup with age. Receptacle concave, smooth, paleate, paleae subtending
the florets, basally folded with elongated apical
appendix. Neuter marginal florets present or absent. Corolla yellow, radiate. Cypselas flattened,
subcordate, with anchor-shaped hairs. Pappus absent. Disc florets perfect. Corolla yellow. Anthers
minutely calcarate, with long tails; endothecial tissue polarized. Style branches with acute sweepinghairs not reaching the furcation. Cypselas quadrangular, ribbed, with anchor-shaped hairs; epidermis
with elongated crystals. Pappus absent. 2n = 14.
Two species, Morocco to Iran.
1089. Lifago Schweinf. & Muschl.
Lifago Schweinf. & Muschl., Bot. Jahrb. 45: 429 (1911).
Niclouxia Battand. (1915).
Annual herb. Leaves spathulate, white-woolly.
Capitula heterogamous, radiate, falling as a unit.
Receptacle paleate; paleae subulate, villous.
Fig. 73. Compositae-Inuleae. Anvillea garcinii. A Habit,
subsp. garcinii. B Habit, subsp. radiata. C Ray floret. D Disc
floret. E Stamen. F Style. G Palea, subsp. garcinii. H Palea,
subsp. radiata. (Anderberg 1982)
Marginal florets female. Corolla yellow, radiate.
Cypselas triquetrous, villous. Pappus absent. Disc
florets perfect. Corolla yellow. Anthers minutely
calcarate, with branched tails; endothecial tissue
polarized. Style branches with acute sweepinghairs not reaching the furcation. Cypselas flattened
or tetragonous, glabrous but distally with long,
brown hairs; epidermis with elongated crystals.
Pappus absent. One species, L. dielsii Schweinf. &
Muschl., Algeria and Morocco.
382
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
1090. Ighermia Wikl.
Ighermia Wikl., Nordic J. Bot. 3: 445 (1983).
Shrub. Leaves alternate, simple, narrowly linear,
needle-like. Capitula solitary, heterogamous, radiate. Receptacle smooth, paleate; paleae folded. Involucre hemispherical; involucral bracts hard and
cartilaginous. Marginal florets female. Corolla yellow, radiate. Disc florets perfect. Corolla yellow.
Anthers long-calcarate, with short tails; endothecial tissue polarized. Style branches short, blunt
with subacute sweeping-hairs not reaching the furcation. Cypselas triquetrous, glabrous; sclerenchymatic tissue more or less confluent; epidermis with
elongated crystals. Pappus of large hard scales.
2n = 14. One species, I. pinifolia (Maire & Wilczek)
Wikl., Morocco.
1091. Asteriscus Tourn. ex Mill.
Asteriscus Tourn. ex Mill., Gard. Dict. 1, ed. 4 (1754);
Wiklund (as Nauplius), Nordic J. Bot. 7: 1–23. (1987), rev.;
Halvorsen & Borgen, Sommerfeltia 3: 1–103 (1986), rev.;
Greuter, Flora Mediterranea 7: 41–48 (1997) nomencl.;
Francisco-Ortega et al., Biol. J. Linn. Soc. 72: 77–97 (2001),
phylog.
Bubonium J. Hill (1761).
Nauplius Cass. (1822).
Odontospermum Necker ex Schultz-Bip. (1844).
Shrublets, perennial or annual herbs. Leaves
alternate, sessile, sometimes fleshy, generally
entire to somewhat lobed, hairy. Capitula solitary,
heterogamous, radiate. Receptacle convex to flat,
paleate; paleae subtending the florets, folded and
crested with secretory cavities under the crest.
Marginal florets female. Corolla yellow, radiate;
epidermis crested. Disc florets perfect, yellow.
Corolla yellow. Anthers long-calcarate, with short
tails; endothecial tissue polarized. Style branches
with subacute sweeping-hairs not reaching the
furcation. Cypselas more or less triquetrous, with
confluent sclerenchymatic tissue and chambered
secretory cavities; epidermis with elongated
crystals. Pappus of broad scales in c. two rows.
2n = 14, 16, 18. Eight species, southern Europe,
North Africa, Middle East, Macaronesia.
1092. Pallenis (Cass.) Cass.
Pallenis (Cass.) Cass. in Cuvier, Dict. Hist. Nat. 23: 566
(1822), nom. cons.; Wiklund (as Asteriscus), Nordic J. Bot. 5:
299–314 (1985), rev.; Greuter, Flora Mediterranea 7: 41–48
(1997), nomencl.; Francisco-Ortega et al., Biol. J. Linn. Soc.
72: 77–97 (2001), phylog.
Subshrubs, perennial or annual herbs. Leaves alternate, simple, not lobed, hairy. Capitula heterogamous, radiate, solitary, hygrochastic or not. Receptacle smooth, paleate, paleae folded and crested.
Involucre broadly campanulate with subtending
leaves. Marginal florets female. Corolla yellow, radiate; epidermis crested. Cypselas oblong-obovate,
terete or dorsiventrally compressed and winged;
epidermis with elongated crystals. Pappus of large
scales or absent. Disc florets perfect. Corolla yellow.
Anthers long-calcarate, with short tails; endothecial tissue polarized. Style branches short with subacute sweeping-hairs not reaching the furcation.
Cypselas somewhat triquetrous; sclerenchymatic
ribs few and inconspicuous; epidermis with elongated crystals. Pappus of large scales. 2n = 10, 12.
Three species, Mediterranean region, Middle East,
one species introduced elsewhere.
1093. Buphthalmum L.
Buphthalmum L., Sp. Pl.: 903 (1753).
Xerolekia A. Anderb. (1991).
Perennial herbs. Leaves alternate, entire or dentate, hairy. Capitula heterogamous, radiate, solitary. Receptacle paleate, paleae folded, subtending
the florets. Involucre hemispherical, phyllaries linear, herbaceous. Marginal florets female. Corolla
yellow, radiate. Cypselas triquetrous to flattened.
Pappus absent or a rim of scales. Disc florets perfect. Corolla yellow. Anthers long- or minutely calcarate, with very short tails; endothecial tissue polarized. Style branches with acute sweeping-hairs
not reaching the furcation. Cypselas flattened to
triquetrous, glabrous or hairy along the edges; epidermis with elongated crystals. Pappus a rim of
irregularly incised scales. 2n = 20. Three species,
southern and central Europe, Sardinia.
1094. Inula L.
Inula L., Sp. Pl.: 881 (1753).
Bojeria DC. (1836).
Codonocephalum Fenzl (1843).
Perennial or annual herbs. Leaves alternate, simple, hairy or glabrous. Capitula solitary or few in
corymb, heterogamous radiate to minutely radiate, heterogamous disciform, or homogamous discoid. Receptacle smooth, epaleate. Marginal florets female. Corolla yellow, radiate, minutely radiate or rarely tubular. Disc florets perfect. Corolla
yellow. Anthers ecalcarate, with branched tails; endothecial tissue radial. Style branches with acute
Compositae
sweeping-hairs not reaching the furcation. Cypselas ellipsoid, cylindrical or angled, ribbed; epidermis with elongated crystals. Pappus of barbellate,
capillary bristles in one or several rows. 2n = 16,
18, 24, 32, 36. About 100 species, Old World.
Inula is heterogeneous and not monophyletic
as now circumscribed. Genera such as Telekia
Baumg., Pentanema Cass., Amblyocarpum Fisch.
& Mey. and Chrysophthalmum Sch. Bip. have their
closest relatives within Inula (Anderberg et al.
2005).
1095. Amblyocarpum Fisch. & Mey.
Amblyocarpum Fisch. & Mey., Index Sem. Hort. Bot. Petrop.
3 (1837).
Annual herb. Leaves alternate, simple, entire, hairy.
Capitula heterogamous, disciform, subtended by
radiating leaves. Receptacle smooth, epaleate.
Marginal florets female. Corolla yellow, minutely
radiate. Disc florets perfect. Corolla yellow. Anthers ecalcarate with long tails; endothecial tissue
radial. Style branches with acute sweeping-hairs
not reaching the furcation. Cypselas cylindrical,
longer than the florets, glandular; epidermis with
elongated crystals. Pappus absent. One species,
A. inuloides Fisch. & Mey., Caspian Sea region.
383
flowering. Involucral bracts herbaceous, cordate.
Marginal florets female. Corolla yellow, radiate;
ray very long, linear. Disc florets perfect. Corolla
yellow. Anthers minutely calcarate, with branched
tails; endothecial tissue polarized. Style branches
with acute sweeping-hairs not reaching the furcation. Cypselas linear, flattened; epidermis with
elongate crystals. Pappus a rim of minute scales, or
absent. 2n = 20. One species, T. speciosa Baumg.,
Europe, Caucasus, Asia Minor.
1098. Chrysophthalmum Sch. Bip.
Chrysophthalmum Sch. Bip., Walpers Repert. 2: 955 (1843)
[as Chrysophtalmum]; Aytec & Anderberg, Bot. J. Linn. Soc.
137: 211–214 (2001).
Perennial herbs or subshrubs. Leaves simple, alternate, hairy. Capitula heterogamous, disciform. Receptacle smooth, paleate; paleae linear, somewhat
canaliculate. Marginal florets female. Corolla yellow, minutely radiate. Disc florets perfect. Corolla
yellow. Anthers ecalcarate, with branched tails; endothecial tissue radial. Style branches with acute
sweeping-hairs not reaching the furcation. Cypselas elongated; epidermis with elongated crystals.
Pappus a short rim of scales, or absent. Three
species, Turkey, Syria, Iraq.
1096. Carpesium L.
Carpesium L., Sp. Pl.: 859 (1753).
Perennial or annual herbs. Leaves alternate, often
in a rosette, hairy, serrate. Capitula heterogamous,
disciform, solitary or few, or many in spiciform
racemes. Receptacle smooth, epaleate. Marginal
florets female, tubular to minutely radiate, in
many rows. Corolla yellow. Disc florets perfect.
Corolla yellow. Anthers ecalcarate, with branched
tails; endothecial tissue radial. Style branches with
acute sweeping-hairs not reaching the furcation.
Cypselas ellipsoid, bottle-necked with conspicuous
glands, longer than the corolla, ribbed; epidermis
cells with elongated crystals. Pappus absent.
2n = 20, 36, 40. Twenty-five species, mainly Asia
but extending into Europe and Australia.
1097. Telekia Baumg.
Telekia Baumg., Enum. Stirp. Transsilv. 3: 149 (1816).
Perennial herb. Leaves large, alternate, cordate,
dentate, hairy. Capitula heterogamous, radiate,
large, in loose corymbs. Receptacle paleate; paleae
subulate with a widened base, persistent after
1099. Pentanema Cass.
Pentanema Cass., Bull. Sci. Soc. Philom. Paris 1818: 74
(1818).
Vicoa Cass. (1825).
Shrublets or annual herbs. Leaves alternate, oblong to lanceolate, entire to serrate, often villous.
Capitula generally heterogamous, radiate, but in
some species heterogamous, disciform or homogamous discoid. Overwintering young capitula often present. Receptacle epaleate, smooth. Marginal
florets female. Corolla yellow, radiate, minutely radiate or more or less tubular. Disc florets perfect.
Corolla yellow. Anthers ecalcarate, with branched
tails; endothecial tissue radial. Style branches with
acute sweeping-hairs not reaching the furcation.
Cypselas ellipsoid, shorter than the corolla; epidermis with elongated crystals. Pappus of barbellate,
capillary bristles in one row. 2n = 18, 27. About 20
species, Turkey through central Asia, India and Sri
Lanka to Africa.
Pentanema is heterogeneous and it seems
doubtful that it is monophyletic with its present
circumscription.
384
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
1100. Varthemia DC.
Varthemia DC., Prodr. 5: 473 (1836).
Shrublet. Leaves alternate, lanceolate to narrowly
oblong, sparsely hairy. Capitula homogamous,
discoid, in loose corymbs. Receptacle epaleate,
smooth. Florets perfect. Corolla yellow. Anthers
ecalcarate, with branched tails; endothecial tissue
radial. Style branches with acute sweeping-hairs
not reaching the furcation. Cypselas shorter
than the corolla, ellipsoid, hairy; epidermis with
elongated crystals. Pappus of barbellate, capillary
bristles in one row. 2n = 16. One species, V. persica
DC., Afghanistan, Iran, Pakistan.
1101. Stenachaenium Benth.
Stenachaenium Benth. in Benth. & Hook. f., Gen. Pl. 2: 289
(1873).
Herbs. Leaves alternate, dentate to serrate, decurrent, tomentose. Capitula heterogamous,
disciform, terminal, solitary or few. Receptacle
epaleate. Involucral bracts in several rows. Florets
white, yellow or purple. Marginal florets female;
corolla filiform. Disc florets perfect. Anthers
tailed; endothecial tissue polarized. Style branches
with obtuse sweeping-hairs reaching below the
furcation. Cypselas ellipsoid, either dark brown,
smooth, glossy, laterally compressed, or pale
brown, terete, ribbed, with straight hairs. Pappus
of numerous free, stiff, barbellate, capillary bristles
in one row; each bristle with patent teeth. Five
species, South America (Brazil, Uruguay, Paraguay,
Argentina).
1102. Antiphiona Merxm.
Antiphiona Merxm., Mitt. Bot. Staatssamml. München 9–
10: 432 (1954).
Shrubs or subshrubs. Leaves alternate, pinnatisect
or bi- to tripinnatisect. Capitula homogamous, discoid, solitary or few. Receptacle smooth, epaleate.
Florets perfect. Corolla purple, corolla lobes with
acute one-celled hairs. Anthers minutely calcarate
with long tails; endothecial tissue polarized. Style
branches with acute sweeping-hairs not reaching
the furcation. Cypselas ellipsoid, hairy; epidermis
without elongated crystals. Pappus of barbellate,
capillary bristles. Two species, Namibia.
1103. Calostephane Benth.
Calostephane Benth., Hooker’s Ic. Pl. 12: 1111 (1872).
Mollera O. Hoffm. (1890).
Herbs. Stem winged or not. Leaves alternate, simple, with scattered hairs. Capitula heterogamous,
radiate. Receptacle smooth, epaleate. Marginal florets female. Corolla yellow, radiate. Disc florets
perfect. Corolla yellow. Anthers minutely calcarate,
tailed; endothecial tissue polarized. Style branches
with acute sweeping-hairs not reaching the furcation. Cypselas broadly elliptic, with glands alternating with ribs; epidermis without elongated
crystals. Pappus of short obtuse scales in two rows
often with protruding, bristle-like mid-portion, or
absent. Six species, tropical Africa, Madagascar.
1104. Pegolettia Cass.
Pegolettia Cass., Dict. Sci. Nat. 38: 230 (1825); Anderberg,
Cladistics 2: 158–186 (1986), rev., phylog.
Shrubs or herbs. Leaves alternate, simple, dentate
or entire, or pinnatifid, glandular, hairy. Capitula
homogamous, discoid. Receptacle epaleate. Florets
perfect. Corolla yellow or purple-tinged; corolla
lobes with acute hairs. Anthers minutely calcarate,
with branched tails; endothecial tissue polarized.
Style branches with acute sweeping-hairs not
reaching the furcation. Cypselas cylindrical, with
sclerenchymatous ribs; epidermis with crystal
sand. Pappus of barbellate or plumose capillary
bristles in one row, with an outer row of small
narrow, entire to lacerate or almost bristle-like
scales. 2n = 20. Nine species, southern Africa, one
species extending into North Africa and Middle
East.
1105. Geigeria Griess.
Geigeria Griess., Linnaea 5: 411 (1830); Merxmüller, Mitt.
Bot. Staatssamml. München 1: 239–316 (1953), rev.
Herbs or shrublets. Stem winged. Leaves alternate,
sessile, simple, linear to lanceolate, sometimes
decurrent, hairy. Capitula heterogamous, radiate
(sometimes homogamous, discoid). Receptacle
epaleate but with numerous bristles. Marginal
female florets present or absent. Corolla yellow,
radiate. Disc florets perfect. Corolla yellow; corolla
deeply lobed. Anthers long-calcarate, with short
tails; endothecial tissue polarized. Style branches
with acute sweeping-hairs reaching below the
furcation. Cypselas ellipsoid or turbinate, villous;
epidermis without crystals. Pappus of large scales
in two rows, often with a long bristle-shaped
mid-portion. 2n = 20. Twenty-eight species,
Africa, mainly in the southern part.
Compositae
1106. Ondetia Benth.
Ondetia Benth., Hooker’s Ic. Pl. 12: t. 1112 (1872).
Perennial herb. Stem winged. Leaves alternate, linear. Capitula solitary, heterogamous, radiate. Receptacle paleate; paleae flattened. Marginal florets
female. Corolla yellow, radiate. Disc florets perfect.
Corolla yellow; corolla deeply lobed. Anthers longcalcarate, with short tails; endothecial tissue polarized. Style branches with acute sweeping-hairs
not reaching below the furcation. Cypselas villous;
epidermis without crystals. Pappus of short bristlelike scales. One species, O. linearis Benth., Namibia.
1107. Rhodogeron Griseb.
Rhodogeron Griseb., Cat. Pl. Cuba 151 (1866); Liu et al., Intl
J. Pl. Sci. 165: 209–217 (2004), phylog.
Perennial herb. Stem scapose. Leaves rosulate, dentate or pinnatifid, hairy. Capitula few in a corymb,
heterogamous, radiate. Receptacle epaleate. Florets purple. Marginal florets female; corolla radiate. Disc florets functionally male. Anthers tailed;
endothecial tissue polarized. Style undivided, with
acute sweeping-hairs reaching below the furcation.
Cypselas ellipsoid. Pappus of free, barbellate, capillary bristles in one row. One species, R. coronopifolius Griseb., Cuba.
1108. Sachsia Griseb.
Sachsia Griseb., Cat. Pl. Cuba 150 (1866); Liu et al., Intl J. Pl.
Sci. 165: 209–217 (2004), phylog.
Perennial herbs. Stem scapose. Leaves rosulate,
dentate, hairy. Capitula few in a corymb, heterogamous, disciform. Receptacle epaleate. Florets
white. Marginal florets female; corolla filiform.
Disc florets functionally male. Anthers tailed;
endothecial tissue polarized. Style undivided, with
acute sweeping-hairs reaching below the furcation.
Cypselas ellipsoid. Pappus of free, barbellate,
capillary bristles in one row. 2n = 20. Four species,
West Indies, southern USA.
1109. Iphionopsis A. Anderb.
Iphionopsis A. Anderb., Nordic J. Bot. 5: 52 (1985), rev.
Shrubs. Leaves alternate, entire or dentate, sparsely
hairy. Capitula homogamous, discoid, in corymbs.
Receptacle epaleate. Involucral bracts broad with
conspicuous median secretory duct. Florets perfect. Corolla white to yellowish, sometimes with
red tinge, with conspicuous longitudinal secretory
385
ducts. Anthers minutely calcarate, with short tails;
endothecial tissue radial. Style branches with subacute to obtuse sweeping-hairs not reaching the furcation. Cypselas hairy, without sclerenchymatic tissue; epidermis without crystals. Pappus of barbellate capillary bristles in several rows. Three species,
East Africa, Madagascar.
1110. Cratystylis S. Moore
Cratystylis S. Moore, J. Bot. 43: 138 (1905).
Stera Ewart (1912).
Dioecious shrubs. Leaves alternate, entire, hairy or
glabrous. Capitula solitary, narrowly cylindrical,
heterogamous but functionally homogamous, discoid. Receptacle epaleate. Florets white. Anthers
ecalcarate, shortly tailed; endothecial tissue polarized. Style branches almost smooth, with minute
acute sweeping-hairs reaching the furcation.
Cypselas ellipsoid, glabrous. Pappus of barbellate,
capillary bristles. 2n = 20. Four species, Australia.
1111. Pterocaulon Ell.
Fig. 74
Pterocaulon Ell., Sketch Bot. S.-Carolina Georgia 2: 323
(1823); Cabrera & Ragonese, Darwiniana 21: 185–257
(1978), rev.
Monenteles Labill. (1825).
Herbs. Leaves alternate, dentate to serrate, decurrent into long wings. Capitula heterogamous,
disciform, terminal, forming dense glomerules
or long, more or less dense spikes or racemes of
capitula. Receptacle epaleate. Florets purplish.
Marginal florets female; corolla filiform. Disc
florets perfect, purple. Anthers tailed with radial
endothecial tissue. Style branches with acute
sweeping-hairs not reaching the furcation. Cypselas hairy. Pappus of free, barbellate, capillary
bristles, in one to several rows; each bristle with
patent teeth. 2n = 20. Eighteen species, North and
South America, Australia and adjacent areas.
1112. Cylindrocline Cass.
Cylindrocline Cass., Bull. Sci. Soc. Philom. Paris 1817: 11
(1817).
Shrubs or small trees. Leaves elliptic, on tips of
branches, not decurrent, marginally dentate, whitewoolly below. Capitula heterogamous, disciform, in
dense terminal corymbs. Florets mauve. Receptacle
paleate; paleae hairy. Involucral bracts with a tuft of
hairs. Marginal florets female; corolla filiform. Disc
florets perfect. Anthers tailed; endothecial tissue
386
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
1113. Epaltes Cass.
Epaltes Cass., Bull. Sci. Soc. Philom. Paris 1818: 139 (1818).
Sphaeromorphaea DC. (1837).
Ethuliopsis F. Muell. (1861).
Herbs. Leaves alternate or subopposite, dentate,
sparsely hairy or glabrous. Capitula heterogamous,
solitary or few in terminal clusters. Receptacle
epaleate. Florets white or purple. Marginal florets
female; corolla filiform. Disc florets functionally
male. Anthers tailed; cells of filament collar mammillose; endothecial tissue radial. Style undivided,
with obtuse sweeping-hairs reaching far down
the shaft. Cypselas glandular, or sparsely hairy.
Pappus absent or present as a short rim or, as in
E. cunninghamii Benth., a few elongated, flattened
scale-like bristles. 2n = 20. Fourteen species,
pantropical.
Epaltes is heterogeneous and likely to be polyphyletic. The genus is diagnosed only by reduction
or loss of pappus.
1114. Litogyne Harv.
Litogyne Harv., Fl. Cap. 3: 48 (1865).
Subshrub. Leaves alternate, entire, decurrent, hairy.
Capitula heterogamous, disciform, few in terminal clusters. Receptacle epaleate, dome-shaped and
centrally depressed with a marginal ridge. Florets purple. Marginal florets female, situated on
the vertical outer face of the marginal receptacle ridge; corolla filiform. Disc florets functionally
male, in the central depression of the receptacle.
Anthers tailed; endothecial tissue radial. Style undivided, with obtuse sweeping-hairs reaching far
down the shaft. Cypselas hairy. Pappus absent. One
species, L. gariepina (DC.) Anderb., tropical Africa,
Namibia.
1115. Pechuel-loeschea O. Hoffm.
Pechuel-loeschea O. Hoffm., Bot. Jahrb. Syst. 10: 274 (1888).
Fig. 74.
Compositae-Inuleae. Pterocaulon virgatum.
A Flowering branch. B Capitulum. C Female floret.
D Hermaphrodite floret. E Stamen. F Style. (Cabrera 1978)
radial. Style branches with acute sweeping-hairs
reaching below the furcation. Cypselas terete to
triquetrous, sparsely hairy on the edges. Pappus
of a few free, stiff, scale-like bristles; each bristle
with patent teeth. Two species, Mauritius.
Shrub. Leaves alternate, lanceolate, sericeous.
Capitula heterogamous, disciform, in loose
corymbs. Receptacle epaleate. Marginal florets
female. Corolla purple, filiform. Disc florets perfect. Corolla purple. Anthers minutely calcarate,
with branched tails; endothecial tissue polarized;
cells of filament collar longer than wide. Style
branches with acute sweeping-hairs not reaching
the furcation. Cypselas ellipsoid, hairy; epidermis
without crystals. Pappus of one row of basally
Compositae
387
connate, barbellate, capillary bristles. One species,
P. leubnitziae O. Hoffm., Namibia.
1116. Adelostigma Steetz
Adelostigma Steetz in Peters, Naturwissensch. Reise
Mossambique 6, Bot.: 428 (1864).
Herbs. Leaves alternate, bipinnatifid. Capitula
heterogamous, disciform, solitary. Receptacle epaleate. Marginal florets female; corolla filiform. Disc
florets functionally male. Anthers very shortly
tailed; endothecial tissue radial. Style almost
undivided, with obtuse sweeping-hairs reaching
far down the shaft. Cypselas sparsely hairy. Pappus
of free, barbellate, capillary bristles in one row;
each bristle with adpressed teeth. Two species,
tropical Africa.
This description is based on A. senegalensis
Benth., one of two species of the genus. The identity of the generic type (A. athrixioides Steetz, destroyed in Berlin) is unclear and its description
differs from that above. Possibly the two taxa represent different genera, in which case A. senegalensis
should be treated under a new generic name.
1117. Pluchea Cass.
Fig. 75
Pluchea Cass., Bull. Sci. Soc. Philom. Paris 1817: 31 (1817);
King-Jones, Englera 23: 1–136 (2001), rev., phylog.
Berthelotia DC. (1836).
Tecmarsis DC. (1836).
Eyrea F. Muell. (1852).
Eremohylema A. Nels. (1924).
Shrubs or herbs. Leaves alternate or subopposite,
dentate to serrate or entire, generally not decurrent,
hairy. Capitula heterogamous, disciform, solitary
or few to many in corymbs. Receptacle epaleate.
Florets purple. Marginal florets female; corolla filiform. Disc florets functionally male. Anthers tailed;
cells of filament collar mammillose or flattened; endothecial tissue radial. Style entirely or almost undivided or sometimes divided; style branches with
obtuse sweeping-hairs reaching below the furcation. Cypselas sometimes indistinct or entirely reduced to a small carpopodium, hairy. Pappus of
free, barbellate, capillary bristles in one row; each
bristle with patent or sometimes with adpressed
teeth. 2n = 20, 30. About 80 species, pantropical.
Pluchea is heterogeneous and not monophyletic. It is composed of several entities which
have been given generic status. Genera such as
Coleocoma F. Muell. and Streptoglossa Steetz have
their closest relatives within Pluchea (Anderberg
Fig. 75. Compositae-Inuleae. Pluchea microcephala.A Flowering branch. B Capitulum. C Marginal floret. D Disc floret.
E Stamen. F Style. (Cabrera 1978)
et al. 2005). Other genera may also prove to be part
of the Pluchea complex, e.g. Monarrhenus Cass.,
Karelinia Less. and Tessaria Ruiz & Pavon.
1118. Karelinia Less.
Karelinia Less., Linnaea 9: 187 (1834).
Perennial herb. Leaves alternate, elliptic-oblong,
entire, sparsely hairy. Capitula heterogamous,
disciform, solitary. Receptacle epaleate. Involucral
bracts broadly oblong. Florets purple. Marginal
florets female; corolla filiform. Disc florets functionally male. Anthers tailed; cells of filament
collar mammillose; endothecial tissue radial. Style
branches with obtuse sweeping-hairs reaching
below the furcation. Cypselas hairy. Pappus of
free, barbellate, capillary bristles, in one row; each
388
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
bristle with patent teeth. 2n = 20. One species,
K. caspia Less., Caspian Sea region.
1119. Tessaria Ruiz & Pavon
Tessaria Ruiz & Pavon, Prod. Fl. Peruv. Chil. (1794).
Trees. Leaves alternate, entire, glabrous or sparsely
hairy. Capitula heterogamous, disciform, few in
loose corymbs. Receptacle epaleate. Florets pink.
Marginal florets female; corolla filiform. Disc florets functionally male, usually one; corolla lobes
longer than tube. Anthers tailed, conspicuously
calcarate; cells of filament collar mammillose; endothecial tissue radial. Style undivided, with obtuse sweeping-hairs reaching far down the shaft.
Cypselas of marginal female florets small, ellipsoid, glabrous. Pappus of free, barbellate, capillary
bristles in one row; each bristle with patent teeth.
2n = 20. One species, T. integrifolia Ruiz & Pavon,
South America.
1120. Monarrhenus Cass.
Monarrhenus Cass., Bull. Soc. Philom. Paris 1817: 31
(1817).
Shrubs. Leaves on the distal portion of the
branches, ovate-lanceolate, subentire, hairy. Capitula heterogamous, disciform, small, in dense
terminal clusters. Receptacle epaleate. Florets
purplish. Marginal florets female; corolla filiform.
Disc florets functionally male. Anthers tailed; endothecial tissue radial. Style branches with obtuse
sweeping-hairs reaching the furcation. Cypselas
of marginal florets small, ellipsoid, distally hairy.
Pappus of free, barbellate capillary bristles, in one
row; each bristle with patent teeth. Two species,
Mauritius, La Réunion.
1122. Coleocoma F. Muell.
Coleocoma F. Muell., Hooker’s J. Bot. Kew Gard. Misc. 9: 19
(1857).
Small rigid herb. Leaves alternate, glabrous, dentate. Capitula heterogamous, disciform, solitary,
subsessile, subterminal, ovoid. Receptacle epaleate.
Involucral bracts broadly ovate, obtuse. Marginal
florets female; corolla filiform. Disc florets functionally male. Anthers tailed; endothecial tissue radial. Style undivided, with obtuse sweeping-hairs
reaching far down on the shaft. Cypselas quadrangular with several narrow ribs. Pappus of scale-like
bristles forming a narrow tube surrounding the
corolla base; each bristle with patent teeth. One
species, C. centaurea F. Muell., Australia.
1123. Delamerea S. Moore
Delamerea S. Moore, J. Bot. 38: 457 (1900).
Procumbent herb. Leaves alternate, spathulate,
shallowly dentate, not decurrent, greyishtomentose. Capitula heterogamous, disciform,
solitary, subsessile, subterminal. Receptacle
epaleate. Involucral bracts acute, slightly squarrose. Florets purple. Marginal florets female;
corolla filiform. Disc florets perfect. Style branches
with obtuse sweeping-hairs reaching below the
furcation. Anthers tailed; endothecial tissue
radial. Cypselas terete or angled. Pappus of a few
scale-like bristles connate into a tube surrounding
the corolla; each bristle with patent teeth. One
species, D. procumbens S. Moore, East Africa.
1124. Neojeffreya Cabr.
1121. Thespidium F. Muell. ex Benth.
Thespidium F. Muell. ex Benth., Fl. Austral. 3: 534 (1867).
Perennial herb, often densely tufted. Leaves alternate, marginally dentate, hirsute. Capitula heterogamous, disciform, small, more or less sessile at the
base of the stems. Receptacle epaleate. Involucral
bracts few, acute. Marginal florets female; corolla
filiform. Disc florets perfect. Anthers tailed; endothecial tissue radial. Style branches with obtuse sweeping-hairs reaching below the furcation.
Cypselas obconical, ribbed, hairs uncinate. Pappus
of a few, free, rather stiff, scale-like bristles in one
row; each bristle with patent teeth. One species,
T. basiflorum F. Muell., Australia.
Neojeffreya Cabr., Hickenia 1: 160 (1978).
Annual herb. Leaves alternate, denticulate to
sinuate, decurrent into long wings, woolly-hairy.
Capitula heterogamous, disciform, small, aggregated in dense terminal glomerules. Receptacle
paleate; paleae marginally plumose. Marginal
florets female; corolla filiform. Disc florets perfect.
Anthers tailed; endothecial tissue radial. Style
branches with obtuse sweeping-hairs reaching
below the furcation. Cypselas laterally compressed.
Pappus of free, barbellate, capillary bristles in one
row; each bristle with patent teeth. One species,
N. decurrens (L.) A. L. Cabrera, Madagascar, East
Africa.
Compositae
1125. Triplocephalum O. Hoffm.
Triplocephalum O. Hoffm., Engler & Prantl, Nat. Pflanzenfam. 4, 5: 389 (1894).
Shrub. Leaves alternate, shallowly dentate, hairy.
Capitula heterogamous, disciform, small, in
globose, terminal secondary heads. Receptacle
epaleate. Florets purplish. Marginal florets female;
corolla filiform. Disc florets functionally male.
Anthers tailed; endothecial tissue radial. Style
undivided, with obtuse sweeping-hairs reaching
far down on the shaft. Cypselas hairy. Pappus
absent, or a minute rim of scales. One species, T.
holstii O. Hoffm., East Africa.
1126. Sphaeranthus L.
Sphaeranthus L., Sp. Pl.: 927 (1753); Ross-Craig, Hooker’s
Ic. Pl. 6: 1–117 (1955), rev.
Tisserantia Humbert (1927).
Herbs. Leaves alternate, dentate, decurrent into
wings. Capitula heterogamous, disciform, small,
congested in globose or ovoid to cylindrical,
terminal secondary heads. Receptacle epaleate.
Florets purple. Marginal florets female; corolla
filiform. Disc florets functionally male. Anthers
tailed; endothecial tissue radial. Style undivided,
with obtuse sweeping-hairs reaching far down
on the shaft. Cypselas often reduced, often with
uncinate or anchor-shaped hairs. Pappus absent.
2n = 20. Forty-one species, Old World tropics.
1127. Pseudoblepharispermum Lebrun & Stork
Pseudoblepharispermum Lebrun & Stork, Adansonia 4: 419
(1981); Beentje & Hind, Kew Bull. 57: 214 (2002), key.
Shrubs. Leaves alternate, entire, densely whitetomentose, ovate or subglabrous, linear. Capitula
heterogamous, disciform, small, congested into
terminal secondary heads. Receptacle paleate;
paleae hairy. Marginal florets female; corolla
filiform. Disc florets functionally male. Anthers
without tails or with short tails; endothecial
tissue radial (type). Style undivided, with obtuse
sweeping-hairs reaching far down on the shaft.
Cypselas with straight or uncinate hairs. Pappus
absent. Two species, East Africa.
Pseudoblepharispermum is heterogeneous, and
the tribal position for the two rare and little-known
species placed in this genus remains to be tested.
1128. Pseudoconyza Cuatr.
Pseudoconyza Cuatr., Ci. Mexico 21: 30 (1961).
389
Herb. Leaves alternate, dentate and sometimes
basally shallowly lobed, slightly stem-clasping.
Capitula heterogamous, disciform, terminal,
solitary or few together. Receptacle epaleate.
Florets greenish-white. Marginal florets female;
corolla filiform. Disc florets perfect. Anthers tailed;
endothecial tissue radial. Style branches with obtuse sweeping-hairs reaching below the furcation.
Cypselas hairy. Pappus of free, barbellate, capillary
bristles in one row; each bristle with adpressed
teeth. One species, P. viscosa (Mill.) D’Arcy, Central
America, Africa, Asia.
1129. Blumeopsis Gagnep.
Blumeopsis Gagnep., Bull. Mus. Hist. Nat. Paris 26: 75
(1929).
Herb. Leaves alternate, sharply dentate, sparsely
hairy. Capitula heterogamous, disciform, rather
small, few, in terminal corymb. Receptacle
epaleate. Florets yellow. Marginal florets female;
corolla filiform. Disc florets perfect. Anthers
without tails; endothecial tissue radial. Style
branches with obtuse sweeping-hairs reaching
below the furcation. Cypselas hairy. Pappus of
free, barbellate, capillary bristles in one row; each
bristle with adpressed teeth. One species, B. falcata
(D. Don) Merr., eastern Asia.
1130. Laggera Sch. Bip. ex Koch
Laggera Sch. Bip. ex Koch, Linnaea 19: 391 (1847).
Herbs. Leaves alternate, dentate, decurrent, hairy.
Capitula heterogamous, disciform, terminal, few to
many. Receptacle epaleate. Florets purple. Marginal
florets female; corolla filiform. Disc florets perfect.
Anthers without tails; endothecial tissue radial.
Style branches with obtuse sweeping-hairs reaching below the furcation. Cypselas hairy. Pappus of
free, barbellate, capillary bristles in one row; each
bristle with adpressed teeth. 2n = 20. Seventeen
species, tropical Africa, Arabia, Asia.
1131. Nicolasia S. Moore
Nicolasia S. Moore, J. Bot. 38: 458 (1900); Merxmüller, Mitt.
Bot. Staatssamml. München 2: 1–10 (1954), rev.
Herbs or subshrubs. Leaves alternate, entire to dentate, generally decurrent, sparsely hairy. Capitula
heterogamous, disciform, solitary or few together.
Receptacle epaleate. Florets purple. Marginal florets female; corolla filiform. Disc florets perfect.
Anthers without tails; endothecial tissue radial.
390
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Style branches with obtuse or sometimes acute
sweeping-hairs reaching below the furcation.
Cypselas hairy. Pappus of few, free, barbellate,
capillary bristles in one row; each bristle with
adpressed teeth. Seven species, southern tropical
Africa.
filament collar mammillose; endothecial tissue radial. Style branches with obtuse sweeping-hairs
reaching below the furcation. Cypselas villous. Pappus of free, rather stiff, barbellate bristles in one
row; each bristle with patent teeth. 2n = 20. Eight
species, Australia.
1132. Doellia Sch. Bip.
1135. Allopterigeron Dunlop
Doellia Sch. Bip., Walpers Repert. 2: 953 (1843); Anderberg,
Willdenowia 25: 19–24 (1995).
Allopterigeron Dunlop, J. Adelaide Bot. Gard. 3, 2: 183
(1981).
Herbs. Leaves alternate, dentate, hairy. Capitula
heterogamous, disciform, solitary or few together.
Receptacle epaleate. Florets purple. Marginal
florets female; corolla filiform. Disc florets perfect.
Anthers tailed; endothecial tissue radial. Style
branches with obtuse sweeping-hairs reaching
below the furcation. Cypselas with conspicuous
red longitudinal resin ducts, hairy. Pappus of
free, barbellate, capillary bristles in one row; each
bristle generally with adpressed teeth. Two species,
Arabia, Africa.
Annual herb. Leaves alternate, fleshy, entire.
Capitula heterogamous, minutely radiate or
disciform, solitary. Receptacle epaleate. Florets
white. Marginal florets female; corolla minutely
radiate. Disc florets functionally male, three-lobed.
Anthers three, without tails. Style undivided, with
obtuse sweeping-hairs reaching far down the shaft.
Cypselas ellipsoid, hairy. Pappus of several free,
stiff, barbellate, capillary bristles. One species, A.
filifolius (F. Muell.) Dunlop, Australia.
1133. Porphyrostemma Benth. ex Oliv.
Porphyrostemma Benth. ex Oliv., Trans. Linn. Soc. London
29: 96 (1873).
Herbs. Leaves alternate, narrowly linear, entire,
glabrous. Capitula heterogamous, radiate or
disciform, solitary. Receptacle epaleate. Involucral
bracts in many rows. Florets purple. Marginal
florets female; corolla minutely radiate. Disc florets
perfect; corolla with conspicuous secretory canals.
Anthers tailed; endothecial tissue radial. Style
branches with obtuse sweeping-hairs reaching
below the furcation. Cypselas with conspicuous,
red, two- or three-celled secretory cavities in
longitudinal rows, hairy. Pappus of free, barbellate,
capillary bristles in one row each with adpressed
teeth, and a short rim of connate scales, sometimes
rim-shaped or absent. Four species, tropical Africa.
1134. Streptoglossa Steetz
Streptoglossa Steetz, Trans. Proc. Bot. Soc. Edinburgh 7: 491
(1863); Dunlop, Adelaide Bot. Gard. 3: 167–182 (1981), rev.
Pterigeron (DC.) Benth. (1867).
Herbs or subshrubs. Leaves alternate, dentate, often
decurrent, hairy. Capitula heterogamous, radiate
to disciform, solitary or few. Receptacle epaleate.
Florets purple. Marginal florets female, radiate or
minutely radiate to almost tubular. Disc florets perfect or functionally male. Anthers tailed; cells of
Genera Incertae Sedis
1136. Nanothamnus Thoms.
Nanothamnus Thomson, J. Linn. Soc. 9: 342, t. 3 (1867).
Annual herb. Leaves lanceolate, serrate, sericeous.
Capitula heterogamous, disciform, few-flowered,
in dense terminal clusters. Receptacle epaleate.
Involucre narrowly cyathiform, bracts cartilaginous, acute, in few rows. Florets yellow. Outer
florets functionally female, with vestigial stamens,
pseudobilabiate (1/4). Disc florets perfect, bilabiate. Anthers long-calcarate, with short tails, apical
appendage soft. Endothecial tissue radial. Style
branches with semi-acute sweeping-hairs reaching
below the furcation. Cypselas ellipsoid, ribbed;
epidermis with elongated crystals. Pappus absent.
One species, N. sericeus Thoms., India.
The large crystals in the cypsela epidermis
would indicate a position in Inuleae s.str., possibly
near Blumea DC., but the tribal position for this
genus is uncertain. Its bilabiate corolla, calcarate
anthers and soft anther appendage are character
states that poorly match the other taxa of the
tribe.
1137. Feddea Urb.
Feddea Urb., Repert. Spec. Nov. Regni Veg. 21: 73 (1925).
Scandent shrub. Leaves alternate, elliptic, glabrous, coriaceous. Capitula homogamous, discoid.
Compositae
Involucral bracts in several rows, imbricate, cartilaginous, rather broad with a median secretory
duct. Receptacle flat, epaleate. Florets deeply
lobed, perfect. Anthers ecalcarate, caudate. Style
branches obtuse-rounded, dorsally and apically
with obtuse sweeping-hairs; stigmatic area in
two marginal bands, confluent apically. Cypselas
oblong, glabrous. Pappus of barbellate capillary
bristles in several rows. One species, F. cubensis
Urb., Cuba.
A stigmatic area with two lateral bands confluent at the apex is a typical character state in Inuleae. The median secretory ducts resemble those
of Iphionopsis A. Anderb. If it belongs in Inuleae,
then Feddea may be another odd member of the
Pluchea Cass. complex.
Key to the Tribes
of the Heliantheae Alliance
J.L. Panero
The common denominator in the history of the
classification of the Heliantheae alliance has
been the subjective delimitation and selective
recognition of the classical tribes Heliantheae and
Helenieae. Cassini (1819) erected tribe Heliantheae
to accommodate most New World sunflowers with
key characters including yellow corollas, opposite
leaves, and uniseriate or biseriate involucres. He
placed Eupatorieae in the proximity of Vernonieae but drew attention to their connection to
Heliantheae. In his final classification of the family
(Cassini 1829), he recognized Helenieae as a group
within Heliantheae and separated HeliantheaeAmbrosiinae and Tageteae-Pectidinae as tribes
(Ambrosieae and Tageteae respectively). Bentham
(1873b) recognized Helenieae at the tribal level,
in which he placed most helianthoid genera
with epaleate receptacles. This implied that the
paraphyletic nature of Helenieae was a necessary condition to have a monophyletic (paleate)
Heliantheae. These decisions were to dominate
the classification of the genera of these tribes for
more than a century. Robinson (1981) published
a revolutionary classification of the group in
which he placed Helenieae in the synonymy
of Heliantheae, as done earlier by Cronquist
(1955), and recognized 35 subtribes. His system
benefited from the use of floret morphological
features previously not used in classification and
391
termed microcharacters, most being visible only
with the aid of a light microscope. Subsequent
classifications (Karis and Ryding 1994a, b), although accepting some of the findings reported
by Robinson (1981), reverted to a bipolar system
in which Heliantheae were deemed as probably
monophyletic and Helenieae as paraphyletic.
Eupatorieae was seen as sister tribe to these. The
advent of molecular comparative studies of these
groups of sunflowers has clarified the phylogenetic
position of Eupatorieae and, if Eupatorieae are to
be maintained at tribal rank, forces the recognition
of several tribes as hypothesized by some previous
authors.
Readers will notice that the tribal classification
outlined in the Introduction to Compositae in
this volume differs from that of the authors of
tribes XVIII–XXX of the so-called Heliantheae
alliance (cf. below). This discrepancy centres on
the recognition of Eupatorieae, a tribe of more
than 2,000 species and 190+ genera, well-defined
by both morphological and molecular features,
and long established in the taxonomic literature
(Cassini 1819; King and Robinson 1987; Bremer
et al. 1994b). To maintain recognition and use of
Eupatorieae, and at the same time a classification
which recognizes only monophyletic groups,
Heliantheae can no longer be recognized in the
traditional, broad sense of Robinson (1981). This
is because all molecular evidence to date (Kim
and Jansen 1995; Baldwin et al. 2002; Panero and
Funk 2002) indicates that Eupatorieae are not
a basal member of subfamily Asteroideae (King
and Robinson 1987), and not sister to the classical
Heliantheae-Helenieae as previously thought
(Karis and Ryding 1994a, b) but rather embedded
within that lineage (Fig. 76). Eupatorieae are now
identified as one of at least 13 major lineages
(Panero and Funk 2002) recognized at tribal rank
by the authors of tribal accounts XVIII–XXX.
These 13 groups are accepted in the Introduction
to Compositae but recognized at the rank of
supersubtribe (Jeffrey 2004), with Eupatorieae
included in an expanded Heliantheae of more
than 5,200 species. For ease of discussion, these
13 tribes (Heliantheae s.l. of the Introduction)
hereafter are referred to as the Heliantheae alliance
(Fig. 76). Comparative molecular studies provided
the phylogenetic hypotheses adopted in tribes
XVIII–XXX (Baldwin et al. 2002; Panero and Funk
2002; Panero, unpubl. data). Among the most
important conclusions from these studies is the
recognition of a more exclusive tribe Heliantheae
392
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
s.s., the elucidation of the phylogenetic position of
tribe Helenieae, and the sister-group relationship
of Eupatorieae and Perityleae within the clade
containing tribes Millerieae and Madieae (Fig. 76).
An equally interesting, if not perplexing, result
is the relatively basal position of Coreopsideae
and the sister-group relationship of Neurolaeneae
to the epaleate tribes Tageteae, Bahieae and
Chaenactideae (Fig. 76). Equally exciting is the
realization that the tremendous diversity of the
Heliantheae alliance in the New World is the
result of a single introduction from the southern
hemisphere, most probably Africa. This assertion
Fig. 76. Summary of phylogenetic relationships among
tribes recognized in the Heliantheae alliance, based
on chloroplast DNA sequence analysis. Numbers along
branches indicate bootstrap support. Numbers after tribal
names indicate approximate number of species. (Modified
from Panero and Funk 2002)
is supported by the fact that the mostly African
tribe Athroismeae was found to be sister to the
rest of the Heliantheae alliance. In addition,
the genus Callilepis, also from southern Africa
and traditionally classified in Gnaphalieae, is
sister to the Heliantheae alliance (Anderberg
et al. 2005; Panero, unpublished chloroplast
data). Apparently, the only unifying feature of
the Heliantheae alliance within Asteraceae is
the presence of a phytomelanin layer in the
cypselae. This layer renders the fruit black or dark
brown, hence the term carbonized cypselae or
cypselae with carbonized walls. This character
is sometimes difficult to observe because some
cypselae are either densely pubescent, covered
by epidermal outgrowths in the form of wings
(Heliantheae-Ecliptinae), or phyllaries are fused to
the fertile cypselae (Millerieae-Melampodiinae).
Tribe Helenieae and subtribes Anisopappinae and
Centipedinae of Athroismeae lack phyomelanin
in their cypselae, and could be confused with
other groups of Asteraceae. Most members of tribe
Helenieae are distinctive by having crystals in each
of the epidermal cells of the cypsela, and lacking
a phytomelanin layer. Elsewhere in Asteraceae,
this character is seen only in some 30 genera
of tribe Inuleae. Helenieae are endemic to the
New World and, for the most part, have a scaly
pappus whereas those Inuleae with crystals in
their cypselae are found only in the Old World
and tend to have a pappus of bristles. Subtribe
Anisopappinae contains taxa with an Old World
distribution which differ from most Inuleae in
having a scaly pappus and obtuse style papillae.
Subtribe Centipedinae is distinctive for having
mostly disciform capitula with several rows of
tubular florets, central florets with tetramerous
corollas, and anthers with very reduced or obsolete
appendages.
The key below facilitates identification of the
tribes and distinctive genera of the Heliantheae alliance. Most members of this group are endemic
to the American continent where they constitute
approximately 40–50% of its sunflower diversity.
The key uses characteristics which sometimes are
not included in the descriptions. My aim in constructing this key has been to use characters which
can be readily observed in the field and in herbarium specimens. Nevertheless, a number of leads
are based on characters which require the use of
a microscope.
Compositae
Key to the Tribes
1. Cypsela walls with 1–3 crystals per cell, rarely none,
not carbonized; plants from the New World
XIX. Helenieae (p. 400)
– Cypsela walls without crystals, carbonized, rarely not
carbonized; plants mostly from America, rarely fromq
the Old World
2
2. Capitula with 1, rarely 2 florets and fused phyllaries,
central ribs of phyllaries swollen and wing-like; shrubs
with opposite, glaucous leaves, growing at the edge of
saline marshes surrounding La Paz, Baja California,
Mexico
XXII. Tageteae-Coulterellinae (p. 422)
– Capitula otherwise, if with 1–2 florets, then phyllaries
without wing-like central ribs; plants various, if from
La Paz, Baja California, then capitula with more than
2 florets
3
3. Cypselae not carbonized
4
– Cypselae carbonized
5
4. Capitula radiate; corollas yellow; receptacles mostly
paleate; cypselae mostly pappose
XVIII. Athroismeae-Anisopappinae (p. 397)
– Capitula disciform, rarely radiate; corollas mostly
white or greenish white, rarely yellow; receptacles
epaleate; cypselae epappose
XVIII. Athroismeae-Centipedinae (p. 399)
5. Capitula arranged in compact, glomerule-like, compound cymes; plants of Africa and Asia, with white
corollas and alternate leaves
XVIII. Athroismeae-Athroisminae (p. 398)
– Capitula otherwise, if arranged in glomerule-like compound cymes, then female florets enclosed in perigynia; plants mostly from America, if from Africa or
Asia, then corolla colour various and leaves opposite 6
6. Capitula paleate
7
– Capitula epaleate, sometimes paleae present as a single
row between ray florets and first row of disc florets or
in discoid capitula paleae fused and resembling an
involucre
26
7. Anther appendages linear, flat, without basal constrictions, reduced or lacking; capitula always discoid;
corollas white, pink, blue or purple; style arm appendages longer than stigmatic lines
XXX. Eupatorieae (p. 510)
– Anther appendages lanceolate to oval, rarely linear,
shallowly to strongly concave, constricted at base; capitula discoid, disciform or radiate; corollas mostly
yellow or yellow-orange, sometimes white, rarely pink
or purple; style arm appendages usually as long as or
shorter than stigmatic lines
8
8. Capitula with dimorphic involucres, innermost phyllaries translucent, chartaceous, normally dull orange
or yellow, outer herbaceous, green, mostly reflexed,
rarely capitula ligulate and all phyllaries herbaceous,
gradate (Fitchia); cypselae obcompressed with resin
canals; anther appendages mostly with resin canals
and without glandular trichomes
XX. Coreopsideae-Chrysanthellinae (p. 407),
Coreopsidinae (p. 409)
– Capitula without dimorphic involucres or, if dimorphic, innermost series of phyllaries rarely chartaceous
and translucent; cypselae various, rarely with resin
canals; anther appendages without resin canals, either
glabrous or with glandular trichomes
9
393
9. Capitula with 4 deeply trilobed yellow florets subtended by 4 scarious phyllaries enclosed by two, broad
suborbicular phyllaries
XX. Coreopsideae 1186. Staurochlamys
– Capitula and phyllaries otherwise
10
10. Phyllaries of innermost series fused to ray cypselae
XXVII. Millerieae-Melampodiinae (p. 487)
– Phyllaries of innermost series not fused to ray cypselae
11
11. Innermost phyllaries partially or fully covering ray floret cypselae; ray corollas trilobed; plants of the Pacific
coast of the USA and extreme north-western Mexico,
Hawaii XXVIII. Madieae-Madiinae in part (p. 497)
– Innermost phyllaries rarely covering ray cypselae; if
covering ray cypselae, then edge of phyllaries translucent or ray corollas reduced to a tube; plants of tropical
America
12
12. Anther filaments papillose, thecae green
XXVII. Millerieae-Guardiolinae (p. 486)
– Anther filaments glabrous, if papillose, then thecae
black or brown, never green
13
13. Leaves with expanded leaf bases or more commonly
forming a cupule (invaginated) around stem; plants
forming persistent rosettes with woody stems (acaulior caulirosulae) or sometimes shrub- or tree-like with
leaves aggregated at distal ends of stems
XXVII. Millerieae-Espeletiinae (p. 482)
– Leaves without expanded bases; plants otherwise, if
forming rosettes, then stems not woody, if shrubs or
trees, then leaves evenly spaced along branches
14
14. Capitula with functionally staminate disc florets;
cypselae without wings, epappose, broadly biconvex
or trigonal; phyllaries and herbage with long stipitate
glandular trichomes
15
– Capitula mostly with bisexual disc florets, if with
functionally staminate disc florets, then either cypselae strongly flattened, mostly winged and pappose
or cypselae broadly biconvex and peripheral floret
corollas tubular with very reduced limbs and corollas
white or light yellow; phyllaries and herbage rarely
with long stipitate glandular trichomes
16
15. Anther thecae hyaline or pale yellow; phyllaries subequal, outermost series never reflexed nor patent
XXV. Polymnieae (p. 439)
– Anther thecae brown or black; phyllaries gradate with
outermost series much larger than inner series, sometimes reflexed XXVII. Millerieae-Milleriinae (p. 487)
16. Plants aquatic with fistulose stems and axillary capitula
XXI. Neurolaeneae-Enydrinae (p. 418)
– Plants otherwise
17
17. Disc florets with very reduced or undivided styles and
functionally staminate
18
– Disc florets bisexual
20
18. Ray corollas conspicuous, yellow or greenish yellow
XXVI. Heliantheae-Dugesiinae (p. 446),
Engelmanniinae in part (p. 461),
Ecliptinae in part (p. 446),
1360. Zinnia in part
– Ray corollas inconspicuous, mostly reduced to a tubular corolla or ring, sometimes missing, if protruding beyond phyllaries, then these white and clearly
trilobed
19
19. Anthers mostly light yellow or hyaline, if light brown,
then paleae of adjacent disc florets fused to ray cypsela;
394
–
20.
–
21.
–
22.
–
23.
–
24.
–
25.
–
26.
–
27.
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
styles of disc florets absent or with 1 (rarely 2) vascular traces; anthers normally free and with numerous
minute thickenings surrounding endothecial cells
XXVI. Heliantheae-Ambrosiinae (p. 443)
Anthers black or brown; styles with 2 vascular traces,
anthers normally fused, endothecial thickenings 1–4,
polar
XXVI. Heliantheae-Ecliptinae (p. 446)
1261. Clibadium,
1275. Lantanopsis,
XXVII. Millerieae-Milleriinae (p. 487)
1386. Ichthyothere
Disc floret cypselae moderately to strongly compressed, rarely quadrate or the exocarp juicy and
cypselae drupe-like; pappus elements, when present,
mostly disposed in a narrowly oval to linear arrangement on cypsela neck; cypselae sometimes winged;
ray cypselae never enclosed in perigynia
XXVI. Heliantheae (p. 440)
Disc floret cypselae mostly terete or subterete, if
quadrate, then cypselae either with 4 small horns
(Greenmaniella) or prismatic (Lycapsus), without
juicy exocarps; pappus elements mostly present and
in radial arrangement on cypsela neck; cypselae never
winged; ray cypselae sometimes tightly wrapped by
phyllaries or enclosed in perigynia
21
Disc corolla throats with resinous areas between
vascular strands just below lobe sinuses; style arms
canaliculate on adaxial surface; carpopodia missing,
base of cypsela appearing invaginated
XXII. Tageteae-Varillinae (p. 430)
Disc corolla throats otherwise; style arms not canaliculate; carpopodia present
22
Leaves alternate; ray florets fertile
23
Leaves opposite, if alternate, then ray florets neuter or
plants shrubby and leaves linear (Dyscritothamnus) 25
Capitula discoid; corollas tetramerous
XXIX. Perityleae-Lycapsinae (p. 509)
Capitula radiate, rarely discoid; disc corollas
pentamerous
24
Leaves entire or lobed, never bipinnate
XXI. Neurolaeneae-Neurolaeninae (p. 419)
1189. Calea in part,
1190. Greenmaniella,
1191. Neurolaena
Leaves bipinnate, segments linear or filiform
XXIV. Bahieae 1237. Hymenopappus in part
Corollas with reddish resin in resin ducts
XXI. Neurolaeneae-Neurolaeninae (p. 419)
1189. Calea in part
Corollas with yellow resin in resin ducts or these
colourless, if with reddish resin in resin ducts, then
plants endemic to Africa (Guizotia)
XXVII. Millerieae-Galinsoginae (p. 483),
Desmanthodiinae (p. 479),
Dyscritothamninae (p. 480),
Jaegeriinae (p. 487),
Milleriinae (p. 488)
Leaves and/or phyllaries with embedded or marginal,
pellucid secretory cavities/glands or pustules,
aromatic
XXII. Tageteae-Pectidinae (p. 423)
Leaves and phyllaries without embedded or marginal,
pellucid secretory cavities/glands or pustules, rarely
aromatic
27
Capitula with one row of paleae between ray florets and
first row of disc florets; in discoid capitula phyllaries
–
28.
–
29.
–
30.
–
31.
–
32.
–
33.
–
34.
–
35.
–
36.
–
37.
–
38.
reduced, wanting or missing and these paleae connate
and appearing as an involucre
XXVIII. Madieae-Madiinae in part (p. 497)
Capitula otherwise
28
Capitula with 1 broadly biconvex, obcompressed
cypsela
XXVI. Heliantheae-Ecliptinae (p. 446)
1275. Lantanopsis,
1292. Riencourtia
Capitula otherwise
29
Anther appendages linear, flat, without basal constrictions, reduced or lacking; capitula always discoid;
corollas white, pink or purple; style arm appendages
longer than stigmatic lines XXX. Eupatorieae (p. 510)
Anther appendages lanceolate to oval, rarely linear,
shallowly to strongly concave, constricted at base; capitula discoid, disciform or radiate; corollas mostly
yellow or yellow-orange, sometimes white, rarely pink
or purple; style arm appendages usually as long as or
shorter than stigmatic lines
30
Disc corollas tetramerous; cypselae strongly compressed or prismatic with 4 ribs usually ciliate on
margins; styles subulate; herbage of most species
densely glandular, sticky
XXIX. Perityleae-Peritylinae (p. 509)
Disc corollas mostly pentamerous; cypselae various,
rarely strongly compressed or with 4 ribs, pubescence
various; styles various, sometimes subulate, herbage
glabrous or pubescent, rarely glandular
31
Plants forming small rosettes with axillary, solitary
capitula on long peduncles; plants from Cuba
32
Plants otherwise, if forming small rosettes, then
capitula terminal in corymbiform inflorescences,
rarely solitary (Chamaechaenactis), if appearing
axillary, then capitula nearly sessile; plants from
America, rarely Cuba
33
Ray florets without corollas; pappus of 4 accrescent
scales
XX. Coreopsideae-Pinillosiinae
1185. Tetraperone
Ray florets with corollas; pappus of several accrescent,
ciliate scales
XXI. Neurolaeneae-Heptanthinae (p. 418)
Decumbent perennial herbs with ovate to reniform,
opposite leaves; capitula with 4 phyllaries and 4 florets;
ray florets either without corollas or corollas, when
present, deeply bilobed; plants from the Caribbean
XX. Coreopsideae-Pinillosiinae (p. 416)
Plants otherwise; capitula otherwise; ray florets, when
present, always corollate, limbs various, never deeply
bilobed; plants from America, rarely the Caribbean 34
Phyllaries of capitula with black striae; receptacles setose
XXII. Tageteae-Clappinae (p. 421)
Phyllaries without black striae; receptacles rarely setose
35
Leaves glabrous, linear to acicular, glaucous, succulent;
plants of saline environments
36
Leaves otherwise; plants rarely from saline environments
37
Leaves opposite XXII. Tageteae-Jaumeinae (p. 423)
Leaves alternate XXII. Tageteae 1223. Pseudoclappia
Cypselae terete with multiple ribs (8–11)
XXII. Tageteae-Flaveriinae (p. 422)
Cypselae compressed or obcompressed, sometimes
prismatic to terete with 5 or fewer ribs
38
Weak shrubs with alternate, entire, petiolate, ovate to
deltate leaves; phyllaries fused at base forming a bowl-
Compositae
–
39.
–
40.
–
41.
–
42.
–
43.
–
44.
–
45.
–
46.
–
47.
–
48.
–
49.
shaped involucre
XXVIII. Madieae-Venegasiinae (p. 507)
Annual or perennial herbs, if shrubs, then leaves
mostly sessile and linear, if petiolate, then blades
dissected
39
Corollas with long, stipitate, biseriate, glandular trichomes, trichome stalk approximately 20+ times as
long as diameter of glandular terminal group of cells
or head, head slightly depressed in centre and appearing obcordate in cross-section
XXII. Tageteae
1220. Arnicastrum,
1221. Jamesianthus
Corollas glabrous or pubescent, if with glandular trichomes, then these with shorter stalks, approximately
1–5 times as long as diameter of glandular head; head
round, never depressed in centre nor obcordate in
cross-section
40
Ray cypsela with 3 and disc cypselae with 5 triangular,
subulate scales
XXII. Tageteae
1222. Oxypappus
Ray and disc cypselae otherwise
41
Cypselae epappose, mostly trigonal with abaxial face
broader than other two faces, glabrous; plants with
mostly glandular herbage, viscid
XXIX. Perityleae-Galeaninae (p. 401)
Cypselae pappose, rarely epappose, cypselae compressed or obcompressed, subterete to terete,
sometimes pyramidal, if trigonal, then all faces of
equal size; plants glabrous to sparsely to densely
pubescent, sometimes lanate-white, herbage rarely
glandular and sticky
42
Capitula disciform, minute; pappus a lacerate crown;
plants of the northern Caribbean
XXIV. Bahieae 1246. Thymopsis
Capitula discoid or radiate, pappus various, rarely
a lacerate crown; plants mostly from North America,
some from South America, few from Africa, rarely
from the Caribbean
43
Pappus of scales with a thickened median rib sometimes protruding beyond scales as an awn
XXIV. Bahieae in part (p. 433)
Pappus various, if of scales, then scales without a median midrib, rarely absent
44
Pappus of bristles
45
Pappus otherwise or absent
46
Cauline leaves mostly opposite
XXVIII. Madieae-Arnicinae (p. 493)
Cauline leaves alternate
XXIV. Bahieae
1231. Bartlettia,
1242. Peucephyllum,
1244. Psathyrotopsis
Shrubs with petiolate leaves
47
Annual or perennial herbs
48
Capitula discoid; corollas white XXIV. Bahieae (p. 433)
1229. Apostates,
1240. Loxothysanus
Capitula radiate; corollas yellow
XVIII. Madieae-Baeriinae
1398. Constancea
Pappus deciduous as a unit
XIII. Chaenactideae
1224. Dimeresia,
1226. Orochaenactis
Pappus persistent or absent
49
Plants aquatic, glabrous, with fistulose stems; capitula
solitary, axillary on long peduncles; ray florets white
395
with black veins on abaxial surfaces; plants of vernal
pools of western Durango state, Mexico
XVI. Heliantheae-Zinniinae
1359. Trichocoryne
– Plants not aquatic, mostly pubescent; capitula terminal, if appearing axillary, then capitula sessile; ray florets mostly yellow or absent, if white, then inconspicuous and without black veins on abaxial surfaces; plants
from western USA and extreme north-western Mexico
50
50. Capitula radiate, sometimes discoid; leaves mostly sessile; phyllaries in 1–3 series, mostly broadly ovate,
sometimes with a thickened rib
XXVIII. Madieae-Baeriinae (p. 494),
Hulseinae (p. 497)
– Capitula discoid; leaves petiolate; phyllaries in 1 series,
linear to lanceolate
XXIII. Chaenactideae
1225. Chaenactis
XVIII. Tribe Athroismeae Panero
(2002).
J.L. Panero
Annual or perennial herbs, shrubs or small trees,
sometimes aromatic. Leaves simple, alternate or
fasciculate on brachyblasts, petiolate or sessile,
abaxial surfaces with or without sessile glandular
trichomes. Capitula terminal, rarely appearing
axillary, in congested, glomerule-like or open,
paniculiform cymes, or solitary, pedunculate,
sometimes sessile, radiate, disciform or discoid;
phyllaries in 1 to 4 series, sometimes absent, herbaceous, subequal or gradate. Receptacles convex to
conic, sometimes subglobose to globose, paleate
or epaleate. Ray florets pistillate or absent, corollas
yellow. Peripheral and disc florets pistillate, bisexual or functionally staminate, rarely shallowly
zygomorphic, peripheral pistillate florets absent
to several, corollas tubular, disc florets bisexual
or functionally staminate, 2 to many, corollas
campanulate, glabrous or pubescent mostly with
twin trichome glands, lobes 4–5, corollas white,
greenish white, white-yellow, rarely purplish;
anthers 4–5, usually ecalcarate, shortly caudate
or ecaudate, rarely the tails well-developed and
branched, endothecium with polarized thickenings, sometimes radial or both, rarely evenly
thickened, appendages shallowly constricted above
thecae or lacking, sometimes with apical gland;
style arms with parallel stigmatic areas sometimes
meeting at style apices, or style filiform and undivided. Cypselae black or brown, obcompressed
to subterete, sometimes with thickened striations
or ribs, sometimes with a proliferation of spongy
396
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
cells at apical portions of cypselae, glabrous or
with ciliate lateral ridges and glandular trichomes,
sometimes with apically arcuate twin trichomes.
Pappus of fertile floret cypselae a crown of twin
trichomes, sometimes with recurved apices or of
deeply lacerate fused squamellae, sometimes with
two awns at angles of cypselae, a variously lacerate
crown or of a few, free, lacerate scales, or absent.
Six genera in three subtribes and about 59
species mostly in eastern Africa, a few species in
western Africa, India, Madagascar, south-eastern
Asia and Indonesia. Centipeda also occurs in Australia, New Zealand, south-eastern Asia, Madagascar and southern South America. Most members of
Athroismeae are found in a variety of habitats ranging from dry open grassland and thickets to dense
woodland (Eriksson 1992, 1993). Centipeda is unusual in the tribe by growing in habitats prone to
inundation, such as the edges of irrigation canals or
of intermittent ponds or on brackish, muddy tidal
flats.
Eriksson (1991) placed Athroisma, Blepharispermum and Leucoblepharis (as the Blepharispermum group, here subtribe Athroisminae) in tribe
Heliantheae, away from their original placement in
Inuleae, because of their carbonized cypselae and
paleate receptacles. Molecular studies (Kim and
Jansen 1995; Baldwin et al. 2002) do not support
the inclusion of the group in any of the lineages
of the Heliantheae alliance but rather support
its status as their sister taxon. The recognition
of the group as a tribe of Asteraceae is based
primarily on evidence from molecular phylogenetic studies of the family (Panero and Funk
2002). Centipeda has been placed in several tribes
including Anthemideae and Astereae. Bremer
(1994) considered the genus difficult to place but
maintained it in Asteroideae unassigned to tribe.
Nesom (1994c) considered the genus a member
of Astereae. An ITS analysis aimed at identifying
the phylogenetic relationships of endemic New
Zealand Asteraceae by Wagstaff and Breitwieser
(2002) found strong support for the exclusion of
the genus from Astereae and its close relationship
to Athroismeae and the Heliantheae alliance. These
results were recently confirmed using several coding regions of chloroplast DNA (Panero 2005, and
unpubl. data). In consequence, Centipeda is here
included in an expanded tribe Athroismeae. The
monotypic Symphyllocarpus of eastern Asia may
prove to be sister to Centipeda. Symphyllocarpus
shares with Centipeda a similar habit, alternate
phyllotaxy, sessile leaves, terminal capitula with
marginally scarious, subequal phyllaries, several
rows of peripheral, tubular female florets and
central bisexual florets with tetramerous corollas.
The main difference between Symphyllocarpus
and Centipeda lies in their cypsela morphology.
The cypselae of Symphyllocarpus are subterete
and smooth and lack the deep grooves and ribs
found in the cypselae of Centipeda. In addition,
Symphyllocarpus is reported to have paleate
receptacles (epaleate in Centipeda), although the
paleae may represent asymmetrical growth of
the cypsela exocarp. Molecular studies of Inuleae
by Eldenäs et al. (1999) revealed that the genus
Anisopappus is more closely related to Heliantheae
and relatives than to Inuleae. Additional molecular
evidence (Panero et al., unpubl. data) supports the
placement of Anisopappus in tribe Athroismeae.
Welwitschiella has been traditionally viewed as
a member of Helenieae (Karis and Ryding 1994b).
Based on my interpretation of the morphology of
the genus, I believe Welwitschiella is not a member
of the Heliantheae alliance of tribes. For the
purposes of this publication, I have decided to
include the genus in Athroismeae, as Anisopappus
appears to share more characteristics with Welwitschiella than any other genus of the Heliantheae
alliance. Welwitschiella shares with some or all
species of Anisopappus an involucre of several
series of phyllaries, strongly ribbed cypselae,
a pappus of scales fused at the base and forming
a lacerate corona, alternate phyllotaxy, yellow
corollas, and with A. athanasioides, also from
Angola, irregularly lobed, peripheral corollas.
Welwitschiella lacks paleate receptacles present
in Anisopappus, although lacking in A. latifolius.
The three original genera of Athroismeae are now
included in subtribe Athroisminae, Centipeda
in subtribe Centipedinae, and Anisopappus and
Welwitschiella in subtribe Anisopappinae (Panero
2005). The three subtribes are quite distinctive
morphologically, and their genera have never been
grouped together in the past or even hypothesized
to be closely related. Recent molecular studies of
Inuleae and related genera by Anderberg et al.
(2005) support, albeit weakly, a close relationship
of the southern African Callilepis to either Athroismeae or the rest of the tribes of the Heliantheae
alliance. Molecular studies based on several genes
of chloroplast DNA (Panero et al., unpubl. data)
clearly show that Callilepis is neither a member
of Gnaphalieae, Inuleae or Athroismeae nor is it
closely related to other tribes of the Heliantheae
alliance.
Compositae
397
Key to the Subtribes
1. Cypselae black, carbonized, obcompressed
2. Athroisminae
– Cypselae brown or dark brown to black, never carbonized, subterete to compressed
2
2. Capitula disciform; receptacles epaleate; innermost
corollas tetramerous, leaves sessile 3. Centipedinae
– Capitula radiate, very rarely discoid; receptacles
paleate, very rarely epaleate; innermost corollas
pentalobed; leaves mostly petiolate, rarely sessile
1. Anisopappinae
XVIII.1. Subtribe Anisopappinae Panero
(2005).
Annual or perennial herbs. Leaves alternate, petiolate or subsessile, blades linear to ovate, sometimes
subcordate, entire to pinnatifid, triplinerved,
sometimes pinnate. Capitula terminal, in open
paniculiform or rarely subumbelliform cymes,
radiate, rarely discoid or disciform. Involucres
campanulate to mostly hemispheric. Receptacles
convex, rarely shallowly conic, usually paleate.
Ray florets pistillate, sometimes with staminodes,
corollas yellow. Peripheral florets in disciform
capitula with zygomorphic corollas, penta- or
tetramerous. Disc florets bisexual, rarely functionally staminate, corollas 5-lobed, golden-yellow;
stamens 5, anthers shallowly calcarate or ecalcarate, shallowly to sometimes strongly tailed, tails
sometimes branched, appendages ovate; style with
divided stigmatic surfaces, with obtuse to subacute
papillae or sweeping trichomes. Cypselae subterete, narrowly obovate in outline, ribbed, epidermis without crystals. Pappus of small scales fused
at the base and forming a lacerate cup or absent.
Fig. 77. Compositae-Athroismeae. A–M Anisopappus chinensis subsp. chinensis. A Flowering branch. B Different
habits. C, D Leaves. E Capitulum. F–H Involucral bracts
(external, middle, internal). I Anthers. J Style arms of female floret. K Style arms of bisexual floret. L, M Cypselae.
N–S Anisopappus chinensis subsp. buchwaldii. N Flowering
branch. O Capitulum. P–R Involucral bracts (external, middle, internal). S Two forms of cypselae. (Humbert 1962, with
permission from the Muséum National d’Histoire Naturelle,
Paris; taxon names according to Ortiz et al. 1996)
Key to the Genera
1. Capitula radiate; receptacles paleate, rarely epaleate
1138. Anisopappus
– Capitula disciform; receptacles epaleate
1139. Welwitschiella
1138. Anisopappus Hook. & Arn.
Fig. 77
Anisopappus Hook. & Arn., Bot. Beech. Voy. 196 (1837);
Humbert, Fl. Madag. 189, 2: 595–611 (1962), part. rev. as
Epallage; Wild, Kirkia 4: 45–76. (1964), rev.; Eldenäs & Anderberg, Pl. Syst. Evol. 199: 167–192 (1996), phylog.; Ortiz
et al., Anales Jard. Bot. Madrid 54: 378–391 (1996), rev.
Epallage DC. (1837).
Sphacophyllum Benth. (1873).
Astephania Oliver (1886).
Temnolepis Baker (1887).
Eenia Hiern & S. Moore (1899).
Annual or perennial herbs. Leaves alternate, blades
linear to ovate, sometimes subcordate, entire to
pinnatifid, triplinerved, sometimes pinnate. Capitula terminal, in open paniculiform or corymbiform, rarely subumbelliform cymes, radiate, rarely
discoid or disciform. Involucres campanulate to
mostly hemispherical. Receptacles convex, rarely
shallowly conic, usually paleate. Ray florets pistillate, sometimes with staminodes, corollas yellow. Disc florets bisexual, corollas 5-lobed, goldenyellow; stamens 5, anthers shallowly calcarate or
398
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
ecalcarate, tailed, tails rarely branched, appendages
ovate; styles with divided stigmatic surfaces, with
obtuse papillae. Cypselae subterete, ribbed, pappus
of small scales or absent. x = 7. Approximately 20
species, tropical Africa, Madagascar, one species in
south-eastern Asia.
of fertile floret cypselae a crown of twin trichomes,
or of deeply lacerate and fused squamellae, sometimes with two awns at angles of cypsela. Pappus
of functionally staminate floret ovaries a variously
lacerate crown or of a few, free, lacerate scales.
1139. Welwitschiella O. Hoffm.
Key to the Genera
Welwitschiella O. Hoffm. Nat. Pflanzenfam. IV, 5: 390
(1894).
Perennial herbs, stems ridged. Leaves alternate,
sessile, blades narrowly oblong, pinnately veined.
Capitula disciform, terminal, in small paniculiform cymes. Involucres campanulate, phyllaries
in 3–4 series, slightly gradate, innermost longer
and somewhat chartaceous, otherwise herbaceous. Receptacles convex, epaleate. Peripheral
florets tetramerous, rarely pentamerous, fertile,
corollas yellow, some zygomorphic, inner florets
functionally staminate(?), corollas yellow, edge of
lobes thickened, cells slightly sclerified; anthers
hyaline or white/yellow(?), ecalcarate, without
tails, appendages narrowly ovate, glabrous; style
arm apices of peripheral florets acute to acuminate, with divided stigmatic surfaces, glabrous on
abaxial surface, style arms of disc florets densely
papillose, acuminate. Cypselae obcompressed,
mostly 4-ribbed, brown, sparsely pubescent.
Pappus an erose crown or multiple erose scales
fused at base. One species, W. neriifolia O. Hoffm.,
south-central tropical Africa.
XVIII.2. Subtribe Athroisminae Panero
(2005).
Perennial herbs, shrubs or small trees. Leaves
alternate or fasciculate on brachyblasts, petioles
sometimes with a basal spine; blades linear,
lanceolate to ovate or obovate. Capitula disciform
or discoid, homogamous or heterogamous, in
congested glomerule-like cymes; glomerules
solitary or arranged in open to compact, paniculiform cymes. Involucres narrowly campanulate,
phyllaries 0–2. Receptacles paleate. Florets all
actinomorphic, rarely zygomorphic, female florets
2 to several or absent, corollas tubular, bisexual
florets 2–25, corollas campanulate, lobes (4–)5;
anthers (4–)5, ecalcarate, caudate; styles divided or
filiform and undivided in functionally staminate
florets, stigmatic area divided and confluent at
style apices, sweeping hairs present. Cypselae
obcompressed, black, lateral ridges ciliate. Pappus
1. Florets with campanulate corollas functionally staminate, styles filiform
1141. Blepharispermum
– Florets with campanulate corollas perfect, styles divided
2
2. Herbaceous bracts subtending capitula longer than
capitula; apices of twin trichomes on cypselae straight
1142. Leucoblepharis
– Herbaceous bracts subtending capitula shorter than
capitula; apices of twin trichomes on cypselae diverging or recurved
1143. Athroisma
Genera of Athroisminae
1140. Athroisma DC.
Fig. 78
Athroisma DC., Guill. Arch. Bot. 2: 516 (1833); Eriksson,
Bot. J. Linn. Soc. 119: 101–184 (1993), rev.
Perennial herbs or shrubs. Leaves petiolate or sessile, blades linear to ovate or obovate, margins entire to pinnatifid. Capitula heterogamous or homogamous with 0–2 female florets and 4–45 perfect florets. Florets with white to pinkish, glabrous
or glandular corollas, anthers 5, rarely 4, styles divided. Cypselae elliptic to obovate. Pappus a corona
of twin trichomes. Twelve species, Africa, India, Indonesia, Madagascar, south-eastern Asia.
1141. Blepharispermum Wight ex DC.
Blepharispermum Wight ex DC. in Wight, Contrib.: 11
(1834); Eriksson, Pl. Syst. Evol. 182: 149–227 (1992), rev.
Shrubs or small trees. Leaves sometimes on
brachyblasts, petiolate to sessile, petiole sometimes with a spine or spine-like protuberance,
blades linear to ovate or obovate, rarely spathulate,
entire to serrate. Capitula heterogamous with
2 female florets and 2–4 functionally staminate
florets. Functionally staminate florets with greenish-white, creamy white or yellowish-white
corollas, anthers 5, style undivided or dividing late
in anthesis. Cypselae elliptic to obcordate, margins
ciliate to scaly. Pappus of female floret cypselae
of 2–15 scales of equal length or with two lateral
scales or awns longer, pappus of functionally
staminate floret ovaries a lacerate crown or of
few, free, lacerate scales or awns. Fifteen species,
Compositae
399
Africa, Arabian Peninsula, India, Madagascar, Sri
Lanka.
1142. Leucoblepharis Arnott
Leucoblepharis Arnott, Mag. Zool. Bot. 2: 422 (1838); Eriksson, Bot. Jahrb. Syst. 112: 167–191 (1990), rev.
Shrubs. Leaves petiolate, blades lanceolate to obovate, entire to shallowly serrate. Capitula heterogamous with 2 female florets and 10–15 perfect florets. Florets with greenish-white corollas, anthers 5,
styles divided. Cypselae elliptic to obcordate, margins ciliate, twin hairs with straight apical cells.
Pappus of 10–15 flat scales. One species, L. subsessilis (DC.) Arnott, south-western India
XVIII.3. Subtribe Centipedinae Panero
(2005).
Annual or perennial herbs. Leaves alternate,
sessile, blades linear to obovate, variously toothed,
rarely entire. Capitula sessile to shortly pedunculate, terminal, appearing axillary, solitary
or in simple monochasial cymes, disciform or
radiate. Involucres campanulate to hemispherical,
phyllaries in 1–2 series, subequal, herbaceous,
margins scarious. Receptacles shallowly concave
to convex, subglobose, epaleate. Peripheral/radiate
florets in several series, sometimes lobes (3)
extremely reduced and corolla tubular, pistillate,
corollas creamy-white, green, light yellow or purplish. Disc florets bisexual, corollas tetramerous,
creamy-white, green, light yellow or purplish,
glabrous with a few glandular trichomes; stamens
4, anthers hyaline, tailed, ecalcarate, appendages,
when present, ovate; style arms with divided
stigmatic surfaces, concave, urceolate in outline,
apices papillose. Cypselae clavate to subterete,
broadly ribbed, glabrescent on basal end, cells of
distal end with raised apical tips, elongating and
forming biseriate trichomes, increasing in density
at apical end and forming a short, dentate, spongy
corona (“pappus”) around corolla tube, glandular
trichomes normally found between ribs. Pappus
absent.
Only one genus:
Fig. 78. Compositae-Athroismeae. Athroisma boranense.
A Flowering branch. B Basal subtending bract. C Median
subtending bract. D Involucral bract. E Female floret.
F Palea. G Hermaphrodite floret. H Twin trichomes.
I Anther. (Eriksson 1993, with permission from Botanical
Journal of the Linnean Society; artist T. Eriksson)
1143. Centipeda Lour.
Fig. 79
Centipeda Lour., Fl. Cochinchin. 492 (1790); Walsh, Muelleria 15: 33–64 (2001), reg. rev.
Myriogyne Less. (1831).
400
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Fig. 79. Compositae-Athroismeae. A–C Centipeda minima.
A Habit. B Capitulum. C Cypsela. D Centipeda nidiformis,
cypsela. E–G Centipeda elatinoides. E Habit. F Leaf.
G Cypsela. H–J Centipeda cunninghamii. H Habit. I Leaf.
J Cypsela. K–M Centipeda thespidioides. K Habit. L Leaf.
M Cypsela. (Walsh 1999, with permission from the author;
artist Enid Mayfield; species names according to N.G.
Walsh, pers. comm.)
Characters of the subtribe. x = 10. Ten species, Australia, New Zealand, south-eastern Asia, Madagascar, Papua New Guinea, southern South America.
XIX. Tribe Helenieae Lindl. (1829).
J.L. Panero
Annual, biennial, perennial herbs, rarely shrubs.
Leaves alternate, rarely opposite, petiolate, subsessile or sessile, blades entire to pinnatifid,
linear to lanceolate, rarely ovate, glabrous to
densely white lanate-floccose. Capitula radiate or
discoid, terminal, solitary and scapose or in open,
rarely congested paniculiform or corymbiform
cymes. Involucres cylindrical, campanulate or
hemispheric, phyllaries subequal to gradate, in 2
to several series, sometimes reflexed at anthesis.
Receptacles flat to convex or globose, usually
epaleate. Ray florets fertile or neuter, obscurely but
most commonly conspicuously 3–5-lobed, limbs
mostly attenuate to oblique into tube, rarely obtuse
to truncate. Disc florets bisexual, rarely functionally staminate, corollas tubular to campanulate,
glabrescent to densely pubescent, veins meeting
at apices of corolla lobes (except in Plateilema);
anthers shortly caudate to ecaudate, appendages
narrowly ovate to round and strongly carinate,
cells of appendages mostly sclerified, filaments
and endothecium with or without crystals, median
endothecial cells quadratic to fusiform with 1–5
polar bridges, rarely with radial thickenings
confined to distal ends of thecae; style arms with
two stigmatic surfaces parallel to the edges of the
style arm, rarely meeting apically, mostly truncate
with a tuft of papillae, sometimes with a tapered,
vascularized appendage (Balduina, Gaillardia), or
with a proliferation of cells producing an obtuse
apex. Ray and disc cypselae essentially homomorphic, clavate to subterete, walls not carbonized,
mostly with large quadratic or isodiametric
crystals, glabrous to densely sericeous, sometimes
pubescence confined to ridges and alternating
with glandular trichomes. Pappus of several ovate
to quadratic, rarely triangular scales with aristate,
acuminate or obtuse apices, or of multiple bristles
either free or coalescing into scales, rarely absent.
The tribe contains 13 genera and approximately
120 species in North and South America, with most
species in the south-western USA and northern
Mexico. Many species grow in desert or dry regions and produce small rosettes through the wetter winter months, from which flowering shoots
arise rapidly as temperatures increase in the drier
spring months. Species of Balduina, Helenium and
Marshallia are adapted to wetter habitats of eastern North America. Several species of Helenium
and Gaillardia are horticulturally important.
Helenieae were characterized by Cassini
(1819) as having cypselae with scaly pappi. The
concept was modified by Bentham (1873b) to
include all taxa of Heliantheae affinity lacking
paleae. Turner and Powell (1977) concluded that
Helenieae were a paraphyletic assemblage and
disposed of the genera in several tribes including
Senecioneae and Heliantheae. Robinson (1981)
placed Helenieae into a large Heliantheae containing 35 subtribes. His subtribes Gaillardiinae
and Marshalliinae correspond to the concept of
Helenieae subtribe Gaillardiinae of Karis and
Ryding (1994b). Molecular studies of Helenieae
by Baldwin and Wessa (2000b) and Baldwin
et al. (2002) support a narrower interpretation
of Helenieae to include only 13 genera, which
corresponds to the composition of Robinson’s
Compositae
subtribes Gaillardiinae and Marshallinae, showing
that Helenieae is the basalmost lineage of the
classical Helenieae-Heliantheae complex. Most
Heleniae can be distinguished from any other
group in the Heliantheae alliance by their noncarbonized cypselae which rather contain crystals
in the cypsela wall (similarly to some Inuleae).
Additional characters which collectively can be
useful in recognizing Helenieae include mostly
truncate style arms, conspicuously trilobed ray
limb apices, a scaly pappus and mostly alternate
leaves. Baldwin and Wessa (2000b) recognize five
subtribes in tribe Helenieae, namely Gaillardiinae,
Marshallinae, Plateileminae, Psathyrotinae and
Tetraneurinae (as Riddelliinae; see Bierner 2001).
Key to the Subtribes
1. Pappus of bristles sometimes fused at base
4. Psathyrotinae (p. 403)
– Pappus of erose, quadratic to ovate, aristate or acuminate scales or absent
2
2. Receptacles paleate throughout
2. Marshalliinae (p. 402)
– Receptacles epaleate or paleate near margins
3
3. Veins of corolla lobe margins not merging at corolla
lobe apices
3. Plateileminae (p. 403)
– Veins of corolla lobe margins merging at corolla lobe
apices
4
4. Phyllaries reflexed at anthesis, if erect, then anther appendages with glandular trichomes on abaxial surface
1. Gaillardiinae (p. 401)
– Phyllaries erect (not reflexed) at anthesis
5. Tetraneuriinae (p. 404)
XIX.1. Subtribe Gaillardiinae Less. (1831).
Subtribe Heleniinae Less. (1832).
Annual or perennial herbs, sometimes caespitose,
rhizomatous and producing a thickened root
caudex or short stem. Leaves alternate, entire or
pinnatifid. Capitula scapose, solitary or in open
paniculiform or corymbiform cymes. Ray florets
fertile, rarely neuter. Disc florets bisexual. Cypselae
with a pappus of ovate, acuminate, aristate, rarely
linear scales.
Key to the Genera
1. Leaves without decurrent bases; styles with conic to
linear, conspicuous vascularized appendages, style
papillae with minute mucronate tips
2
– Leaves usually with decurrent bases; styles without
vascularized appendages, normally truncate with papillose tips; style papillae without mucronate tips
1146. Helenium
401
2. Anther appendages and style arms with glandular trichomes; pappus of scales without midrib extending as
an awn or bristle
1144. Balduina
– Anther appendages and style arms without glandular
trichomes; pappus of scales with midribs extending as
an awn or bristle
1145. Gaillardia
Genera of Gaillardiinae
1144. Balduina Nutt.
Balduina Nutt., Gen. N. Amer. Pl. 2: 175 (1818), nom. cons.;
Parker & Jones, Brittonia 27: 355–361 (1975), rev.
Annual or perennial herbs (?). Leaves sessile to
shortly petiolate, entire, linear to oblanceolate. Capitula radiate, terminal, solitary or in open corymbiform cymes. Involucres hemispheric, phyllaries
gradate. Receptacles strongly alveolate, with multiple aristae, growing after anthesis and forming an
aristate basket or cup around cypselae. Ray florets
neuter, 3-lobed, corollas bright to golden yellow.
Disc corollas bright yellow to purple or yellow with
purple lobes, sparsely glandular, without sclerified
cells, lobes densely pubescent on abaxial surfaces;
anthers ecaudate, appendages ovate, with glandular
trichomes, cells sclerified; style arms with tapered,
papillose apices and glandular trichomes, style appendages vascularized. Cypselae obconic, glabrous
to densely sericeous, sometimes glandular. Pappus
of several ovate scales with acuminate apices and
erose margins. x = 18. Three species, south-eastern
USA.
1145. Gaillardia Foug.
Gaillardia Foug., Obs. Phys. 29: 55 (1786); Biddulph, Res.
Stud. State College Wash. 12: 195–256 (1944), rev.
Annual or perennial herbs, rarely caespitose,
low shrubs with xylopodia, sometimes forming
rosettes, rarely strongly aromatic. Leaves petiolate
or sessile, entire to pinnatifid sometimes semisucculent. Capitula terminal or axillary, solitary,
sometimes on long peduncles, radiate or discoid.
Involucres hemispheric to broadly hemispheric,
phyllaries subequal, herbaceous or chartaceous.
Receptacles convex to hemispheric, densely setose.
Ray florets neuter, rarely fertile, corollas yellow
to red or purple, often bicoloured with deeply
trilobed apices. Disc corollas yellow or greenish
yellow, often red or deep purple distally, or purple
throughout, glabrous or sparsely pubescent,
without sclerified cells, lobes densely pubescent
on abaxial surfaces; anthers caudate, appendages
ovate to narrowly ovate, cells sclerified, especially
402
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
distally and on margins; style arms with tapered
papillose apices, style appendages vascularized.
Cypselae obpyramidal to obconic, glabrous to
densely sericeous, sometimes with ridges. Pappus
of 6–12 aristate, rarely linear scales with erose
margins. x = 9, 15, 17, 19. Twenty species, southeastern Canada, northern Mexico, south-western
USA, temperate eastern South America.
1146. Helenium L.
Fig. 80
Helenium L., Sp. Pl. 2: 886 (1753).
Annual or perennial herbs. Leaves alternate,
usually sessile, mostly decurrent, blades linear
to lanceolate, elliptic. Capitula terminal, solitary
or in open paniculiform cymes, sometimes on
swollen peduncles, radiate or discoid. Involucres
rotate to reflexed at anthesis, phyllaries subequal,
herbaceous. Receptacles convex to globose, rarely
with paleae on outermost rows of florets. Ray
florets neuter or pistillate and fertile, corollas
yellow or orange, sometimes brownish orange.
Disc corollas yellow, sometimes obscured by red or
purple pilose lobes, glabrous to sparsely pubescent,
without sclerified cells, lobes densely pubescent
on abaxial surface; anthers ecaudate, appendages
ovate to deltate, cells sclerified; style arms with
truncate papillose apices. Cypselae obpyramidal to
cylindric, sometimes ridged, glabrous or variously
pubescent, sometimes glandular. Pappus absent or
of a few scales with obtuse, acuminate or aristate
tips. x = 17. Thirty species, North and South
America, most species in northern Mexico and
southern USA.
XIX.2. Subtribe Marshalliinae H. Rob.
(1978).
Perennial, mostly glabrous herbs, sometimes rhizomatous and producing a thickened root caudex.
Leaves alternate, petiolate to subsessile, blades linear to narrowly ovate to oval, triplinerved or with
a single midvein. Capitula solitary or in open paniculiform cymes, discoid, paleate throughout. Involucres campanulate to hemispheric, phyllaries
herbaceous in 2 series. Receptacles flat to shallowly
convex. Florets bisexual, corollas deeply dissected,
white to cream, purplish or pale lavender, without sclerified cells, lobes sparsely pubescent with
a few large, round glandular trichomes; anthers
caudate, appendages ovate to round, cells sclerified only on upper margins; style arms with obtuse
papillose apices, papillae extending down to bifurcation. Cypselae obpyramidal, weakly 5-angled,
with tapered trichomes on ridges interspersed with
glandular trichomes. Pappus of 5–6 hyaline to ferrugineous acuminate scales. x = 9.
Only one genus:
1147. Marshallia Schreb.
Fig. 80. Compositae-Helenieae. Helenium mexicanum.
A Flowering branch and young plant. B Capitulum
with denuded receptacle. C Ray floret. D Disc corolla.
E Anthers. F Style. G Cypsela. (Reproduced from Flora
Novo-Galiciana, vol. 12, with permission of the University
of Michigan Herbarium; see McVaugh 1984)
Marshallia Schreb., Gen. Pl. 2: 810 (1791), nom. cons.; Channell, Contr. Gray Herb. 181: 41–132, (1957), rev.; Watson &
Estes, Syst. Bot. 15: 403–414, (1990), phylog.
Characters of the subtribe. Seven species, northeastern Mexico (M. caespitosa Nutt. ex DC., pers.
obs.), south-western and eastern USA.
Compositae
XIX.3. Subtribe Plateileminae B.G. Baldwin
(2000).
Perennial herbs, rosulate. Leaves alternate,
oblanceolate in outline, pinnatifid. Capitula
radiate, terminal, scapose, solitary. Involucres
campanulate. Receptacles flat, epaleate. Ray florets
fertile, corollas 3-lobed, yellow on adaxial surface,
pink to purple on abaxial surface. Disc florets
bisexual, corollas variously yellow, glabrous, without sclerified cells, lobes sparsely pubescent on
abaxial surfaces, trichomes with mucronate tips;
anthers ecaudate, appendages ovate to round, cells
sclerified; style arms with obtuse apices, papillae
with short mucronate tips. Ray cypselae shallowly
obcompressed to obconic, triquetrous, essentially
glabrous. Disc cypselae clavate, quadrate, essentially glabrous. Pappus of 5–8 quadratic or ovate
scales with obtuse, bifid or erose apices.
403
Genera of Psathyrotinae
1149. Pelucha S. Wats.
Pelucha S. Wats., Proc. Amer. Acad. Arts 24: 55 (1889).
Shrubs. Leaves alternate, sessile, blades oblanceolate in outline, pinnatifid with 1–2 pairs of linear
lobes. Capitula in terminal, corymbiform cymes,
discoid. Involucres campanulate. Receptacles flat.
Florets bisexual, corollas yellow, densely pubescent,
without sclerified cells, lobes densely pubescent on
abaxial surfaces; anthers caudate, appendages narrowly ovate, cells sclerified; style arm apices tapered, papillae confined to distal half of style arm.
Cypselae cylindric, densely sericeous. Pappus of
multiple bristles. x = 19. One species, P. trifida S.
Wats., island in Sea of Cortes, Mexico.
1150. Psathyrotes (Nutt.) A. Gray
Only one genus:
Psathyrotes (Nutt.) A. Gray, Smithsonian Contr. Knowl. 5:
100 (1853); Strother & Pilz, Madroño 23: 24–40, (1975), rev.
1148. Plateilema (A. Gray) Cockerell
Annual herbs, dichotomously branched, leaves of
first nodes opposite, soon thereafter producing
three leaves with very short internodes and
appearing whorled with a capitulum terminating
the main axis of the shoot, with three shoots
alternating with the leaves becoming the three
main branches of the plant. Leaves subalternate or
opposite, petiolate, blades ovate to deltate, margins
dentate to crenulate, densely pubescent, glaucous,
semisucculent. Inflorescence a terminal, solitary,
vegetative dichasium in which the central shoot
is a solitary capitulum and the lateral shoots are
vegetative. Capitula discoid. Involucres cylindric
to campanulate. Receptacles flat to convex. Florets
bisexual, corollas golden yellow, becoming purplish with age, glabrescent to sparsely pubescent,
without sclerified cells, lobes densely pubescent
on abaxial surfaces; anthers shallowly caudate,
appendages narrowly ovate, cells sclerified; style
arm apices obtuse, vascular strands conspicuously
expanded at style arm apices. Cypselae cylindric
to obpyramidal, sparsely to densely sericeous,
trichomes on ridges. Pappus of multiple bristles.
x = 17. Three species, north-western Mexico,
western USA.
Plateilema (A. Gray) Cockerell, Bull. Torrey Bot. Club 31:
462 (1904).
Characters of the subtribe. One species, P. palmeri
(A. Gray) Cockerell, north-western Mexico, Texas,
USA.
XIX.4. Subtribe Psathyrotinae B.G. Baldwin
(2000).
Annual or perennial herbs, rarely shrubs. Leaves
alternate or opposite. Capitula in dichasial cymes,
discoid, sometimes with peripheral florets with two
lobes expanded. Receptacles epaleate. Pappus of
multiple bristles, either free or connate into 5 scales.
Key to the Genera
1. Style arms with tapered apices and with papillae on
abaxial surfaces extending approximately halfway
down from apices; capitula in corymbiform cymes;
leaves linear, trilobed. Shrubs endemic to Isla San
Pedro Mártir, Sea of Cortes, Mexico
1149. Pelucha
– Style arms with obtuse to rounded apices with
papillae on abaxial surfaces confined to distalmost
end; capitula solitary in centres of dichotomously
branched cymes, the lateral shoots of which are
vegetative; leaves deltate, ovate or rhombic, variously
lobed, never trilobed. Annuals of the western USA
and Mexico
2
2. Pappus of free bristles
1150. Psathyrotes
– Pappus of bristles fused into 5 distinctive scales at base
1151. Trichoptilium
1151. Trichoptilium A. Gray
Trichoptilium A. Gray, Rep. U.S. Boundary Bot.: 97 (1859).
Annual herbs, dichotomously branched. Leaves
subalternate or opposite, shortly petiolate or ses-
404
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
sile, blades oblanceolate, bases attenuate, margins
dentate distally, white-floccose to lanate on abaxial
surface. Capitula arranged in dichasia in which the
central shoot is a solitary capitulum and the lateral
shoots are vegetative. Capitula discoid. Involucres
campanulate to hemispheric. Receptacles flat to
convex. Florets bisexual, corollas yellow, glabrescent, without sclerified cells, lobes moderately
pubescent on abaxial surfaces; anthers caudate,
appendages ovate, cells sclerified; style arm apices
obtuse. Cypselae cylindric to obpyramidal, densely
sericeous. Pappus of multiple bristles coalesced
into 5(–6) discrete scales. x = 13. One species,
T. incisum (A. Gray) A. Gray, north-western
Mexico, western USA.
XIX.5. Subtribe Tetraneuriinae Rydb.
(1915).
Subtribe Riddelliinae A. Gray (1884), nom illegit.
Subtribe Psilostrophinae B.L. Turner & A.M. Powell (1978).
Annual or perennial herbs, caespitose shrubs.
Leaves alternate, entire or dissected, sometimes
pinnatifid.Capitula scapose, solitary or in open to
congested paniculiform or corymbiform cymes,
radiate or discoid. Receptacles epaleate. Cypselae
epappose or with a pappus of ovate, acuminate or
aristate scales.
Key to the Genera
1. Lobes of disc corollas narrowly deltate, thickened and
without multicellular trichomes on abaxial surfaces;
cypselae prominently ribbed
1152. Amblyolepis
– Lobes of disc corollas deltate with variously shaped
multicellular trichomes on abaxial surfaces; cypselae
not prominently ribbed
2
2. Pappus absent
3
– Pappus present
4
3. Disc florets perfect
1153. Baileya
– Disc florets functionally staminate 1154. Hymenoxys
4. Throats (excluding lobes) of disc corollas with strongly
sclerified cells; rays with obtuse to truncate bases;
plants usually with floccose, whitish stems
1155. Psilostrophe
– Throats of disc corollas without sclerified cells; rays
with attenuate to oblique bases; plants usually without
floccose whitish stems
5
5. Plants caespitose with scapose, solitary capitula;
plants with mostly entire leaves
1156. Tetraneuris
– Plants various and with leaves associated with solitary or cymose capitula; plants with variously dissected leaves, when entire, ovate, sessile and evenly
distributed along flowering scape 1154. Hymenoxys
Genera of Tetraneuriinae
1152. Amblyolepis DC.
Amblyolepis DC., Prodr. 5: 667 (1836); Bierner, Madroño
37: 133–140 (1990), rev.
Annual herbs. Leaves petiolate to sessile, basal
spathulate, blades elliptic to ovate, bluish green
when live, drying green. Capitula terminal, solitary or loosely corymbiform, radiate. Involucres
campanulate to hemispheric, phyllaries subequal.
Receptacles convex to slightly globose with age.
Ray florets fertile, 3-lobed, corollas yellow-orange.
Disc corollas yellow-orange, essentially glabrous,
without sclerified cells; anthers ecaudate, appendages narrowly ovate, cells sclerified; style
arms with truncate papillose apices. Cypselae
obconic, strongly ribbed, densely pubescent,
glandular between ribs. Pappus of 5–6 quadrate,
hyaline scales with obtuse, rarely acuminate apices.
x = 19. One species, A. setigera DC., north-eastern
Mexico, Texas, USA.
1153. Baileya Harv. & A. Gray ex A. Gray Fig. 81
Baileya Harv. & A. Gray ex A. Gray, Not. Milit. Recon. 01.144
(1848).
Annual or perennial herbs, stems conspicuously
floccose. Leaves sessile, basal producing a rosette,
pinnatifid, cauline entire, linear to oblanceolate,
white-floccose especially on abaxial surfaces. Capitula terminal, solitary and scapose or in open
corymbiform cymes, radiate. Involucres campanulate to hemispheric, phyllaries subequal. Receptacles shallowly convex. Ray florets fertile, entire
to mostly 3-lobed, in 2–3 series, corollas bright
yellow, ray erect to patent during anthesis, then
reflexed, persistent. Disc florets bisexual, corollas
bright yellow, sparsely glandular, without sclerified
cells, lobes densely pubescent on abaxial surfaces;
anthers ecaudate, appendages ovate, cells sclerified;
style arms with truncate to slightly obtuse papillose
apices. Cypselae obconic to oblong, conspicuously
ribbed, glabrescent with a few, scattered glandular trichomes and short double trichomes. Pappus
absent. x = 16. Three species, northern Mexico,
western USA.
1154. Hymenoxys Cass.
Hymenoxys Cass., Dict. Sci. Nat., ed. 2, 55: 278 (1828);
Parker, Leaflets W. Bot. 9: 197–224 (1962), part. rev.;
Bierner, Brittonia 26: 385–392 (1974), part. rev.; Bierner,
Compositae
405
ovate, acuminate or caudate to aristate scales,
rarely absent. x = 15. Twenty-eight species (Bierner
2001), eastern and central Mexico, western USA,
temperate South America.
1155. Psilostrophe DC.
Psilostrophe DC., Prodr. 7: 261 (1838); Heiser, Ann.
Missouri Bot. Gard. 32: 265–278 (1944), rev.
Fig. 81.
Compositae-Helenieae. Baileya multiradiata.
A Flowering branch and leaf. B Capitulum past flower,
to show naked receptacle. C Ray floret. D Disc corolla.
E Anthers. F Style arms. G Cypsela. (Reproduced from
Flora Novo-Galiciana, vol. 12, with permission of the
University of Michigan Herbarium; see McVaugh 1984)
Sida 16: 1–8 (1994), part. rev.; Bierner & Jansen, Lundellia
1: 17–26 (1998), phylog.; Bierner, Lundellia 4: 37–63 (2001),
part. rev.
Annual or perennial herbs, sometimes with a thickened caudex. Leaves sessile or petiolate, entire or
mostly trilobed or bipinnate, blades mostly linear
or lanceolate, rarely ovate. Capitula terminal, solitary or in open to congested corymbiform cymes,
radiate, rarely discoid. Involucres campanulate
to hemispheric, phyllaries mostly in two series,
outer phyllaries fused at base. Receptacles flat to
shallowly convex. Ray florets fertile, conspicuously to obscurely 3-lobed, corollas yellow. Disc
florets bisexual or rarely functionally staminate,
corollas yellow, glabrescent to densely pubescent,
sometimes with a few sclerified cells, lobes densely
pubescent on abaxial surfaces; anthers ecaudate,
appendages ovate to round, cells sclerified; style
arms with truncate, rarely obtuse papillose apices. Cypselae obconic, moderately to densely
sericeous with glandular trichomes. Pappus of
Perennial, rarely biennial herbs, stems conspicuously floccose. Leaves petiolate to sessile, entire
to pinnatifid, pinnatifid leaves mostly in basal
rosette, blades linear, lanceolate to oblanceolate or
spathulate, floccose especially on abaxial surfaces.
Capitula in compact to open corymbiform cymes,
radiate. Involucres cylindric to campanulate, innermost phyllaries with membranaceous margins.
Receptacles flat to convex. Ray florets fertile,
3–5-lobed, corollas yellow, persistent and turning
white-stramineous with age. Disc florets bisexual,
corollas yellow, glabrous to sparsely pubescent,
with sclerified cells throughout except lobes, lobes
moderately pubescent on abaxial surfaces; anthers
ecaudate, appendages ovate, cells sclerified; style
arms with truncate apices. Cypselae obconic,
essentially glabrous to densely villous/sericeous,
trichomes uniseriate. Pappus of 4–6 hyaline
quadratic scales with obtuse, erose or laciniate
apices. x = 16. Seven species, northern Mexico,
western USA.
1156. Tetraneuris Greene
Tetraneuris Greene, Pittonia 3: 265 (1898); Bierner & Jansen,
Lundellia 1: 17–26 (1998), phylog.; Bierner & Turner, Lundellia 6: 44–96 (2003), rev.
Annual or perennial herbs, sometimes caespitose,
rarely shrubs. Leaves alternate, mostly tightly clustered on short nodes forming rosettes. Capitula
terminal, solitary on long scapes, rarely in open
paniculiform cymes, radiate. Involucres campanulate to hemispheric. Receptacles flat to convex. Ray
florets fertile, 3-lobed, persistent, corollas yellow.
Disc florets bisexual, corollas yellow, glabrescent
to sparsely pubescent, with a few sclerified cells
on throat, lobes moderately to densely pubescent;
anthers ecaudate, appendages ovate, cells sclerified; style arms with truncate apices. Cypselae obconic, moderately to densely sericeous especially
on ridges. Pappus of several acuminate to aristate
scales. x = 14, 15. Nine species, northern Mexico,
western USA.
406
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
XX. Tribe Coreopsideae Lindl. (1829). Mexico contains the highest generic diversity of
Family Coreopsidaceae Link (1829).
Tribe Bidentideae Godr. (1850).
Tribe Coreopsideae B.L. Turner &
A.M. Powell (1977), isonym.
J.L. Panero
Annual or perennial herbs, shrubs, sometimes
aquatic, rarely trees. Leaves alternate or opposite,
petiolate or sessile, blades mostly ovate or trullate,
commonly dissected, 1–3-pinnatifid, segments
lanceolate or linear sometimes filiform. Capitula
terminal, rarely axillary, solitary or more commonly in open paniculiform, sometimes congested
corymbiform, cymes, discoid or mostly radiate,
rarely disciform or ligulate. Involucres cylindric
to hemispheric, phyllaries mostly dimorphic
and gradate, rarely isomorphic, in 1–6 series,
outermost herbaceous, green, mostly reflexed,
innermost membranaceous, sometimes with
scarious margins. Receptacles flat to conic, usually
paleate. Ray florets with or rarely without corollas,
pistillate or sterile. Disc florets bisexual or functionally staminate, corollas usually pentamerous,
peripheral sometimes zygomorphic; anthers
yellow, brown, black or deep purple, appendages
lanceolate to round, sometimes wanting, glabrous,
rarely with glands, sometimes with prominent
resin canals, endothecial cells mostly quadrate
or oblong, sometimes fusiform with 1–3 polar
thickenings, sometimes with multiple radial and
polar thickenings; style arm apices broadly deltate
or acuminate to subulate, appendages short or very
well developed, appendages penicillate, papillose
and vascularized, papillae apically round, rarely
tapered. Cypselae usually isomorphic, sometimes
with resin canals, rarely striate, walls carbonized,
sometimes corky on edges, sometimes winged.
Pappus of 1–8 awns or absent, rarely of 4–6 round,
lanceolate or obovate scales, awns smooth or with
retrorse or antrorse barbs.
The tribe contains 30 genera and approximately 550 species with a worldwide distribution.
Most species are found in America, especially
North America. The tribe, as opposed to other
helianthoid tribes, has a fair number of genera endemic to the palaeotropics, Oceania and Polynesia.
Several members of the tribe are valued for their
horticultural potential; wild species of Coreopsis,
Cosmos and Dahlia and their hybrids are now
important components of gardens worldwide.
Coreopsideae.
Molecular studies of Heliantheae and related
tribes (Panero and Funk 2002) support the recognition of Coreopsideae at the tribal level. Similar
studies (Panero et al., unpubl. data) support, albeit tentatively, the inclusion of the genera Staurochlamys and Pinillosia in Coreopsideae. Pinillosiinae, as circumscribed by Robinson (1981) and including Pinillosia and Koehneola, are transferred
here to Coreopsideae. The other Cuban endemic
genus Tetraperone (in Heptanthinae sensu Robinson 1981) is also included in Coreopsideae on the
basis of its involucre morphology. Generic relationships within Coreopsideae until recently have been
very difficult to ascertain. The difficulty in separating Coreopsis from Bidens, and the apparent
close relationship of certain lineages of Coreopsis
to other genera in the tribe have increased uncertainty about the monophyly of several genera of
Coreopsideae (Tadesse et al. 2001). Molecular studies using the ITS region of the nuclear ribosomal
DNA by Kimball and Crawford (2004) show that the
traditional morphological features used to circumscribe genera and infer generic relationships (Ryding and Bremer 1992) are labile. Kimball and Crawford (2004) show that Ericentrodea appears to be the
basal lineage of the tribe, followed by a large polytomy including all genera of Coreopsideae sampled
to date. Bidens and Coreopsis are revealed as polyphyletic or paraphyletic assemblages, because distinctive genera such as Coreocarpus, Cosmos and
Thelesperma are derived from within them. Based
on the results by Kimball and Crawford (2004), the
relative position of Chrysanthellinae within Coreopsideae is impossible to ascertain but the distinctive morphology and photosynthesis of this
group provide support for its recognition. Their
data also support the close relationship of Oparanthus to Fitchia. Selleophytum clearly is a distinctive
genus sister to Narvalina, also endemic to Hispaniola. Coreopsis cyclocarpa is here included in Heterosperma. The genus Henricksonia is not placed
in the synonymy of Heterosperma, as it has a series of distinctive morphological features. Future
studies of the Heterosperma complex may support
the recognition of Coreopsis mutica as a distinctive
genus. The polyphyletic nature of Coreopsis, as revealed by molecular studies, supports the views of
Tadesse et al. (1995) that the genus may be treated as
a smaller assemblage of species essentially endemic
to the USA, with the other sections of the genus recognized as closely related genera. My circumscrip-
Compositae
tion of Coreopsis follows scenario 3 of Tadesse et al.
(1995), in which the genus is viewed as endemic
to the New World. I include the African species of
Coreopsis in Bidens. The inclusion of Diodontium
in Coreopsideae is tentative because the original
description states the herbage to be aromatic. Most
species of Coreopsideae have a particular odour
when the herbage is crushed, although this can
hardly be considered aromatic. Therefore, it is possible that Diodontium does not belong in Coreopsideae but rather in another tribe of Asteroideae,
possibly Astereae or Anthemideae. The relationship of Staurochlamys to Coreopsideae is difficult
to ascertain. The genus has a series of morphological features which are unique in the helianthoid alliance. My decision to include this genus in Coreopsideae is based solely on very limited molecular evidence; further analysis may support the inclusion
of Staurochlamys in Neurolaeneae, as suggested by
Robinson (1981), or its recognition as a distinctive lineage. Future molecular studies of Coreopsideae should focus on including Staurochlamys,
Pinillosiinae and members of related tribes such
as Helenieae, Tageteae and Neurolaeneae in an attempt to elucidate the relationships of this complex
lineage of the Heliantheae alliance. Furthermore, it
should be noted that Coreopsideae may be of hybrid origin, as molecular studies based on chloroplast DNA (Panero and Funk 2002) and nuclear
DNA (Goertzen et al. 2003) show widely different
but well-supported phylogenetic positions for the
tribe. Finally, it should be kept in mind that molecular studies and morphological evidence support
the recognition of subtribes Chrysanthellinae and
Pinillosiinae, with the residuum as a large subtribe
Coreopsidinae. It is very probable that Coreopsidinae, as circumscribed here, are not monophyletic.
407
– Leaves opposite, sometimes those associated with
the capitulescences alternate, stigmatic branch
appendages present normally shorter than length of
stigmatic area; plants without Kranz anatomy
2. Coreopsidinae (p. 409)
4. Plants small rosettes or trailing, forming mats; plants
of the Caribbean
3. Pinillosiinae (p. 416)
– Plants bushy; northern Australia 1. Chrysanthellinae
(1158. Diodontium)
XX.1. Subtribe Chrysanthellinae Ryding &
K. Bremer (1992).
Annual or perennial herbs, sometimes forming
rosettes, rarely shrubs. Leaves usually alternate,
blades entire or dissected, 1–2-pinnatifid. Capitula
terminal, solitary, mostly on long peduncles
appearing scapose or in open paniculiform cymes,
radiate, sometimes discoid. Involucre turbinate or
campanulate, sometimes hemispheric, phyllaries
in 2–5 series, dimorphic, outermost herbaceous,
normally much smaller than innermost, inner
erect, scarious. Receptacles paleate. Ray florets
pistillate or sterile, corollas sometimes lacking,
apices bilobed or trilobed. Disc florets bisexual, sometimes functionally staminate, corollas
tetramerous or pentamerous, mostly yellow;
anthers yellow to brown or yellow-purple, appendages ovate, rarely wanting, glabrous, with or
without resin canals; style arm apices subulate,
appendages normally as long as or several times
longer than stigmatic surfaces, densely papillose
and vascularized. Ray cypselae coiled or straight,
clavate to terete, sometimes narrowly obconic.
Disc cypselae obcompressed, rarely compressed,
quadrate, straight or sometimes concave. Pappus
of 2–4 erect or reflexed, smooth or retrorsely
barbed awns, sometimes absent.
Key to the Genera
Key to the Subtribes
and Unplaced Genera
1. Plants without glandular trichomes on anther
appendages, if glandular, then appendages hyaline,
ray corollas never deeply trilobed
2
– Plants with glandular, black anther appendages; ray
corollas deeply trilobed
1186. Staurochlamys
2. Receptacles paleate
3
– Receptacles epaleate
4
3. Leaves alternate, sometimes on short internodes and
restricted to the base of the plant; stigmatic branches
with prominent, penicillate appendages either as long
as or several times longer than stigmatic area; plants
with C4 photosynthesis and Kranz anatomy syndrome
1. Chrysanthellinae (p. 407)
1. Anther appendages with a prominent resin canal; capitula solitary on long peduncles; corollas mostly deep
purple or yellow-purple, sometimes pink; plants of
South America
1161. Isostigma
– Anther appendages without resin canals or these very
reduced; corollas yellow, white, pink or purple; plants
of North America, Oceania and palaeotropics, if in
South America, then corollas yellow and capitula not
scapose
2
2. Cypselae with a pappus of 3 or 4 awns
1162. Trioncinia
– Cypselae with a pappus of 2 awns or absent
3
3. Capitula epaleate
1158. Diodontium
– Capitula paleate
4
4. Ray florets with mostly bifid apices; tubes of disc corollas shorter than the throats
5
408
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
– Ray florets with mostly trifid apices; tubes of disc
corollas as long as throats
1160. Glossocardia·
5. Plants perennial and forming a woody, bulbous root or
base; leaves ampliated at base 1159. Eryngiophyllum
– Plants annual, if perennial, then with a narrow, woody
root, not a bulbous base
1157. Chrysanthellum
Genera of Chrysanthellinae
1157. Chrysanthellum Rich.
Fig. 82
Chrysanthellum Rich., Syn. Pl. 2: 471 (1807); Turner,
Phytologia 64: 410–444 (1988), rev.
Annual or perennial herbs, sometimes with
a woody caudex. Leaves basal in a loose rosette,
cauline mostly alternate, pinnatifid. Capitula
terminal, solitary or in open paniculiform cymes,
radiate. Involucres turbinate to campanulate,
phyllaries in 2–3 series, outermost very reduced,
innermost membranaceous, oval to ovate. Receptacles flat to convex. Ray florets pistillate,
corollas yellow, golden yellow or yellow-white,
narrowly lanceolate to oblong, apices bilobed.
Disc florets bisexual or functionally staminate,
corollas yellow to reddish orange, tetramerous or
pentamerous, sometimes innermost twice as large
as those surrounding them; anther appendages
round to ovate, acuminate or bilobed, without
resin canals; style arm apices subulate, appendages
penicillate, mostly longer than stigmatic surfaces.
Ray cypselae coiled or straight, clavate to terete,
sometimes narrowly obconic, sometimes with
prominent ridges, brown or blackish, glabrescent,
not winged. Disc cypselae like those of the ray
florets or biconvex, compressed or obcompressed,
sometimes winged, black or brown, glabrescent.
Pappus absent or of two obsolete awns on some
cypselae of the disc florets. x = 8, 9, 12. Eleven
species, pantropical, apparently not in Oceania,
most species in Mexico.
1158. Diodontium F. Muell.
Diodontium F. Muell., Hooker’s J. Bot. Kew Gard. Misc. 9:
19 (1857); Veldkamp & Kreffer, Blumea 35: 459–482 (1991),
rev.
Perennial herbs, shrubs (?), aromatic (?). Leaves
opposite (?), basal in tufts, sessile, blades linear. Capitula terminal in subscapose paniculiform cymes,
discoid. Receptacles flat, paleate, (epaleate?).
Disc florets bisexual, corollas yellow, tubular,
pentameous; anthers brownish, appendiculate.
Cypselae obcompressed (?), narrowly winged,
concave. Pappus of 2 smooth or barbed awns.
One species, D. filifolium F. Muell., northern
Australia.
1159. Eryngiophyllum Greenm.
Eryngiophyllum Greenm., Proc. Amer. Acad. Arts 39: 113
(1903).
Fig. 82. Compositae-Coreopsideae. Chrysanthellum filiforme. A Habit. B Flowering capitulum. C Capitulum in
bud showing involucre. D Ray corolla. E Disc corolla. F
Anthers. G Style arms. H Palea. I Cypsela. (Reproduced
from Flora Novo-Galiciana, vol. 12, with permission of the
University of Michigan Herbarium; see McVaugh 1984)
Perennial herbs forming a small rosette. Leaves alternate, sessile, blades linear, sometimes dissected.
Capitula terminal, subscapose, solitary or in open
paniculiform cymes, radiate. Involucres campanulate, phyllaries in 3–4 series, dimorphic, outermost
as long as innermost, innermost membranaceous.
Receptacles flat. Ray florets pistillate, corollas yellow, apices mostly trifid, rarely bifid. Disc florets bisexual, functionally staminate (?), corollas yellow;
anthers yellow, appendages broadly ovate, without
resin canals; style arm apices round to acuminate,
Compositae
appendages penicillate, very conspicuous, approximately 7–10 times as long as stigmatic surfaces,
papillose. Cypselae obcompressed, those of the ray
perhaps obpyramidal. Pappus absent. Two species,
western Mexico.
1160. Glossocardia Cass.
Glossocardia Cass., Bull. Sci. Soc. Philom. Paris 1817: 138
(1817); Veldkamp & Kreffer, Blumea 35: 459–482 (1991),
rev.
Annual or perennial herbs. Leaves alternate, uppermost sometimes opposite, sometimes forming
a loose rosette, sessile or petiolate, trullate or ovate
in outline, simple with trilobed apices or 1–2pinnatifid. Capitula terminal, solitary or in open
paniculiform cymes, radiate. Involucres cylindric
to campanulate, phyllaries in 2–3 series, outermost
filiform, herbaceous, innermost several times
longer, broader, membranaceous. Receptacles flat.
Ray florets pistillate or sterile, corollas yellow,
white, pink or purple, apices trilobed, rarely
bilobed. Disc florets bisexual, sometimes functionally staminate, corollas pentamerous, sometimes
tetramerous, yellow, white or brownish; anther
appendages ovate, sometimes wanting, without
resin canals; style arm apices filiform, appendages
papillose, 2–5 times as long as stigmatic surfaces.
Cypselae obcompressed, sometimes triquetrous,
oblong to linear-lanceolate, black to brown,
glabrous to densely pubescent. Pappus absent
or of two awns. x = 12. Eleven species, eastern
Africa, south-eastern Asia, Australia and Pacific
islands.
1161. Isostigma Less.
Isostigma Less., Linnaea 6: 513 (1831); Sherff, Bot. Gaz. 81:
241–257 (1926), rev.; Peter & Katinas, Austral. J. Bot. 51:
217–226 (2003), anat.
Perennial herbs, sometimes forming rosettes with
well-developed xylopodia or large subterranean
tuber-like roots. Leaves alternate, petiolate or
sessile, blades linear to obovate in outline, entire
or variously dissected, deeply dentate to twice
trifoliolate, segments linear to filiform. Capitula
terminal, solitary, scapose, radiate, rarely discoid.
Involucres campanulate or hemispheric, phyllaries
in 2–5 series, outermost reflexed, innermost membranaceous to scarious, erect. Receptacles flat. Ray
florets pistillate, corollas dull red-purple, pink,
sometimes yellow on adaxial surface, limb oval,
sometimes conspicuously trilobed. Disk florets
409
bisexual, corollas yellow-red or deep purple, pentamerous, rarely tetramerous; anther appendages
broadly ovate, glabrous, with a central resin
canal; style arm apices subulate, strongly arcuate,
appendages as long as or longer than stigmatic
surfaces, cylindric. Cypselae obcompressed, rarely
quadrate, rectangular to obpyramidal, black,
glabrous to sparsely pubescent. Pappus of 2 erect
to reflexed, smooth or barbellate awns, sometimes
with a few minute scales in between, sometimes
wanting. Approximately 15 species, Argentina,
Bolivia, Brazil, Paraguay and Uruguay.
1162. Trioncinia (F. Muell.) Veldkamp
Trioncinia (F. Muell.) Veldkamp, Blumea 35: 480 (1991).
Perennial herbs. Leaves alternate, mainly basal,
blades trullate in outline, simple or 1–2-pinnatifid.
Capitula in terminal, open paniculiform cymes,
radiate. Involucres campanulate (?), phyllaries in
2–3 series. Receptacles flat. Ray florets sterile (?),
corollas yellow (?). Disc florets bisexual, tetramerous. Cypselae obcompressed, brown or black,
tuberculate. Pappus of 3–4 strongly reflexed awns.
One species, T. retroflexa (F. Muell) Veldkamp,
north-eastern Australia.
XX.2. Subtribe Coreopsidinae Cass. ex
Dumort. (1829).
Subtribe Petrobiinae Benth. in Benth. & Hook. f. (1873).
Subtribe Fitchiinae Carlquist (1957).
Annual or perennial herbs, shrubs, sometimes
aquatic, rarely trees. Leaves alternate or opposite, blades mostly ovate or trullate, commonly
dissected, 1–3-pinnatifid, segments lanceolate or
linear. Capitula in terminal, rarely axillary, open
paniculiform or sometimes congested corymbiform cymes, discoid or mostly radiate, rarely
ligulate. Involucres cylindric to hemispheric,
phyllaries mostly dimorphic and gradate, rarely
isomorphic, in 1–6 series, outermost herbaceous,
green, innermost membranaceous. Receptacles
paleate. Ray florets pistillate. Disc florets bisexual
or functionally staminate, corollas pentamerous,
sometimes tetramerous rarely hexamerous, peripheral sometimes zygomorphic, mostly yellow
or purple; anther appendages lanceolate to round,
sometimes wanting, glabrous, sometimes with
glands, sometimes with prominent resin canals;
style arm apices broadly deltate or acuminate to
subulate, appendages short or very well developed
410
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
and sometimes several times longer than stigmatic
surfaces, penicillate, papillose and vascularized.
Cypselae isomorphic or sometimes those of the
ray or the innermost disc florets different in shape
from the other cypselae, sometimes growing after
anthesis, sometimes corky on edges or winged.
Pappus of 1–8 awns or absent, rarely of 4–6 round,
lanceolate or obovate scales, awns smooth or with
retrorse or antrorse barbs.
–
12.
–
Key to the Genera
1. Anther filaments papillose or pubescent
1166. Cosmos
– Anther filaments not papillose nor pubescent
2
2. Abaxial surface of disc corollas with a ring of broad trichomes at base of throat; cypselae with corky sinuate
or pectinate margins
1164. Coreocarpus
– Abaxial surface of disc corollas glabrous or variously
pubescent, pubescence not restricted to a ring at base
of throat; cypselae with or without wings, wings entire,
very rarely sinuate or pectinate
3
3. Phyllaries of innermost series variously fused at base
up to mid-length
1182. Thelesperma
– Phyllaries free
4
4. Capitula with dimorphic cypselae, normally those
of the ray floret obcompressed, broadly oblong and
winged, those of the disc florets or innermost disc
florets obpyramidal to terete, fusiform
5
– Capitula with isomorphic cypselae
8
5. Disc cypselae terete to obpyramidal, striate; pappus of
scales; anther appendages glandular
1173. Henricksonia
– Disc cypselae obcompressed to weakly terete, not striate; pappus absent or of two diverging awns; anther
appendages not glandular
6
6. Involucres with gradate phyllaries, not dimorphic; leaf
bases oblique
1172. Goldmanella
– Involucres with dimorphic phyllaries, outermost
herbaceous, green, reflexed, innermost erect,
membranaceous
7
7. Shrubs or small trees with corymbiform inflorescences
1165. Coreopsis
– Annual or perennial herbs, rarely woody with capitula
solitary or in open paniculiform inflorescences
1174. Heterosperma
8. Leaf bases oblique; capitula in umbelliform cymes
1172. Goldmanella
– Leaf bases various, never oblique; inflorescences various, never umbelliform cymes
9
9. Capitula ligulate, pendulous; shrubs or small trees of
Polynesia
1171. Fitchia
– Capitula various, never ligulate, erect; worldwide, if
shrubs or trees from Polynesia, then capitula not ligulate
10
10. Plants aquatic with dimorphic leaves; capitula solitary
1176. Megalodonta
– Plants terrestrial, if aquatic without dimorphic leaves;
capitula variously arranged
11
11. Cypselae urceolate in outline; pappus of several (3 or
more) retrorsely barbed awns arising from shoulders
13.
–
14.
–
15.
–
16.
–
17.
–
18.
–
19.
–
20.
–
21.
–
of cypsela; vines of tropical Andean South America
1170. Ericentrodea
Cypselae various, pappus absent or, if retrorsely
barbed, then awns mostly 2, one on each side of
cypsela; annual or perennial herbs, shrubs or trees, if
vines, then not from Andean South America
12
Shrubs or trees with broadly ovate to round, very thick,
coriaceous leaves and densely pectinate foliar venation; style branches with continuous stigmatic surfaces, disc corollas tetramerous; plants of Rapa Island,
French Polynesia
1179. Oparanthus
Plants various, without thick, broadly ovate or round
leathery leaves; style branches with parallel stigmatic
surfaces; disc corollas mostly pentamerous, rarely
tetramerous; worldwide, not from Rapa Island
13
Small, dioecious trees; disc corollas tetramerous,
lobes narrowly lanceolate; plants from St. Helena
island
1180. Petrobium
Annual, perennial herbs or shrubs, never trees; disc
corollas pentamerous, if tetramerous, then mostly yellow and without narrowly lanceolate lobes; worldwide,
not from St. Helena island
14
Capitula with winged cypselae, wings corky, stramineous
15
Capitula rarely with winged cypselae, wings, if present,
very narrow, not corky
16
Leaves simple; cypselae with a pappus of two awns;
plants from Hispaniola
1178. Narvalina
Leaves compound; cypselae with a pappus of two awns
arising from a cup or corona; plants of Ecuador
1167. Cyathomone
Capitula with all disc florets functionally staminate 17
Capitula with bisexual disc florets, sometimes a few of
the innermost florets functionally staminate
19
Disc florets 3–4; limbs of ray florets very reduced and
mostly covered by phyllaries; cypselae with a pappus
of broadly divergent awns
1169. Dicranocarpus
Disc florets 10–many; limbs of ray florets conspicuous,
protruding and spreading beyond phyllaries; cypselae
epappose or with two broad awns
18
Perennial herbs or shrubs; ray florets with conspicuous trilobed apices; cypselae oval and plump, broadly
convex in cross-section; plants of southern India and
Sri Lanka
1177. Moonia
Vines; ray florets with minute and inconspicuous
bilobed or trilobed apices; cypselae broadly oval,
flattened, lenticular in cross-section; plants of Mexico
and Central America
1175. Hidalgoa
Style branches with large, conspicuous papillose appendages, appearing plumose to the naked eye; capitula nodding
1168. Dahlia
Style branches without large, conspicuous, papillose
appendages, capitula erect
20
Cypselae with antrorsely barbed awns; plants of America and surrounding islands
21
Cypselae with retrorsely barbed awns, when plants
from Africa, cypselae may be antrorsely awned
1163. Bidens
Shrubs with sessile, subcoriaceous, lanceolate to ovate
leaves; plants from Hispaniola
1181. Selleophytum
Annual or perennial herbs, if shrubs, then leaves otherwise; plants mostly from continental America
1165. Coreopsis
Compositae
411
Genera of Coreopsidinae
1163. Bidens L.
Bidens L., Sp. Pl. 2: 831 (1753); Sherff, Field Mus. Nat. Hist.,
Bot. 16: 1–709 (1937), rev.; Tadesse, Kew Bull. 48: 437–516
(1993), part. rev.
Annual or perennial herbs, vines or shrubs,
sometimes aquatic. Leaves opposite, sometimes
alternate towards inflorescence, rarely whorled,
simple to pinnately compound, blades deltate to
ovate in outline. Capitula in terminal, open to
congested corymbiform to paniculiform cymes,
radiate or discoid. Involucres campanulate to
hemispheric, phyllaries dimorphic, outer herbaceous, inner membranaceous. Receptacles flat
to convex. Ray florets sterile, rarely pistillate,
corollas yellow, orange, white with black nerves on
abaxial surface, rarely purple or pink. Disc florets
bisexual, corollas mostly yellow to orange, rarely
greenish, pink, white or purple; anther appendages
ovate, glabrous, with or without resin canals; style
arm apices deltate, densely papillose, appendages
cylindric, sometimes not vascularized. Cypselae
obcompressed, triquetrous, quadrate, obovoid
to oblong, linear or fusiform in outline, black to
reddish brown, glabrous to densely pubescent,
sometimes tuberculate, rarely winged. Pappus
absent or mostly 2 to few awns or scales, awns
usually retrorsely barbed, antrorsely barbed in
some African species, rarely glabrous. x = 10, 11
12, 17, 18. Approximately 280 species, worldwide,
most species in America.
1164. Coreocarpus Benth.
Coreocarpus Benth., Bot. Voy. Sulphur 28, pl. 16 (1844);
Smith, Syst. Bot. 14: 448–472 (1989), rev.; Kimball et al.,
Evolution 57: 52–61 (2003), syst.
Annual or perennial herbs or weak shrubs. Leaves
opposite, blades ovate to trullate in outline,
1–2-pinnate, segments linear to lanceolate. Capitula in open paniculiform, rarely congested
corymbiform cymes, radiate, rarely discoid.
Involucre campanulate to hemispheric, phyllaries
in 2–3 series, dimorphic, outer 1–3 very reduced,
linear, herbaceous, membranaceous, appressed.
Receptacles flat to convex. Ray florets sterile, rarely
pistillate, corollas yellow or white with pink or
purple veins. Disc florets bisexual or sometimes
functionally staminate, sometimes only 1 floret
fertile, corollas yellow, with a ring of broadly
lanceolate, glandular trichomes on abaxial surface
Fig. 83. Compositae-Coreopsideae. Coreopsis petrophiloides.
A Flowering branch. B Flowering capitulum with some ray
florets removed. C Ray corolla. D Disc corolla. E Anthers.
F Style arms. G Palea. H Cypsela. (Reproduced from Flora
Novo-Galiciana, vol. 12, with permission of the University
of Michigan Herbarium; see McVaugh 1984)
at junction of tube and throat; anther appendages
ovate to suborbicular, glabrous, resin canals
poorly developed or wanting; style arm apices
subulate, appendages as long as or longer than
stigmatic lines, subulate to cylindric. Cypselae
obcompressed, somewhat concave or cucullate,
black, margins corky, stramineous, wings sinuate
or tuberculate to shallowly crenate, essentially
glabrous. Pappus of 1–2 retrorsely barbed awns
or absent. x = 12. Approximately eight species,
western North America.
1165. Coreopsis L.
Fig. 83
Coreopsis L., Sp. Pl. 2: 907 (1753); Sherff, Field Mus. Nat.
Hist., Bot. (1936), rev.; Crawford, Amer. J. Bot. 58: 361–367
(1971), part. rev.; Smith, Brittonia 25: 200–208 (1973), part.
rev.; Crawford, Brittonia 28: 329–336 (1976), part. rev.;
412
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Smith, Sida 123–215 (1976), part. rev.; Jansen et al., Pl. Syst.
Evol. 157: 73–84 (1987) phylog.; Tadesse et al., Brittonia 47:
61–91 (1995), anat., phylog.
x = 12, 17. Approximately 28 species, Mexico, Central America and Andean South America.
Annual or perennial herbs, shrubs or small trees.
Leaves opposite to alternate towards inflorescence, blades simple or 1–3 pinnately compound,
triplinerved, pinnate. Capitula solitary or in open
paniculiform or corymbiform cymes, discoid
or radiate. Involucres cylindric to campanulate, phyllaries in 2–3 series, dimorphic, outer
herbaceous, spreading, inner membranaceous to
scarious, erect. Receptacles flat to convex. Ray
florets sterile, rarely pistillate, corollas yellow
to orange, sometimes bicoloured with purplish
bases. Disc florets bisexual, sometimes innermost
functionally staminate, corollas pentamerous,
rarely tetramerous, yellow, orange to purplish
brown; anther appendages ovate, glabrous, with
or without resin canals; style arm apices deltate,
densely papillose, appendages cylindric, sometimes not vascularized. Cypselae obcompressed,
oblong, elliptic, fusiform, sometimes with corky,
entire or pectinate margins or wings, glabrous to
moderately pubescent. Pappus absent or of two
awns, sometimes a short crown of scales, awns
antrorsely barbed. x = 12, 13, 14. Approximately
70 species, America.
1167. Cyathomone S.F. Blake
1166. Cosmos Cav.
Cosmos Cav., Icon. 1: 9, pl. 14 (1791).
Annual or tuberous perennial herbs. Leaves opposite, blades deltate, sagittate or ovate in outline, 1–3-pinnatifid or pinnate, rarely simple. Capitula terminal, solitary or in very open paniculiform cymes, discoid or radiate. Involucres cylindric
to hemispheric, phyllaries dimorphic, outermost
herbaceous, shortly fused, inner membranaceous
or scarious. Receptacles flat. Ray florets neuter,
corollas pink, magenta, maroon to black-purple,
white, rarely red or orange. Disc florets bisexual,
corollas yellow, green, white, brownish yellow, purple or bicoloured; anther filaments pubescent, appendages broadly ovate, with a prominent resin
canal; style arm apices deltate-aristate, appendages
prominent, as long as or longer than stigmatic surfaces, deltate/aristate. Cypselae fusiform to linear
or narrowly lanceolate, apices conspicuously tapered, usually quadrate in cross-section, black to
dark brown, glabrescent to hispid, curved outwards
with age. Pappus of 1–8 retrorsely barbed and either erect or divergent awns, sometimes absent.
Cyathomone S.F. Blake, J. Wash. Acad. Sci. 13: 105 (1923).
Shrubs (?). Leaves opposite, ternate, biternate or
pinnate-ternate. Capitula in terminal, paniculiform cymes. Involucre campanulate (?), phyllaries
in 2–3 series, outermost about 5, herbaceous,
innermost longer and membranaceous. Receptacles convex. Cypselae obcompressed, obovate
in outline, black, wings stramineous, erose and
fused to two fragile awns, squamellae around neck
fused to awns and forming a cup. Pappus of two
fragile awns fused to a ciliolate cup. One species,
C. sodiroi (Hieron.) S.F. Blake, Ecuador.
1168. Dahlia Cav.
Dahlia Cav., Icon. 1: 56 (1791); Sorensen, Rhodora 71:
309–365, 367–416 (1969), rev.; Gatt, Hammett & Murray,
Bot. J. Linn. Soc. 133: 229–239 (2000), phylog.; Saar et al.,
Syst. Bot. 28: 627–639. (2003), phylog.
Tuberous perennial herbs or shrubs, one species
epiphytic, sometimes rupicolous. Leaves opposite,
sometimes whorled, sometimes semisucculent,
blades simple to 1–3-pinnatifid, ovate to deltate
in outline, rarely cordate. Capitula solitary or
loosely aggregated in paniculiform cymes, radiate,
nodding. Involucres cylindric to campanulate,
hemispheric, phyllaries in 2–3 series, strongly
dimorphic, outermost herbaceous, reflexed at
anthesis, innermost broad, scarious, translucent.
Receptacles flat. Ray florets sterile, corollas in
a wide variety of shades of pink and lavender,
yellow, orange, red or dark maroon sometimes
appearing black, white. Disc florets bisexual,
innermost often functionally staminate, corollas
yellow to orange, red or purple, sometimes
bicoloured; anther appendages ovate to deltate,
glabrous or sometimes with a few short trichomes
or papillae; style arms densely papillose abaxially, apices narrowly deltate, subulate, as long
as or longer than stigmatic surfaces, cylindric,
densely papillose. Cypselae obcompressed, linear
to spathulate, sometimes shallowly tuberculate,
glabrous to sparsely pubescent, black or greyish
black. Pappus absent or of 2(5) small teeth or of
two weak, filiform, caducous bristles. x = 12, 16,
17, 18. Approximately 40 species, Mexico, Central
America, Colombia, widely cultivated.
Compositae
1169. Dicranocarpus A. Gray
Dicranocarpus A. Gray, Mem. Amer. Acad. Arts n.s. 5: 322
(1854).
Annuals. Leaves opposite, blades ovate in outline,
1–2-pinnatifid, segments linear. Capitula terminal,
solitary, sometimes nodding and appearing axillary, radiate. Involucre turbinate to campanulate,
phyllaries in 2–3 series, dimorphic, outer narrowly
lanceolate to linear, herbaceous, innermost scarious. Receptacle flat. Ray florets pistillate, corollas
very reduced and covered by phyllaries, yellow. Disc
florets functionally staminate, style undivided, papillose, corollas yellow, throat abruptly narrowed
into tube; anther appendages translucent, sometimes suffused with black, glabrous, without resin
canals; style arm apices of disc corollas deltate to
acuminate, of ray corollas rounded to subulate.
Cypselae obcompressed to shallowly terete, linear,
moderately pubescent, growing after anthesis and
sometimes up to 2–3 times length of capitulum.
Pappus of two diverging, arcuate, very conspicuous, smooth awns, growing after anthesis. n = 10.
One species, D. parviflorus A. Gray, Mexico, USA.
1170. Ericentrodea S.F. Blake
Ericentrodea S.F. Blake, J. Wash. Acad. Sci. 13: 104 (1923).
Weak shrubs or vines. Leaves opposite, blades
ovate to deltate in outline, 2–3-pinnate, segments linear to lanceolate. Capitula in terminal,
corymbiform cymes, discoid, rarely radiate.
Involucres campanulate to hemispheric, phyllaries
in 2–4 series, dimorphic, outermost 2 series
reflexed, green, herbaceous, innermost series
erect, membranaceous. Receptacles convex. Ray
florets pistillate, corollas white (?) or yellow. Disc
florets bisexual, corollas yellow or white, throats
broad and abruptly narrowed into tube; anther
appendages black, glabrous, with a prominent
central resin canal; style arm apices deltate,
stigmatic surfaces very broad, dense, touching
and appearing continuous, confluent at apices,
stigmatic papillae sometimes black. Cypselae
obcompressed, flatly biconvex, obovate, with two
minute or well-developed shoulders flanking neck,
ciliate, brown. Pappus of 3 to numerous, retrorsely
barbed awns on each shoulder of cypselae, awns
spreading, reflexed or arched inwards. Six species,
Andes, Colombia to Bolivia.
1171. Fitchia Hook. f.
Fitchia Hook. f., London J. Bot. 4: 640 (1845).
413
Shrubs or small trees. Leaves opposite, blades
entire, broadly ovate to cordate, sometimes
elliptic, entire to crenulate. Capitula axillary,
pendulous on lax or recurved peduncles, ligulate.
Involucres hemispheric, phyllaries in 5–6 series,
outermost woody, innermost with scarious margins. Receptacles flat to shallowly convex. Florets
bisexual, corollas yellow-orange, inrolled forming
a navicular structure; anthers with prominent,
lanceolate appendages, without resin canals; style
arms reduced, acute to round, stigmatic surfaces
continuous. Cypselae obcompresed, oblong,
densely pubescent. Pappus of 2 barbellate awns.
Seven species, Polynesia.
1172. Goldmanella Greenm.
Goldmanella Greenm., Bot. Gaz. 45: 198 (1908).
Goldmania Greenm. (1907), nom. illegit.
Prostrate, perennial herbs rooting at nodes. Leaves
alternate, blades asymmetrically ovate to elliptic,
bases oblique. Capitula terminal in umbelliform
cymes, radiate. Involucres campanulate, phyllaries
in 3–4 series, gradate with conspicuous brown
striae, membranaceous. Receptacles conic, paleate.
Ray florets pistillate, corollas white to pale yellow.
Disc florets bisexual, corollas yellow-orange, pentamerous, rarely hexamerous; anther appendages
deltate, sides involute, glabrous, without resin
canals; style arm apices subulate, tapered, papillae
sometimes extending below bifurcation point,
appendages long, cylindric, papillose. Cypselae
shallowly obcompressed, innermost subterete,
cylindric, brown or reddish brown, corky. Pappus
absent or of 2–4 minute protuberances. One
species, G. sarmentosa Greenm., Mexico, Belize,
Guatemala.
1173. Henricksonia B.L. Turner
Henricksonia B.L. Turner, Amer. J. Bot. 64: 78 (1977).
Weak, rupicolous shrubs. Leaves opposite, blades
twice tripinnate, segments lanceolate. Capitula
terminal, solitary on long peduncles, radiate.
Involucres campanulate to hemispheric, phyllaries in 2 series, dimorphic, outer herbaceous,
patent to reflexed, inner chartaceous, striate, each
subtending a ray floret. Receptacles shallowly
convex. Ray florets pistillate, corollas yellow with
conspicuous reddish veins. Disc florets bisexual, corollas yellow; anther appendages ovate,
glandular abaxially, glands also on distal part of
connective, without resin canals; style arm apices
414
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
acuminate, appendages cylindric. Ray cypselae obcompressed, oval, broadly winged, wings smooth,
corky, of even width throughout circumference
of cypsela, glabrous, black, wings stramineous.
Disc cypselae obpyramidal, 3–5-sided, brown,
striate, some striae thickened (carinate), brown,
sparsely pubescent. Pappus of ray cypselae of two
small projections or awns, disc cypselae with 3–5
obovate squamellae. One species, H. mexicana B.L.
Turner, northern Mexico.
1174. Heterosperma Cav.
Fig. 84
Heterosperma Cav., Icon. 3: 34 (1795/1796).
Annual or perennial herbs, weak shrubs. Leaves
opposite, blades simple to 1–2-pinnatifid, segments linear. Capitula terminal, solitary, radiate.
Involucres cylindric to campanulate or hemi-
spheric, phyllaries in 2–3 series, dimorphic, outer
herbaceous, linear, inner membranaceous to
chartaceous, ovate. Receptacles flat to convex.
Ray florets pistillate, corollas yellow to orange,
sometimes conspicuously trilobed. Disc florets
bisexual, fertile, corollas yellow to orange; anther
appendages ovate to deltate with shallowly to
strongly developed resin canals; style arm apices
deltate, papillose, appendages cylindrical. Cypselae obcompressed, those of the ray and sometimes
some or nearly all except for two to four of the
innermost disc florets broad, oval, with corky wide
margins, flatly biconvex, the innermost growing
after anthesis, terete, protruding above the capitula, and slightly arched. Pappus absent or of
2–4 retrorsely barbed awns, mostly perpendicular
to the cypsela, mostly restricted to innermost
cypselae. x = 11, 12, 13. Approximately seven
species, neotropical, introduced elsewhere.
1175. Hidalgoa La Llave & Lex.
Hidalgoa La Llave & Lex., Nov. Veg. Descr. 1: 15 (1824).
Vines or lianas. Leaves opposite, petioles twining
at their bases, blades suborbicular in outline, pedate or pinnate with 3–5 or more leaflets. Capitula
terminal or axillary, solitary or in simple cymes
on long peduncles, radiate. Involucres campanulate, phyllaries dimorphic, outer fleshy, herbaceous, spreading, inner membranaceous. Receptacles flat. Ray florets pistillate, corollas bright yellow
to orange or red. Disc florets functionally staminate, corollas yellow to orange; anther appendages
shortly acute, brown, glabrous, sometimes with one
gland, without resin canals; style arms of disc florets fused, penicillate, of ray florets slender, subulate, spreading, stigmatic surfaces parallel, confluent at apices. Cypselae obcompressed, biconvex,
fusiform, brown to black. Pappus absent, wings
projecting as thickened, lacerate awns above neck.
x = 15, 16. Five species, Mexico, Central America.
1176. Megalodonta Greene
Megalodonta Greene, Pittonia 4: 271 (1901).
Fig. 84. Compositae-Coreopsideae. Heterosperma pinnatum. A Habit. B Fruiting capitulum. C Ray corolla. D Disc
corolla. E Anthers. F Style arms. G Ray cypsela. H Inner disc
cypsela. (Reproduced from Flora Novo-Galiciana, vol. 12,
with permission of the University of Michigan Herbarium;
see McVaugh 1984)
Aquatic annual or perennial herbs. Leaves opposite,
submerged leaves finely dissected, filiform, those
above water simple, lanceolate, serrate. Capitula
terminal, solitary, radiate. Involucres hemispheric,
phyllaries in 2–3 series, shallowly gradate, outermost few, herbaceous, reflexed, innermost scarious. Receptacles flat. Ray florets neuter, corollas
yellow. Disc florets bisexual, corollas yellow; anther
Compositae
appendages hyaline to yellow, glabrous; style arm
apices deltate, densely papillose. Cypselae obcompressed, linear to narrowly fusiform, brown, essentially glabrous. Pappus of approximately 6, easily
caducous, retrorsely barbed awns, sometimes one
of the awns reflexed. One species, M. beckii (Torr.
ex Spreng.) Greene, northern USA, Canada.
1177. Moonia Arn.
Moonia Arn., Nova Acta Phys.-Med. Acad. Caes. Leop.Carol. Nat. Cur. 18: 348 (1836); Stuessy, Brittonia 27: 97–102
(1975), rev.
Perennial herbs, shrubs (?). Leaves opposite, blades
trilobed, lobes ovate to elliptical, serrate, terminal
lobe deeply to strongly trilobed. Capitula terminal,
solitary on long peduncles, radiate. Involucres
campanulate to hemispheric, phyllaries in 4 series,
outermost 2 series lanceolate, herbaceous, reflexed,
innermost series appressed, densely and minutely
striate. Receptacles flat. Ray florets pistillate,
corollas yellow with conspicuous reddish veins,
deeply trilobed, outer lobes wider than central and
overlapping it. Disk florets functionally staminate,
corollas yellow; anther appendages short, ovate,
black or sometimes basal half black, distal half
stramineous, with a weak resin canal in the centre;
styles of disc florets undivided, apices deltate,
papillose, styles of ray florets subulate, spreading.
Cypselae obcompressed, broadly biconvex, slightly
concave with abaxial surface larger than adaxial
surface, apices with a short annular neck, greenish
black, glabrous. Pappus absent. One species,
M. heterophylla Arn., south-eastern India, Sri
Lanka.
1178. Narvalina Cass.
Narvalina Cass., Dict. Sci. Nat. 38: 17 (1825).
Shrubs. Leaves opposite, blades obovate or elliptic,
semisucculent, with a few, evenly spaced, conspicuous teeth on each side of lamina. Capitula
in terminal, open paniculiform cymes, radiate.
Involucres turbinate to campanulate, phyllaries in
3 series, gradate, outermost herbaceous, spreading, innermost membranaceous to coriaceous.
Receptacles flat. Ray florets pistillate, corollas
yellow. Disc florets functionally staminate, corollas
yellow; anther appendages ovate, glabrous with
a conspicuous resin canal; style arm apices deltate,
densely papillose. Cypselae obcompressed, oval in
outline, black, glabrous, with a thick, stramineous,
corky wing all around cypsela. Pappus of diverg-
415
ing, retrorsely barbed awns, stramineous, fragile.
One species, N. domingensis Cass., Hispaniola.
1179. Oparanthus Sherff
Oparanthus Sherff, Occas. Pap. Bernice P. Bishop. Mus. 12: 9
(1937); Stuessy, Fieldiana, Bot. 38: 63–70 (1977), rev.; Shannon & Wagner, Allertonia 7: 273–295 (1997), rev.
Shrubs or trees. Leaves opposite, blades oval to
round, entire, pinnately veined, sometimes very
thick, leathery. Capitula in terminal, solitary, simple, open or congested cymes, radiate. Involucres
cupuliform, campanulate, phyllaries in 2–3 series,
coriaceous, striate. Receptacles convex. Ray florets
pistillate, corollas white, yellow to yellow-green,
apices trilobed, rarely bilobed or tetralobed, style
arms with continuous stigmatic surfaces. Disc florets functionally staminate, corollas white or yellow; anthers black, styles of disc florets undivided,
penicillate. Cypselae obcompressed, linear to narrowly fusiform, narrowly winged on one or both
sides, wings extending as awns, glabrous. Pappus
of 2–3 smooth awns. Approximately six species,
Rapa Island, Marquesas Islands, French Polynesia.
1180. Petrobium R. Br.
Petrobium R. Br., Trans. Linn. Soc. London 12: 113 (1817).
Shrubs or small trees, dioecious. Leaves opposite,
blades ovate to elliptic, dentate. Capitula terminal in simple cymes, discoid. Involucres campanulate, phyllaries dimorphic in 2–4 series, outermost herbaceous in 1–2 series, green, innermost
membranaceous. Receptacles flat. Florets bisexual
or functionally staminate or functionally pistillate,
corollas tetramerous, white or brownish white; anther appendages ovate. Cypselae obcompressed to
triquetrous, linear, shallowly winged, margins ciliate. Pappus of 2–3 awns. One species, P. arboreum
R. Br., St. Helena island.
1181. Selleophytum Urb.
Selleophytum Urb., Repert. Spec. Nov. Regni Veg. 13: 483
(1915).
Shrubs. Leaves opposite, blades lanceolate to
narrowly ovate, coriaceous, apparently semisucculent. Capitula terminal, solitary or in simple
cymes, radiate. Involucres hemispheric, phyllaries
in 2–3 series, dimorphic, outermost herbaceous,
reflexed, innermost erect, membranaceous to
coriaceous. Receptacles flat. Ray florets pistillate,
corollas yellow. Disc florets bisexual, corollas
416
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
yellow; anther appendages ovate, yellow, glabrous,
with a resin canal in the centre; style arm apices
deltate, densely papillose. Cypselae obcompressed
to shallowly quadrate, edges flattened (wings
wanting), black, sparsely to moderately pubescent.
Pappus of 2 antrorsely barbed awns. One species,
S. buchii Urb., Hispaniola.
1182. Thelesperma Less.
Thelesperma Less., Linnaea 6: 511 (1831); Blake, Proc. Biol.
Soc. Wash. 41: 145–150 (1928), syn.; Shinners, Field Lab. 18:
14–24 (1950), part. rev.
Perennial herbs or weak shrubs. Leaves opposite,
blades linear or ovate to trullate in outline, 1–2pinnatifid, segments linear. Capitula terminal, solitary on long peduncles or in open paniculiform
cymes, radiate or discoid. Involucres campanulate
to hemispheric, phyllaries dimorphic in 2 series,
outer herbaceous, free, linear, inner fused in basal
half, membranaceous. Receptacles convex to conic.
Ray florets neuter, corollas yellow or sometimes
bicoloured with distal half yellow and basal half
brown or deep purple. Disc florets bisexual, corollas yellow, in discoid capitula outermost disc corollas zygomorphic, with one or two lobes shorter
than other 3; anther appendages ovate, glabrous,
resin canals prominent or wanting; style arm apices
deltate to aristate, appendages aristate. Cypselae
terete to slightly obpyramidal, quadrate, strongly
incurved, brown, tuberculate, glabrescent. Pappus
of 2–3 retrorsely barbed awns or absent. x = 9, 10,
11, 12. Approximately 15 species, south-western
USA, north-eastern Mexico, Argentina, southern
Brazil, Uruguay.
XX.3. Subtribe Pinillosiinae H. Rob. (1978).
Prostrate perennial or small rosette herbs, sometimes forming dense mats and appearing caespitose. Leaves opposite or alternate, blades sometimes semisucculent. Capitula axillary, solitary, radiate or disciform. Involucres turbinate to hemispheric, phyllaries either 4 in 1 series, subequal,
free, herbaceous or more than 4 in 3 series and
gradate, shortly fused at base, outermost filiform,
herbaceous, innermost lanceolate to oval, membranaceous. Receptacles flat to convex, epaleate.
Ray florets with or without corollas, 2–10, pistillate,
corollas when present yellow. Disc florets 2, or 9–11,
bisexual, fertile or functionally staminate, corollas
tetramerous or pentamerous, lobes long, lanceolate and spreading or very short and erect; an-
thers hyaline to white or black, sometimes broadly
quadrate or oval, appendages present or wanting
and reduced to a tuft of trichomes; style arm apices
broadly deltoid or subulate, sometimes broadly
expanded and densely papillose, sometimes appendages tapered, penicillate. Cypselae obconic,
terete, sometimes triquetrous, essentially glabrous.
Pappus of either 4 short, distally retrorsely barbed
awns or 4 scales which grow after anthesis, sometimes absent.
Key to the Genera
1. Phyllaries 4, subequal; prostrate herbs, sometimes
forming mats or growing in between short vegetation;
capitula with 4 florets
2
– Phyllaries 6–9, dimorphic; plants forming small
rosettes, caespitose with age; capitula with more than
4 florets
1185. Tetraperone
2. Capitula radiate
1183. Koehneola
– Capitula disciform, outermost florets without corollas
1184. Pinillosia
Genera of Pinillosiinae
1183. Koehneola Urb.
Koehneola Urb., Symb. Antill. 2: 463 (1901); Liogier, Fl.
Cuba 5: 184–185 (1962), rev.
Prostrate perennial herbs. Leaves opposite, blades
cordate to deltate. Capitula radiate. Involucres
turbinate to campanulate, phyllaries 4 in 1 series,
lanceolate to elliptic. Receptacles flat. Ray florets
2, pistillate, corolla apices deeply bilobed. Disc
florets 2, bisexual, corollas yellow (?), tetramerous,
throat short, abruptly narrowed into tube, as
long as lobes, lobes spreading, lanceolate; anther
appendages black, glabrous; style arm apices subulate, with a small papillose, tapered appendage.
Cypselae obpyramidal to obconic, triquetrous,
brownish black, densely pubescent. Pappus absent.
One species, K. repens Urb., Cuba.
1184. Pinillosia Ossa ex DC.
Pinillosia Ossa ex DC., Prodr. 5: 528 (1836); Liogier, Fl.
Cuba 5: 184 (1962), rev.
Prostrate perennial herbs, sometimes forming
mats. Leaves opposite, blades cordate to deltate,
sometimes reniform or elliptic. Capitula disciform. Involucres turbinate to campanulate,
phyllaries 4 in 1 series, obtuse, somewhat connate
at base. Receptacles flat. Florets 4, outermost 2
without corollas, pistillate, inner 2 functionally
Compositae
staminate, corollas white or greenish yellow,
pentamerous, rarely hexamerous, throat short,
as long as lobes, abruptly narrowed into tube,
lobes spreading, lanceolate; anthers broadly
quadrate to oval, appendages brown, reduced
to a tuft of cells, glabrous; style arms of inner
florets broadly deltoid, very conspicuous, styles
of ecorollate florets slender, subulate. Cypselae
obconic with mostly 4 retrorsely barbed awns
when young, which develop into short knobs or
protuberances in fully developed cypselae, brown,
slightly tuberculate. Pappus of 4 very short, distally
retrorsely barbed awns, the awns soon absorbed
by developing cypsela. One species, P. berteri Urb.,
Cuba, Hispaniola.
1185. Tetraperone Urb.
Tetraperone Urb., Symb. Antill. 2: 462 (1901); Liogier, Fl.
Cuba 5: 184 (1962), rev.
Small rosulate herbs with rhizomes, becoming
caespitose with age. Leaves alternate, blades elliptic
to reniform, semisucculent. Capitula disciform. Involucre campanulate to hemispheric, phyllaries in
3 series, gradate, outermost filiform, herbaceous,
innermost lanceolate to oval, membranaceous.
Receptacles convex. Outer florets without corollas,
approximately 8–10, pistillate. Disc florets 9–11,
functionally staminate, corollas white, greenish
white, tetramerous, lobes short; anther appendages
wanting; style arm apices subulate. Cypselae terete,
tuberculate, brownish black, glabrous. Pappus of 4
triangular, thickened stramineous scales, growing
after anthesis. One species, T. bellioides Urb.,
Cuba.
Genus Unassigned to Subtribe
1186. Staurochlamys Baker
Staurochlamys Baker, Hooker’s Icon. Pl. t. 1825 (1889).
Annual herbs. Leaves opposite, blades entire,
lanceolate. Capitula in terminal, open paniculiform cymes, radiate. Involucres hemispheric,
globose, phyllaries 8 in 3 series, phyllaries of first
series 2, broadly ovate, flat, of second series 2,
erect, round, herbaceous, opposite, striate, of third
series enclosed by outer phyllaries, 4, scarious,
ovate, each subtending a ray floret. Receptacles
conic, paleate. Ray florets 4, pistillate, corollas
yellow, apices deeply trilobed. Disc florets bisexual,
corollas yellow, lobes conspicuously rimmed by
orange-red resin canals, throat extremely short,
417
lobes long, spreading; anthers black, glandular;
style arms short, apices acute, papillose. Cypselae
terete, slightly concave, with a brownish, glandular
epidermis, black underneath, glabrous. Pappus absent. One species, S. burchellii Baker, north-central
Brazil.
XXI. Tribe Neurolaeneae Rydb.
(1927).
J.L. Panero
Annual or perennial herbs sometimes forming
rosettes, shrubs, rarely trees. Leaves alternate or
opposite, rarely whorled, blades various, most
commonly linear, ovate or trullate. Capitula
terminal or axillary, solitary on long peduncles or
more commonly in paniculiform or corymbiform
cymes, discoid or radiate. Involucres cylindric,
campanulate or hemispheric, 4 or 6 to many in
1–8 series, subequal to gradate, rarely dimorphic.
Receptacles flat to conic, usually paleate, pales
chartaceous or sometimes indurate and wrapping
tightly around cypselae. Ray florets pistillate,
limbs conspicuous or sometimes barely protruding beyond paleae, rarely corolla tubular. Disc
florets bisexual or sometimes functionally staminate, (4–)5(–6)-lobed, corollas with conspicuous
reddish or orange resin canals; anthers ecalcarate,
without tails, appendages deltate to ovate, rarely
broadly ovate, abaxially glandular, endothecium
of 1–3 polar bridges; styles with two vascular
strands with parallel stigmatic areas, usually
not confluent at apex, apices acute to acuminate
with a short tuft of papillae. Cypselae obconic to
obpyramidal, sometimes obcompressed, rarely
quadrate, walls carbonized, glabrous to densely
pubescent, sometimes glandular. Pappus present
or absent, persistent, of multiple bristles or scales
of subequal or uneven length, scales sometimes
fused at the base forming a cup, rarely a shallow
crown with 4 minute awns, sometimes growing
after anthesis. x = 11, 19.
The tribe contains five genera and approximately 150 species, with most species in tropical
areas of Mexico and South America, Heptanthus
endemic to Cuba, and a few species of Enydra endemic to the Old World tropics.
Molecular studies of Heliantheae and relatives
(Panero et al. 2001c; Panero and Funk 2002) support recognition of the group as a tribe of the He-
418
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
liantheae alliance sister to Bahieae, Chaenactideae
and Tageteae. Neurolaeneae include only five genera with Neurolaena sister to Greenmaniella, and
these two sister to Calea. The genus Enydra of Enydrinae is sister to the latter three genera. Preliminary molecular studies of the genus Heptanthus
(Panero et al., unpubl. data) support, albeit weakly,
the inclusion of Heptanthus in Neurolaeneae. The
genus Tetraperone, the second member of Heptanthinae (Robinson 1981), is treated as part of Coreopsideae. The genera Tetrachyron, Staurochlamys
and Unxia, included in Neurolaeninae by Robinson (1981), are treated under Heliantheae, Coreopsideae and Millerieae respectively. The limits of the
tribe are preliminary, and will probably change as
future molecular phylogenetic studies of members
of the Heliantheae alliance include broader sampling and more characters.
hemispherical, phyllaries 4–6 in 2–3 series, the outermost larger, herbaceous, striate. Receptacles convex to minutely conical, paleate, pales wrapping
tightly around florets. Ray florets pistillate, corollas white, yellow, rarely brownish, limbs minute or
wanting, trilobed. Disc florets bisexual or functionally staminate, corollas pentamerous, white, yellow or pale brown, corollas barely protruding beyond paleae, with a few large, glandular trichomes;
style arm apices acute. Cypselae obcompressed to
subterete, shallowly stipitate, black, essentially glabrous. Pappus absent.
Key to the Subtribes
Characters of the subtribe. Ten species, pantropical.
1. Pales wrapping tightly around cypselae; limbs of ray
floret very reduced or wanting and barely protruding
beyond paleae or phyllaries; capitula solitary, axillary
and sessile to subsessile; cypselae epappose; stems fistulose, rooting at nodes; plants of aquatic or marshy
areas
1187. Enydrinae (p. 418)
– Pales never wrapping around cypselae, sometimes
lacking; limbs of ray florets, when present, conspicuous; capitula mostly in paniculiform or corymbiform
cymes, if solitary and axillary, then capitula on long
peduncles; cypselae with a pappus of multiple bristles
or scales, rarely wanting or reduced to a shallow
crown; stems not fistulose nor plants rooting at nodes;
plants of various habitats, rarely aquatic or of marshy
areas
2
2. Plants forming a small rosette; disc florets functionally
staminate, style arms densely covered with broadly
acute to round papillae; capitula axillary on long peduncles; endothecium of numerous radial and polar,
minute thickenings
1188. Heptanthinae (p. 418)
– Plants various, never forming a rosette, disc florets
bisexual; style arms sparsely to moderately papillose,
papillae tapered and confined to distal half and abaxial surface, capitula terminal, solitary or in compound
thyrsoid, corymbiform or paniculiform cymes; endothecium of 1–3 polar thickenings
3. Neurolaeninae (p. 419)
XXI.1. Subtribe Enydrinae H. Rob. (1979).
Perennial herbs, aquatic or of very wet areas, with
fistulose stems, rooting at nodes. Leaves opposite,
blades lanceolate to ovate or obovate, entire to
broadly serrate. Capitula axillary, solitary, sessile,
radiate or disciform. Involucres campanulate to
Only one genus:
1187. Enydra Lour.
Enydra Lour., Fl. Cochinchin. 510 (1790); Lack, Willdenowia
10: 3–12 (1980), part. rev.
XXI.2. Subtribe Heptanthinae H. Rob. (1978).
Perennial herbs, forming rosettes, sometimes persistent on a thickened single caudex. Leaves alternate, blades lanceolate to ovate or cordiform
on long petioles, entire or deeply lobed, sometimes trilobed. Capitula axillary, solitary on long
peduncles, radiate. Involucres campanulate, phyllaries in 1–2 series, subequal, herbaceous to chartaceous. Receptacles convex, epaleate. Ray florets
pistillate, corollas yellow or creamy yellow, limbs
trilobed. Disc florets functionally staminate, (4–
)5(–6)-lobed, corollas yellow with conspicuous reddish resin canals; disc floret style arms densely papillose, glandular (?), style of ray florets with divided stigmatic surfaces, tapered, essentially without papillae. Ray cypselae obconical, terete, black
or dark brown, with a few glandular trichomes, otherwise glabrous, rugose, especially on basal half.
Pappus of several, subequal, ciliate scales, shorter
than disc corollas, growing after anthesis.
Only one genus:
1188. Heptanthus Griseb.
Heptanthus Griseb., Cat. Pl. Cub. 148 (1866); Liogier, Fl.
Cuba 5: 187–189 (1962), reg. rev.
Characters of the subtribe. Seven species, Cuba.
Compositae
419
XXI.3. Subtribe Neurolaeninae (Rydb.)
Stuessy, B.L. Turner & A.M. Powell (1977);
Robinson, Smithsonian Contrib. Bot. 51:
58–60 (1981), rev.
Annual or perennial herbs, shrubs or small trees.
Leaves alternate or opposite, sometimes whorled.
Capitula in terminal, paniculiform to corymbiform
cymes, discoid or radiate. Involucres cylindrical to
hemispherical, phyllaries in 2–8 series, subequal
to gradate, rarely dimorphic. Receptacles flat to
conical, paleate, rarely epaleate. Ray florets pistillate. Disc florets bisexual, corollas with prominent
resin canals along veins of throat and lobes; anthers ecalcarate, without tails, appendages deltate
or ovate, with glandular trichomes on abaxial surface, sometimes glabrous, endothecium of 1–3 polar thickenings; style arms with parallel stigmatic
areas, not confluent, acute to acuminate, apices in
ray florets tapered, resin canals outside the vascular
traces. Cypselae obconical to obpyramidal, rarely
quadrate or urceolate in outline. Pappus persistent,
of numerous bristles or scales of unequal or equal
size, rarely a shallow crown. x = 11, 19.
Key to the Genera
1. Leaves alternate, blades ovate, trullate, or weakly
trilobed, never linear
2
– Leaves opposite, if alternate, then blades linear
1189. Calea
2. Cypselae with a pappus reduced to a shallow crown
with 4 minute awns at angles of cypsela
1190. Greenmaniella
– Cypelae with a pappus of multiple bristles
1191. Neurolaena
Genera of Neurolaeninae
1189. Calea L.
Fig. 85
Calea L., Sp. Pl., ed. 2, 1179 (1763); Wussow et al., Syst. Bot.
10: 241–267 (1985); Urbatsch et al., Syst. Bot. 11: 501–504
(1986), reg. rev.
Tyleropappus Greenm. (1931).
Brasilia G.M. Barroso (1962).
Perennial herbs sometimes with woody xylopodia
and tuberous roots, shrubs, sometimes scandent to
vine-like or small trees. Leaves opposite, rarely alternate, whorled or basal, blades linear to ovate.
Capitula solitary or in variously thyrsoid, paniculiform or corymbiform cymes, discoid or radiate. Involucres cylindrical to hemispherical, phyllaries in 2–8 series, subequal to gradate, some-
Fig. 85. Compositae-Neurolaeneae. Calea crocinervosa.
A Habit. B Capitulum. C Disc floret. D Ray floret. (Wussow
et al. 1985, with permission from Systematic Botany; artist
J.K. Sullivan)
times dimorphic. Receptacles flat to conical, usually paleate. Ray florets pistillate, corollas yellow,
rarely whitish. Disc florets bisexual, corollas yellow,
less commonly white or purplish; style arm apices
acute. Cypselae obconical or obpyramidal, black or
brown, glabrous to densely pubescent, sometimes
glandular. Pappus of multiple unequal or subequal,
linear or tapered, rarely obovate, ciliate or erose
scales, much shorter than or as long as the disc
corolla, sometimes an erose crown. x = 19. Approximately 125 species, neotropical.
1190. Greenmaniella W.M. Sharp.
Greenmaniella W.M. Sharp, Ann. Missouri Bot. Gard. 22:
141 (1935).
420
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Perennial herbs or weak shrubs. Leaves alternate,
blades ovate. Capitula in terminal, paniculiform
cymes, radiate. Involucres hemispherical, phyllaries in 2–3 series, dimorphic. Receptacles shallowly
to conspicuously conical with age. Ray florets
pistillate, corollas yellow. Disc florets bisexual,
corollas yellow to yellow-orange; style arm apices
acute. Cypselae quadrate, with minute wings on
each angle, black, sometimes with a greenish tinge,
glabrous. Pappus a shallow crown with 4 minute
awns at angles of cypselae, the crown enveloping
corolla tube when young and expanding outwards
as cypsela develops. One species, G. resinosa W.M.
Sharp, north-eastern Mexico.
1191. Neurolaena R. Br.
Neurolaena R. Br., Trans. Linn. Soc. London 12: 120 (1817);
Turner, Pl. Syst. Evol. 140: 119–139. (1982), reg. rev.
Annual or perennial herbs, shrubs or small trees.
Leaves alternate, blades lanceolate to ovate, trullate, sometimes shallowly trilobed. Capitula in terminal, paniculiform, monochasial cymes, discoid
or radiate. Involucres cylindrical to hemispherical,
phyllaries in 2–5 series, subequal. Receptacles convex to conical. Ray florets pistillate, corollas yellow.
Disc florets bisexual, corollas yellow or greenish
white; style arm apices acute to acuminate. Cypselae obconical to urceolate in outline, terete to obscurely obpyramidal with 4–5 shallow ribs, black,
glabrous or pubescent especially on apices, sometimes with glandular trichomes, apices flat, bases
tapered. Pappus of numerous bristles in 1–2 series,
sometimes sigmoid at base. x = 11. Eleven species,
neotropical.
XXII. Tribe Tageteae Cass. (1819).
J.L. Panero
Annual or perennial, sometimes aquatic herbs,
shrubs, rarely small trees, sometimes with C4
photosynthesis(Flareria, Pectis). Leaves alternate
or opposite, blades filiform to broadly ovate,
simple, pinnate or bipinnate, often with marginal
and scattered pellucid glands. Capitula in open,
terminal, paniculiform cymes, sometimes solitary
and scapose, peduncles sometimes fistulose, capitula radiate, rarely discoid. Involucres cylindric,
turbinate, campanulate or hemispheric, sometimes
with a calyculus, phyllaries in 1–5 series, subequal
to gradate, free or variously fused, with pellucid,
oval glands or inflated pustules, sometimes with
linear glands, sometimes eglandular. Receptacles
flat to conical, mostly epaleate, sometimes with
minute scales or bristles, sometimes foveolate. Ray
florets pistillate. Disc florets bisexual, rarely functionally staminate, corollas actinomorphic, rarely
zygomorphic, pentamerous, rarely tetramerous or
hexamerous; anther appendages usually without
glandular trichomes, usually strongly sclerified
rarely without sclerification, endothecium cells
with 1–2, sometimes with 2–4 polar thickenings,
rarely evenly thickened or with radial thickenings;
style arms acute to narrowly tapered, with narrow
to broad (sometimes touching) stigmatic surfaces,
usually not confluent at apices, appendages
wanting or well developed, rarely style arms
short and round and essentially fused except
for distalmost apices and style shaft papillose.
Cypselae cylindric to narrowly fusiform or obpyramidal, rarely compressed and narrowly convex,
walls carbonized, black, rarely brown, striate,
sparsely to densely pubescent, most commonly
with trichomes concentrated on distal and basal
ends, carpopodium usually well developed and
conspicuous. Pappus of few to multiple scales
or bristles or sometimes reduced to a crown of
scales, rarely absent, persistent, rarely caducous,
scales variously dissected into sometimes slightly
flattened bristles, sometimes pappus of alternating
scales and bristles.
The tribe contains 32 genera and approximately
270 species found mostly in the south-western
USA and Mexico. Temperate and tropical South
America contain an important number of species;
Schizotrichia is the only genus of Tageteae endemic
to South America. A few species occur in the
Caribbean, with the monotypic genera Harnackia
and Lescaillea being endemic to Cuba. One species
of Flaveria is native to Australia. Members of
the genus Tagetes (marigolds) are widely used in
horticulture as well as a source of edible orange
pigments. Some species of Tagetes are important
components in the religious rituals of the peoples
of Mesoamerica.
Tribe Tageteae was named by Cassini (1819)
to accommodate a group of plants, previously
placed in his Heliantheae. Strother (1977) provided
the first modern account of the tribe. He viewed
Tageteae as having two main phyletic lines, subtribe Pectidinae containing the distinctive genus
Pectis, and subtribe Tagetinae containing the other
15 genera he recognized. Robinson (1981) viewed
Tageteae as another subtribe of Heliantheae, and
Compositae
placed Tagetinae in the synonymy of Pectidinae.
A similar interpretation of Pectidinae was provided
by Karis and Ryding (1994a). Molecular studies of
Heliantheae and related tribes (Baldwin et al. 2002;
Panero and Funk 2002) support the recognition of
Pectidinae at the tribal level as tribe Tageteae. In
addition, Baldwin’s studies support, albeit weakly,
a broader circumscription of Tageteae which
‘dilutes’ the convenient original circumscription of
the tribe including only those taxa with glandular
leaves and/or phyllaries. My decision to include
the eglandular taxa in Tageteae follows Baldwin
et al. (2002) and morphological considerations.
The eglandular genera of Tagetae, for the most
part, have glabrous, strongly sclerified anther
appendages, striate cypselae with well-developed
carpopodia, and a tendency to have cypselae
with a pappus of bristles or dissected scales.
These morphological characteristics are shared
by most members of the traditional Tageteae
(herein subtribe Pectidinae), and are unusual
among the epaleate tribes with carbonized cypselae. The traditional view of two main phyletic
lines within tribe Tageteae-Pectidinae has been
recently challenged by results from molecular
studies (Loockerman et al. 2003). These data
support the monophyly of the subtribe and the
dismemberment of Dyssodia into seven genera,
as proposed by Strother (1986). The endemic
Cuban genera Harnackia and Lescaillea are sister
to Boeberastrum and, along with Chrsyactinia,
they represent the basal lineages of the tribe. The
other genera of the subtribe are placed in two
clades, each with an equivalent number of genera.
The findings of Loockerman et al. (2003) support
the redefinition of the genus Porophyllum, by
removing Porophyllum tridentatum and related
species from Baja California and placing these
in the new genus Bajacalia. Poropyllum is sister
to Pectis and collectively to Nicolletia, Bajacalia,
Urbinella and Leucactinia. The other clade contains Dyssodia and segregates, sister to Strotheria
and Gymnolaena and collectively sister to the
clade containing Tagetes and Hydropectis. Their
findings also support the placement of Vilobia and
Adenopappus in Tagetes.
Key to the Subtribes
and Genera Unassigned to Subtribe
1. Capitula with 1(2) florets and fused phyllaries
2. Coulterellinae (p. 422)
421
– Capitula normally with 3 or more florets, if fewer than
3, then phyllaries free
2
2. Receptacles paleate
6. Varillinae (p. 430)
– Receptacles epaleate , sometimes with minute scales
or bristles
3
3. Leaves and/or phyllaries with pellucid glands or pustules, aromatic
5. Pectidinae (p. 423)
– Leaves and phyllaries without glands
4
4. Leaves alternate, filiform to acicular, succulent
5
– Leaves opposite, if alternate towards the inflorescences, then leaves not succulent
6
5. Receptacles setose
1. Clappiinae (p. 421)
– Receptacles not setose
1223. Pseudoclappia
6. Leaves on stem and inflorescences alternate
1222. Oxypappus
– Leaves opposite throughout
7
7. Plants stoloniferous, rooting at the nodes, forming
mats; phyllaries succulent; plants of brackish, coastal
environments
4. Jaumeinae (p. 423)
– Plants not stoloniferous, nor forming mats nor rooting
at the nodes; phyllaries not succulent, plants found
mostly in the interior of continents
8
8. Corolla tubes with long, stipitate, glandular trichomes;
anther endothecial cells with 1–2 polar thickenings;
style arms acute to deltate; pappus of disc florets easily
caducous
9
– Corolla tubes glabrous or with a few scattered, short
glandular trichomes; anther endothecial cells with 2–4
polar thickenings; style arms truncate; pappus of disc
florets, when present, persistent
3. Flaveriinae (p. 422)
9. Capitula solitary or in simple cymes; disc cypselae
with a pappus of numerous bristles; plants of western
and southern Mexico
1220. Arnicastrum
– Capitula in open paniculiform cymes; disc cypselae
with a pappus of 6–12 bristles; plants of the SE USA
1221. Jamesianthus
XXII.1. Subtribe Clappiinae H. Rob. (1978).
Compact, perennial herbs or weak shrubs. Leaves
alternate, sessile, blades linear, rarely shallowly
lobed at base, succulent, mucronate, not aromatic.
Capitula terminal, solitary, on long, distally fistulose peduncles, radiate. Involucres hemispheric,
phyllaries in 3 series, subequal. Receptacles
convex, epaleate, setose. Ray florets pistillate,
corollas yellow. Disc florets bisexual, corollas
yellow, slightly zygomorphic, lobes acuminate,
edges thickened, shallowly sclerified, with two
resin canals along edges; anther appendages
deltate, glabrous; style arms tapered, with broad,
parallel stigmatic surfaces, appendages papillose,
not vascularized. Cypselae cylindric, 10-ribbed,
glabrous, slightly stipitate. Pappus of multiple,
flattened, scabrid bristles, stramineous. x = 16.
Only one genus:
422
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
1192. Clappia A. Gray
Clappia A. Gray, Rep. U.S. Mex. Bound.2:93 (1859).
Characters of the subtribe. One species, C. suaedifolia Wooton & Standl., south-western USA, northeastern Mexico.
XXII.2. Subtribe Coulterellinae H. Rob.
(1978).
Brittle shrubs with whitish stems. Leaves opposite, sessile, blades ovate, dentate, concave, succulent, glaucous. Capitula with 1(2) florets, aggregated in terminal, umbelliform cymes, discoid.
Involucres cylindric to rhombic, phyllaries 2–4,
fused, subtended by a bract, ribs of each phyllary thickened after anthesis. Receptacles flat. Florets bisexual, corollas deep golden yellow, lobes
as long as or longer than throat, corollas sometimes zygomorphic with 2 lobes fused; anther appendages ovate, with abaxial glandular trichomes;
style arms acute to deltate, without appendages,
with very broad, parallel stigmatic surfaces confluent at apices, papillae covering completely their
abaxial surfaces. Cypselae cylindric to narrowly
obconic, striate, black, distal end stramineous, indurate, glabrous. Pappus absent.
Only one genus:
1193. Coulterella Vasey & Rose
Coulterella Vasey & Rose, Contr. U.S. Natl Herb. 1: 71, pl. 1
(1890).
Characters of the subtribe. One species, C. capitata Vasey & Rose, southernmost Baja California,
Mexico.
XXII.3. Subtribe Flaveriinae Less. (1832).
Annual or perennial herbs, shrubs, rarely trees.
Leaves opposite, blades linear to elliptic, sometimes semisucculent. Capitula in terminal, open to
congested corymbiform cymes, radiate or discoid.
Involucres cylindric to hemispheric, phyllaries in
1–3 series, normally 5–6, subequal. Receptacles
flat to conic, epaleate, sometimes with setae. Ray
florets pistillate, corollas yellow, apices shallowly
bilobed or trilobed. Disc florets bisexual, corollas yellow; anther appendages linear to ovate or
deltate, glabrous; style arms truncate, with narrow,
parallel stigmatic surfaces, appendages very short
or wanting, not vascularized. Cypselae cylindric to
narrowly obovate, rarely shallowly compressed, 8–
11-ribbed, black, glabrous to sparsely pubescent,
sometimes with a prominent, conic carpopodium.
Pappus absent or of a few to numerous scales or
bristles. x = 18.
Key to the Genera
1. Heads discoid, if radiate, then with only one or two ray
florets
1194. Flaveria
– Heads radiate, with 3 or more ray florets
2
2. Disc cypselae with a pappus of numerous bristles
1195. Haploësthes
– Disc cypselae with a pappus of 4–6 scales alternating
with 4–6 bristles
1196. Sartwellia
Genera of Flaveriinae
1194. Flaveria Juss.
Flaveria Juss., Gen. Pl. 186 (1789); Powell, Ann. Missouri
Bot. Gard. 65: 590–636 (1978), rev.
Annual or perennial herbs, shrubs, rarely small
trees. Leaf blades linear to elliptic. Capitula in terminal or axillary congested corymbiform cymes,
tightly aggregated into glomerules with peripheral
heads having one ray floret, radiate or discoid. Involucres cylindric, phyllaries in 1–2 series, subequal, navicular, herbaceous to corky. Receptacles
flat to convex, paleae sometimes present as setae.
Ray florets 1–2 per capitulum, corollas yellow. Disc
floret anther appendages ovate to deltate. Cypselae shallowly compressed, obovate, 8–11-ribbed,
glabrous, carpopodium wanting, base of cypsela
sometimes invaginated. Pappus absent or rarely either a crown of minute scales, or 2–4 minute, free
squamellae. x = 18. Twenty-two species, America,
one species in Australia; introduced elsewhere.
1195. Haploësthes A. Gray
Haploësthes A. Gray, Mem. Amer. Acad. Arts n.s. 4: 109
(1849); Turner, Wrightia 5: 108–115 (1975), rev.
Multistemmed, annual or perennial herbs or weak
shrubs. Leaf blades linear, semisucculent. Capitula
in terminal, corymbiform cymes, radiate. Involucres campanulate to hemispheric, phyllaries 5–6, in
2–3 series, subequal, broadly ovate to suborbicular.
Receptacles convex to conic, epaleate. Disc floret
anther appendages lanceolate. Cypselae cylindric,
9–15-ribbed, sparsely to densely pubescent. Pappus
of multiple bristles. x = 18. Three species, southwestern USA, north-eastern Mexico.
Compositae
1196. Sartwellia A. Gray
XXII.5. Subtribe Pectidinae Less. (1830).
Sartwellia A. Gray, Smithsonian Contr. Knowl. 3: 122
(1852); Turner, Sida 4: 265–273 (1971), rev.
Subtribe Tagetinae Less. (1831).
Annual herbs. Leaf blades filiform, flat to terete,
semisucculent, glabrous. Capitula in terminal,
congested corymbiform cymes, radiate. Involucres
campanulate to hemispheric, phyllaries 5 in 2 series, subequal, herbaceous, yellowish. Receptacles
conic, epaleate. Ray floret apices shallowly bilobed
or trilobed. Disc floret anther appendages narrowly ovate to linear. Cypselae narrowly elliptic,
9–11-ribbed, glabrous, with a conic carpopodium.
Pappus of (4–)5(–6) erose, isodiametric scales
alternating with (4–)5(–6) capillary bristles,
sometimes united into a shallow crown, rarely
united to corolla and deciduous with it, ray
florets sometimes with a pappus of erose, minute,
obovate scales, rarely pappus absent. x = 18.
Four species, south-western USA, north-eastern
Mexico.
XXII.4. Subtribe Jaumeinae Benth. & Hook. f.
(1873).
Perennial herbs or prostrate shrubs of estuarine
sandy areas or salty mud flats. Leaves opposite,
blades linear to terete, succulent. Capitula terminal, appearing axillary, solitary, radiate or discoid.
Involucres cylindrical to campanulate, rarely
hemispheric, phyllaries in 3–4 series, gradate,
herbaceous, succulent. Receptacles conic, epaleate.
Ray florets pistillate, corolla apices shallowly
trilobed. Disc florets bisexual, corollas yellow, glabrous, edge of lobes thickened; anther appendages
narrowly ovate/deltate, glabrous; style arms with
parallel, broad stigmatic surfaces, apices deltate,
appendages short, papillose, not vascularized.
Cypselae cylindric, conspicuously striate, abruptly
narrowed into a short cylindric carpopodium,
black to brown, glabrous. Pappus persistent,
of multiple awns, stramineous, or absent.
x = 19.
Only one genus:
1197. Jaumea Pers.
Jaumea Pers., Syn. Pl. 2: 397 (1807).
Characters of the subtribe. Two species, coastal
western USA, north-western Mexico, coastal, central Argentina, Uruguay.
423
Annual or perennial herbs, sometimes shrubs,
rarely vines, sometimes aquatic herbs, aromatic.
Leaves opposite or alternate, entire or dissected,
with pustules or pellucid glands, either embedded
in the leaf, marginal or terminal on the tips of
lobes, sometimes with conspicuous setose bristles
at ends of lobes or scattered along margins of
leaves, sometimes bristles restricted to basal
portions of leaves. Capitula terminal, solitary or
in open paniculiform cymes, rarely congested
corymbiform cymes, radiate or discoid. Involucres
cylindric, turbinate, campanulate or hemispheric,
phyllaries normally in 1–5 series, sometimes with
a calyculus, free, partially or completely fused,
usually with oval or linear glands. Receptacles
flat to conic, epaleate, sometimes setose, scaly or
with minute projections, sometimes foveolate. Ray
florets pistillate. Disc florets bisexual, rarely functionally staminate, corollas sparsely to moderately
covered with glandular trichomes, actinomorphic
or zygomorphic, sometimes with one lobe or
two either smaller or longer than rest; anther
appendages ovate to lanceolate or very reduced,
without glandular trichomes, normally densely
sclerified; style arms acute to narrowly tapered,
with narrow to broad, parallel stigmatic surfaces
never confluent at apices, appendages wanting to
well developed and sometimes as long as length
of stigmatic arms, acute to deltate, sometimes
cylindric and densely papillose, rarely arms
very reduced, knob-like and style shaft densely
papillose (Pectis). Cypselae cylindric to narrowly
fusiform or obpyramidal, rarely compressed and
narrowly biconvex, black or brown, sparsely to
densely pubescent, with well-developed carpopodia. Pappus of a few to many scales or bristles,
scales normally dissected into many bristles,
bristles barbellulate, stramineous, rarely purple or
red, sometimes scales alternating with bristles.
Key to the Genera
1. Vines or scandent shrubs; genera endemic to Cuba 2
– Annual or perennial herbs, erect or wiry shrubs, never
scandent; genera endemic to America, some species in
Cuba
3
2. Leaves reduced to minute scales, eglandular; capitula
discoid
1209. Lescaillea
– Leaves twice trilobed, lobes glandular; capitula radiate
1207. Harnackia
424
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
3. Styles with minute, erect, rounded, style arms, papillae
densely covering distal 1/2–2/3 of style 1212. Pectis
– Styles with spreading or arcuate style arms, papillae
rarely found below style bifurcation point
4
4. Capitula with 4 or 5 disc florets
1215. Strotheria
– Capitula with more than 5 disc florets
5
5. Ray corollas white or pink, rarely white with a yellow
base
6
– Ray corollas yellow, greenish-yellow, orange, red, if
white, then phyllaries connate
9
6. Plants perennial, rhizomatous; leaves tightly clustered
and forming a loose, rosette-like structure; capitula
solitary, on long peduncles, scapose; plants of northeastern Mexico
1210. Leucactinia
– Plants annuals, leaves evenly spaced and never
forming rosette-like structures; capitula in open
paniculiform cymes, rarely solitary, plants of the
south-western USA and north-western Mexico
7
7. Pappus of numerous bristles, sometimes fused at base
1217. Thymophylla
– Pappus of entire or dissected scales
8
8. Style arms long, acuminate, appendages well developed, nearly as long as length of stigmatic surface,
cylindric; pappus of dissected scales nearly as long as
disc corollas
1211. Nicolletia
– Style arms short, appendages very short, acute to
deltate; pappus of 5–6 obovate scales, much shorter
than disc corolla tube
1218. Urbinella
9. Plants aquatic, stems fistulose
1208. Hydropectis
– Plants not aquatic, if in marshy areas, then stems not
fistulose, sometimes along rivers and stems shallowly
fistulose, then capitula with prominent, orange ray
corollas
10
10. Receptacles with scales or setae, scales or setae as long
as or longer than the cypselae
1203. Comaclinium
– Receptacles without scales or setae or if setose, then
setae minute and much shorter than cypselae
11
11. Phyllaries free
12
– Phyllaries or innermost series of phyllaries partially
to nearly completely connate
16
12. Calyculus of 5–8 bracts, herbaceous not scarious, normally narrower than phyllaries, reflexed
13
– Calyculus absent or reduced to 1 bract, sometimes
phyllaries in several series with outermost ending in
an inflated pustule
14
13. Annual or perennial herbs; plants of North America
1205. Dyssodia
– Shrubs, plants of Andean Peru
1214. Schizotrichia
14. Annuals
1200. Boeberastrum
– Shrubs
15
15. Phyllaries in 1–3 series, outermost phyllaries ending
in an inflated pustule; plants of Baja California, Mexico
1199. Bajacalia
– Phyllaries in 1 series, phyllaries with a pellucid gland,
gland not inflated nor at the very tips of the phyllaries;
plants of Mexico and the south-western USA, not in
Baja California
1202. Chrysactinia
16. Phyllaries in 4–5 series (or calyculus of several gradate,
acute bracts), innermost series fused at base
1201. Boeberoides
– Phyllaries in one series, variously fused, calyculus, if
present, of bracts or bracteoles with setose apices 17
17. Calyculus of 5 to many setose bracts, sometimes as
long as phyllaries
18
– Involucres without a calyculus, sometimes with 1–3
bracts
19
18. Leaves linear; style arm apices deltate with a narrow,
cylindric appendage
13. Dysodiopsis
– Leaves lanceolate or compound; style arm apices long,
acuminate or subulate, if deltate, then without cylindric appendages
1198. Adenophyllum
19. Capitula discoid
20
– Capitula radiate
21
20. Pappus of multiple bristles
1213. Porophyllum
– Pappus of 5 scales alternating with 5 bristles
1217. Thymophylla
21. Capitula solitary, on peduncles held above foliage, if
not above foliage, then herbage densely pubescent,
lanate; leaves opposite below, alternate above; capitula
with 1–3 bracts
1217. Thymophylla
– Capitula in open paniculiform or corymbiform cymes,
rarely solitary, herbage never densely pubescent so
as to appear lanate, leaves opposite; capitula without
bracts
22
22. Pappus of scales dissected into multiple bristles
1206. Gymnolaena
– Pappus of a few scales, apices acute to acuminate, never
dissected into multiple bristles
1216. Tagetes
Genera of Pectidinae
1198. Adenophyllum Pers.
Adenophyllum Pers., Syn. Pl. 2: 458 (1807); Strother, Sida 11:
371–378 (1986), rev.
Annual or perennial herbs, shrubs. Leaves opposite, sometimes alternate distally, blades pinnatifid, deltate to lanceolate-ovate in outline with ovate,
obovate, linear or filiform lobes, teeth usually setose or bristly, glands marginal, on bases of lobes
and subterminal in lobe tips. Capitula terminal,
solitary, or in open paniculiform cymes, discoid or
radiate. Involucres cylindric-campanulate to hemispheric, phyllaries in 1–2 series, weakly connate
1/3–2/3 of their length. Receptacles flat to shallowly
convex, setose. Ray floret corollas yellow, orange,
vermillion or bright scarlet. Disc floret corollas
yellow to orange, sometimes zygomorphic, lobes
lanceolate to subulate; anther appendages lanceolate to ovate; style arms either narrowly lanceolate, tapered with a prominent, partly vascularized,
papillose, cylindric appendage, or acute to deltate
with a reduced appendage. Cypselae linear to narrowly obpyramidal, dark brown, glabrous to moderately pubescent. Pappus of multiple squamellae,
aristate or dissected into multiple bristles. x = 7,
13. Approximately 10 species, USA, Mexico, Central
America, Cuba.
Compositae
1199. Bajacalia Loockerman, B.L. Turner & R.K.
Jansen
Bajacalia Loockerman, B.L. Turner & R.K. Jansen, Syst.
Bot. 28: 204 (2003).
Shrubs. Leaves alternate, blades broadly acicular
to linear, sometimes apices tridentate, in the
shape of a T, semisucculent or succulent, each
lobe terminated by a gland. Capitula terminal,
solitary or in simple cymes, discoid. Involucres
campanulate, phyllaries in 1–3 series, subequal,
herbaceous, semisucculent, apices of outermost
ending in a gland, innermost with a gland embedded in central region. Receptacles convex. Corollas
light yellow with conspicuous brown or dark
orange resin canals along vascular strands; anther
appendages ovate; style arms acuminate, with vascularized, tapered, papillose appendages, papillae
confined to distal half of style arms. Cypselae terete
to narrowly obconic, black, sparsely to moderately
pubescent. Pappus of multiple, unequal, capillary
bristles, stramineous. x = 15. Three species, Baja
California, Mexico.
1200. Boeberastrum (A. Gray) Rydb.
Boeberastrum (A. Gray) Rydb., N. Amer. Fl. 34: 161 (1915);
Strother, Sida 11: 371–378 (1986), rev.
Dyssodia Cav. sect. Boeberastrum A. Gray (1883).
Annual herbs. Leaves alternate, blades oblong in
outline, pinnatifid, leaflets linear, semisucculent,
with a few, oval glands scattered throughout.
Capitula terminal, solitary or in open paniculiform cymes, on fistulose peduncles, radiate.
Involucres hemispheric, phyllaries free, in 2
series, membranaceous, suborbicular, with a few
scattered, oval glands. Receptacles convex to conic,
minutely scaly. Ray floret corollas yellow. Disc
floret corollas yellow, zygomorphic, one lobe
shorter than other 4, lobe apices thickened, with
a mucronate projection; anther appendages ovate;
style arms acute to deltate, appendages acuminate,
papillose, not vascularized. Cypselae narrowly
obconic, black, sparsely pubescent, pubescence
denser on the base. Pappus of multiple scales each
dissected into multiple bristles. x = 7. Two species,
Baja California, Mexico.
1201. Boeberoides (DC.) Strother
Boeberoides (DC.) Strother, Sida 11: 373 (1986); Strother,
Sida 11: 371–378 (1986), rev.
Dyssodia Cav. sect. Boeberoides DC. (1836).
425
Stout, perennial herbs or weak shrubs. Leaves opposite, blades shallowly trullate to elliptic, or obovate, minutely serrate, with numerous small glands,
scattered throughout. Capitula terminal, solitary
or a few in decussate, long-pedunculate paniculiform cymes, radiate. Involucres broadly cylindric
to campanulate, bracts of calyculus free, phyllaries
connate, in 4–5 series, last series connate from base
to mid section, without glands. Receptacles convex. Ray floret corollas orange to vermillion. Disc
floret corollas yellow, zygomorphic with two lobes
appearing longer than others because of deeper
sinuses; anther appendages lanceolate, glabrous;
style arms long, acuminate, appendages acuminate,
papillose, not vascularized, appendages approximately 1/4–1/5 the length of the stigmatic surfaces.
Cypselae obpyramidal, sparsely pubescent. Pappus of multiple scales each dissected into multiple
bristles. One species, B. grandiflora (DC.) Strother,
Mexico.
1202. Chrysactinia A. Gray
Chrysactinia A. Gray, Mem. Amer. Acad. Arts n.s. 4: 93
(1849); Strother, Madroño 24: 129–139 (1977), rev.
Compact, erect or weak trailing shrubs, sometimes
hanging from crevices in rocky canyons. Leaves alternate, simple or compound, blades simple, acicular to linear or compound and ovate in outline, 1-pinnate, segments filiform to linear, sometimes cuneate. Capitula terminal, solitary, on long
peduncles, radiate. Involucres campanulate, phyllaries in 1 series, subequal, with a single gland
on central, apical region. Receptacles flat to convex. Ray floret corollas lemon yellow to golden
yellow, drying greenish or cherry red. Disc floret corollas yellow; anther appendages narrowly
ovate; style arms acuminate, with narrow, parallel stigmatic surfaces, appendages acute. Cypselae cylindric, black, glabrous except for distalmost
end, carpopodium broad, conic. Pappus of multiple
bristles, white to stramineous. x = 15. Six species,
south-western USA, northern Mexico.
1203. Comaclinium Scheidw. & Planch.
Comaclinium Scheidw. & Planch., Fl. Serres Jard. Eur. 8: 19,
t. 756 (1852); Strother, Sida 11: 371–378 (1986), rev.
Perennial herbs. Leaves opposite, alternate along
peduncles, blades simple, rarely trifoliolate, lanceolate, glands scattered between secondary veins and
in a single row along edge of leaf. Capitula solitary, on long peduncles, radiate. Involucres cam-
426
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
panulate, phyllaries in 2 series, subequal, herbaceous with pellucid glands. Receptacles conic, scaly
with 3–4 caducous scales surrounding each cypsela,
scales as long as or longer than cypselae. Ray floret corollas orange. Disc floret corollas yellow to
orange, lobes lanceolate to linear, subulate with
oval, slightly expanded apices; anther appendages
lanceolate, glabrous; style arms acute to deltate, appendages shortly deltate, not vascularized. Cypselae obpyramidal, weakly 4–5-angled, pubescent.
Pappus of approximately 20 scales in 2 series, the
outer shorter with each scale dissected into 5–10
bristles. x = 13. One species, C. montanum (Benth.)
Strother, Mexico to Panama.
1204. Dysodiopsis (A. Gray) Rydb.
Dysodiopsis (A. Gray) Rydb., N. Amer. Fl. 34: 170 (1915);
Strother, Sida 11: 371–378 (1986), rev.
Annual or perennial herbs. Leaves alternate, with
a few, randomly spaced, filiform or setaceous serrations, setaceous at base, with scattered glands.
Capitula terminal, in open paniculiform cymes, radiate. Involucres cylindric to turbinate, phyllaries
in 2 series, connate with a reflexed calyculus of
a few, deeply serrate bracts, phyllaries and bracts
with several oval glands. Receptacles flat to convex,
minutely setose. Ray floret corollas yellow. Disc floret corollas yellow; anther appendages lanceolate;
style arms acute, appendages acuminate, vascularized only at the base. Cypselae obpyramidal, black,
sparsely pubescent. Pappus of several scales, scale
apices sometimes trifid. x = 13. One species, D.
tagetoides Rydb., south-central USA.
1205. Dyssodia Cav.
Fig. 86
Dyssodia Cav., Descr. Pl. 202 (1801) [1802]; Strother, Univ.
Calif. Publ. Bot. 48: 1–88 (1969), rev.; Strother, Sida 11:
371–378 (1986), rev.
Annual or perennial herbs. Leaves opposite or
alternate above, blades pinnatifid, segments
linear to oblanceolate with scattered, pellucid
glands. Capitula terminal, solitary, or in open
paniculiform cymes, sometimes syncephalous
and the syncephalia resembling single, radiate
capitula, discoid or radiate. Involucres cylindric to
hemispheric, phyllaries dimorphic in 1–3 series,
outer herbaceous (calyculus), inner herbaceous to
membranaceous, with pellucid oval or narrowly
oblong glands. Receptacles flat to conic. Ray floret
corollas yellow to orange. Disc floret corollas
yellow to dull orange, sometimes with purple tips;
Fig. 86. Compositae-Tageteae. Dyssodia papposa. A Habit.
B Flowering capitulum. C Ray corolla. D Disc corolla. E Anthers. F Style arms. G Cypsela. H Pappus scale. (Reproduced
from Flora Novo-Galiciana, vol. 12, with permission of the
University of Michigan Herbarium; see McVaugh 1984)
anther appendages ovate to lanceolate; style arms
acute to deltate, appendages deltate, not vascularized. Cypselae obpyramidal to obconical, black,
glabrous to moderately pubescent. Pappus of 15–20
squamellae of various lengths, each squamella
composed of 5–10 basally connate bristles, stramineous to burgundy. x = 13. Approximately five
species, USA, Mexico, Guatemala.
1206. Gymnolaena (DC.) Rydb.
Gymnolaena (DC.) Rydb., N. Amer. Fl. 34: 160 (1915);
Strother, Sida 3: 110–114 (1967), rev.
Compositae
Shrubs. Leaves opposite, blades lanceolate to
ovate, conspicuously serrate, pinnately nerved,
with small, round glands scattered throughout.
Capitula terminal, solitary or in simple cymes,
radiate. Involucres cylindric, phyllaries uniseriate,
connate to tips, with pellucid glands. Receptacles
convex, epaleate. Ray floret corollas yellow to red.
Disc floret corollas yellow to red, sometimes bicoloured; anther appendages lanceolate; style arms
strongly acuminate, with broad, parallel stigmatic
surfaces, nearly touching at apices, appendages
filiform, papillose, partly vascularized. Cypselae
obpyramidal, moderately pubescent. Pappus of
multiple scales, each comprised of several connate
bristles. x = 13. Three species, Mexico.
1207. Harnackia Urb.
Harnackia Urb., Repert. Spec. Nov. Regni Veg. 21: 72 (1925);
Liogier, Fl. Cuba 5, 224 (1962), rev; Strother, Madroño 24:
129–139 (1977), rev.
Scandent shrubs or vines. Leaves opposite,
trilobed, lobes cuneate to acuminate, apices entire
to trilobed, each lobe with an oval, inflated gland
at its apex, sometimes central lobe without glands.
Capitula in terminal, simple to compound cymes,
each on a long peduncle, radiate. Involucres
turbinate, sometimes with a calyculus of 2 bracts,
phyllaries few, free, in one series. Receptacles
flat to slightly convex. Ray floret corollas yellow.
Disc floret corollas yellow, with 8–9 vascular
traces; anther appendages ovate, glabrous; style
arms tapered, with narrow, parallel stigmatic
surfaces, appendages lanceolate, papillose, not
vascularized. Cypselae cylindric, black-striate,
glabrous with a few apical trichomes. Pappus of
multiple, barbellulate, subequal bristles, weakly
united at base. One species, H. bisecta Urb., eastern
Cuba.
1208. Hydropectis Rydb.
Hydropectis Rydb., N. Amer. Fl. 34: 216 (1914); Turner,
Phytologia 78: 211–213 (1995), rev.
Hydrodyssodia B.L. Turner (1988).
Aquatic annuals. Leaves opposite, blades linear,
entire or pinnate, sometimes with pellucid, oval
glands. Capitula terminal, sometimes appearing
axillary, solitary or in open simple cymes, discoid
or radiate. Involucres turbinate or hemispheric,
phyllaries in 1–2 series, herbaceous to membranaceous, sometimes with linear glands. Receptacles
convex to conic, epaleate. Ray floret corollas
427
brownish, limbs very reduced. Disc floret corollas
tetramerous or pentamerous, yellow or brownish; anther appendages lanceolate; style arms
acute, without appendages. Cypselae cylindric
or shallowly obpyramidal, sparsely pubescent,
pubescence denser on basal area and around
carpopodium. Pappus of multiple scales or bristles, scales sometimes trifid or divided into a few
bristles. x = 9. Three species, western Mexico.
1209. Lescaillea Griseb.
Lescaillea Griseb., Cat. Pl. Cub. 156 (1866); Liogier, Fl. Cuba
5, 224 (1962), rev.; Strother, Madroño 24: 129–139 (1977),
rev.
Scandent shrubs or vines. Leaves opposite, blades
reduced to minute scales. Capitula in terminal, simple cymes, discoid. Involucres turbinate, phyllaries few, in one series, each with an oval gland at
the apex. Receptacles flat. Corollas yellow, with 8–9
vascular traces; anther appendages ovate, glabrous;
style arms tapered, with narrow, parallel stigmatic
surfaces, appendages lanceolate, papillose, not vascularized. Cypselae cylindric, black, glabrous with
a few apical trichomes. Pappus of multiple, barbellulate, subequal bristles, weakly united at base. One
species, L. equisetiformis Griseb., western Cuba.
1210. Leucactinia Rydb.
Leucactinia Rydb., N. Amer. Fl. 34: 180 (1915).
Perennial herbs, rhizomatous, each shoot forming
a small tuft of leaves. Leaves alternate, on short
internodes, blades filiform with a few, scattered,
oval glands. Capitula terminal, solitary on long
peduncles well above foliage, radiate. Involucres
campanulate, phyllaries in 1 series, subequal,
herbaceous with a few, oval glands, edges and
distal apices scarious. Receptacles convex, with
a few golden, caducous trichomes surrounding
bases of cypselae. Ray floret corollas white, apices
shallowly trilobed. Disc floret corollas yellow with
the base turning purplish after anthesis, with
conspicuous brown or dark orange resin canals
along vascular strands; anther appendages narrowly ovate, glabrous; style arms acute to deltate,
appendages short, papillose, not vascularized.
Cypselae subterete to narrowly obconic, black,
sparsely pubescent, pubescence denser on distal
and basalmost areas. Pappus of multiple, slender,
unequal, capillary bristles, stramineous. x = 16.
One species, L. bracteata Rydb., north-eastern
Mexico.
428
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
1211. Nicolletia A. Gray
Nicolletia A. Gray, Rep. Exped. Rocky Mts 315 (1845);
Strother, Sida 7: 369–374 (1978), rev.
Annual herbs. Leaves alternate, blades linear or filiform, trilobed or with several, filiform projections
on each side. Capitula terminal, solitary on essentially all branches of the plant, radiate. Involucres
turbinate to campanulate, phyllaries in 2 series,
free, subequal, with a few smaller, phyllaries/bracts
on peduncle just below capitula. Receptacles flat to
convex. Ray floret corollas white suffused with pink
or deep pink. Disc floret corollas yellow; anther
appendages ovate, strongly sclerified, glabrous;
style arms long-acuminate, with densely papillose,
vascularized, filiform appendages nearly as long
as the stigmatic surfaces. Cypselae cylindric,
moderately to densely pubescent. Pappus of a few
broad scales, edges soon dissected into multiple
bristles, stramineous. x = 10. Three species,
south-western USA, north-western Mexico.
1212. Pectis L.
Fig. 87
Pectis L., Syst. Nat., ed. 10: 1189, 1221, 1376 (1759); Keil,
Madroño 23: 181–191 (1975), part. rev.; Keil, Rhodora 79:
32–78 (1977), part. rev.; Keil, Rhodora 80: 135–146 (1978),
part. rev.
Annual or perennial herbs, sometimes stoloniferous. Leaves opposite, with pairs of setaceous
bristles, blades linear to narrowly ovate, abaxial
surfaces with large glands. Capitula terminal, solitary or in paniculiform cymes, radiate. Involucres
cylindric to campanulate, phyllaries in 1 series,
free, calyculus absent. Receptacles flat to convex.
Ray florets rarely functionally staminate, corollas
yellow or reddish yellow. Disc floret corollas
actinomorphic or zygomorphic with one lobe
shorter and more deeply cut, yellow; anther appendages ovate to round, sometimes reduced and
shallowly obcordate, not sclerified; style arms very
short, knob-like or rounded, sometimes fusiform,
densely papillose, papillae extending well below
bifurcation, appendages in most species reduced
to an aggregation of minute papillae. Cypselae
cylindric to clavate, sometimes obpyramidal,
black or brown, sparsely to moderately pubescent,
sometimes densely pubescent on angles. Pappus
of few to multiple scales or bristles or sometimes
reduced to a crown of scales, stramineous to
burgundy red. x = 12. Approximately 85 species,
tropical and subtropical America.
Fig. 87.
Compositae-Tageteae. Pectis decemcarinata.
A Flowering branch. B Involucrate cluster of capitula.
C Flowering capitulum. D Ray corolla. E Disc floret.
F Anthers. G Style. H Ray cypselae. (Reproduced from
Flora Novo-Galiciana, vol. 12, with permission of the
University of Michigan Herbarium; see McVaugh 1984)
1213. Porophyllum Adans.
Porophyllum Adans., Hist. Acad. Roy. Sci. Mem. Math. Phys.
(Amsterdam) 1750: 377 (1754); Johnson, Univ. Kansas Sci.
Bull. 48: 225–267 (1969), rev.
Annual or perennial herbs or densely branched
shrubs, sometimes weak stoloniferous shrubs.
Leaves opposite or alternate, blades linear to
ovate, with embedded, pellucid glands, herbage
strongly aromatic. Capitula in terminal, open
corymbiform to paniculiform cymes, discoid.
Involucres cylindric to campanulate, phyllaries in
1 series, free, calyculus absent. Receptacles flat or
slightly convex. Corollas actinomorphic to slightly
zygomorphic, green-brown, purple-brown, purple, green, white-yellow or yellow; style arms
long, tapered, appendages cylindric, papillose.
Cypselae fusiform or narrowly cylindric, brown or
black, sparsely to moderately pubescent. Pappus
of numerous bristles. x = 11, 12. Thirty species,
tropical and subtropical America.
Compositae
1214. Schizotrichia Benth. in Benth. & Hook. f.
Schizotrichia Benth. in Benth. & Hook. f., Gen. Pl. 2: 202,
410 (1873).
Shrubs. Leaves opposite, blades narrowly ovate to
ovate or elliptic, entire to deeply serrate, with scattered orange glands. Capitula in terminal, simple
or open paniculiform cymes, radiate. Involucres
campanulate to hemispheric, phyllaries in 3 series,
gradate, outermost reflexed. Receptacles flat to convex, epaleate. Ray floret corollas yellow, apices shallowly trilobed. Disc floret corollas yellow; anther
appendages ovate, glabrous; style arms acuminate
to deltate, with parallel, broad stigmatic surfaces,
appendages short, not vascularized. Cypselae subterete, black to silvery black, moderately pubescent.
Pappus of multiple, soft scales soon divided into
several bristles of unequal length, stramineous.
Two (to five?) species, Peru.
1215. Strotheria B.L. Turner
Strotheria B.L. Turner, Amer. J. Bot. 59: 180 (1972).
Graciela Rzed. (1975).
Low shrubs forming small mounds. Leaves alternate, few nodes below capitula with opposite
leaves, blades linear, navicular, succulent, not glandular. Inflorescences terminal, solitary, subtended
by two bracts, neighbouring capitula tightly
clustered into small congested, corymbiform to
umbelliform cymes, capitula discoid. Involucres
cylindric to narrowly campanulate, phyllaries 4–6,
fused except for acute apices, subequal, with one to
several glands on apices. Receptacles flat. Florets
4 or 5, corollas yellow to greenish yellow, pentalobed or hexalobed; anther appendages ovate,
strongly sclerified, glabrous; style arms acute to
deltate, with narrow, parallel stigmatic surfaces,
appendages short, acuminate, not vascularized.
Cypselae obpyramidal, obscurely 3- or 4-angled,
black, shiny, moderately sericeous. Pappus of
multiple lanceolate, erose, overlapping scales
with a thickened midrib, sometimes protruding as
a short, erose awn. x = 8. One species, S. gypsophila
B.L. Turner, north-eastern Mexico.
1216. Tagetes L.
Tagetes L., Sp. Pl. 2: 887 (1753); Soule, Ph.D. Dissertation,
University of Texas, Austin (1993), rev.
Adenopappus Benth. (1840).
Vilobia Strother (1968).
Annual or perennial herbs, sometimes shrubs.
Leaves opposite, simple or pinnately dissected,
429
blades linear to lanceolate or ovate in outline,
segments sometimes reduced to bristles, glands
scattered throughout blade. Capitula terminal,
solitary or more commonly in open (rarely congested) paniculiform cymes, sometimes on long,
fistulose peduncles, radiate, rarely discoid. Involucres cylindric, fusiform or campanulate, phyllaries
in 1 series, fused except for apices, with 2 or more
rows of glands, calyculus absent. Receptacles flat
to conic. Ray floret corollas white, yellow, orange
or sometimes bicoloured. Disc floret corollas
yellow, orange, purplish or white, sometimes
zygomorphic with one lobe longer than the rest;
anther appendages narrowly lanceolate to ovate;
style arms acute to deltate, appendages short,
acuminate to deltate, sometimes fully vascularized.
Cypselae clavate, linear or narrowly oblanceolate,
sometimes compressed, biconvex or obpyramidal,
black, sparsely pubescent. Pappus of a few, acute,
acuminate or subulate scales or a combination
thereof, sometimes of 1 or 2 very narrow awns and
a few very short scales in between, rarely of scales
subdivided into bristles, rarely absent. x = 11, 12.
Approximately 50 species, tropical and subtropical
America, a few species adventive and widespread
in temperate and tropical regions of the Old World.
1217. Thymophylla Lag.
Thymophylla Lag., Gen. Sp. Pl. 25 (1816); Strother, Sida 11:
371–378 (1986), rev.
Dyssodia Cav. sect. Acyphyllaea DC. (1836).
Dyssodia Cav. sect. Aurantiacae Strother (1969).
Annual or perennial herbs, sometimes persisting
as weak shrubs. Leaves opposite below or alternate
throughout, blades linear or filiform, spathulate,
entire or pinnatisect, segments linear or filiform,
essentially without setae, glands scattered. Capitula
terminal, pedunculate or sessile, solitary or in open
paniculiform cymes, usually radiate, rarely discoid.
Involucres turbinate to hemispheric, phyllaries in
1 series, membranaceous, connate 1/2–2/3 of their
length, glands small, round, scattered, calyculus
mostly of 1–3 bracts. Receptacles flat to convex,
epaleate. Ray floret corollas yellow, golden yellow,
rarely white. Disc floret corollas yellow to yelloworange; anther appendages ovate, glabrous; style
arms acute to deltate, appendages short, not vascularized. Cypselae narrowly obconic, cylindric or
obpyramidal, black. Pappus of multiple scales, aristate or dissected into several bristles, sometimes
obovate scales alternating with bristles. x = 8. Ap-
430
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
proximately 18 species, south-western USA, Mexico and north-western Argentina.
Genera Unassigned to Subtribe
1220. Arnicastrum Greenm.
Arnicastrum Greenm., Proc. Amer. Acad. 39: 115 (1903).
1218. Urbinella Greenm.
Urbinella Greenm., Proc. Amer. Acad. 39: 117 (1903).
Small, dainty annual herbs. Leaves opposite below, alternate distally, blades filiform, with a few,
scattered, oval glands. Capitula terminal, solitary
or in open paniculiform cymes, radiate. Involucres campanulate, phyllaries in 1 series, subequal,
medial area with a few oval glands. Receptacles
convex to conic, epaleate. Ray floret corollas bicoloured, white, base of limb yellow. Disc floret
corollas yellow, with a very narrow tube; anther appendages reduced to a minute deltate tip or wanting; style arms acute to deltate, appendages deltate,
papillose, not vascularized. Cypselae subterete to
terete, black, glabrous with a few trichomes on distal and basalmost areas. Pappus of a few, broadly
obovate, minute scales, hyaline. x = 8. One species,
U. palmeri Greenm., north-western Mexico.
Perennial herbs or weak, trailing subshrubs.
Leaves opposite, blades broadly lanceolate to ovate.
Capitula solitary or few capitula in open, paniculiform cymes, radiate. Involucres hemispheric,
phyllaries in 3 series, subequal to gradate, ovate
with attenuate tips. Receptacles flat to convex,
epaleate. Ray floret corollas yellow. Disc florets
fertile (?), corollas yellow, with long, stipitate
glandular trichomes; style arms acuminate to
deltate, without appendages, with broad, parallel
stigmatic surfaces. Cypselae narrowly obconic,
striate, black, sparsely pubescent. Pappus of multiple barbellulate bristles, fragile, easily caducous,
absent in ray cypselae. Two species, northern and
southern Mexico.
1221. Jamesianthus S.F. Blake & Sherff
Jamesianthus S.F. Blake & Sherff, Publ. Field Mus. Nat.
Hist., Bot. Ser. 22: 399 (1940).
Only one genus:
Perennial herbs. Leaves opposite, blades narrowly
ovate to trullate, acuminate. Capitula terminal,
in open paniculiform cymes, radiate. Involucres
campanulate to hemispheric, phyllaries in 2–3
series, gradate. Receptacles flat to convex, epaleate.
Ray floret corollas yellow, apices shallowly trilobed,
with distinctive, stipitate, glandular trichomes.
Outermost disc florets (those alternating with
ray florets) bisexual, otherwise functionally
staminate, corollas yellow, with long, stipitate
glandular trichomes; anther appendages ovate,
glabrous; style arms long, arcuate, curling, acute
to deltate, with narrow, parallel stigmatic surfaces,
appendages wanting. Cypselae elliptic, black to
brown, sparsely pubescent, the trichome bases
broad and appearing as glands with age, ray cypselae slightly curved inwards, carpopodium broad,
bowl-shaped. Pappus a shallow crown or ring
with 6–12 caducous, slender bristles, ray cypselae
apparently lacking bristles and the ring slightly
more thickened. One species, J. alabamensis S.F.
Blake & Sherff, south-eastern USA.
1219. Varilla A. Gray
1222. Oxypappus Benth.
Varilla A. Gray, Mem. Amer. Acad. Arts n.s. 4: 106 (1849);
Turner, Phytologia 68: 4–13 (1990), rev.
Oxypappus Benth., Bot. Voy. Sulphur 118 (1845).
XXII.6. Subtribe Varillinae B.L. Turner &
A.M. Powell (1978).
Shrubs. Leaves opposite, sometimes alternate along
peduncles, blades lanceolate or linear, sometimes
succulent. Capitula terminal, in corymbiform
cymes or solitary on long peduncles, discoid.
Involucres campanulate to hemispheric, phyllaries in 2–3 series, subequal. Receptacles conic,
paleate. Disc floret corollas yellow, pubescent with
glandular trichomes and dense resinous areas between vascular strands; anther appendages ovate,
glabrous or sometimes with minute glandular
trichomes abaxially; style arms truncate to broadly
acute, short and canaliculate, appendages acute,
with numerous, minute papillae, not vascularized.
Cypselae cylindric, 9–11-ribbed, without carpopodia, black, glabrous to sparsely pubescent. Pappus
mostly absent, sometimes pappus of a few awns.
Characters of the subtribe. x = 18. Two species,
south-western USA, northern Mexico.
Annual or perennial herbs. Leaves opposite below, alternate above, those at the base ovate and
much larger than the linear, cauline blades, herbage
Compositae
with stipitate glands. Capitula terminal, in open
paniculiform cymes, radiate. Involucres campanulate, phyllaries in 1 series, subequal. Receptacles
flat to convex, epaleate. Ray floret corollas yellow, apices shallowly trilobed. Disc floret corollas
yellow; anther appendages ovate, glabrous; style
arms long-acuminate, with an apical short tuft of
papillae, without appendages. Cypselae subterete,
black, glabrous, carpopodium conspicuous, bowlshaped. Pappus of narrowly triangular scales prolonged into barbellate awns, 3 in the ray cypselae, 5
in the disc cypselae. One species, O. scaber Benth.,
western and southern Mexico.
1223. Pseudoclappia Rydb.
Pseudoclappia Rydb., J. Wash. Acad. Sci. 13: 288 (1923).
Low, open shrubs. Leaves alternate, blades terete to
subacicular, succulent. Capitula terminal, solitary
on long peduncles, radiate. Involucres cylindric to
turbinate, phyllaries in 2–3 series, without glands.
Receptacles flat to shallowly convex, epaleate. Ray
floret corollas yellow. Disc floret corollas yellow;
anther appendages lanceolate to narrowly lanceolate; style arms acute to tapered, with parallel,
broad stigmatic surfaces, confluent at very apex,
appendages short, papillose. Cypselae cylindric,
black, moderately pubescent. Pappus of multiple,
barbellulate bristles, white to stramineous. x =
18. Two species, south-western USA, north-eastern
Mexico.
XXIII. Tribe Chaenactideae
B.G. Baldwin (2002).
J.L. Panero
Annual, often ephemeral, or perennial herbs, rarely
alpine subshrubs. Leaves alternate, sometimes subopposite or opposite, in some species the basal
leaves forming a rosette, petiolate or sessile, blades
entire or trilobed to 1–3 times pinnate, oval to obovate or trullate in outline, variously pubescent. Capitula discoid, terminal, subscapiform and solitary
or usually in open corymbiform or paniculiform
cymes, rarely reduced and congested in the centre
of a rosette. Involucre cylindrical, campanulate or
subhemispherical, phyllaries in 1–2 series. Receptacle flat to convex, epaleate. Florets bisexual, cells
not sclerified, peripheral corollas sometimes with
expanded lobes, sparsely to moderately pubescent
with short to long stipitate glandular trichomes,
431
lobes 5, sometimes densely papillose on adaxial
surfaces; anther thecae white to white-pink, rarely
yellow or purple-pink, anther appendages ovate to
round, rarely oval, weakly to strongly carinate, partially to fully sclerified, with or without glandular
trichomes on abaxial surfaces, median endothecial
cells with one polar thickening, stigmatic surfaces
marginal, rarely meeting at the distal end of the
style arm, style arm apices obtuse to rounded, most
commonly narrowly tapered. Cypselae cylindrical,
terete to shallowly convex, usually with a shallow to prominent neck, walls carbonized, glabrous
to densely pubescent but without glandular trichomes. Pappus of 5–10 ovate, narrowly oval to
round, rarely bilobed, erose scales or rarely multiple, erose or setose, linear to obovate scales fused
at the base, deciduous and dispersing as a unit.
Three genera and approximately 29 species of
the western USA and north-western Mexico.
Tribe Chaenactideae was named by Baldwin
et al. (2002) to accommodate most genera of classical Helenieae having a pappus of scales without
thickened bases or midribs and striate cypselae,
and revealed in molecular studies as a distinctive
lineage of the Heliantheae alliance. Chaenactideae
have traditionally been included in a broad concept of Helenieae. Dimeresia has been placed in Inuleae and Senecioneae (Karis and Ryding 1994b).
Robinson (1981) placed Chaenactis and Orochaenactis in his Chaenactidinae whereas Dimeresia was
viewed as an unusual element of obscure affinity
and, therefore, placed in its own subtribe Dimeresiinae. Karis and Ryding (1994b) were also unable
to clarify the affinities of Dimeresia. Recent molecular studies of the helenioid members of the Heliantheae alliance by Baldwin et al. (2002) resulted
in the recognition of tribe Chaenactideae to include only three genera – Chaenactis, Dimeresia
and Orochaenactis. These results have been confirmed by chloroplast DNA data (Panero and Funk
2002). The latter studies also show that the members of the tribe are sister to Bahieae (sensu Baldwin
et al. 2002).
Key to the Genera
1. Capitula with one or two florets
1224. Dimeresia
– Capitula with more than two florets
2
2. Cypselae epappose or, if pappose, then with persistent
and free pappus elements
1225. Chaenactis
– Cypselae with obovate to pandurate pappus elements
fused at the base and collectively deciduous
1226. Orochaenactis
432
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Genera of Chaenactideae
1224. Dimeresia A. Gray
Dimeresia A. Gray, Syn. Fl. N. Am., ed. 2, 1(2): 448 (1886).
Prostrate annuals with taproot with multiple, short,
congested shoots, forming a tiny rosette-like structure subtending the congested inflorescence. First
pair of leaves opposite to subopposite, subsequent
leaves alternate on short internodes, sessile, blades
oval to obovate, those subtending the heads narrowly spathulate, with a few floccose trichomes
at the base, triplinerved. Capitula with only one
or two florets, in terminal, congested cymes. Involucre cylindrical, phyllaries one or two enclosing
each floret. Receptacle flat. Florets bisexual, corollas white, reddish purple on abaxial surface when
old or dried, white on adaxial surface, glabrous to
sparsely pubescent with widely scattered glandular trichomes; anthers whitish with thecal arc in
connective purplish, appendages ovate to round;
style arms with obtuse papillose apices, style bases
without trichomes. Cypselae cylindrical, terete to
shallowly biconvex, glabrous. Pappus of multiple,
setose, linear scales fused at the base, deciduous
and dispersing as a unit. x = 7. One species, D. howellii A. Gray, western USA.
1225. Chaenactis DC.
Fig. 88
Chaenactis DC., Prodr. 5: 659 (1836); Stockwell, Contr.
Dudley Herb. 3: 89–168 (1940), rev.
Annual or perennial herbs, rarely subshrubs.
Leaves alternate, distalmost sometimes subopposite or opposite, sometimes basal producing
a rosette, petiolate, entire, trilobed, pinnatifid or
2–3-pinnatifid, blades rarely linear or trilobed,
most commonly oval or trullate in outline,
sometimes densely pubescent. Capitula terminal,
solitary and subscapose or in open corymbiform
or paniculate cymes. Involucres campanulate
to subhemispherical. Receptacle flat to convex.
Florets bisexual, corollas white, creamy white, pink
to purple or yellow, peripheral florets sometimes
with expanded lobes; anthers whitish to pale
pink or light yellow, appendages ovate to round
or rarely narrowly oval; style arms with obtuse
or narrowly tapered papillose apices, style bases
with papillae and some multicellular trichomes.
Cypselae terete, glabrous to densely pubescent,
with a shallow to prominent cyathiform neck.
Pappus absent or more commonly of 5–10 ovate,
narrowly oval to round, rarely bilobed or obovate,
Fig. 88. Compositae-Chaenactideae. Chaenactis douglasii
var. achilleifolia. A Habit. B Involucre. C Floret. (Reproduced from Vascular Plants of the Pacific Northwest, with
permission of University of Washington Press; see Hitchcock et al. 1959; artist John H. Rumely)
Compositae
erose scales, sometimes pappus of peripheral
florets not uniform, with scales facing phyllaries
much shorter. x = 5, 6, 7, 8. Twenty-seven species,
western USA, north-western Mexico.
1226. Orochaenactis Coville
Orochaenactis Coville, Contr. U.S. Natl Herb. 4: 134 (1893).
Annual herbs. Leaves alternate, distalmost subopposite, blades linear, sparsely pubescent and
glandular. Capitula in open paniculiform cymes.
Involucres campanulate. Receptacle flat to convex.
Corollas creamy white, without sclerified cells,
sparsely pubescent with scattered glandular
trichomes, lobes papillose on adaxial surface,
veins of most lobes meeting at lobe apices; anthers
whitish, appendages ovate; style arms with obtuse
to tapered papillose apices, style bases with
trichomes. Cypselae cylindrical, terete, sparsely
pubescent with short and broad twin trichomes.
Pappus of multiple, erose, obovate scales fused at
the base, deciduous and dispersing as a unit. x = 9.
One species, O. thysanocarpha Coville, western
USA.
XXIV. Tribe Bahieae B.G. Baldwin
(2002).
J.L. Panero
Annual, biennial or perennial herbs, rarely
shrubs, some subalpine or dry grassland species
forming caespitose woody caudices, herbage
usually glandular. Leaves alternate or opposite,
sessile to petiolate, blades linear to cordiform,
entire to 1–4-pinnate. Capitula terminal, scapose
to subscapose or in open paniculiform, rarely
congested corymbiform cymes, radiate or discoid,
rarely disciform. Involucres cylindric, turbinate
to hemispheric, phyllaries in 1–4 series, subequal,
sometimes gradate, mostly with abundant glandular trichomes on abaxial surfaces. Receptacles
flat to convex, usually epaleate. Ray florets pistillate. Disc florets bisexual, pentamerous, rarely
tetramerous, corollas mostly actinomorphic, in
a few genera zygomorphic; anther appendages
ovate, rarely cochleate, with or without abaxial
glandular trichomes, endothecium of quadrate
or fusiform cells with 1–4 polar thickenings,
rarely radial (Chamaechaenactis); style bases
sometimes papillose or pubescent, arms with
parallel, narrow or broad and sometimes touching
433
stigmatic surfaces, not confluent at apices, with
or without appendages. Cypselae obpyramidal
to subterete, obscurely to strongly 4–5-sided,
rarely biconvex and obcompressed, usually striate,
walls carbonized. Pappus persistent, of oblong to
obovate, erose scales with or without prominent
midribs, midribs usually protruding as an awn or
bristle, sometimes absent or of multiple bristles.
The tribe contains 20 genera and approximately
83 species found mostly in the south-western USA
to central Mexico, with a few species in temperate
and Andean South America. The monotypic
genus Apostates is endemic to Rapa Island, French
Polynesia whereas Hypericophyllum is found in
tropical Africa.
Tribe Bahieae was named by Baldwin (Baldwin et al. 2002) to accommodate most genera of
classical Helenieae having a pappus of scales with
thickened midribs and striate cypselae, and revealed in molecular studies as a distinctive lineage of the Heliantheae alliance. Molecular studies of Asteraceae (Panero and Funk 2002) show
that Bahieae are a derived lineage sister to Chaenactideae, and collectively sister to Tageteae and
Neurolaeneae. The tribe is characterized by its pappus morphology, epaleate receptacles, a tendency
to have large glandular trichomes on the anther
appendages and corollas, and cuneate or obpyramidal, striate cypselae. Only Loxothysanus and Hymenoppapus lack striate cypselae, and these occupy
a basal position in the tribe, along with the genera Thymopsis, Chamaechaenactis and Bartlettia
(Baldwin et al. 2002). These five genera are sister
to three clades informally recognized by Baldwin
et al. (2002) as the Bahia, Peucephyllum and Chaetymenia clades. The circumscription of Bahieae recognized in this treatment follows mostly results
from molecular studies (Baldwin et al. 2002). The
genera Apostates and Welwitschiella were the only
genera of the classical Helenieae allied to Bahieae
not sampled by Baldwin (Baldwin et al. 2002), and
therefore we lack molecular information as to their
relationships. Apostates shares most of the characteristics of Bahieae, and it is accepted here as
a member of the tribe following Karis (1998). Welwitschiella differs from most members of Bahieae
by having obovate cypselae with a pappus of scales
fused at the base, the cypsela walls are apparently
not carbonized, the cypselae lack striations, the
main 4 ribs of the cypsela are strongly thickened
and well developed even in immature cypselae,
the endothecium is composed of uniformly thickened cells, the involucre is composed of 4–5 se-
434
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
ries of lanceolate phyllaries which are somewhat
indurate at base with herbaceous apices, the disc
florets appear to be functionally staminate, and
the peripheral florets are zygomorphic. The geographical distribution and morphology of Welwitschiella provide support for its placement in
Athroismeae.
Key to the Genera
1. Capitula discoid or disciform
2
– Capitula radiate
17
2. Style bases papillose; style arms with prominent appendages as long as or longer than stigmatic surfaces;
endothecium with radial thickenings
1233. Chamaechaenactis
– Style bases smooth, without papillae or trichomes;
style arms without appendages, if appendiculate, then
appendages much shorter than length of stigmatic surface; endothecium with polar thickenings
3
3. Corollas tetramerous or tetramerous and pentamerous in the same capitulum
4
– Corollas pentamerous
6
4. Pappus of a few scales fused at base and forming a cup;
capitula disciform
1246. Thymopsis
– Pappus of approximately 8–10 free scales; capitula discoid
5
5. Leaves ovate, entire; anthers dark brown to black
1236. Holoschkuhria
– Leaves filiform or linear, 1–3-pinnate; anthers hyaline
to yellow
1245. Schkuhria
6. Leaves opposite
7
– Leaves alternate below inflorescence branches, opposite only on basal nodes
10
7. Leaves distinctly petiolate; capitula in corymbiform
cymes
8
– Leaves sessile to shortly petiolate, capitula solitary or
in open paniculiform cymes
9
8. Leaves ovate to trilobed, triplinerved, densely glandular, abaxial surfaces densely pubescent, sometimes
appearing or drying white; plants from eastern, tropical Mexico
1240. Loxothysanus
– Leaves broadly ovate to elliptical, palmately veined,
sparsely pubescent, not glandular, concolorous; plants
endemic to Rapa Island, French Polynesia
1229. Apostates
9. Pappus of obovate scales with a prominent midrib,
midrib sometimes protruding beyond scale as
a thickened, straight awn, never uncinate; corollas
with a very short throat, approximately 1/5 the
length of the lobes; corolla tubes thickened and
with expanded bases; plants of Mexico and Central
America
1234. Espejoa
– Pappus of capillary bristles or scales with a prominent
midrib extending as a bristle, rarely of bristles alternating with small scales, bristle apices mostly uncinate, corollas with throat as long or longer than lobes;
corolla tubes not thickened, nor with expanded bases;
plants of tropical Africa
1239. Hypericophyllum
10. Anther appendages glabrous
11
– Anther appendages with glandular trichomes on abaxial surfaces
14
11. Style arms acuminate, without appendages; papillae
covering abaxial surfaces of style arms and reaching
below bifurcation point
1241. Palafoxia
– Style arms with strongly acuminate to deltate apices
and sometimes with minute to prominent, tapered
appendages; papillae on abaxial surfaces of style arms
confined mostly to distalmost end
12
12. Corollas of all florets or of only the peripheral florets
zygomorphic
13
– Corollas actinomorphic
1245. Schkuhria
13. Corollas with two lobes longer than other three
1235. Florestina
– Corollas with one or all lobes of unequal length, sometimes with one sinus much longer (more deeply cut)
than others
1237. Hymenothrix
14. Shrubs
1242. Peucephyllum
– Annual or perennial herbs
15
15. Corollas with strongly recurved lobes
1237. Hymenopappus
– Corollas without strongly recurved lobes
16
16. Plants with most leaves confined to the base and
scapose inflorescences; pappus of multiple bristles;
leaves ovate, entire
1244. Psathyrotopsis
– Plants with leaves evenly spaced along stems, inflorescences not scapose; pappus of approximately 8–10
obovate to fusiform scales, leaves filiform or dissected
1245. Schkuhria
17. Ray and disc corollas deep pink or purple
1241. Palafoxia
– Ray corollas white, yellow or yellow-orange, if pink,
then disc corollas not pink
18
18. Anther appendages without glandular trichomes on
abaxial surfaces
19
– Anther appendages with glandular trichomes on abaxial surfaces
25
19. Ray corollas white
1227. Achyropappus
– Ray corollas yellow
20
20. Leaves opposite, sessile
1232. Chaetymenia
– Leaves alternate, sometimes confined to base and inflorescences subscapose
21
21. Cypselae obcompressed, biconvex
1231. Bartlettia
– Cypselae obpyramidal to subterete, 4–5-sided
22
22. Leaves entire, narrowly elliptic to ovate or suborbicular
1243. Platyschkuhria
– Leaves filiform or linear, 1–4-pinnate
23
23. Capitula in congested, corymbiform cymes; pappus of
multiple capillary bristles
1237. Hymenothrix
– Capitula solitary or in open paniculiform cymes; pappus of obovate or fusiform scales sometimes with
midrib extending as a bristle
24
24. Capitula with more than 5 ray florets
1228. Amauriopsis
– Capitula with 1–3, rarely 5 ray florets
1245. Schkuhria
25. Lobes of disc corollas strongly recurved
1237. Hymenopappus
– Lobes of ray corollas spreading or straight, not strongly
recurved
26
26. Capitula with more than 5 ray florets
1230. Bahia
– Capitula with 1–3, rarely 5 ray florets
1245. Schkuhria
Compositae
Genera of Bahieae
1227. Achyropappus Kunth
Achyropappus Kunth in Humb., Bonpl. & Kunth, Nov. Gen.
Sp., folio ed. 4: 202 (1818).
Annual herbs. Leaves opposite, blades deltate to
ovate in outline, once or twice pinnate, segments
linear. Capitula in terminal, open paniculiform
cymes, radiate. Involucres hemispheric, phyllaries
in 1–2 series, subequal. Receptacles convex. Ray
florets pistillate, corollas white. Disc florets bisexual, corollas yellow, throats abruptly narrowed into
tube; anther appendages cochleate, without glandular trichomes; style arms acute to deltate with
a minute aristate appendage, stigmatic surfaces
narrow. Cypselae obpyramidal, 3–5-sided, black,
moderately sericeous, ray cypselae triquetrous.
Pappus of approximately 8 erose scales, acute or
bilobed without thickened midribs. x = 10. Two
species, Mexico.
435
deltate, with narrow stigmatic surfaces. Cypselae
narrowly obpyramidal, quadrate, pale brown,
sparsely sericeous. Pappus of multiple, erose awns.
One species, A. rapae (F. Brown) N.S. Lander, Rapa
Island, French Polynesia.
1230. Bahia Lag.
Fig. 89
Bahia Lag., Gen. Sp. Pl.: 30 (1816); Ellison, Rhodora 66:
65–86, 177–215, 281–311 (1964), rev.
Annual, biennial or perennial herbs, sometimes
rhizomatous. Leaves opposite, blades linear to
ovate, entire to 1–3-pinnate, segments filiform
to linear. Capitula terminal, solitary or in open
paniculiform cymes, radiate. Involucres campanulate to hemispherical, phyllaries in 2–3 series,
subequal, herbaceous. Receptacles flat to convex.
Ray florets pistillate, corollas yellow or white. Disc
1228. Amauriopsis Rydb.
Amauriopsis Rydb., N. Amer. Fl. 34: 37 (1914).
Annual herbs. Leaves alternate, blades deltate to
broadly ovate in outline, 1–4-pinnate, segments filiform to linear. Capitula in terminal, open paniculiform cymes, radiate. Involucres broadly campanulate to hemispheric, phyllaries in 2 series, subequal.
Receptacles flat to shallowly convex. Ray florets pistillate, corollas yellow-orange. Disc florets bisexual, rarely functionally pistillate and with staminodes, corollas yellow-orange, sometimes zygomorphic; anther appendages ovate, without glandular
trichomes; style arms acute to deltate, with narrow stigmatic surfaces. Cypselae obpyramidal, obscurely angled, sparsely to moderately pubescent.
Pappus of approximately 8–15 erose scales, midrib
rarely protruding beyond scale, sometimes absent.
x = 12, 18. Five species, south-western USA, northern Mexico
1229. Apostates N.S. Lander
Apostates N.S. Lander, Austral. Syst. Bot. 2: 129 (1989);
Karis, Bot. Jahrb. Syst. 120: 131–135 (1998), syst.
Shrubs. Leaves opposite, blades ovate. Capitula in
terminal, congested corymbiform cymes, discoid.
Involucres campanulate, phyllaries in 2–3 series,
subequal. Receptacles flat to shallowly convex.
Florets bisexual, corollas white; anther appendages
with glandular trichomes; style arms acute to
Fig. 89. Compositae-Bahieae. Bahia absinthifolia. A Habit.
B Capitulum with rays partly removed. C Ray corolla. D Disc
corolla. E Anthers. F Style arms. G Cypsela. (Reproduced
from Flora Novo-Galiciana, vol. 12, with permission of the
University of Michigan Herbarium; see McVaugh 1984)
436
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
florets bisexual, corollas yellow; anther appendages
glandular; style arms acute to deltate, stigmatic
surfaces narrow. Cypselae obpyramidal, 4-sided,
black, sparsely to moderately pubescent. Pappus
of approximately 6–10 obovate or lanceolate scales
with thickened midribs sometimes excurrent as
an awn or bristle. x = 8, 11, 10, 12. Approximately
ten species, western USA, Mexico, Chile.
1231. Bartlettia A. Gray
Bartlettia A. Gray, Mem. Amer. Acad. Arts 5: 323 (1855);
Powell, Southw. Naturalist 8: 117–120 (1963), rev.; Strother,
N. Amer. Fl. ser. II, 10: 146 (1978), rev.
Annuals. Leaves alternate, blades narrowly trullate,
deltate or trilobed, subentire to deeply mucronate,
serrate. Capitula in terminal, open paniculiform
cymes, radiate. Involucres hemispherical, phyllaries in 2 series, subequal. Receptacles flat to
shallowly convex. Ray florets pistillate, corollas
yellow. Disc florets bisexual, corollas yellow;
anther appendages ovate, without glandular
trichomes; style arms acute to blunt with a tuft
of papillae, stigmatic surfaces narrow. Cypselae
obcompressed, narrowly biconvex with one weak
rib on centre of each tangential face, shiny black,
pubescence denser on edges and tangential ribs,
moderately sericeous. Pappus of multiple capillary
bristles. x = 11. One species, B. scaposa A. Gray,
northern Mexico, south-western USA.
1232. Chaetymenia Hook. & Arn.
Chaetymenia Hook. & Arn., Bot. Beechey Voy. 298 (1838).
Perennial herbs or small, multi-stemmed shrubs
of riparious areas. Leaves opposite, linear to narrowly lanceolate, semisucculent. Capitula terminal,
solitary or in open paniculiform simple cymes, radiate, on long, distally fistulose peduncles. Involucres turbinate to campanulate, phyllaries gradate.
Receptacles convex. Ray florets pistillate, corollas
bright yellow, drying orange. Disc florets bisexual, corollas yellow; anther appendages without
glands; style arms acute to deltate, with broad stigmatic surfaces, touching at the very tip. Cypselae narrowly obpyramidal, 4-angled, strigose, especially on angles. Pappus of multiple, awn-like,
subplumose bristles. x = 9. One species, C. peduncularis Hook. & Arn., Mexico.
1233. Chamaechaenactis Rydb.
Chamaechaenactis Rydb., Bull. Torrey Bot. Club 33: 155
(1906); Preece & Turner, Madroño 12: 99–103 (1953).
Small perennial herbs with a long taproot growing
in exposed rocky areas, rosette apices densely
lanate. Leaves alternate, on short internodes, forming a rosette, blades broadly ovate to subcordate on
long petioles, semisucculent, glaucous. Capitula
terminal, solitary, scapose on long, leafless peduncles, discoid. Involucres campanulate, phyllaries
in 2 series, subequal. Receptacles flat to convex.
Florets bisexual, corollas white, turning brownmustard with age, sometimes drying purple, with
conspicuous glandular trichomes; anther appendages broadly ovate, glandular; style arms with
long, thickened appendages, stigmatic surfaces
narrow, base of style with numerous trichomes and
elongated cells. Cypselae obpyramidal, shallowly
quadrate, densely sericeous. Pappus of 7–10,
obovate, erose scales with thickened midribs,
acute or sometimes bilobed. x = 16. One species,
C. scaposa (Eastw.) Rydb., south-western USA.
1234. Espejoa DC.
Espejoa DC., Prodr. 5: 660 (1836).
Annual herbs. Leaves opposite, semisucculent. Capitula terminal, solitary or in open paniculiform
cymes, discoid, peduncles shallowly fistulose. Involucres turbinate to campanulate, phyllaries in
3 series. Receptacles flat to convex. Florets bisexual, corolla tubes yellow-green, tube base conspicuously broad and as wide as open throat, throats
yellow-purple, edges of lobes thickened; anther appendages narrowly oblong, without glands; style
arms short, acute to acuminate, stigmatic surfaces
thin, purple. Cypselae narrowly obpyramidal, 4–5angled, densely sericeous. Pappus of multiple ovate,
erose scales sometimes with the thickened midrib
extending as an awn. x = 9. One species, E. mexicana DC., Mexico, Central America.
1235. Florestina Cass.
Florestina Cass., Bull. Sci. Soc. Philom. Paris 1817: 11
(1817); Turner, Brittonia 15: 27–46 (1963), rev.
Annual or perennial herbs. Leaves opposite
proximally, alternate distally, blades linear to
ovate-elliptic, cordiform, entire or crenate. Capitula in terminal, open paniculiform cymes, discoid.
Involucres obconic to subhemispherical, phyllaries
in 1–2 series, equal to subequal. Receptacles flat
to shallowly convex. Florets bisexual, corollas
white or purple, asymmetrical with two lobes
appearing longer than others due to deeper
sinuses; anther appendages ovate, without glan-
Compositae
dular trichomes; style arms narrowly acuminate,
tapered, stigmatic surfaces thin, purple. Cypselae
obconic to obpyramidal-quadrate, black, sparsely
to moderately pubescent. Pappus of 8 obovate or
fusiform, erose scales, sometimes with thickened
midribs protruding as a bristle or awn, sometimes
with alternating obovate, awn-less scales. x = 10,
12. About eight species, southern USA, Mexico,
Central America.
1236. Holoschkuhria H. Rob.
Holoschkuhria H. Rob., Comp. Newslett. 38: 48–50, f. 1
(2002).
Annual or perennial herbs, weak shrubs. Leaves
opposite, blades simple, ovate to elliptic. Capitula in terminal, open paniculiform cymes, sometimes decussate, capitula discoid. Involucres hemispheric, phyllaries in 1–2 series, subequal. Receptacles flat. Florets bisexual, corollas white, tetramerous; anther appendages ovate, with glandular trichomes; style arms acute to deltate, with narrow
stigmatic surfaces. Cypselae obpyramidal, 4-sided,
black, moderately pubescent. Pappus of approximately 8 erose scales with a weak midrib. One
species, H. tetramera H. Rob., southern Ecuador,
northern Peru.
1237. Hymenopappus L’Hér.
Hymenopappus L’Hér., Hymenopappus 1 (1788); Turner,
Rhodora 58: 163–186, 208–269, 295–308 (1956), rev.
Annual or perennial herbs, sometimes biennial, rosulate or most leaves basal. Leaves alternate, blades
ovate in outline, 2-pinnate, segments filiform to
linear. Capitula in terminal, subscapose, open paniculiform cymes, capitula discoid, rarely radiate.
Involucres campanulate to hemispheric, phyllaries in 3 series, subequal to slightly gradate. Receptacles convex, rarely paleate. Ray florets pistillate, corollas white, rarely suffused with pink, shallowly trilobed. Disc florets bisexual, corollas white
or yellow, abruptly narrowed into tube, glandular,
lobes reflexed soon after anthesis, corollas spreading to slightly reflexed well above the involucre;
anther appendages with glandular trichomes; style
arms acute to acuminate, stigmatic surfaces narrow. Cypselae shallowly obcompressed, obpyramidal, shallowly quadrate, sometimes concave, not
striate, glabrous to moderately pubescent, black.
Pappus of approximately 8–10 erose, obovate scales
with a thickened midrib. x = 17. Approximately 14
species, south-western USA, northern Mexico.
437
1238. Hymenothrix A. Gray
Hymenothrix A. Gray, Mem. Amer. Acad. Arts 4: 102 (1849);
Turner, Brittonia 14: 101–120 (1962).
Annual or perennial herbs. Leaves alternate,
deltate to ovate in outline, 1–4-pinnate, segments
filiform to linear. Capitula in terminal, congested
corymbiform or open paniculiform cymes, discoid or radiate. Involucres turbinate to broadly
campanulate, phyllaries in 2 series, subequal.
Receptacles flat. Ray florets pistillate, corollas
yellow. Disc florets bisexual, corollas yellow, white
or lavender, deeply dissected, lobes reflexed,
especially the peripheral ones, irregularly lobed;
anther appendages without glandular trichomes;
style arms acute to deltate with narrow stigmatic
surfaces. Cypselae obpyramidal, 4-sided, sparsely
to moderately pubescent. Pappus of multiple
bristles or narrowly fusiform scales with strong
midribs, midribs protruding as awns or bristles.
x = 12. Four species, south-western USA, Mexico.
1239. Hypericophyllum Steetz
Hypericophyllum Steetz in Peters, Naturwissensch. Reise:
498 (1864).
Perennial herbs. Leaves opposite, blades ovate, oblong, trullate or obovate, entire to shallowly serrate. Capitula terminal, solitary or in open paniculiform cymes, discoid. Involucres broadly campanulate, phyllaries in 3–5 series, gradate. Receptacles
convex. Florets bisexual, corollas mostly yelloworange or red, rarely white or greenish white; anther appendages without glands; style arms acute
to deltate, with broad stigmatic surfaces. Cypselae narrowly obconic to obovate, obscurely 4–5sided, black or brown, glabrescent to moderately
pubescent. Pappus of multiple aristate scales or
bristles, sometimes bristles alternating with narrow scales, awns straight but most commonly uncinate, golden copper in colour. Seven species, tropical Africa.
1240. Loxothysanus B.L. Rob.
Loxothysanus B.L. Rob., Proc. Amer. Acad. Arts 4: 43 (1907);
Turner, Wrightia 5: 45–50 (1974), rev.
Shrubs. Leaves opposite. Capitula in terminal,
small corymbiform cymes, discoid. Involucres
campanulate to hemispheric, phyllaries in 2 series,
subequal. Receptacles convex. Florets bisexual,
corollas abruptly narrowed into tube, white,
glandular, especially those of peripheral florets
438
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
zygomorphic; anther appendages with a few
glandular trichomes; style arms acute to deltate,
with broad stigmatic surfaces. Cypselae narrowly
obconic to shallowly quadrate, especially with
age, shallowly striate, black, sparsely pubescent.
Pappus of multiple, erose scales, asymmetrical,
scales awn-like and longer on adaxial side of
cypsela. x = 15. Two species, Mexico.
1241. Palafoxia Lag.
Palafoxia Lag., Gen. Sp. Pl.: 26 (1816); Shinners, Field Lab.
20: 92–102 (1952); Turner & Morris, Rhodora 78: 567–628
(1976), rev.
Annual or perennial herbs, rarely shrubs. Leaves
opposite proximally, alternate distally, blades linear
to ovate. Capitula in terminal, open paniculiform
cymes, discoid or radiate. Involucres turbinate
or campanulate, phyllaries in 2 series, subequal,
herbaceous, linear. Receptacles flat. Ray florets
pistillate, corollas pink or pink-purple, deeply
trilobed. Disc florets bisexual, corollas white, pink,
magenta, actinomorphic to slightly zygomorphic;
anther appendages without glands; style arms
acuminate, without appendages, papillae covering
entire length of abaxial surface of style branches
and extending slightly below point of bifurcation.
Cypselae narrowly obpyramidal, 4-sided, black or
brown, glabrous to moderately pubescent. Pappus
of 4–10 obovate or more commonly fusiform or
linear scales with prominent midribs, midribs
sometimes protruding as awns, ray cypselae
sometimes with very reduced scales. x = 10, 11,
12. Twelve species, south-western USA, northern
Mexico.
1242. Peucephyllum A. Gray
Peucephyllum A. Gray, Rep. U.S. Mex. Bound., Bot. 74
(1859); Strother, N. Amer. Fl. ser. II, 10: 170–171 (1978), rev.
Shrubs. Leaves alternate, filiform to linear. Capitula terminal, solitary or in simple corymbiform
cymes, discoid. Involucres campanulate, phyllaries
in 1 series. Receptacles flat. Florets bisexual,
corollas yellow-green, lobes purplish; anther
appendages ovate with a few glandular trichomes;
style arms acute to deltate, with broad stigmatic
surfaces, papillae on abaxial surface reaching
style bifurcation. Cypselae narrowly obconic,
black, moderately pubescent. Pappus of multiple,
narrowly linear scales with prominent midribs
protruding as bristles, or of capillary bristles,
hyaline to stramineous. x = 10. One species,
P. schottii A. Gray, western USA, Mexico.
1243. Platyschkuhria Rydb.
Platyschkuhria Rydb., Bull. Torrey Bot. Club 33: 154 (1906);
Ellison, Brittonia 23: 269–279 (1971), rev.
Perennial herbs with a woody caudex. Leaves alternate, blades narrowly ovate, succulent. Capitula
few, subscapose, in terminal, open, corymbiform
cymes, radiate. Involucres hemispheric, phyllaries
in 2 series, subequal. Receptacles convex. Ray florets pistillate, sometimes with staminodes or additional lobes, corollas golden yellow. Disc florets
bisexual, corollas yellow; anther appendages glabrous; style arms acute to deltate, with broad stigmatic surfaces. Cypselae obpyramidal, shallowly
quadrate, with multiple striations, black, sparsely
pubescent. Pappus of multiple obovate scales with
prominent midribs, midribs sometimes protruding
as awns, hyaline to slightly purplish. x = 12. One
polymorphic species, P. integrifolia Rydb., southwestern USA.
1244. Psathyrotopsis Rydb.
Psathyrotopsis Rydb., N. Amer. Fl. 34: 360 (1927); Strother
& Pilz, Madroño 23: 24–40 (1975), rev.
Pseudobartlettia Rydb. (1927).
Annual or perennial herbs. Leaves tightly clustered
at base of plant, opposite at first nodes, alternate
above, blades ovate to trullate, sometimes broadly
ovate to suborbicular, entire to serrate-crenulate.
Capitula terminal, scapose, solitary or in open
paniculiform cymes, discoid. Involucres campanulate to subhemispheric, phyllaries in 2 series,
subequal. Receptacles convex. Florets bisexual,
corollas yellow or white, sometimes tinged with
pink; anther appendages glandular on abaxial
surface; style arms long, acuminate, sometimes
coiled, with broad stigmatic surfaces. Cypselae
narrowly obconic, with multiple striations or
shallow ribs, black or sometimes golden copper
in colour because of dense pubescence. Pappus of
multiple, unequal bristles, golden stramineous.
x = 19. Three species, south-western USA,
northern Mexico.
1245. Schkuhria Roth
Fig. 90
Schkuhria Roth, Catal. Bot. 1: 116 (1797), nom. cons.; Heiser,
Ann. Missouri Bot. Gard. 32: 265–278 (1945), rev.; Turner,
Phytologia 79: 364–368 (1996), part. rev.
Cephalobembix Rydb. (1914).
Compositae
439
tetramerous and pentamerous in the same capitulum; anther appendages with or without glandular
trichomes; style arms acute, with narrow stigmatic
surfaces and a minute aristate appendage. Cypselae obpyramidal and strongly quadrate, sparsely
pubescent sometimes only on angles. Pappus of 8–
10 obovate or fusiform, sometimes aristate scales,
stramineous, sometimes flecked with purple. x = 8,
10, 11. Six species, North and South America, introduced elsewhere.
1246. Thymopsis Benth.
Thymopsis Benth. in Benth. & Hook. f., Gen. Pl. 2: 201, 407
(1873).
Neothymopsis Britt. & Millsp. (1920).
Annual or perennial herbs. Leaves opposite, blades
trullate or ovate. Capitula terminal or axillary,
solitary or in simple cymes, sessile, disciform.
Involucres narrowly campanulate, phyllaries in
1–2 series, subequal. Receptacles flat. Florets
dimorphic; peripheral florets pistillate, lacking
stamens, corollas cylindric, with 4 minute lobes,
innermost florets bisexual with white, tetramerous
corollas with prominent glandular trichomes; anther appendages without glands; style arms deltate,
narrow stigmatic surfaces with a short, aristate
appendage. Cypselae obcompressed to subterete,
essentially glabrous to sparsely pubescent, black.
Pappus of a few erose scales forming a cup, the
scales fused at base and without a thickened
midrib. Two species, West Indies.
XXV. Tribe Polymnieae (H. Rob.)
Panero (2002).
Fig. 90. Compositae-Bahieae. Schkuhria pinnata. A Habit.
B Flowering capitulum. C Ray corolla. D Disc corolla.
E Anthers. F Style arms. G Cypsela. (Reproduced from
Flora Novo-Galiciana, vol. 12, with permission of the
University of Michigan Herbarium; see McVaugh 1984)
J.L. Panero
Annual herbs. Leaves alternate, filiform or linear,
ovate to trullate in outline, 1–3-pinnate, segments
linear. Capitula terminal, solitary or in open paniculiform cymes, discoid or radiate. Involucres obovate to obtrullate in outline, phyllaries in 1–3 series,
gradate or subequal. Receptacles flat. Ray florets
1–3, rarely 5, pistillate, corollas sometimes irregular with an additional, small limb opposite the
main limb, apices trilobed or acute. Disc florets
bisexual, corollas yellow, pentamerous, sometimes
Perennial herbs. Leaves opposite, petiolate, blades
cordate to deltate in outline, entire or deeply pinnatifid. Capitula in terminal, paniculiform cymes,
radiate. Involucres campanulate to hemispheric,
phyllaries in 2–3 series, subequal, herbaceous,
densely glandular abaxially. Receptacles convex
to shallowly conical, paleate. Ray florets pistillate,
corollas white to creamy-white, limbs trilobed with
central lobe longer and wider than flanking lobes,
abaxially pubescent. Disc florets functionally
staminate, corollas 5-lobed, yellow to white-yellow,
with multicellular trichomes, without glandular
Subtribe Polymniinae H. Rob. (1978).
440
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
trichomes, throats broadly campanulate; ovaries
very reduced; anthers yellow to brown, ecalcarate, ecaudate, appendages deltate, glandular
on abaxial surface, endothecium of fusiform cells
with 1–3 polar bridges; disc floret style arms
fused except for deltate apices, style arms of ray
florets tapered, without abaxial papillae, stigmatic
surface a continuous line at edges of style arms.
Cypselae obcompressed, without striations, walls
carbonized, broadly obovate, biconvex, with 3(–5)
sutures or ribs, sutures on angles and median
adaxial surface of cypsela, rarely with two additional sutures on abaxial, stipitate, dark brown to
greenish brown, glabrous or sometimes with a few
twin trichomes on apices. Pappus a shallow crown
or absent.
Polymnia was circumscribed by Robinson
(1978a) to include only two (now three, Pittman
and Bates 1989) closely related species endemic to
temperate forests of the eastern USA. The other
species of Polymnia, mostly large shrubs or small
trees of tropical America, were placed by him
(Robinson 1978b) in the genus Smallanthus. He
considered Polymnia to be a distinctive taxon
not closely related to members of Melampodiinae
and, therefore, placed the genus in subtribe
Polymniinae (Robinson 1978a). He considered
Polymniinae to be closely related to subtribes
Melampodiinae and Milleriinae in his circumscription (Robinson 1981). Molecular studies of
the family using chloroplast DNA (Panero et al.
2001c; Panero and Funk 2002), and of members of
subtribe Espeletiinae and related members using
nrDNA (Rauscher 2002) confirm that Polymniinae
are not related to Espeletiinae, Melampodiinae or
Milleriinae sensu Robinson (1981). Polymniinae
represent a distinctive helianthoid lineage splitting
off just above Helenieae and Coreopsideae (Panero
and Funk 2002). Molecular results support the
recognition of Polymniinae as tribe Polymnieae.
Only one genus:
1247. Polymnia L.
Polymnia L., Sp. Pl. 2: 926 (1753); Wells, Brittonia 17: 144–
159 (1965), reg. rev.
Polymniastrum Lam. (1798).
Characters of the tribe. x = 15. Three species, eastern USA.
XXVI. Tribe Heliantheae Cass.
(1819).
J.L. Panero
Annual or perennial herbs, shrubs, scandent
shrubs, lianas or trees. Leaves alternate or opposite, blades generally simple, rarely divided,
mostly ovate, triplinerved, adaxial surfaces usually scabrous, abaxial surfaces not as scabrous.
Capitula in terminal, rarely axillary, open
paniculiform cymes, sometimes congested, or
corymbiform cymes, sometimes solitary on long
peduncles, rarely sessile, sometimes arranged
in second-order capitula. Involucres cylindric to
hemispheric, sometimes reflexed or patent. Phyllaries subequal to gradate, in 1–7 series, normally
innermost smaller than outermost and resembling
paleae. Receptacles flat to convex, conic or sometimes columnar, paleate (epaleate in Tetranthus,
Trichocoryne, Riencourtia, and some functionally
staminate capitula in Ambrosiinae), paleae conduplicate, rarely wrapping around and shed as a unit
with cypselae (perigynia), usually persistent. Capitula radiate, discoid, rarely disciform. Ray florets
5–13(–21+), rarely 1 or 2, sometimes none, pistillate and fertile, styliferous and sterile or neuter,
corollas sometimes without a tube, apices trilobed,
sometimes bilobed. Disc florets bisexual or
functionally staminate, rarely functionally pistillate, corollas actinomorphic, rarely zygomorphic,
throats urceolate or campanulate, sometimes tubular, throats gradually or abruptly expanding above
tube, lobes 5, rarely 3 or 4, erect or spreading, rarely
coiled. Stamens 5, rarely 4 or 3, filaments glabrous,
rarely papillose, anthers connate or free, rarely
shallowly tailed, mostly sagittate or with acute to
acuminate lobes, ecalcarate, rarely shallowly calcarate, appendages mostly ovate, endothecial cells
fusiform or isodiametric, rarely quadrate to round,
with 1–5 polar thickenings, sometimes with radial
thickenings or encircled by uniform thickenings,
filaments glabrous or rarely with papillae. Pollen
echinate, caveate, exines with internal foramina
(‘Helianthoid’ pattern). Styles with 2, rarely one,
vascular strand, papillae increasing in density on
median to distal portions of abaxial surface of style
arms, stigmatic surfaces divided or fused, style
arms normally with a tuft of papillae on distal ends
or with non-vascularized, cylindric appendages.
Cypselae normally compressed, rarely obcompressed, biconvex, rarely shallowly quadrate,
narrowly to broadly obovate to suborbicular in
Compositae
outline, rarely terete, walls carbonized, prismatic
or ribbed, those of the ray triquetrous and obcompressed, walls smooth or striate, with uniseriate or
biseriate trichomes, sometimes with juicy exocarp
and resembling a drupe, winged or wingless; pappus of persistent awns and squamellae, disposed
in a narrowly oval to linear pattern on cypsela
neck, rarely in a radial pattern, pappus elements
sometimes reduced to an erose crown, rarely
caducous, or lacking. x = 19 (7–18, dysploid).
The tribe contains 113 genera and approximately 1,500 species. Its distribution is pantropical, with most species in Mexico and seasonally
dry regions of tropical and temperate South America. The tribe contains an important proportion of
the shrubs and trees of the family, most of these
found in the open pine-oak forests of western and
southern Mexico. The tribe is important economically, with Helianthus annuus, the commercial sunflower, being cultivated worldwide for oil production and cut flowers. Several species of Echinacea,
Rudbeckia and Zinnia are important components
of summer gardens across the world.
Tribe Heliantheae was named by Cassini (1819)
to include sunflowers with mostly radiate capitula,
paleaceous receptacles, opposite leaves and yellow
corollas. In 1829, Cassini presented his final
classification for Heliantheae which contained five
major divisions and excluded Ambrosiinae and
Tageteae. Bentham (1873b) adopted many of the
findings and taxonomic conclusions of Cassini. He
included in Helenieae most Heliantheae having
epaleate receptacles. He believed Helenieae to be
closely related to Heliantheae but probably not
natural. This decision was to dominate the debate
as to the phylogenetic relationships and limits
of Heliantheae for decades to come. With minor
revisions, Hoffmann (1894) adopted Bentham’s
concept for Die natürlichen Pflanzenfamilien.
Stuessy (1977) outlined the characteristics of
Heliantheae and provided a careful revision of the
historic decisions leading to the circumscription of
the Heliantheae-Helenieae alliance. His taxonomic
decisions were based on morphological features
but influenced greatly by chromosome number
information. His tribe Heliantheae contained 15
subtribes and, like Cronquist (1955) but with some
reservations, he believed Heliantheae to be the
basalmost lineage of the family. Robinson (1981)
dismissed this idea and instead placed Heliantheae
at the base of Asteroideae as the next lineage to
split from its ancestral stock, Eupatorieae. He
believed Helenieae to be a derived group within
441
the alliance, with subtribe Madiinae bridging the
morphological gap between the two tribes. His
treatment includes 35 subtribes, and is not a mere
merger of traditional tribes Heliantheae and
Helenieae and their corresponding subtribes but
a meaningful comparison of salient morphological
features which he used deftly to accommodate
genera in his classification scheme. Karis (1993a)
summarized the literature of the HeliantheaeHelenieae alliance, providing interesting and
important insight on the phylogenetic relationships and morphology of the group. His studies
were later formalized in a new classification for
tribe Heliantheae (Karis and Ryding 1994a) and
the recognition of a paraphyletic tribe Helenieae
(Karis and Ryding 1994b), with Eupatorieae and
the Athroisma group (Eriksson 1991) basal to
a series of sequentially splitting Helenieae lineages
ending in a monophyletic Heliantheae. These
conclusions were partly supported by the studies
of Kim and Jansen (1995) using sequence data of
the chloroplast gene ndhF. The classification of the
Heliantheae-Helenieae alliance was significantly
improved recently with the publication by Baldwin
et al. (2002) of a nuclear ribosomal ITS phylogeny
Fig. 91. Phylogenetic relationships of subtribes of Heliantheae based on chloroplast DNA sequences (after
Panero et al. 2001c)
442
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
which includes representatives of nearly all genera
of tribe Helenieae and outgroups. This study
supported the basal position of tribe Helenieae in
the strict sense (Gaillardiinae and Marshalliinae
sensu Robinson 1981) and provided evidence for
the recognition of several groups carved out of
the classical Helenieae. The impetus to implement
this classification was not the evidence about the
strength of the support for the groups themselves
but rather the finding that Eupatorieae were
derived from a helenioid lineage. This finding inevitably leads to the dismemberment of Helenieae.
The new classification system outlined by Baldwin
(Baldwin et al. 2002) included six tribes carved
out from classical Helenieae, namely Bahieae,
Chaenactideae, Helenieae, Madieae, Perityleae and
Tageteae. The concept of tribe Heliantheae adopted
here is based primarily on results from comparative studies of chloroplast DNA data (Panero et al.
2001c, and unpubl. data). These studies support
the conclusions of Baldwin et al. (2002), and a more
exclusive concept of tribe Heliantheae. Taxa with
mostly glandular herbage, functionally staminate
disc florets, cypselae somewhat terete with radial pappi of bristles and formerly included in
subtribes Galinsoginae, Melampodiinae and Milleriinae sensu Robinson (1981) are treated under
tribe Millerieae and excluded from Heliantheae. As
presently circumscribed, Heliantheae include 14
subtribes, namely Ambrosiinae, Chromolepidinae,
Dugesiinae, Ecliptinae, Enceliinae, Engelmanniinae, Montanoinae, Helianthinae, Rojasianthinae,
Rudbeckiinae, Spilanthinae, Verbesininae, Zaluzaniinae and Zinniinae (Fig. 91). Members of
these subtribes for the most part share paleate
receptacles, triplinerved blades, yellow corollas, radiate capitula, moderately to strongly
conduplicate paleae, herbaceous, sometimes
foliaceous phyllaries, strongly flattened cypselae
and, therefore, pappi arranged in a narrowly oval
to linear disposition on the neck of the cypselae,
and scabrous herbage. Our understanding of
the phylogenetic relationships of members of
Heliantheae has improved considerably since the
last treatment of the tribe by Karis and Ryding
(1994a). This has allowed me to redefine the
concept of certain subtribes and to propose the
recognition of new ones which are for the first
time used here. It can be argued that the taxonomy
of the tribe has benefited enormously from the use
of floral microcharacters, the main force behind
the overhauling of the classification of the tribe
by Robinson (1981). Most of the characters used
by him to circumscribe the major subtribes of
Heliantheae have apparently evolved several times
when compared with phylogenies produced by
DNA studies. For example, the fibres flanking or
embedding the vascular strands of the throats of
the disc corollas are present in several groups of
Ecliptinae and all members of Engelmanniinae.
These groups are not closely related to each other,
nor do they grow under similar environmental
conditions. The adaptive significance of this and
other characters exhibited by the floral parts of
the members of Heliantheae remains unclear.
Therefore, I stress results based on comparative
studies of DNA as the major component in the
circumscription of the main lineages of the tribe.
The subtribes and their associated genera are
organized alphabetically. The circumscription of
certain taxa continues to be problematic and no
doubt will change as more data become available.
This is the case for the limits and phylogenetic
relationships within Melanthera, Viguiera, and
Wedelia and relatives.
Key to the Subtribes
1.
–
2.
–
3.
–
4.
–
5.
–
6.
–
7.
–
8.
–
Paleae accrescent after anthesis
2
Paleae never accrescent after anthesis
3
Stamen filaments papillose 9. Rojasinthinae (p. 470)
Stamen filaments without papillae
8. Montanoinae (p. 469)
Phyllaries with black markings; ray corollas white with
black veins on abaxial surfaces; plants aquatic, forming loose rosettes
2. Chromolepidinae (p. 446)
Phyllaries without black markings; ray corollas otherwise; plants terrestrial, if aquatic, then not forming
rosettes
4
Disc florets bisexual, fertile; style arms of disc florets
with continuous stigmatic surfaces
5
Disc florets bisexual, fertile or functionally staminate;
style arms of fertile disc florets with parallel stigmatic
surfaces, sometimes these meeting at the apices 10
Disc corollas with prominent fibres embedding vascular strands
6. Engelmanniinae (p. 461)
Disc corollas without prominent fibres embedding
vascular strands
6
Ray corollas fused to cypselae, if ray florets sterile,
then disc floret corollas deeply lobed and lobes densely
papillose on adaxial surface or ray ovary with 3 minute
projections and receptacles columnar
14. Zinniinae (p. 475)
Ray corollas not fused to cypselae
7
Receptacles conspicuously conic or columnar; paleae
with decurrent bases; rarely capitulum with 4 florets
with phyllaries enclosing the cypselae (Tetranthus)
11. Spilanthinae (p. 471)
Receptacles flat to convex, rarely conic; paleae without
decurrent bases; capitula otherwise
8
Ray florets fertile
13. Zaluzaniinae (p. 474)
Ray florets sterile
9
Compositae
9. Paleae deciduous after anthesis; cypselae with long
sericeous trichomes
5. Enceliinae (p. 460)
– Paleae persistent, rarely deciduous, if deciduous, then
tightly enclosing cypselae (perigynia)
7. Helianthinae (p. 464)
10. Receptacles epaleate; disc florets fertile
14. Zinniinae (Trichocoryne)
– Receptacles paleate, if epaleate, then disc florets sterile
11
11. Disc florets functionally staminate and styles with
a single vascular strand, if with two vascular strands,
then ray cypselae fused to adjacent 2–4 disc florets and
their associated paleae or capitula with 2 series of ray
florets
12
– Disc florets bisexual, fertile, if functionally staminate,
then styles with two vascular strands and ray cypselae
not fused to adjacent disc florets and associated paleae
nor with capitula with 2 series of ray florets
14
12. Disc corollas with slender to prominent fibres embedding the vascular strands 6. Engelmanniinae (p. 461)
– Disc corollas without fibres embedding the vascular
strands
13
13. Ray corollas prominent, yellow with greenish veins on
abaxial surfaces
3. Dugesiinae (p. 446)
– Ray corollas absent, if present, then limbs wanting or, if
conspicuous, then ray cypselae fused to adjacent 2 disc
florets and associated paleae 1. Ambrosiinae (p. 443)
14. Receptacles conic to columnar with age
10. Rudbeckiinae (p. 470)
– Receptacles flat to convex, rarely conic, never columnar with age
15
15. Ray corollas white, disc corollas yellow; cypselae narrowly stipitate
12. Verbesininae (Podachaenium)
– Ray corollas mostly yellow or yellow-orange, if white,
then disc corollas also white (Steiractinia, Verbesina
or Wedelia); cypselae rarely stipitate
16
16. Ray cypselae mostly compressed, rarely obpyramidal,
stipitate with a pappus of several squamellae; shrubs or
small trees of the highlands of Mexico and Guatemala
12. Verbesininae (Squamopappus)
– Ray cypselae and corresponding pappus otherwise;
annual or perennial herbs, shrubs or trees of America
17
17. Resin in the ducts of disc corollas red; cypselae obpyramidal, quadrate; pappus of 4 scales or sometimes of 4
awns at angles of cypselae alternating with 4 squamellae; plants of Mexico and Mesoamerica
12. Verbesininae (Tetrachyron)
– Resin in the ducts of disc corollas yellow or orange,
never red; cypselae and pappus otherwise; plants of
America
18
18. Disc cypselae strongly compressed, obovate in outline
with two, stramineous, equivalent (rarely asymmetrical) wings at angles of cypselae, wings fused to pappus; pappus of 2 awns each at angles of cypsela, rarely
of 1 awn or missing; stigmatic surfaces of disc floret
style arms narrow and not touching each other, florets
never functionally staminate; disc corollas without fibres embedding the vascular strands
12. Verbesininae (Verbesina)
– Disc cypselae compressed to quadrate in crosssection, pappus various, cypselae of various shapes,
winged or wingless, if with two wings at angles of
cypselae and a pappus of two awns, then either
443
wings strongly asymmetrical and cypselae narrowly
oblanceolate (Otopappus) or cypsela oblanceolate
with narrow wings and disc corollas with fibres
(Calyptocarpus, Lasianthaea, Tuxtla), without fibres
(Oblivia) or wings forming a cup around neck,
surrounding the disc corolla tubes (Dimerostemma),
or paleae filiform (Eclipta)
4. Ecliptinae (p. 446)
XXVI.1. Subtribe Ambrosiinae Less. (1830);
Bolick, Adv. Clad. 2: 125–141 (1983);
Karis, Syst. Bot. 20: 40–54 (1995); Miao
et al., Amer. J. Bot. 82: 924–932 (1995).
Tribe Ambrosieae Cass. (1819).
Family Ambrosiaceae Link (1829).
Annual or perennial herbs, shrubs, rarely small
trees. Leaves alternate or opposite, blades simple,
unlobed to deeply dissected, venation mostly
triplinerved. Capitula in terminal, paniculiform or
spiciform cymes, sometimes unisexual with female
capitula at base of spicate male inflorescence, discoid, disciform or radiate, mostly wind-pollinated,
capitula mostly nodding. Involucres campanulate
to hemispheric, phyllaries in 1–3 series, subequal
or gradate, free or fused. Receptacles flat, convex
to minutely conical, usually paleate. Ray florets
absent or in bisexual capitula peripheral florets
female, corollas tubular or sometimes reduced
to a shallow ring, sometimes minutely radiate
with bilobed limbs, yellow or white. Disc florets
functionally staminate, mostly pentamerous,
rarely tetramerous, corollas hyaline or light yellow,
without fibres embedding vascular strands, sometimes ovaries absent; stamens 5, anthers yellow
to hyaline rarely with dark connectives or light
brown, filaments free or fused, anthers free or
partially connate, endothecium cells quadrate with
multiple polar and radial thickenings, styles with
one or two vascular strands, style arms of fertile
florets with divided stigmatic surfaces, style arms
of sterile florets fused, terete with only one, rarely
two, vascular strands. Cypselae of unisexual heads
enclosed in a conceptacle in which phyllaries are
fused to the fruits, cypselae of bisexual corollas
obcompressed, sometimes winged or forming
a unit associated with the subtending phyllary and
the adjacent disc florets/paleae. Pappus absent,
a minute crown of squamellae or trichomes, or 2–3
awns or scales. Eight genera, 70 species, America,
introduced elsewhere.
Preliminary chloroplast DNA studies show that
Ambrosiinae may be a paraphyletic assemblage
(see Fig. 91).
444
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Key to the Genera
1. Ray cypselae obcompressed and associated with 2
functionally staminate disc florets and their corresponding paleae
2
– Ray cypselae compressed or obcompressed or enclosed in a conceptacle but never associated with adjacent 2 disc florets and their corresponding paleae 3
2. Ray florets always 5; ray corollas with a small limb
1254. Parthenium
– Ray florets 6–8; ray corollas without a limb
1253. Parthenice
3. Capitula unisexual; plants monoecious
4
– Capitula bisexual; if unisexual, plants dioecious or
population gynodioecious
5
4. Phyllaries of male capitula connate 1248. Ambrosia
– Phyllaries of male capitula free
1255. Xanthium
5. Staminal filaments connate up to anther collars; inner
phyllaries accrescent after anthesis
1249. Dicoria
– Staminal filaments free; inner phyllaries not accrescent
6
6. Capitula or pair of capitula subtended by bracts as
long as or longer than capitulum
1252. Iva
– Capitula not subtended by bracts or, if bracts present,
then these minute and subtending inflorescence
branch of 5 or more capitula
7
7. Corollas of peripheral florets tubular, sometimes reduced to a minute ring around style 1250. Euphrosyne
– Corollas absent and replaced by a few glandular or
moniliform trichomes around style 1251. Hedosyne
Genera of Ambrosiinae
1248. Ambrosia L.
Fig. 92
Ambrosia L., Sp. Pl. 2: 987 (1753); Payne, J. Arnold Arb. 45:
401–430 (1964), rev.; Strother & Baldwin, Madroño 49: 143–
144 (2002), incl. Hymenoclea.
Franseria Cav. (1793) [1793–1794], nom. cons.
Hymenoclea Torr. & A. Gray (1848).
Annual or perennial herbs, shrubs. Leaves opposite or upper alternate, blades lobed or dissected,
deltate, ovate to subrotundiform in outline, triplinerved. Capitula discoid, unisexual, sessile, pistillate capitula solitary or in clusters at base of spiciform cymose inflorescences of functionally staminate capitula; phyllaries of functionally staminate
capitula connate. Involucres campanulate to hemispheric. Receptacles in staminate capitula paleate.
Pistillate florets without corollas and stamens, commonly more than 1 floret aggregated into a conceptacle, staminate corollas 4- or 5-lobed, light yellow or hyaline; filaments free, rarely connate, anthers free; style arms of staminate florets fused,
terete, with only one, rarely two, vascular strands.
Disseminule a conceptacle in which phyllaries are
fused to the fruits, smooth or variously with transverse scarious bracts in one plane or spirally ar-
Fig. 92. Compositae-Heliantheae. Ambrosia canescens.
A Habit. B Anther in abaxial and lateral views. C Staminate
corolla. D Palea. E Involucre. F Staminate capitulum.
G Apex of stylar remnant in staminate floret. Ambrosia
polystachya. H Involucre. (Reproduced from Flora NovoGaliciana, vol. 12, with permission of the University of
Michigan Herbarium; see McVaugh 1984)
ranged or more commonly with hooked or spine
projections. Pappus absent. x = 12, 17, 18. Thirty
species, America, introduced elsewhere.
1249. Dicoria Torr. & A. Gray
Dicoria Torr. & A. Gray, Rep. U.S. Mex. Bound. 86 (1859).
Annual or perennial herbs, shrubs. Leaves alternate, petiolate, blades simple, narrowly lanceolate
to ovate, suborbicular. Capitula in terminal,
open, ebracteate paniculiform cymes, disciform,
sometimes unisexual. Involucres campanulate,
phyllaries in 2 series, accrescent. Receptacles convex. Peripheral florets pistillate, corollas a shallow
ring or absent, other florets functionally staminate,
Compositae
5-lobed, corollas yellow; filaments connate up to
anther collars; styles with one vascular strand.
Cypselae obcompressed, sometimes with prominent pectinate or smooth wings, glabrous. Pappus
absent or of a few hyaline trichomes. x = 18. Four
species, northern Mexico, south-western USA.
1250. Euphrosyne DC.
Euphrosyne DC., Prodr. 5: 530 (1836).
Cyclachaena Fres. (1836).
Oxytenia Nutt. (1848).
Chorisiva Rydb. (1922).
Leuciva Rydb. (1922).
Annual or perennial herbs, sometimes semiaquatic. Leaves alternate or opposite, petiolate,
blades oval to ovate in outline, unlobed to lobed
or 2–3 times pinnate. Capitula in terminal,
ebracteate, paniculiform cymes, disciform. Involucres hemispheric, phyllaries in 1–2 series,
subequal, herbaceous. Receptacles usually paleate.
Peripheral florets pistillate, corollas tubular or
a shallow ring, yellow, other florets functionally
staminate, 5-lobed, corollas yellow; filaments
free, anthers free; styles with one vascular strand.
Cypselae obcompressed, sometimes winged at
maturity, glabrous or with glandular or setose
trichomes. Pappus a minute crown. x = 18. Five
species, Mexico, USA.
1251. Hedosyne (A. Gray) Strother
Hedosyne (A. Gray) Strother, Madroño 47: 204 (2000).
Annual herbs. Leaves alternate, blades ovate in outline, 1–2 times pinnate, segments lobed. Capitula in terminal, mostly ebracteate, paniculiform
cymes, disciform. Involucres hemispheric, phyllaries in 2 series. Receptacles paleate. Peripheral
florets fertile, corollas missing, other florets functionally staminate, corollas 5-lobed, yellow; filaments free. Cypselae obcompressed, oblong, black,
glabrous. Pappus absent. One species, H. ambrosiifolia (A. Gray) Strother, northern Mexico, southwestern USA.
1252. Iva L.
Iva L., Sp. Pl. 2: 988 (1753); Jackson, Univ. Kansas Sci. Bull.
41: 793–876 (1960), rev.; Bolick, Taxon 34: 81–84 (1985),
phylog.
Annual or perennial herbs. Leaves alternate or opposite. Capitula in terminal, bracteate paniculiform cymes or congested spicate cymes, disciform.
445
Involucres campanulate to hemispheric, phyllaries
in 1–2 series. Receptacles convex. Peripheral florets pistillate, corollas tubular, yellow, other florets
functionally staminate, 5-lobed, corollas yellow; filaments free, anthers free; styles with one vascular
strand. Cypselae obcompressed to terete, glabrous
to sometimes completely covered with glandular
trichomes. Pappus absent. x = 16, 17. Ten species,
North America, introduced elsewhere.
1253. Parthenice A. Gray
Parthenice A. Gray, Smithsonian Contr. Knowl. 5: 85 (1853).
Annual herbs. Leaves alternate, petiolate, blades
ovate to deltate or subcordate, triplinerved. Capitula in terminal, paniculiform cymes, disciform.
Involucres hemispheric, phyllaries in 2–3 series,
dimorphic. Receptacles convex. Peripheral florets
pistillate, 6–8, corollas tubular, white or greenwhite. Disc florets functionally staminate, corollas
white or creamy white; filaments and anthers free.
Cypselae obcompressed, deeply concave, wings
involute, associated with subtending phyllary
and adjacent two disc florets and paleae. Pappus
absent. x = 18. One species, P. mollis A. Gray,
Mexico, south-western USA.
1254. Parthenium L.
Parthenium L., Sp. Pl. 2: 988 (1753); Rollins, Contr. Gray
Herb. II, 172: 1–72 (1950), rev.
Annual or perennial herbs, shrubs, rarely small
trees. Leaves alternate, petiolate, blades unlobed
or lobed sometimes 2 times pinnate, ovate to
subsagittate. Capitula in terminal, simple, open or
congested paniculiform cymes, radiate. Involucres
hemispheric, phyllaries in 2 series, mostly 5
in each series, dimorphic. Receptacles flat to
convex. Ray florets pistillate, 5, corollas with small
bilobed limbs, white or green-white. Disc florets
functionally staminate, 5- or 4-lobed, corollas
white or creamy white, styles with one vascular
strand; filaments and anthers free. Cypselae obcompressed, shallowly concave or flat, associated
with subtending phyllary and paleae and florets
of adjacent two disc florets, paleae enclosing the
disc florets, forming a corky disseminule. Pappus
of 2–3 awns or scales or absent. x = 9. Sixteen
species, America.
1255. Xanthium L.
Xanthium L., Sp. Pl. 2: 987 (1753); Millspaugh & Sherff, Field
Mus. Nat. Hist. Publ. Bot. ser. IV, 204: 9–49 (1919), reg. rev.
446
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Annual herbs, sometimes spiny. Leaves alternate,
petiolate, blades ovate to deltate, simple or lobed,
triplinerved. Capitula discoid, unisexual, sessile,
pistillate solitary or in clusters at base of terminal
or axillary spiciform cymose inflorescences composed of functionally staminate heads; phyllaries
of functionally staminate capitula free. Involucres
campanulate to hemispheric. Receptacles in staminate heads paleate. Pistillate florets without corollas
and stamens, commonly more than 1 floret aggregated into a conceptacle; staminate corollas light
yellow or hyaline; anthers free, filaments connate;
styles of staminate florets wanting. Disseminule
a conceptacle in which phyllaries are fused to the
fruits, with hooked or spine projections. Pappus
absent. x = 18. Three species of disturbed habitats,
warm and temperate regions of the world.
XXVI.2. Subtribe Chromolepidinae Panero
(2005).
Rosulate perennial semi-aquatic herbs. Leaves
alternate, petiolate, semi-succulent, ovate to
narrowly lanceolate, entire or pinnatifid. Capitula axillary, solitary, long-pedunculate, radiate.
Involucres campanulate, phyllaries dimorphic
in 2–3 series, subequal, membranaceous with
black markings. Receptacles convex, paleate. Ray
florets pistillate, fertile, corollas white. Disc florets
bisexual, fertile, corollas pentamerous, yellow,
without fibres embedding vascular strands; stamens 5, anthers black, appendages ovate to deltate,
without glandular trichomes; styles with two
vascular strands, style arms with divided stigmatic
surfaces, apices broadly acute to shallowly deltate.
Ray cypselae obcompressed, triquetrous to weakly
quadrate, glabrous. Disc cypselae obpyramidal,
quadrate, sparsely pubescent, conspicuously
smaller than ray cypselae. Pappus a crown of
minute awns and squamellae, absent in ray
cypselae.
Only one genus:
1256. Chromolepis Benth.
Chromolepis Benth., Pl. Hartw. 40 (1840).
Characters of the subtribe. One species, C. heterophylla Benth., Mexico.
XXVI.3. Subtribe Dugesiinae Panero (2005).
Prostrate, rhizomatous, stoloniferous perennial
herbs. Leaves alternate, petiolate, blades obovate
to oval in outline, runcinate to pinnatifid. Capitula
axillary, solitary or in small paniculiform cymes,
radiate. Involucres hemispheric, phyllaries in 2–3
series, subequal, outermost foliaceous. Receptacles
flat to slightly convex. Ray florets pistillate, in
two series, corollas lemon to golden yellow with
greenish to black veins on abaxial sides, rays
deeply 2- rarely 3-lobed. Disc florets functionally
staminate, corollas pentamerous, yellow, without
fibres embedding vascular strands; stamens 5,
anthers black, endothecium of fusiform cells with
1–2 polar bridges or evenly thickened; styles with
two vascular strands, style arms of disc florets
narrowly tapered and papillose, style arms of ray
florets spreading with broad, divided stigmatic
surfaces. Ray cypselae obcompressed, biconvex,
glabrous with a few trichomes on the neck, margins
with shallowly lacerate wings. Pappus absent or
a minute crown.
Only one genus:
1257. Dugesia A. Gray.
Dugesia A. Gray, Proc. Amer. Acad. Arts 17: 215 (1882).
Characters of the subtribe. x = 18. One species,
D. mexicana A. Gray, central Mexico.
XXVI.4. Subtribe Ecliptinae Less. (1831);
Villaseñor & Strother, Syst. Bot. 14: 529–
540 (1989); Strother, Syst. Bot. Monogr.
33: 1–111 (1991), rev.; Panero et al.,
Amer. J. Bot. 86: 413–427 (1999), phylog.
Subtribe Clibadiinae H. Rob. (1978).
Annual or perennial herbs, shrubs, lianas or trees.
Leaves usually opposite, blades lanceolate to ovate,
mostly unlobed. Capitula terminal, rarely axillary,
solitary or in open paniculiform or rarely congested corymbiform cymes, radiate, rarely discoid
or disciform. Involucres campanulate to hemispheric, rarely cylindrical, phyllaries in 1–7 series,
indurate at base with herbaceous, sometimes foliaceous apices, innermost sometimes larger, broader
and chartaceous. Receptacles usually flat to convex.
Ray florets pistillate or sometimes neuter. Disc
florets bisexual, sometimes functionally staminate,
corollas usually pentamerous, with or without
fibres embedding the vascular strands; stamens
5, rarely 4; styles with two vascular strands, style
arms with divided stigmatic surfaces, apices with
or without papillae, sometimes with a papillose
Compositae
appendage, in functionally staminate disc florets
style arms fused. Ray cypselae obcompressed and
triquetrous, disc cypselae compressed, biconvex,
sometimes shallowly quadrate, usually black,
mostly winged on angles, faces glabrous or moderately pubescent, sometimes tuberculate, with or
without elaiosomes. Pappus of easily caducous or
persistent awns, normally as many as angles of
cypsela with or without intermediate squamellae,
squamellae sometimes fused with awns and
forming a crown, sometimes this elevated on
a rostrum, rarely a minute crown or absent.
13.
–
14.
–
15.
–
Key to the Genera
1. Capitula with 1 round phyllary; involucres planocompressed
1263. Delilia
– Capitula without round phyllaries; involucres campanulate to hemispheric
2
2. Capitula nodding; anthers free; plants with tuberculate
cypselae
1267. Eleutheranthera
– Capitula erect; anthers connate, rarely free; plants with
or without tuberculate cypselae
3
3. Disc florets functionally staminate
4
– Disc florets bisexual, fertile
10
4. Ray corollas yellow; corolla limbs expanded or patent
and protruding above involucres
5
– Ray corollas white; corolla reduced to a tube or corolla
limbs minute and not expanded above involucres 8
5. Anthers yellow to pale brown; Australia
1286. Pentalepis
– Anthers brown or black; neotropical
6
6. Cypselae without wings; pappus a crown of minute
scales
1258. Baltimora
– Cypselae winged; wings fused to a pappus of two awns
7
7. Shrubs; style arms of disc florets fused except for distal end; cypsela wings stramineous; Central America
1291. Rensonia
– Annual herbs; style arms of disc florets not fused;
cypsela wings black; Ecuador
1293. Schizoptera
8. Capitula with more than one pistillate, fertile floret
1261. Clibadium
– Capitula with one pistillate, fertile floret
9
9. Leaves alternate; capitula with only 2 functionally
staminate florets; Cuba and Hispaniola
1257. Lantanopsis
– Leaves opposite; capitula with more than 2 functionally staminate florets; mostly eastern, tropical South
America
1292. Riencourtia
10. Ray florets absent or, if present, then neuter or pistillate
and sterile
11
– Ray florets present, pistillate and fertile
31
11. Capitula with 1–6 disc florets
12
– Capitula with more than 6 disc florets
14
12. Capitula with 4 disc florets; cypselae obcompressed,
linear to narrowly rhombic in cross-section, with
lacerate, stramineous wings; Argentina, Bolivia and
Paraguay
1297. Synedrellopsis
– Capitula with 5 or 6 disc florets; cypselae obcompressed or terete, biconvex to circular in cross-section,
16.
–
17.
–
18.
–
19.
–
20.
–
21.
–
22.
–
23.
–
24.
–
447
without lacerate, stramineous wings; Philippines or
western tropical Africa
13
Plants with two sterile ray florets; phyllaries 5 or more
in 2 series; Philippines
1269. Fenixia
Plants without ray florets; phyllaries 2 in 1 series; western tropical Africa
1270. Hoffmanniella
Plants dioecious, capitula discoid, receptacles globose;
Chile
1290. Podanthus
Plants not dioecious, capitula discoid or radiate, receptacles rarely globose, mostly flat to convex or conical;
America, rarely of Chile or pantropical
15
Cypselae with a pappus of several, unequal, caducous
awns on a circular or oval rostrum, rarely in combination with two awns at angles of cypselae or at base of
rostrum
16
Cypselae epappose or with a pappus of persistent awns
or squamellae
18
Cypselae greyish black or black and always mottled,
not tuberculate; phyllaries of innermost series broadly
obovate or sometimes with orbicular apices, chartaceous, translucent and amber in colour; Colombia,
Ecuador and Venezuela
1295. Steiractinia
Cypselae black, rarely mottled, sometimes tuberculate; phyllaries of innermost series ovate to lanceolate,
apices never orbicular nor broadly obovate, chartaceous, translucent or amber in colour
17
Cypselae with two slightly wider, longer, caducous
awns; scandent shrubs of eastern Mexico, corollas yellow
1287. Perymeniopsis
Cypselae awns equivalent in width, without two
slightly wider, longer, caducous awns; pantropical
herbs or weak shrubs with yellow or white corollas,
those in Mexico with only white corollas
1280. Melanthera
Cypselae tuberculate
19
Cypselae not tuberculate
22
Ray corollas deep orange or red; Venezuela
1300. Tuberculocarpus
Ray corollas, when present, mostly yellow or yelloworange; pantropical
20
Leaves linear; Galápagos Islands 1299. Trigonopterum
Leaves mostly ovate, never linear
21
Pappus of a few lacerate squamellae or minute awns
on a rostrum
1268. Exomiocarpon
Pappus a lacerate crown, not constricted into a rostrum
1267. Eleutheranthera
Cypselae with a pappus on a rostrum or separated
from the main body of the cypselae by a small constriction, the pappus of 0–3 awns fused to an erose cup
of free or partially fused squamellae or small awns 23
Cypselae epappose or with pappus various but never
on a rostrum, continuous with the main body of the
cypselae
26
Cypselae with a pappus on an eccentric, half sigmoid
or geniculate rostrum; disc corollas white, ray corollas
absent; central Yucatan, Mexico 1289. Plagiolophus
Cypselae with a pappus separated from the cypsela by
a small constriction or on a straight rostrum; disc
corollas yellow, if white, then ray corollas present,
white; pantropical
24
Cypselae with a pappus on a rostrum 1303. Wedelia
Cypselae with a pappus not on a rostrum but separated
from the body of the cypsela by a small constriction
25
448
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
25. Plants from southern Mexico and adjacent Belize
1306. Zyzyxia
– Plants from Panama and tropical South America
1284. Oyedaea
26. Cypselae winged, wings either lacerate or smooth and
normally of a lighter colour than cypselae, rarely concolorous, sometimes wings reduced to a smooth, corky
rim
27
– Cypselae never winged
29
27. Cypselae broadly rhombic in cross-section, drupaceous when fresh; wings concolorous with the body
of the cypsela
1298. Tilesia
– Cypselae linear to biconvex, rarely narrowly rhombic
in cross-section, not drupaceous when fresh; wings
not concolorous with the body of the cypsela
28
28. Wings of cypselae asymmetrical, distal end of one
of the wings smoothly to abruptly broader than the
corresponding wing on the other side of the cypsela
1283. Otopappus
– Wings fused at neck of cypselae producing a crown
around the tube of the corolla 1264. Dimerostemma
29. Throats of disc corollas with fibres embedding the
vascular strands
1266. Elaphandra
– Throats of disc corollas without fibres embedding the
vascular strands or these wanting
30
30. Capitula discoid, corolla abruptly expanding above
tube into a campanulate or broadly urceolate throat,
cypselae fusiform; Ecuador and Peru 1281. Monactis
– Capitula radiate, corolla gradually expanding into
a narrowly campanulate throat, cypselae obcuneate
to obovate; central Chile
1277. Leptocarpha
31. Paleae fused to adjacent paleae at base on lateral sides
(removing the cypsela and its associated palea on
a single pull brings off half of the paleae of the contiguous cypselae; removing the cypsela before it is very
mature and the capitulum very dry normally brings
with it its associated palea)
1279. Lundellianthus
– Paleae not fused laterally nor to the cypsela
32
32. Disc corollas tetramerous
33
– Disc corollas pentamerous
34
33. Paleae filiform, ray floret limbs linear or narrowly
lanceolate
1265. Eclipta
– Paleae not filiform, ray floret limbs oval
1278. Lipochaeta
34. Pappus of 2 to several caducous awns
35
– Pappus various, awns, when present, persistent
36
35. Leaves mostly lobed or compound, sometimes simple;
cypselae sometimes tuberculate; Hawaii
1278. Lipochaeta
– Leaves simple; cypselae minutely tuberculate and appearing as if having striped faces; neotropical
1288. Perymenium
36. Cypselae tuberculate
37
– Cypselae not tuberculate
40
37. Involucres with outermost phyllaries resembling small
foliage leaves
1273. Jefea
– Involucres with outermost phyllaries not resembling
foliage leaves
38
38. Cypselae with two divergent awns
1260. Calyptocarpus
– Cypselae without awns
39
39. Leaves linear, cypselae obpyriform with a pappus
a small crown on a small rostrum; Galápagos Islands
1299. Trigonopterum
– Leaves lanceolate to trullate, never linear, cypselae
broadly fusiform with a pappus an erose crown continuous with cypselae; pantropical 1294. Sphagneticola
40. Capitula axillary
41
– Capitula terminal
42
41. Ray cypselae with lacerate wings, pappus of two slender awns fused to wings
1296. Synedrella
– Ray cypselae without wings, sometimes margins irregular, pappus of two divergent awns
1260. Calyptocarpus
42. Wings of cypsela fused around the neck forming
a cup-like structure surrounding the tube of the
corolla
1264. Dimerostemma
– Wings of cypselae not fused around the neck
43
43. Leaves alternate
44
– Leaves opposite, those associated with the capitula
sometimes subopposite or alternate
46
44. Cypselae with a pappus on a rostrum 1259. Blainvillea
– Cypselae without a rostrum
45
45. Capitula with only 1–3 ray florets
1281. Monactis
– Capitula with more than three ray florets
1274. Kingianthus
46. Cypsela epappose
1262. Damnxanthodium
– Cypselae pappose, pappus sometimes reduced to
a minute crown
47
47. Limbs of ray corollas narrowly oblanceolate to filiform
and involute
48
– Limbs of ray corollas oval or oblong, sometimes obovate, rarely oblanceolate, never involute
49
48. Disc corolla throats with prominent fibres embedding the vascular strands, leaf-blades elliptic, awns
of cypsela slightly curved outwards; Mexico and Costa
Rica
1301. Tuxtla
– Disc corolla throats without fibres embedding the
vascular strands, leaf-blades lanceolate-elliptic, awns
of cypsela not curved; Panama and northern South
America
1282. Oblivia
49. Disc corolla throats with prominent fibres embedding
the vascular strands
50
– Disc corolla throats without fibres
52
50. Cypselae prominently winged
1293. Schizoptera
– Cypselae not winged or with a narrow hyaline margin
along cypsela edges
51
51. Disc cypselae compressed, pappus of two awns, rarely
with squamellae in between
1276. Lasianthaea
– Disc cypselae weakly quadrate, pappus of two awns
and a few squamellae in between
1272. Iogeton
52. Involucres with outermost phyllaries resembling small
foliage leaves
1273. Jefea
– Involucres with outermost phyllaries not resembling
foliage leaves
53
53. Cypselae with a rostrum
54
– Cypselae without a rostrum
56
54. Cypselae without elaiosomes
55
– Cypsela with elaiosomes
1303. Wedelia
55. Cypselae winged
1305. Zexmenia
– Cypselae not winged
1259. Blainvillea
56. Cypselae without wings, disc cypselae shallowly
quadrate, rarely pentagonal
57
– Cypselae with asymmetrical wings 1283. Otopappus
57. Cypselae weakly stipitate and shallowly navicular;
small trees of central Ecuador
1271. Idiopappus
– Cypselae not weakly stipitate nor shallowly navicular; annual or perennial herbs, shrubs of Mexico and
Compositae
58.
–
59.
–
60.
–
Central America or south-eastern South America and
Chile
58
Cypselae with a pappus a minute crown; annual or
perennial herbs of south-eastern South America and
central Chile
1285. Pascalia
Cypselae with a pappus of a few awns or squamellae
free or fused at the base
59
Leaves triplinerved
60
Leaves pinnately veined
1272. Iogeton
Cypselae obpyramidal; Mesoamerica
1302. Wamalchitamia
Cypselae oblong or obovate; coastal sand dunes of
Indo-Pacific region
1304. Wollastonia
Genera of Ecliptinae
1258. Baltimora L.
Baltimora L., Mant. Pl. 2: 158, 288 (1771), nom. cons.;
Stuessy, Fieldiana Bot. 36: 31–50 (1973), rev.
Annual erect herbs. Leaves opposite, petiolate,
blades ovate, triplinerved. Capitula in terminal,
open paniculiform cymes, radiate. Involucres
cylindrical to campanulate, phyllaries in 2 series,
subequal to shallowly gradate. Receptacles flat to
convex. Ray florets pistillate, corollas yellow to
yellow-orange. Disc florets functionally staminate,
corollas yellow, without fibres embedding vascular strands, lobes deeply papillose on adaxial
surface; anthers black, appendages with glandular trichomes; styles arms of ray florets with
divided stigmatic surfaces, spreading. Cypselae
triquetrous, obpyramidal, the apices with three
protruding horns, black, strongly to shallowly
tuberculate, puberulent. Pappus a crown of minute
scales. x = 12, 13. Two species, neotropics.
1259. Blainvillea Cass.
Blainvillea Cass., Dict. Sci. Nat., ed. 2, 29: 493–494 (1823).
Annual or perennial erect herbs, weak shrubs.
Leaves opposite or alternate, petiolate, blades
lanceolate to broadly ovate, triplinerved. Capitula
in terminal, open paniculiform cymes, radiate.
Involucres subcylindrical, campanulate to oblong,
sometimes hemispheric, phyllaries in 2–4 series,
chartaceous with green longitudinal striae. Receptacles flat to minutely convex. Ray florets pistillate,
corollas yellow or white. Disc florets bisexual,
corollas yellow or white, without fibres embedding
vascular strands, tubes long and slender; anthers
black, appendages with or without glandular
trichomes; style arms narrowly acuminate. Ray
cypselae obcompressed, obpyramidal, black,
glabrous. Disc cypselae compressed, biconvex,
449
sometimes shallowly triquetrous to quadrate,
black. Pappus on a rostrum, of 2–3 awns and
sometimes with a few scales fused at the base.
x = 17. Ten species, pantropical.
1260. Calyptocarpus Less.
Calyptocarpus Less., Syn. Gen. Comp. 221 (1832); McVaugh
& Smith, Brittonia 19: 268–272 (1967), rev.
Prostrate to weakly erect perennial herbs or weak
shrubs. Leaves opposite, petiolate, blades ovate
to lanceolate, triplinerved. Capitula terminal and
axillary, solitary or in simple cymes, radiate.
Involucres campanulate, phyllaries in 1–2 series,
mostly 5, herbaceous. Receptacles convex. Ray florets pistillate, corollas yellow. Disc florets bisexual,
corollas yellow, with fibres embedding vascular
strands; anthers black, appendages with glandular
trichomes; style arms narrowly acuminate. Cypselae compressed, narrowly obovate or obcuneate,
often tuberculate, sometimes striate. Pappus of 2
divergent or reflexed awns. x = 12. Two species,
southern USA, Mexico, Central America, Cuba.
1261. Clibadium L.
Clibadium L., Mant. Pl. 2: 161, 294 (1771); Arriagada,
Brittonia 55: 245–301 (2003), rev.
Trichapium Gilli (1983).
Shrubs or trees. Leaves opposite, sessile to petiolate,
blades lanceolate to broadly cordiform, triplinerved. Capitula in terminal open or congested paniculiform to corymbiform, sometimes glomerulelike cymes, disciform, rarely epaleate. Involucres
cylindric, campanulate, hemispherical, phyllaries
in 2–6 series, subequal, membranous, scarious, the
inner enclosing the marginal cypselae, greenish
to purple, sometimes white. Receptacles flat to
shallowly convex. Pistillate florets in 1–2 series,
corollas inconspicuous, white, greenish white
or white-yellow, 2–5-lobed, sometimes weakly
zygomorphic. Disc florets functionally staminate,
corollas white or greenish white, 4- or 5-lobed; anthers black, appendages with glandular trichomes;
style arms acuminate. Cypselae obcompressed,
obovoid to obpyriform, black to blackish purple,
sometimes drupe-like with juicy exocarp, glabrous
to variously pubescent especially at apices. Pappus
absent. x = 16. About 24 species, neotropics.
1262. Damnxanthodium Strother
Damnxanthodium Strother, Syst. Bot. 12: 41 (1987).
450
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Erect to slightly decumbent perennial herbs. Leaves
opposite, blades ovate to elliptic, 3–5-plinerved.
Capitula solitary on long peduncles or in simple
dichasial cymes, radiate. Involucres campanulate
to hemispherical, phyllaries in 2 series, subequal,
herbaceous. Receptacles convex. Ray florets pistillate, corollas golden yellow to yellow-orange. Disc
florets bisexual, corollas yellow, with fibres embedding vascular strands, lobes densely papillose on
adaxial surfaces; anthers black, appendages without glandular trichomes; style arms tapered, apices
papillose. Cypselae obliquely pyriform to oblanceolate in outline, black to brownish black, glabrous
with a few trichomes around neck. Pappus absent.
x = 12. One species, D. calvum (Greenm.) Strother,
western Mexico.
3–4 series. Receptacles convex. Ray florets neuter
or pistillate, corollas golden to bright yellow. Disc
florets bisexual, corollas yellow without fibres embedding vascular strands; anthers black or brown,
appendages ovate with glandular trichomes;
style arms tapered, apices papillose. Cypselae
compressed, biconvex (ray cypselae when present
triquetrous), obovate in outline, brown to black,
tuberculate, shallowly to prominently winged,
rarely without wings. Pappus of 2–3 awns fused
with the wings and forming a shallow cup around
the corolla tube. Approximately 26 species, Bolivia,
Paraguay, Argentina, Brazil.
1263. Delilia Spreng.
Erect or decumbent annual or perennial herbs.
Leaves opposite, blades lanceolate, narrowly ovate
Delilia Spreng., Bull. Sci. Soc. Philom. Paris 10: 54 (1823);
Delprete, Pl. Syst. Evol. 194: 111–122 (1995), rev.
Elvira Cass. (1824).
1265. Eclipta L.
Fig. 93
Eclipta L., Mant. Pl. 2: 157, 286 (1771), nom. cons.
Erect or decumbent annual herbs. Leaves opposite,
blades ovate to lanceolate, triplinerved. Capitula in
terminal and axillary umbelliform cymes, radiate.
Involucres flattened, disc-like, plano-compressed,
phyllaries 2–4, herbaceous, one orbicular the
others suborbicular, ovate to obovate. Ray florets
1(–3), pistillate, corollas yellow. Disc florets
1(–4), functionally staminate, corollas yellow,
with fibres embedding vascular strands; anthers
black, appendages with glandular trichomes;
style arms of ray floret spreading with divided
stigmatic surfaces. Cypselae triquetrous, obovate
in outline, glabrous with a few trichomes around
apices. Pappus absent. x = 12. Two species, one
widespread in tropical America, the other endemic
to the Galápagos Islands.
1264. Dimerostemma Cass.
Dimerostemma Cass., Bull. Sci. Soc. Philom. Paris 11
(1817); Robinson, Proc. Biol. Soc. Wash. 97: 961–969
(1984), circumscr.; Robinson, Proc. Biol. Soc. Wash. 97:
618–626 (1984), new spp.; Dias de Moraes, Ph.D. Thesis,
Universidade Estadual Campinas (2004), rev.
Angelphytum Barroso (1980).
Erect annual or perennial herbs, shrubs. Leaves
opposite or alternate, petiolate or subsessile,
blades narrowly lanceolate to ovate, sometimes
subcordate, triplinerved. Capitula solitary or in
paniculiform cymes, discoid or radiate. Involucres
broadly campanulate to hemispheric, phyllaries in
Fig. 93.
Compositae-Heliantheae. Eclipta prostrata.
A Habit. B Capitulum in flower. C Capitulum in fruit.
D Capitulum past flower with persistent palea on receptacle.
E Cypsela. F Ray corolla. G Disc corolla. H Anthers. I Style
arms. J Palea. (Reproduced from Flora Novo-Galiciana,
vol. 12, with permission of the University of Michigan
Herbarium; see McVaugh 1984)
Compositae
or oval, pinnately veined or triplinerved. Capitula
axillary or terminal, solitary or in simple cymes,
radiate. Involucres campanulate to hemispherical,
phyllaries in 2–3 series, subequal. Receptacles
shallowly convex, paleae filiform. Ray florets
pistillate, corollas white to pale yellow, linear,
apices bilobed or entire. Disc florets 4(–5)-lobed,
bisexual, corollas white or yellow, without fibres
embedding vascular strands; anthers yellow or
black, appendages with or without glandular
trichomes; style arms acuminate. Cypselae compressed (those of the ray sometimes triquetrous
and obcompressed), shallowly 3–4-angled, tuberculate to rugose, dark brown, glabrous, sometimes
broadly winged or with corky margins, with a few
simple trichomes around neck. Pappus a minute
corona sometimes with 2 teeth at angles of cypsela
or of 2 awns fused to the wings. x = 11. Five
species, America, Australia, introduced elsewhere.
1266. Elaphandra Strother
Elaphandra Strother, Syst. Bot. Monogr. 33: 17 (1991);
Robinson, Phytologia 72: 144–151 (1992), new comb.;
Pruski, Novon 6: 404–418 (1996), reg. rev.
Perennial herbs or shrubs. Leaves opposite,
petiolate, blades lanceolate to ovate, triplinerved.
Capitula in terminal, simple dichasial or open
paniculiform cymes, radiate, rarely discoid.
Involucres hemispherical, phyllaries in 2–3 series,
subequal to gradate. Receptacles flat to convex. Ray
florets neuter, corollas yellow to yellow-orange.
Disc florets bisexual, corollas yellow or blackish
green with yellow lobes, with fibres embedding
vascular strands; anthers black, appendages rarely
with glandular trichomes; style arms tapered,
apices papillose. Cypselae compressed, shallowly
quadrate, sometimes base narrowed or stipitate
and then cypselae narrowly obpyriform, dark
brown to black or reddish brown, glabrescent to
moderately pubescent. Pappus absent or a bicornute, minute crown. x = 13, 14. Fourteen species,
Panama, tropical Andes of South America.
1267. Eleutheranthera Poit. ex Bosc.
Eleutheranthera Poit. ex Bosc., Bull. Sci. Soc. Philom. Paris
3: 137 (1802); Lawalrée, Bull. Jard. Bot. Etat Bruxelles 17: 59
(1943), reg. rev.
Gymnolomia Kunth (1818).
Erect annual herbs. Leaves opposite, petiolate,
blades ovate to trullate, triplinerved. Capitula
axillary, simple, nodding, radiate or discoid.
451
Involucres campanulate, phyllaries in 1 series,
herbaceous. Receptacles flat. Ray florets neuter,
corollas yellow to yellow-orange. Disc florets
bisexual or functionally staminate (?), corollas
yellow, without fibres embedding vascular strands,
sometimes only 2 or 3 florets per head producing
fertile cypselae; anthers black or golden brown,
appendages with glandular trichomes; style arms
tapered, apices papillose. Cypselae shallowly
compressed, shallowly quadrate to terete, black,
tuberculate. Pappus a small lacerate, tubular
crown. x = 10, 15. Two species, neotropical,
adventive in tropical regions of the Old World.
1268. Exomiocarpon Lawalrée
Exomiocarpon Lawalrée, Bull. Jard. Bot. Etat Bruxelles 17:
62 (1943).
Erect annuals. Leaves opposite, petiolate, triplinerved. Capitula axillary and terminal, solitary or
in paniculiform cymes, discoid, rarely radiate.
Involucres campanulate, phyllaries in 1–2 series,
herbaceous. Receptacles flat to shallowly convex.
Ray florets when present neuter, corollas yellow.
Disc florets 5(–4)-lobed, bisexual, corollas yellow,
with fibres embedding vascular strands at base of
throat; anthers black, appendages ovate with glandular trichomes; style arms deltate with densely
papillose apices. Cypselae obcompressed (outermost) or compressed (innermost), black or greyish
black and mottled, obscurely winged, wings shallowly tuberculate, moderately pubescent. Pappus
a small crown of minute, lacerate, filiform scales.
One species, E. madagascariense (Humbert)
Lawalrée, Madagascar.
1269. Fenixia Merr.
Fenixia Merr., Philipp. J. Sci., Bot. 12: 119 (1917).
Erect to prostrate annual herbs. Leaves opposite, petiolate, triplinerved. Capitula terminal,
solitary or in simple dichasial cymes, radiate.
Involucres campanulate, phyllaries in 2 series,
herbaceous. Receptacles flat, epaleate. Ray florets
2, sterile, corollas yellow. Disc florets 5, bisexual,
corollas tetramerous, yellow; anthers 4, black,
appendages ovate; style arms tapered. Cypselae
shallowly obcompressed to shallowly quadrate,
winged, obovate, rugose, moderately pubescent.
Pappus absent. One species, F. pauciflora Merr.,
Philippines.
452
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
1270. Hoffmanniella Schltr ex Lawalrée
Hoffmanniella Schltr ex Lawalrée, Westafr. KautschukExped. 325 (1900); Bull. Jard. Bot. Etat Bruxelles 17: 59
(1943).
Prostrate to erect annual herbs. Leaves opposite, petiolate, triplinerved. Capitula terminal,
appearing axillary, solitary, sessile, discoid. Involucres cylindrical with only 2 opposite phyllaries.
Receptacles flat. Florets 6, bisexual, corollas
(3–)4-lobed, yellow, those subtended by phyllaries
sometimes slightly zygomorphic, without fibres
embedding vascular strands, sometimes a few
fibrous cells flanking veins on distal area of throat;
anthers black, free, appendages without glandular
trichomes; style arms with round to subacute,
smooth apices. Cypselae obcompressed, broadly
obovate, depressed obovate in cross-section,
brown to black, edges thickened, stramineous,
glabrous except for densely pubescent apices.
Pappus a minute, pubescent crown. One species,
H. sylvatica Schltr ex Lawalrée, tropical west
Africa.
1271. Idiopappus H. Rob. & Panero
Idiopappus H. Rob. & Panero, Syst. Bot. 19: 359–362 (1994).
Small trees. Leaves opposite, petiolate, broadly
ovate to subcordate, triplinerved. Capitula in
terminal, open paniculiform to thyrsoid cymes,
radiate. Involucres hemispherical to shallowly
patent with age, phyllaries in 3 series, subequal.
Receptacles conical to subglobose with age. Ray
florets pistillate, corollas yellow. Disc florets bisexual, corollas yellow, without fibres embedding
vascular strands, the throats expanding conspicuously above tube, subcampanulate, lobes coiled;
anthers black, appendages without glandular
trichomes; style arm apices deltate. Cypselae
compressed, sometimes those of the ray trigonous,
those of the disc shallowly quadrate, somewhat
navicular, shallowly stipitate with curved bases,
black, glabrous. Pappus of 1 or mostly 2, rarely 3
squamellae at main angles of cypsela. x = 31–33.
One species, I. quitensis H. Rob. & Panero, Ecuador.
1272. Iogeton Strother
Iogeton Strother, Syst. Bot. Monogr. 33: 20 (1991).
Stoloniferous perennial herbs or weak shrubs.
Leaves opposite, petiolate, blades narrowly
lanceolate, pinnately nerved. Capitula terminal,
solitary or in lax dichasial cymes, radiate. In-
volucres narrowly obconic, phyllaries subequal.
Receptacles conical or hemispherical. Ray florets
pistillate, corollas yellow. Disc florets bisexual,
corollas yellow; anthers brown, appendages ovate;
style arms tapered. Immature ray cypselae and
outermost disc cypselae triquetrous, brown to
black. Innermost disc cypselae quadrate, brownish
to black, hispidulous on the angles. Pappus of 2–4
slender, persistent, minutely barbellulate awns
sometimes with 1–3 minute scales. One species,
I. nowickeanus (D’Arcy) Strother, Panama.
1273. Jefea Strother
Jefea Strother, Syst. Bot. Monogr. 33: 22 (1991).
Basally woody perennials or shrubs. Leaves
opposite or alternate, petiolate, blades deltate,
ovate or lanceolate, triplinerved, sometimes
densely tomentose beneath. Capitula terminal,
solitary, radiate. Involucres broadly campanulate
to hemispheric, phyllaries dimorphic, outermost
leaf-like and spreading. Receptacles strongly convex to conic. Ray florets pistillate, corollas yellow
to yellow-orange. Disc florets bisexual, corollas
yellow to orangish, without fibres embedding the
vascular strands; anthers yellow, brown or black,
appendages lanceolate with glands; style arms
tapered, apices papillose. Ray cypselae triquetrous,
disc cypselae compressed, oblanceolate, wings
continuous or shallowly lacerate, often corky.
Pappus of 2–3 slender, unequal, caducous or
persistent awns and 2–10 or more much shorter,
free or basally connate squamellae, all inserted on
apices of cypselae, not raised on a rostrum. x = 14.
Five species, Mexico, Guatemala.
1274. Kingianthus H. Rob.
Kingianthus H. Rob., Phytologia 38: 415 (1978); Robinson,
Phytologia 44: 70–78 (1979), new spp.
Shrubs. Leaves alternate, petiolate, blades ovate,
triplinerved. Capitula in terminal, paniculiform
cymes of congested corymbiform cymes, radiate. Involucres campanulate, phyllaries in 1–2
series, subequal. Receptacles convex. Ray florets
pistillate, corollas light to golden yellow. Disc
florets bisexual, corollas yellow to orange-yellow,
without fibres embedding the vascular strands;
anthers black, appendages with glands, sometimes
glandular trichomes scattered lengthwise along the
connective; style branch apices deltate. Cypselae
quadrate, slightly navicular, shallowly stipitate
with curved bases, black, glabrous. Pappus of 1
Compositae
squamella or absent. x = 15, 16, 17. Two species,
Ecuador, Peru.
1275. Lantanopsis C. Wright ex Griseb.
Lantanopsis C. Wright ex Griseb., Mem. Amer. Acad. Arts
n.s. 8: 513 (1862).
Annual or perennial herbs, sometimes weak
shrubs. Leaves opposite, petiolate, blades lanceolate to ovate, triplinerved. Capitula in terminal,
congested corymbiform or pseudoscorpioid
cymes, disciform. Involucres cylindrical to campanulate, phyllaries 4, in 2 series, subequal.
Receptacles flat, epaleate. Florets 3, unisexual and
dimorphic, the single pistillate floret tubular and
the corolla with 2 or 3 lobes, the two functionally
staminate florets with a tetramerous corolla, corollas white, without fibres embedding the vascular
strands; anthers black, appendages with glandular
trichomes; style arms tapered and at least 10 times
as long as style. Cypselae obcompressed, obovoid
to obpyriform, black or brown, densely covered
with glandular and a few tapered trichomes
especially on distal ends. Pappus absent. Three
species, Cuba and Hispaniola.
1276. Lasianthaea DC.
Lasianthaea DC., Prodr. 5: 607 (1836); Becker, Mem. New
York Bot. Gard. 31: 1–64 (1979), rev.
Perennial herbs or shrubs, rarely small trees.
Leaves opposite, petiolate or subsessile, blades
lanceolate to ovate or elliptic, triplinerved. Capitula
terminal, solitary or in open to congested corymbiform or paniculiform cymes, radiate. Involucres
cylindric to hemispheric, phyllaries in 3–5 series,
subequal to imbricate, distal apices sometimes
chartaceous, red, purple or yellow. Receptacles
convex. Ray florets pistillate, corollas yellow,
orange, vermillion, purple, sometimes lighter in
colour abaxially. Disc florets bisexual, corollas
yellow, orange, reddish-orange, purple, with fibres
embedding the vascular strands; anthers brown or
black, appendages without glandular trichomes;
style arms tapered, apices papillose. Ray cypselae
obcompressed, triquetrous, those of disc compressed, cuneate to slightly oblong in outline,
black with hyaline or greenish edges, glabrous.
Pappus of 2–3 awns continuous with cypsela edges,
sometimes with a few free or connate squamellae.
x = 8, 10, 11. Twelve species, south-western USA,
Mexico, Central America, Venezuela.
453
1277. Leptocarpha DC.
Leptocarpha DC., Prodr. 5: 495 (1836).
Perennial herbs or weak shrubs. Leaves opposite
or alternate, blades ovate, triplinerved. Capitula
in terminal, simple dichasia or open paniculiform
cymes, radiate. Involucres hemispherical to patent
with age, phyllaries in 2 series. Receptacles convex with linear paleae. Ray florets neuter or pistillate but sterile, corollas yellow. Disc florets bisexual, corollas yellow, without fibres embedding
the vascular strands; anthers brown, appendages
with minute glandular trichomes; style arms small,
apices deltate, not papillose. Cypselae compressed,
biconvex to triquetrous, obovate, black, sparsely
pubescent. Pappus of 2–4 small squamellae. One
species, L. rivularis DC., Chile.
1278. Lipochaeta DC.
Lipochaeta DC., Prodr. 5: 610 (1836); Gardner, Rhodora 81:
291–343 (1979), reg. rev.; Wagner & Robinson, Brittonia 53:
539–561 (2002), delim.
Annual or perennial herbs, weak shrubs. Leaves
opposite, petiolate or sessile, rarely ternate, blades
ovate to elliptic, sometimes filiform, mostly
triplinerved. Capitula terminal, solitary or in
open paniculiform cymes, radiate. Involucres
campanulate to hemispherical, phyllaries in 2
series, normally five in each. Receptacles convex,
paleae sometimes deciduous. Ray florets pistillate,
corollas yellow. Disc florets bisexual, corollas 4or 5-lobed, yellow, without fibres embedding the
vascular strands; anthers brown, appendages with
glandular trichomes; style arms deltate, apices
shallowly papillose. Ray cypselae obcompressed,
triquetrous, black, tuberculate, with lacerate
wings. Disc cypselae compressed, biconvex,
oblong to obcuneate, otherwise as ray cypselae.
Pappus of connate scales forming a corona,
sometimes with caducous awns and intervening
squamellae. x = 15, 26. Twenty species, Hawaii.
1279. Lundellianthus H. Rob.
Lundellianthus H. Rob., Wrightia 6: 41 (1978); Strother,
Syst. Bot. 14: 544–548 (1989), delim.
Shrubs. Leaves opposite, petiolate to subsessile,
blades ovate-elliptic to ovate, triplinerved. Capitula
terminal, solitary or in simple dichasia, sometimes
in open paniculiform cymes, radiate. Involucres
campanulate to hemispherical, phyllaries in 2–3
series, outermost foliaceous. Receptacles convex,
454
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
tuberculate, paleae fused laterally. Ray florets
pistillate, corollas yellow to orange-yellow. Disc
florets bisexual, corollas yellow to yellow-orange,
with or without fibres embedding the vascular
strands; anthers black, appendages with or without
glandular trichomes; style arms tapered, apices
papillose. Ray cypselae triquetrous, oblanceolate
in outline, black or stramineous, glabrous or
sparsely pubescent, shallowly to prominently
winged, wings corky, stramineous, in some species
edges of wings fused and producing a tuberculate,
stramineous cover on faces of cypsela. Disc cypselae compressed, primarily biconvex but with small
ridges on faces rendering them shallowly quadrate
and rhombic in outline. Pappus of 2 or 3 broad,
lacerate awns at main angles of cypsela, with or
without squamellae in between, awns sometimes
fused with wings. x = 16. Eight species, Mexico,
Central America.
1280. Melanthera Rohr
Melanthera Rohr, Skr. Naturhist.-Selsk. 2: 213 (1792); Wild,
Kirkia 5: 1–17 (1965), reg. rev.; Parks, Rhodora 75: 169–210
(1973), reg. rev.; Wagner & Robinson, Brittonia 53: 539–561
(2002), delim.
Annual or perennial herbs, scandent or erect
shrubs. Leaves opposite, blades narrowly obovate
to linear-lanceolate, ovate, triplinerved, unlobed,
trilobed or pinnately lobed. Capitula in terminal,
open paniculiform cymes, rarely solitary, discoid
or radiate. Involucres campanulate to hemispherical, phyllaries in 1–2 series, subequal. Receptacles
convex. Ray florets pistillate but sterile or neuter,
corollas yellow. Disc florets bisexual, corollas
yellow or white, without fibres embedding the vascular strands, sometimes with a few sclerified cells
along vascular strands; anthers black, appendages
with glandular trichomes; style arms tapered to
acute, apices shallowly papillose. Ray cypselae
obcompressed, triquetrous to quadrate, obovate
to obpyramidal, black, apices truncate, sometimes
winged on angles of cypsela, glabrescent. Disc
cypselae compressed, obconical, somewhat quadrangular, otherwise as ray cypselae. Pappus of
several caducous awns. x = 15, 16. Twenty species,
pantropical.
1281. Monactis Kunth
Monactis Kunth in Humb., Bonpl. & Kunth, Nov. Gen. Sp.
folio ed. 4: 225 (1818); Robinson, Phytologia 34: 33–45
(1976), notes; Robinson, Phytologia 44: 70–78 (1979), new
spp.
Shrubs or small trees. Leaves alternate, petiolate,
blades lanceolate to broadly ovate or trullate,
pinnately veined or triplinerved. Capitula in
terminal, congested to open corymbiform cymes,
radiate or discoid, only one or two ray florets per
head, ray florets towards the outside of congested
corymbiform cymes. Involucres cylindrical to
campanulate, phyllaries in 1–2 series, gradate.
Receptacles flat to shallowly convex. Ray florets pistillate, corollas yellow to orange-yellow.
Disc florets bisexual, corollas white, greenish
white, yellow or orange-yellow, without fibres
embedding the vascular strands, the throats
expanding conspicuously above tube, campanulate to subhemispheric, lobes coiled; anthers
black, appendages with glandular trichomes,
sometimes these scattered lengthwise along the
connective; style arm apices deltate. Cypselae shallowly triquetrous or quadrate, fusiform, slightly
asymmetrical, sometimes geniculate at apices,
black, glabrous. Pappus of 1 rarely 2 squamellae
or absent. x = 27, 28, 30, 32. About 10 species,
Ecuador and Peru.
1282. Oblivia Strother
Oblivia Strother, Syst. Bot. 14: 541 (1989); Anderson et al.,
Syst. Bot. 4: 44–56 (1979), phylog.; Villaseñor & Strother,
Syst. Bot. 14: 529–540 (1989), syst.; Robinson, Phytologia
76: 24–26 (1994), new spp.
Scandent shrubs. Leaves opposite, petiolate, blades
lanceolate-elliptic to ovate, 3–5-plinerved. Capitula in terminal, congested corymbiform cymes,
radiate. Involucres campanulate, phyllaries in 2–
3 series, subequal. Receptacles convex. Ray florets
pistillate, corollas dull yellow. Disc florets bisexual, corollas yellow, without fibres embedding the
vascular strands; anthers black, appendages with
glandular trichomes; style arms tapered. Cypselae
obcompressed, triquetrous, narrowly oblanceolate
in outline, shallowly winged. Disc cypselae compressed, otherwise as ray cypselae. Pappus of 2(–4)
awns and a few squamellae in between. x = 16.
About three species, Panama, tropical Andes.
1283. Otopappus Benth.
Otopappus Benth. in Benth. & Hook. f., Gen. Pl. 2: 196, 380
(1873); Hartman & Stuessy, Syst. Bot. 8: 185–210 (1983),
rev.; Villaseñor & Strother, Syst. Bot. 14: 529–540 (1989),
new spp.
Shrubs or small trees. Leaves opposite, petiolate,
blades lanceolate to ovate, 2–7-plinerved. Capit-
Compositae
ula terminal, solitary or in small paniculiform to
corymbiform cymes, radiate or discoid. Involucres campanulate to hemispherical, phyllaries in
3–7 series, gradate. Receptacles convex to conic.
Ray florets pistillate, rarely neuter, corollas yellow to yellow-orange. Disc florets bisexual, corollas yellow, rarely white, without fibres embedding
the vascular strands; anthers black or deep purple, appendages with glandular trichomes; style
arms tapered. Cypselae compressed, peripheral triquetrous, otherwise oblanceolate to obovate in outline, black or brown, glabrous to puberulent, especially at apices, with 1–3 wings of unequal width.
Pappus of 1–3 awns with connate or free minute
squamellae in between, awns fused to wings. x =
16. Fourteen species, Mexico, Central America.
1284. Oyedaea DC.
Oyedaea DC., Prodr. 5: 576 (1836); Blake, Contr. U.S. Natl
Herb. 20: 411–422 (1921), rev.; Robinson, Wrightia 6: 43–48
(1979), reg. rev.; Pruski, Novon 6: 404–418 (1996), reg. rev.
Shrubs or small trees. Leaves opposite, petiolate,
blades elliptic to ovate, pinnately veined or
triplinerved. Capitula terminal, in simple or open
paniculiform cymes, radiate. Involucres campanulate to hemispherical, phyllaries in 2–5 series.
Receptacles shallowly convex to convex. Ray florets
neuter, corollas yellow, rarely white. Disc florets
bisexual, corollas yellow, rarely white, without
fibres embedding the vascular strands; anthers
black, appendages with or without glandular
trichomes; style arms tapered, apices papillose
with appendages. Cypselae oblong, compressed,
winged at maturity. Pappus of 2 subequal awns
on a rostrum, sometimes with intermediate
squamellae. x = 14. Eighteen species, Central and
South America.
1285. Pascalia Ortega
Pascalia Ortega, Novarum, aut Rariorum Plantarum Horti
Reg. Bot. Matrit. 39 (1797); Strother, Syst. Bot. Monogr. 33:
1–111 (1991), rev.
Stoloniferous perennial herbs. Leaves opposite,
sessile to shortly petiolate, blades lanceolate
to broadly ovate or oval, triplinerved. Capitula
terminal, solitary or in simple dichasia, radiate.
Involucres hemispherical, phyllaries in 2–3 series.
Receptacles convex to hemispherical. Ray florets
pistillate, sometimes with staminodes, corollas
yellow-orange. Disc florets bisexual, corollas
yellow-orange, without fibres embedding the
455
vascular strands; anthers yellow or brown, appendages without glandular trichomes; style arms
acute, apices papillose. Cypselae compressed,
broadly biconvex to quadrate, broadly obcuneate
in outline, neck broad and flat, rhombic in outline,
edge sinuate, brown, glabrous. Pappus a minute
crown. x = 33. Two species, eastern South America,
north-central Chile.
1286. Pentalepis F. Muell.
Pentalepis F. Muell., Trans. Bot. Soc. Edinburgh 7: 496
(1863); Karis et al., Austral. Syst. Bot. 6: 149–153 (1993),
re-establ.
Erect annuals or shrubs. Leaves opposite, blades
lanceolate to narrowly ovate, triplinerved. Capitula
in terminal, congested corymbiform cymes or
open paniculiform cymes, radiate. Involucres
campanulate, phyllaries in 2 series. Receptacles
shallowly convex. Ray florets pistillate, corollas
yellow. Disc florets functionally staminate, corollas
yellow, without fibres embedding the vascular
strands; anthers yellow to brown, appendages
without glandular trichomes; style arms of disc
florets fused, style arms of ray florets with divided
stigmatic surfaces, without appendages. Cypselae
obcompressed, obovate, shallowly winged, with
a few tuberculae on edge of wings, brown-black,
glabrous with a few trichomes on neck. Pappus
a minute crown. About three species, Australia.
1287. Perymeniopsis H. Rob.
Perymeniopsis H. Rob., Phytologia 40: 495 (1978).
Scandent shrubs. Leaves opposite, petiolate, blades
oblong, triplinerved. Capitula in terminal, open
paniculiform cymes, radiate. Involucres campanulate to hemispherical, phyllaries in 2–3 series. Receptacles convex. Ray florets neuter, corollas golden
bright yellow. Disc florets bisexual, corollas yellow, without fibres embedding the vascular strands;
anthers brown or black, appendages ovate with
minute glandular trichomes; style arms tapered,
apices papillose. Cypselae compressed, obovate,
black or brown, shallowly winged, wings wider
at distal apices, cypsela bases narrowed, essentially glabrous. Pappus of 2 easily caducous awns
at angles of cypsela and a small rostrum bearing numerous minute and easily caducous awns.
One species, P. ovalifolia (A. Gray) H. Rob., eastern
Mexico.
456
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
1288. Perymenium Schrad.
Perymenium Schrad., Index Sem. (Göttingen) 1830: 4
(1830); Fay, Allertonia 1: 235–296 (1978), part. rev.
Perennial herbs, shrubs or small trees. Leaves
opposite, blades lanceolate to ovate sometimes
cordiform, triplinerved, rarely pinnately veined.
Capitula in terminal, simple dichasia or open paniculiform cymes, radiate. Involucres hemispherical,
phyllaries in 2–4 series. Receptacles convex. Ray
florets pistillate, corollas golden yellow. Disc
florets bisexual, corollas yellow, without fibres
embedding the vascular strands, some sclerified
cells scattered throughout throat; anthers brown or
black, appendages with glandular trichomes; style
arms deltate to tapered, apices shallowly papillose.
Ray cypselae obcompressed, triquetrous, obovate
to oblong, those of the disc compressed, biconvex,
obovate, black, variously pubescent, shallowly to
broadly winged, minutely tuberculate, tuberculae
collectively appearing as shiny lines. Pappus of 0–3
awns at angles of cypselae and a rostrate crown of
numerous unequal, easily caducous awns. x = 15.
About 43 species, neotropics.
1289. Plagiolophus Greenm.
Plagiolophus Greenm., Publ. Field Columb. Mus., Bot. Ser.
3: 125 (1904).
Erect annuals. Leaves opposite, blades ovate,
triplinerved. Capitula in terminal, open paniculiform cymes, discoid. Involucres hemispheric,
phyllaries in 2 series, outermost 5, spreading,
innermost appressed. Receptacles shallowly
convex. Florets bisexual, corollas white, without
fibres embedding the vascular strands; anthers
black, appendages with glandular trichomes; style
arms tapered, acuminate. Cypselae compressed,
obovate, with geniculate, eccentric necks, winged
at maturity. Pappus of 2 subequal awns and a few
squamellae in between on a crown at apex of
neck. One species, P. millspaughii Greenm., central
Yucatan, Mexico.
1290. Podanthus Lag.
Podanthus Lag., Gen. Sp. Pl. 24 (1816), nom. cons.
Dioecious shrubs, sometimes with purplish stems.
Leaves opposite, blades ovate to trullate. Capitula
terminal, solitary, or in simple or leafy paniculiform cymes, discoid. Involucres patent or reflexed,
phyllaries in 1–2 series, subequal. Receptacles convex to globose. Corollas yellow or green-yellow,
without fibres embedding the vascular strands, dimorphic, those of functionally pistillate florets approximately half the size of those of functionally
staminate florets and with much shorter tubes; anthers dark yellow or brown with black connectives and appendages, appendages without glandular trichomes; style arms small, apices deltate
to acute, not papillose. Cypselae compressed to
quadrate, obpyramidal, bases slender to narrowed.
Pappus a minute crown of squamellae or absent.
Two species, central Chile.
1291. Rensonia S.F. Blake
Rensonia S.F. Blake, J. Wash. Acad. Sci. 13: 144 (1923).
Shrubs. Leaves opposite, petiolate, blades ovate to
oval, triplinerved. Capitula in terminal, paniculiform cymes, radiate. Involucres campanulate, phyllaries in 2 series, subequal. Receptacles flat to shallowly convex. Ray florets pistillate, corollas golden
yellow. Disc florets functionally staminate, corollas
golden yellow with fibres embedding the vascular
strands; anthers black, sometimes shallowly calcarate, appendages without glandular trichomes;
style arms of disc florets fused except for distal end.
Cypselae obovate, glabrous, conspicuously winged.
Pappus of 2 awns fused to wings. x = 17. One
species, R. salvadorica S.F. Blake, Mexico, Central
America.
1292. Riencourtia Cass.
Riencourtia Cass., Dict. Sci. Nat. 45: 466 (1827).
Annual or perennial herbs. Leaves opposite,
petiolate or sessile, blades linear to elliptic or
ovate, 1–5-veined. Capitula in terminal, open paniculiform cymes of tightly clustered glomerule-like
cymes, disciform. Involucres cylindrical, phyllaries
in 3–4 series, subequal. Receptacles flat, epaleate.
Ray/tubular floret 1, pistillate, corolla white,
tubular or with a very reduced limb. Disc florets
functionally staminate and campanulate, the lobes
with conspicuously long and stiff trichomes,
without fibres embedding the vascular strands;
anther appendages without glandular trichomes;
style arms tapered. Cypselae broadly ellipsoid,
biconvex, black or brown with a broad stramineous
carpopodium on adaxial side, glabrous to sparsely
pubescent especially on apices. Pappus absent.
x = 16. About six species, neotropics, eastern
South America.
Compositae
1293. Schizoptera Turcz.
Schizoptera Turcz., Gen. Pl. 2: 191, 349 (1873).
Erect annual herbs. Leaves opposite, petiolate,
blades ovate to oval, apices acuminate, triplinerved.
Capitula in paniculiform cymes with subumbellate paracladia, on pedicels of various lengths,
radiate. Involucres campanulate, phyllaries in
2 series, subequal. Receptacles flat to shallowly
convex. Ray florets pistillate, in two rows, corollas
golden yellow. Disc florets functionally staminate,
rarely perfect, corollas golden yellow with fibres
embedding the vascular strands; anthers black,
appendages ovate; style arms of disc florets with
no or very few stigmatic papillae, style arms of ray
florets spreading. Ray cypselae broadly obovate,
obcompressed, black, concave, glabrous, margins
winged. Disc cypselae when present similar to ray
cypselae. Pappus of two awns fused to wings. One
species, S. peduncularis S.F. Blake, Ecuador.
1294. Sphagneticola O. Hoffm.
Sphagneticola O. Hoffm., Notizbl. Konigl. Bot. Gart. BerlinDahlem 3: 36 (1900); Strother, Syst. Bot. Monogr. 33: 1–111
(1991), rev.; Pruski, Novon 6: 404–418 (1996), reg. rev.
Thelechitonia Cuatrec. (1954).
Complaya Strother (1991).
Perennial herbs, prostrate, rooting at nodes. Leaves
opposite, blades ovate to trullate, trilobed, triplinerved. Capitula terminal, appearing axillary, solitary, radiate. Involucres turbinate, phyllaries in 2
series, subequal, apices foliaceous, expanded. Receptacles convex to conical. Ray florets pistillate,
corollas golden yellow to orange. Disc florets bisexual, corollas yellow to orange, with long multicellular trichomes on adaxial surface of lobes,
without fibres embedding the vascular strands; anthers black, appendages with glandular trichomes;
style arms tapered. Ray cypselae trigonal, clavate
to obpyriform, winged, wings corky, stramineous,
thicker at the base of cypselae, smooth to strongly
tuberculate, glabrous. Disc cypselae compressed,
biconvex to rounded-quadrate, otherwise as ray
cypselae. Pappus an erose or lacerate corona on
a stout rostrum, the rostrum and pappus sometimes covered by a corky collar continuous with
the cypsela body. x = 15. Four species, pantropical.
1295. Steiractinia S.F. Blake
Steiractinia S.F. Blake, J. Bot. 53: 153 (1915); Díaz-Piedrahita
& Vélez-Nauer, Asteraceae-Heliantheae 1. Steiractinia, Fl.
Colomb. 13: 1–65 (1990), reg. rev.
457
Shrubs or small trees. Leaves opposite, blades
lanceolate to ovate, pinnate or triplinerved.
Capitula in terminal, simple or open paniculiform cymes, radiate. Involucres hemispherical,
phyllaries in 2–4 series. Receptacles convex.
Ray florets neuter, corollas golden yellow. Disc
florets bisexual, corollas yellow with a few fibres
embedding the vascular strands; anthers brown
or black, appendages with or without glandular
trichomes; style arms tapered, apices papillose
with appendages. Cypselae compressed or obcompressed, shallowly biconvex to quadrate, oblong or
obovoid, shallowly to broadly rhombic in outline,
black to greyish black and mottled or striped,
mostly with 2, rarely 3 or 4 wings. Pappus of
numerous bristles disposed radially on neck of
cypsela. x = 14. Twelve species, Colombia, Ecuador
and Venezuela.
1296. Synedrella Gaertn.
Synedrella Gaertn., Fruct. Sem. Pl. 2: 456 (1791), nom.
cons.; McVaugh & Smith, Brittonia 19: 268–272 (1967);
Turner, Phytologia 76: 39–51 (1994), rev.
Erect to slightly decumbent annual or perennial
herbs. Leaves opposite, shortly petiolate, blades
ovate to elliptic, triplinerved. Capitula in axillary,
simple cymes, radiate. Involucres cylindrical,
phyllaries in 2 series. Receptacles flat. Ray florets
pistillate, corollas yellow. Disc florets bisexual,
corollas yellow with fibres embedding the vascular strands; anthers black, appendages without
glandular trichomes; style arms tapered, with papillose appendages. Ray cypselae obcompressed,
biconvex, oblong, black, winged, wings deeply
lacerate, stramineous, glabrous. Disc cypselae obcompressed, biconvex, obcuneate, black, slightly
tuberculate, glabrous. Pappus of 2 awns, in ray
cypselae fused to wings, in disc cypselae stout.
x = 18, 19, 20. One species, S. nodiflora Gaertn.,
neotropics, adventive in other tropical regions.
1297. Synedrellopsis Hieron. & Kuntze
Synedrellopsis Hieron. & Kuntze, Rev. Gen. 3: 180 (1898);
Cabrera, Fl. Prov. de Jujuy, Rep. Argent. 10 (1978), rev.
Prostrate to decumbent annual or perennial herbs
rooting at the nodes. Leaves opposite, petiolate,
blades ovate to trullate, triplinerved. Capitula
axillary, solitary, discoid. Involucres cylindrical,
phyllaries 2. Receptacles flat, epaleate. Florets
4, those subtended by the opposing phyllaries
tubular, pistillate, corollas 3- or 4-lobed, the
458
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
internal bisexual, corollas 4-lobed, yellow and
uniformly thickened with sclerified cells; anthers
black, appendages without glandular trichomes;
style arms tapered, without papillose appendages.
Cypselae obcompressed, shallowly rhombic in
cross-section, with a shallow ridge on each
face, broadly oval to elliptic, black, wings corky,
stramineous, deeply lacerate, sparsely pubescent,
trichomes uniseriate. Pappus of 2 awns fused
to wings. One species, S. grisebachii Hieron. &
Kuntze, Argentina, Bolivia, Paraguay.
1298. Tilesia G. Mey.
Tilesia G. Mey., Prim. Fl. Esseq. 251 (1818); Pruski, Novon
6: 404–418 (1996), reg. rev.
Wulffia Neck. (1790), nom. non rite public.
Wulffia Neck. ex Cass. (1823), nom. cons. prop.
Perennial herbs, mostly scandent shrubs, stems
striate. Leaves opposite, blades lanceolate, ovate
or oval, pinnately veined or triplinerved. Capitula
in terminal, simple or paniculiform cymes, discoid or radiate. Involucres hemispherical, phyllaries in 2–3 series. Receptacles shallowly convex. Ray
florets neuter or pistillate but sterile, corollas yellow to orange. Disc florets bisexual, corollas yellow,
rarely red, with a few fibres embedding the vascular
strands; anthers black, appendages ovate with glandular trichomes; style arms tapered. Cypselae compressed, obpyriform to obovate, rhombic in crosssection, fleshy at maturity, exocarp drying brown,
black beneath, glabrous to sparsely pubescent at the
apices. Pappus absent or a minute crown. x = 30.
Three species, tropical, humid forests of America.
1299. Trigonopterum Hook. f.
Trigonopterum Hook. f., Q. J. Sci. Lit. Arts ser. 2, 1: 104
(1828); Harling, Acta Horti Bergiani 20: 63–120 (1962),
rev.; Eliasson, Bot. J. Linn. Soc. 88: 253–256 (1984), cytol.;
Wagner & Robinson, Brittonia 53: 539–561 (2002), tax.
Shrubs. Leaves opposite, blades linear and slightly
involute. Capitula terminal, solitary, radiate or
discoid. Involucres campanulate to hemispherical,
phyllaries in 2 series. Receptacles convex. Ray
florets pistillate, corollas yellow. Disc florets
bisexual, corollas yellow without fibres embedding
the vascular strands; anthers black, appendages
without glandular trichomes; style arms tapered,
apices papillose. Cypselae of ray floret triquetrous,
otherwise biconvex, narrowly obpyriform, glabrescent, tuberculate, with 1 wing and 1 or 2 additional
weakly developed wings or commissures. Pappus
a small, rostrate crown. x = 14. One species,
T. laricifolium (Hook. f.) W.L. Wagner & H. Rob.,
Galápagos Islands.
1300. Tuberculocarpus Pruski
Tuberculocarpus Pruski, Novon 6: 415 (1996); Pruski,
Novon 6: 404–418 (1996), rev.
Perennial herbs or subshrubs. Leaves opposite,
blades lanceolate, pinnately veined. Capitula terminal, solitary or in simple cymes, radiate. Involucres
campanulate to hemispherical, phyllaries in 2–3
series. Receptacles flat to shallowly convex. Ray
florets neuter, corollas red. Disc florets bisexual,
corollas orange-red; anthers black, appendages
with glandular trichomes, apices papillose; style
arms tapered. Cypselae compressed to subterete,
with small wings, black, tuberculate, glabrous
with a few trichomes at apices. Pappus a minute
crown. One species, T. ruber (Aristeg.) J.F. Pruski,
southern Venezuela.
1301. Tuxtla Villaseñor & Strother
Tuxtla Villaseñor & Strother, Syst. Bot. 14: 537 (1989).
Lianas or scandent shrubs. Leaves opposite, blades
elliptic, 3–5-plinerved. Capitula in terminal,
corymbiform cymes, radiate. Involucres campanulate, phyllaries in 2–3 series. Receptacles convex.
Ray florets pistillate, corollas yellow. Disc florets
bisexual, corollas yellow with fibres embedding
the vascular strands; anthers brown or black,
appendages with minute glandular trichomes
extending to distal parts of connective; style arms
tapered, apices shortly papillose. Ray cypselae
triquetrous, brown or black with stramineous
wings, glabrous. Disc cypselae compressed,
biconvex, otherwise as ray cypselae. Pappus of
2–3 awns continuous with wings and body of
cypsela. x = 17. One species, T. pittieri (Greenm.)
Villaseñor & Strother, Mexico, Costa Rica.
1302. Wamalchitamia Strother
Wamalchitamia Strother, Syst. Bot. Monogr. 33: 30 (1991).
Shrubs. Leaves opposite, blades ovate to lanceolate, triplinerved. Capitula terminal, solitary or in
open dichasial cymes, radiate. Involucres broadly
cylindric to hemispheric, phyllaries in 2–4 series.
Receptacles flat to convex. Ray florets pistillate,
corollas yellow to red-orange. Disc florets bisexual,
corollas yellow to orange without fibres embedding
the vascular strands; anthers black, appendages
Compositae
without glandular trichomes; style arms tapered,
apices with a long papillose appendage. Cypselae
prismatic, narrowly cuneate to linear in outline,
those of the ray triquetrous, otherwise compressed,
quadrate, rarely 5-angled, not winged, dark reddish
brown to black, glabrous. Pappus of 1–5 unequal,
fragile or persistent awns plus 0–6 minute, free or
strongly connate squamellae, all inserted directly
on apex of cypsela body and not raised on a rostrum. n = 89–96, 100–108, 110. Four species, Mexico and Central America.
1303. Wedelia Jacq.
Wedelia Jacq., Enum. Syst. Pl. 8: 28 (1760), nom. cons.;
Strother, Syst. Bot. Monogr. 33: 1–111 (1991), rev.
Aspilia Thouars (1806).
Erect to prostrate annual or perennial herbs,
shrubs. Leaves opposite, blades lanceolate to ovate,
oval or elliptic, subcordate, usually triplinerved,
sometimes trilobed, rarely deeply serrate. Capitula
terminal, solitary or in open paniculiform cymes,
radiate, rarely discoid. Involucres campanulate to
hemispherical, phyllaries in 2–4 series. Receptacles convex. Ray florets pistillate, fertile or neuter,
corollas yellow to yellow-orange, sometimes white,
pink or burgundy. Disc florets bisexual, corollas
yellow to orange, without fibres embedding the
vascular strands; anthers light to dark brown or
black, appendages with glandular trichomes; style
arms tapered to deltoid, apices papillose. Cypselae
of ray floret obcompressed, triquetrous, obovate or
oblong, grey, brown or black, sometimes mottled,
glabrous to moderately pubescent, winged at
angles of cypselae or without wings, with an
elaiosome. Disc cypselae like those of the ray,
biconvex. Pappus on a rostrum, an erose cup or
corona with 0–3 unequal, persistent awns. x = 11,
12, 13. Approximately 110 species, pantropical,
most species in America.
1304. Wollastonia DC. ex Decne
Wollastonia DC. ex Decne, Nouv. Ann. Mus. Hist. Nat. 3:
414 (1834); Fosberg & Sachet, Smithsonian Contr. Bot. 45:
1–40 (1980), rev.; Wagner & Robinson, Brittonia 53: 539–561
(2002), rev.
Perennial herbs or weak shrubs. Leaves opposite,
blades oval to ovate, triplinerved. Capitula terminal, solitary or in open paniculiform cymes, radiate. Involucres hemispherical, phyllaries in 2 series. Receptacles convex. Ray florets pistillate, yellow. Disc florets bisexual, corollas yellow or green-
459
ish yellow, without fibres embedding the vascular strands; anthers brown to black. Ray cypselae
cuneiform, triquetrous, with truncate, flat apices,
hirtellous on distal end. Disc cypselae compressed
and shallowly quadrate, shallowly rhombic in outline, otherwise as ray cypselae. Pappus absent or
of a single awn. One species, W. biflora (L.) DC.,
Indo-Pacific coastal regions.
1305. Zexmenia La Llave & Lex.
Zexmenia La Llave & Lex., Nov. Veg. Descr. 1: 13 (1824);
Strother, Syst. Bot. Monogr. 33: 1–111 (1991), rev.
Scandent to erect perennial herbs, shrubs or lianas.
Leaves opposite, blades ovate-elliptic to lanceolate, pinnately nerved or triplinerved. Capitula in
terminal, simple dichasia or umbelliform cymes,
radiate. Involucres broadly campanulate to hemispheric, phyllaries in 2–3 series. Receptacles convex. Ray florets pistillate, corollas yellow to yelloworange. Disc florets bisexual, corollas yellow to orange with or without a few fibres embedding the
vascular strands; anthers black, appendages ovate
with a few glandular trichomes; style arms tapered,
apices papillose. Cypselae compressed, those of the
ray triquetrous, oblanceolate to narrowly cuneate,
winged, stramineous to dark brown or blackish,
sparsely pubescent distally, the wings thin to corky.
Pappus on a distinct rostrum, of 2–3 unequal awns
plus 4–10 minute awns often connate to form a lacerate cup. Two species, Mexico and Central America.
1306. Zyzyxia Strother
Zyzyxia Strother, Syst. Bot. Monogr. 33: 91 (1991).
Shrubs or small trees. Leaves opposite, blades
lanceolate to narrowly elliptic, pinnately nerved.
Capitula terminal, solitary or in simple dichasia, radiate. Involucres broadly campanulate to
hemispheric, phyllaries in 2–3 series, gradate.
Receptacles convex. Ray florets neuter, corollas
yellow. Disc florets bisexual, corollas yellow with
fibres embedding the vascular strands; anthers
brown, appendages without glandular trichomes;
style arms tapered, apices papillose. Cypselae
compressed, oblong, brownish, sparsely ciliate
distally and on angles. Pappus of 2–3 persistent,
unequal awns and 6–10 free or variously connate
bristles between the awns, all borne on a rostrum.
One species, Z. lundellii (H. Rob.) Strother, Mexico
and Belize.
460
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
XXVI.5. Subtribe Enceliinae Panero (2005).
Annual or perennial herbs, shrubs, rarely trees.
Leaves usually alternate, petiolate, sometimes with
resinous exudates, blades linear to ovate or trullate,
sometimes laciniate. Capitula terminal, solitary
and scapose or in paniculiform or corymbiform
cymes, radiate or discoid. Involucres turbinate,
campanulate or hemispheric, phyllaries in 2–5
series, subequal, rarely gradate, herbaceous, rarely
chartaceous, sometimes with resinous exudates.
Receptacles flat to convex, paleae deciduous.
Ray florets neuter or rarely pistillate but sterile,
corollas sometimes with biseriate trichomes on
tube, rarely on limb. Disc florets bisexual, corollas
5-lobed, mostly without fibres embedding the
vascular strands, lobes sometimes with thickened
cells or with glandular or multicelular trichomes
on lobe abaxial surfaces; stamens 5, yellow, brown
or black, appendages linear to ovate, sometimes
with glandular trichomes, endothecium cells
fusiform rarely quadrate with 1–3 polar bridges;
styles with two vascular strands, style arms with
fused stigmatic surfaces, some species with two
stigmatic surfaces which fuse slightly above style
bifurcation point, densely papillose below, apices
acute to broadly acute. Cypselae compressed,
rarely thickened or terete, obovate to oblong,
densely sericeous, rarely glabrous, sometimes with
conspicuous wings or bands/margins at cypselae
edges and neck. Pappus of two slender awns with
or without squamellae in between, rarely absent,
rarely of awns fused into a crown surrounding the
neck.
Key to the Genera
1. Plants with mostly solitary, large capitula on long,
leafless peduncles
1308. Enceliopsis
– Plants otherwise
2
2. Shrubs or small trees
3
– Annual or perennial herbs
4
3. Cypselae strongly flattened with glabrous or sparsely
pubescent faces and ciliate margins; leaves mostly
canescent, never resinous or glutinous; disc corollas
yellow or purple, rarely white
1307. Encelia
– Cypselae biconvex, not strongly flattened, with
sparsely to densely pubescent faces and mostly
without ciliate margins; leaves mostly green, resinous
or glutinous or membranaceous; disc corollas yellow
1309. Flourensia
4. Phyllaries conspicuously ciliate; cypselae without
a white or stramineous edge or this wanting; cypselae
apices with a conspicuous crown surrounding neck of
cypsela and fused to awns
1310. Geraea
– Phyllaries glabrescent to variously pubescent,
rarely ciliate; cypselae mostly with a conspicuous
white/yellow (when fresh) or stramineous (when dry)
wing surrounding cypselae; cypselae apices without
a crown surrounding neck
5
5. Cypselae with a pappus of two awns and intervening
squamellae, rarely squamellae absent, cypselae mostly
without conspicuously ciliate margins
1311. Helianthella
– Cypselae epappose or with a pappus of two awns and
without intervening squamellae, cypselae mostly with
conspicuously ciliate margins
1307. Encelia
Genera of Enceliinae
1307. Encelia Adans.
Encelia Adans., Fam. Pl. 2: 128 (1763); Blake, Proc. Amer.
Acad. Arts 49: 346–396 (1913), rev.
Annual or perennial herbs, shrubs. Leaves alternate, blades linear to ovate sometimes laciniate,
triplinerved or with a single vein. Capitula terminal, solitary or in open paniculiform cymes, rarely
in a corymbiform cyme, radiate or discoid. Involucres campanulate to hemispherical, phyllaries in
2–4 series, subequal. Receptacles shallowly convex.
Ray corollas yellow, rarely white, sometimes with
biseriate trichomes on tube and along veins. Disc
corollas yellow or deep purple; anthers yellow or
black, appendages ovate with glandular trichomes;
style arms with fused stigmatic surfaces, sometimes divided at very base, apices acute. Cypselae broadly obovate to oblong, margins deeply ciliate, sparsely to densely pubescent. Pappus of two
awns, rarely of one or absent. x = 17, 18. Fifteen
species, Mexico, western USA, Galápagos Islands,
Peru, Chile.
1308. Enceliopsis (A. Gray) A. Nelson
Enceliopsis (A. Gray) A. Nelson, Bot. Gaz. 47: 432 (1909).
Perennial herbs or shrubs (caulirosula). Leaves alternate, petiolate, leaf internodes very short above
taproot producing a rosette, blades ovate to oblong, sometimes glaucous, bluish when fresh, semisucculent, triplinerved. Capitula terminal, solitary
(rarely with 2–3 capitula) on long leafless scapes,
radiate or discoid. Involucres hemispheric, phyllaries in 2–4 series, subequal. Receptacles shallowly
convex. Ray corollas yellow, sometimes with biseriate trichomes on tube. Disc corollas yellow; anthers yellow, appendages ovate with glandular trichomes; style arms with stigmatic surfaces continuous, sometimes divided at very base, apices acute.
Cypselae broadly obovate to oblong with yellow-
Compositae
ish and strongly ciliate or glabrous margins, sometimes with trichomes on centre of cypselae. Pappus
of two awns or small shoulders with few squamellae in between. x = 17, 18. Four species, western
USA.
1309. Flourensia DC.
Flourensia DC., Prodr. 5: 592 (1836); Dillon, Fieldiana Bot.
n.s. 16: 1–66 (1984), rev.
Shrubs or small trees. Leaves alternate, blades
lanceolate to ovate, pinnately nerved, usually
with resinous exudates, entire to broadly dentate.
Capitula terminal, solitary or in paniculiform or
corymbiform cymes, radiate or discoid. Involucres
turbinate, campanulate to hemispheric, phyllaries
in 2–5 series, subequal to gradate. Receptacles flat
to convex. Ray florets rarely pistillate and sterile,
corollas golden yellow, sometimes with biseriate
trichomes on tube. Disc corollas golden yellow;
anthers yellow or black, appendages sometimes
with glandular trichomes; style arms with continuous stigmatic surfaces, apices acute to broadly
acute. Cypselae thickened, rarely terete, obovate
to oblong, glabrous to densely sericeous with
ciliate margins. Pappus absent or most commonly
of 2 rarely 3 or 4 persistent or caducous awns.
x = 18. About 33 species, Mexico, south-western
USA, south-central Andes and interior of central
Argentina.
1310. Geraea Torr. & A. Gray
Geraea Torr. & A. Gray, Proc. Amer. Acad. Arts 1: 48 (1846)
[1847].
Erect annuals. Leaves alternate, blades ovate to
trullate, triplinerved. Capitula in terminal, open
paniculiform cymes, radiate or discoid. Involucres hemispheric, phyllaries in 2–3 series, subequal, conspicuously ciliate. Receptacles shallowly
convex. Ray corollas golden yellow. Disc corollas
golden yellow; anthers yellow, appendages with few
glandular trichomes; style arms with continuous
stigmatic surfaces, apices acute. Cypselae oblong
to obovate, densely sericeous except for base, with
yellowish wings and crown. Pappus of two awns
fused to a crown projecting from the neck. x = 18.
Two species, north-western Mexico, south-western
USA.
1311. Helianthella Torr. & A. Gray
Helianthella Torr. & A. Gray, Fl. NW Coast 2: 333 (1842);
Weber, Amer. Midl. Naturalist 48: 1–35 (1952), rev.
461
Perennial herbs. Leaves opposite, alternate distally,
blades linear to ovate-lanceolate, triplinerved to
pentanerved. Capitula terminal, solitary or in
corymbiform cymes, radiate. Involucres hemispherical, phyllaries in 2–4 series, subequal, rarely
gradate, the outermost sometimes enlarged and
leaf-like. Receptacles flat or convex. Ray corollas
yellow, sometimes with biseriate trichomes on
tube. Disc corollas yellow, purple or brownishpurple; anthers brown or black, appendages
sometimes with glandular and tapered trichomes;
style arms with continuous stigmatic surfaces,
sometimes at very base with two stigmatic areas
parallel to each other, apices acute. Cypselae
sometimes shallowly winged, cuneate to obcordate, blackish brown, glabrous to variously
pubescent. Pappus absent or of two persistent
awns, sometimes with a crown of short, lacerate
squamellae. x = 15. Eight species, western Canada
and USA, northern Mexico.
XXVI.6. Subtribe Engelmanniinae Stuessy
(1977); Bolick, Adv. Clad. 2: 125–144
(1983), phylog.; Clevinger & Panero,
Amer. J. Bot. 87: 565–572 (2000), phylog.;
Moore & Bohs, Amer. J. Bot. 90: 1653–
1660 (2003), phylog.
Annual or perennial herbs, rarely shrubs. Leaves
alternate or opposite, blades ovate to oblong, trullate, rarely obovate or sagittate, sometimes perfoliate, entire or pinnatifid. Capitula terminal, solitary,
sometimes on long scapes, or in open paniculiform
cymes, radiate, rarely discoid. Involucres mostly
hemispheric, sometimes campanulate, phyllaries
in 2–4 series, usually subequal, sometimes gradate,
ovate to suborbicular, mostly herbaceous, sometimes with indurate bases. Receptacles flat to convex. Ray florets pistillate, fertile, rarely in two series. Disc florets 5-lobed, bisexual or functionally
staminate, corollas with slender to prominent fibres
embedding vascular strands, glabrous to moderately pubescent; stamens 5, anthers rarely shallowly
calcarate, rarely tailed, anther appendages with or
without glandular trichomes; styles with two vascular strands, style arms either fused and filiform,
or slender with parallel stigmatic surfaces fusing
at distal end of style arms, sometimes with continuous stigmatic surfaces, sometimes with glandular trichomes. Ray cypselae obcompressed and in
some species with functionally staminate disc florets associated with the 2(–4) adjacent disc florets
and paleae, or shallowly quadrate or triquetrous.
462
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Disc cypselae when present compressed, narrowly
rhombic in cross-section. Pappus mostly of two
awns at angles of cypselae sometimes associated
with a few scales, sometimes of several minute
scales, a cup of fused squamellae or absent.
Key to the Genera
1. Disc florets bisexual
2
– Disc florets functionally staminate
4
2. Fibre sheaths of disc corollas continuous along edges of
lobes; paleae stiff, lignified, apices becoming acicular
with age, stoloniferous shrubs of coastal marshes of
North America, Caribbean and Peru 1313. Borrichia
– Fibre sheaths of disc corollas restricted to the throats;
paleae flexible, not lignified, apices not acicular with
age; perennial herbs or shrubs, rarely stoloniferous, of
mountainous regions of North America
3
3. Shrubs of north-eastern Mexico; anthers with glandular trichomes on appendages and connectives
1318. Vigethia
– Perennial herbs of western North America; anthers
with glandular trichomes on appendages, not on the
connectives
1319. Wyethia
4. Capitula with 2 series of ray florets; ray cypselae not associated with adjacent 2–4 disc florets and associated
paleae
1317. Silphium
– Capitula with 1 series of ray florets; ray cypselae associated with adjacent 2–4 disc florets and associated
paleae
5
5. Leaves opposite
1314. Chrysogonum
– Leaves alternate
6
6. Leaves entire, ovate, rhombic or trullate, rarely
trilobed
7
– Leaves pinnatisect or lyrate
8
7. Phyllaries suborbicular; styles with glandular
trichomes
1312. Berlandiera
– Phyllaries ovate; styles without glandular trichomes
1316. Lindheimera
8. Disc corollas red; veins of ray corollas green or reddish; phyllaries of outermost series suborbicular; foliage dull green
1312. Berlandiera
– Disc corollas yellow; veins of ray corolla yellow;
phyllaries of outermost series narrowly ovate; foliage
bright green
1315. Engelmannia
Genera of Engelmanniinae
1312. Berlandiera DC.
Fig. 94
Berlandiera DC., Prodr. 5: 517 (1836); Pinkava, Brittonia
19: 285–298 (1967), rev.
Perennial herbs, sometimes with strong, pungent
odour. Leaves alternate, simple, ovate to obovate,
sometimes pinnatifid, serrate to runcinate. Capitula terminal, solitary or in open paniculiform
cymes, radiate. Involucres hemispheric, phyllaries in 2–3 series, subequal, broadly ovate to
suborbicular, bases indurate, herbaceous apices
Fig. 94.
Compositae-Heliantheae. Berlandiera lyrata.
A Habit. B Ray corolla. C Disc corolla. D Ovary of
sterile disc floret and attached palea. E Anthers. F Ray
cypsela. G Inner face of phyllary and two attached paleae.
(Reproduced from Flora Novo-Galiciana, vol. 12, with
permission of the University of Michigan Herbarium; see
McVaugh 1984)
spreading. Receptacles shallowly convex. Ray
florets pistillate, corollas light to golden yellow.
Disc florets functionally staminate, corollas yellow
or deep red or burgundy; anthers black to reddish
black, appendages densely covered with glandular
trichomes on abaxial side; style arms of disc florets
fused and filiform with glandular trichomes, of ray
florets with divided stigmatic surfaces. Cypselae
obcompressed, suborbicular, flat or with a median
rib on each tangential side. Pappus of a few scales
or absent. x = 15. Four species, Mexico, USA.
1313. Borrichia Adans.
Borrichia Adans., Fam. Pl. 2: 130, 527 (1763); Semple, Ann.
Missouri Bot. Gard. 66: 681–693 (1978), rev.
Stoloniferous shrubs of muddy or sandy coastal
marshes. Leaves opposite, blades obovate, succulent, perfoliate, entire to broadly serrate. Capitula
terminal, solitary, radiate. Involucres hemispheric,
Compositae
phyllaries in 2–4 series, subequal. Receptacles
convex, paleae lignified, apices somewhat acicular, capitulum echinate. Ray florets pistillate,
corollas yellow, sometimes persistent and shed
with cypsela, tube lacking. Disc florets bisexual,
fertile, corollas yellow, fibres meeting at each lobe
apex; anthers black, with glandular trichomes
on abaxial side of appendages and connectives;
style arms with continuous stigmatic surfaces,
tapered. Cypselae obcompressed, oblong, rhombic
in outline, with a median rib on each tangential
side. Pappus a cup with an erose, sinuate edge,
sometimes with free squamellae. x = 13, 14. Two
species, tropical and subtropical coasts of North
America and Caribbean, Peru.
1314. Chrysogonum L.
Chrysogonum L., Sp. Pl. 2: 920 (1753); Stuessy, Rhodora 79:
190–202 (1977), rev.
Stoloniferous perennial herbs. Leaves opposite,
petiolate, ovate to suborbicular, shallowly serrate.
Capitula terminal, solitary or in simple dichasial
cymes, radiate. Involucres hemispherical, phyllaries in 2 series, subequal, broadly ovate, dimorphic.
Receptacles shallowly convex. Ray florets pistillate,
corollas yellow. Disc florets functionally staminate,
corollas yellow; anthers black, shallowly tailed,
shallowly calcarate, appendages ovate; style arms
of disc florets filiform, abaxial sides deeply papillose, style arms of ray florets 2–3, spreading,
with a continuous stigmatic surface. Ray cypselae
obcompressed, suborbicular, with a median rib on
each tangential side. Pappus of two small awns or
squamellae. x = 16. One species, C. virginianum
L., south-eastern USA.
1315. Engelmannia Torr. & A. Gray
Engelmannia Torr. & A. Gray, Trans. Amer. Philos. Soc. n.s.
7: 343 (1840).
Perennial herbs. Leaves alternate, blades ovate to
oval, pinnatifid, venation pectinate. Capitula in
terminal, paniculiform cymes, radiate. Involucres
hemispherical, phyllaries in 2–3 series, dimorphic.
Receptacles shallowly convex. Ray florets pistillate,
corollas yellow. Disc florets functionally staminate,
corollas yellow; anthers black; style arms of disc
florets filiform, those of ray florets spreading with
fused stigmatic surfaces. Cypselae obcompressed,
broadly ovate to oblong or suborbicular with
a median rib on each tangential side. Pappus
of 2–4 unequal awns, those at edge of cypselae
463
longer. x = 9. One species, E. peristenia (Raf.) G.J.
Goodman & C.A. Lawson, south-western USA,
Mexico.
1316. Lindheimera A. Gray & Engelm.
Lindheimera A. Gray & Engelm., Proc. Amer. Acad. Arts 1:
47 (1846) [1847]; Hind, Kew Mag. 7: 128–131 (1990), rev.
Erect annuals. Leaves alternate or opposite, blades
ovate to trullate, rarely sagittate, weakly triplinerved, sinuate to broadly dentate. Capitula in terminal, dichasial paniculiform cymes, rarely solitary,
radiate. Involucres campanulate, phyllaries in 2 series, normally 5 in each series, strongly dimorphic.
Receptacles shallowly convex. Ray florets pistillate,
mostly 5, corollas golden yellow. Disc florets functionally staminate, corollas yellow; anthers black,
appendages without glandular trichomes; styles
of disc florets filiform, style arms fused, those of
ray florets spreading with acute apices, stigmatic
surfaces divided. Cypselae obcompressed, broadly
ovate to oblong or suborbicular with a median rib
on each tangential side, wings entire and fused
to pappus. Pappus of two diverging awns. x = 8.
One species, L. texana A. Gray, south-western USA,
northern Mexico.
1317. Silphium L.
Silphium L., Sp. Pl. 2: 919 (1753); Clevinger & Panero, Amer.
J. Bot. 87: 565–572 (2000), phylog.
Perennial herbs, sometimes forming a rosette.
Leaves opposite or alternate, blades ovate to
trullate, sometimes perfoliate, simple or deeply
dissected, venation pectinate, rarely triplinerved.
Capitula terminal, solitary or in paniculiform
cymes, sometimes with very short internodes
and pedicels and capitula congested on a short
axis, radiate. Involucres hemispherical, rarely
campanulate, phyllaries broadly ovate to suborbicular. Receptacles flat to shallowly convex. Ray
florets pistillate, in 2 series, corollas golden yellow,
stramineous or white. Disc florets functionally
staminate, corollas pale to golden yellow or
white; anthers black; style arms of disc florets
fused, filiform, those of ray florets spreading with
fused stigmatic surfaces. Cypselae obcompressed,
broadly obovate to suborbicular or oblong, margins winged. Pappus of two small awns connate to
the wings or absent. x= 7. About 25 species, USA,
Canada.
464
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
1318. Vigethia W.A. Weber
Vigethia W.A. Weber, Madroño 7: 98 (1943).
Shrubs. Leaves alternate, ovate to subcordate,
serrate. Capitula terminal, solitary or in simple
dichasial cymes, radiate. Involucres hemispherical, phyllaries in 5–7 series, subequal, outermost
reflexed. Receptacles convex. Ray florets pistillate,
sometimes with staminodes, corollas light yellow.
Disc florets bisexual, corollas yellow; anthers
black, appendages with glandular trichomes on
abaxial side and on connective; style arms of disc
florets with stigmatic surfaces divided at the base,
fused midway, tapered, papillose, style arms of
ray florets spreading, with acute apices. Cypselae
ovate to oblong, obcompressed to quadrate, ray
cypselae triquetrous to shallowly quadrate. Pappus
of several lacerate, minute scales. x = 19. One
species, V. mexicana (S. Watson) W.A. Weber,
north-eastern Mexico.
1319. Wyethia Nutt.
Wyethia Nutt., J. Acad. Nat. Sci. Philadelphia 7: 39 (1834);
Sharp, Ann. Missouri Bot. Gard. 22: 51–153 (1935), rev.; Weber, Amer. Midl. Naturalist 35: 400–452 (1946), rev.; Moore
& Bohs, Amer. J. Bot. 90: 1653–1660 (2003), phylog.
Balsamorhiza Nutt. (1840).
Scabrethia W.A. Weber (1999).
Agnorhiza W.A. Weber (1999).
Perennial herbs with thickened taproots, sometimes forming large stoloniferous colonies in shifting sand dunes, sometimes resinous. Leaves alternate or opposite, blades linear to deltate-ovate,
spathulate, entire, deeply lobed to serrate. Capitula
terminal, large, very showy, solitary or few in paniculiform cymes, on floral scapes with or without
leaves, radiate or discoid. Involucres broadly campanulate to hemispheric, phyllaries in 2–6 series.
Receptacles broadly convex. Ray florets pistillate,
yellow to light yellow or white. Disc florets bisexual, corollas golden yellow, sometimes fibres wanting; anther appendages with glandular trichomes
on abaxial side; style arms with parallel or continuous stigmatic surfaces, sometimes fused at mid to
distal end. Cypselae ovate to oblong, compressed
to shallowly quadrate, ray cypselae triquetrous to
shallowly quadrate. Pappus absent or a crown of
persistent scales, some species with slender awns
at angles of cypselae. x = 19. About 28 species,
western North America.
XXVI.7. Subtribe Helianthinae Dumort.
(1827).
Subtribe Lagasceinae Benth. & Hook. f. (1873).
Annual or perennial herbs, shrubs, trees. Leaves
alternate or opposite, blades unlobed to dissected,
pinnately veined to mostly triplinerved, rarely pentanerved, entire to deeply serrate. Capitula terminal, rarely axillary, solitary or in simple to large
monochasial or dichasial paniculiform or corymbiform cymes, sometimes with fistulose peduncles,
capitula discoid or radiate, rarely disciform. Involucres cylindrical to hemispherical, phyllaries in
2–7 series, subequal to gradate, most commonly
with an indurate base and herbaceous apices, innermost normally smaller than outermost, outermost often foliaceous. Receptacles flat to conic,
paleae sometimes forming perigynia and shedding with cypselae as a unit. Ray florets neuter,
sometimes with a vestigial style or fertile anther.
Disc florets bisexual, rarely functionally pistillate,
corollas pentamerous, without fibres embedding
the vascular strands, sometimes apices of lobes
with sclerified cells; stamens 5, filaments rarely
papillose; styles with two vascular strands, style
arms with fused stigmatic surfaces, deltate, sometimes with long and tapered apices, sometimes
with a narrow, cylindric appendage. Cypselae compressed, biconvex in cross-section, rarely quadrate
to terete, mostly black, striate, glabrous to densely
pubescent. Pappus absent or more commonly of
two awns with or without squamellae in between,
sometimes a crown of awns.
Key to the Genera
1. Capitula with 1–6(–8) disc florets
2
– Capitula with more than 10 disc florets
4
2. Phyllaries fused except for distal herbaceous apices
1328. Lagascea
– Phyllaries free
3
3. Leaves opposite; cypselae terete, circular in crosssection; pappus of multiple awns
1321. Alvordia
– Leaves alternate; cypselae biconvex to weakly quadrangular, rhombic in cross-section; pappus absent
1323. Calanticaria
4. Capitula with paleae tightly wrapping cypselae (perigynia) and shed with cypsela as a unit
5
– Capitula with paleae not tightly wrapping cypselae, if
deciduous, then generally not shed with cypselae as
a unit
6
5. Disc florets with black papillose trichomes on adaxial
surface of throat; anthers mostly yellow
1333. Sclerocarpus
– Disc florets without black papillose trichomes on
adaxial surface of throat; anthers black 1320. Aldama
Compositae
6. Plants forming weak rosettes with scapose capitula;
cypselae shallowly quadrate, broadly rhombic in outline; corollas pink, purple or yellow, rarely white
1327. Iostephane
– Plants otherwise with capitula in paniculiform
or corymbiform cymes, rarely scapose; cypselae
biconvex; corollas yellow to golden yellow
7
7. Cypselae with a well-developed elaiosome; pappus of
two broad awns and mostly 4 intervening squamellae;
perennial herbs or shrubs of the western Andes in
central Peru
1336. Syncretocarpus
– Cypselae without elaiosomes; pappus various; plants
from America
8
8. Leaves alternate, capitula in monochasial cymes,
rarely solitary
9
– Leaves opposite, capitula in dichasial cymes, rarely
solitary
19
9. Shrubs or trees of the Galápagos Islands; corollas
mostly white, rarely greenish
1332. Scalesia
– Annual or perennial herbs, shrubs or trees of continental America, Caribbean; corollas yellow, very rarely
white (Viguiera)
8
10. Peduncles shallowly to strongly fistulose
1337. Tithonia
– Peduncles never fistulose
11
11. Pappus caducous, rarely absent
12
– Pappus persistent
17
12. Pappus absent
13
– Pappus of two awns and sometimes a few intervening
squamellae, mostly easily caducous
14
13. Abaxial leaf surfaces with glandular trichomes; cypselae glabrous
1325. Heliomeris
– Abaxial leaf surfaces without glandular trichomes;
cypselae pubescent
1326. Hymenostephium
14. Plants with tuberous roots
15
– Plants without tuberous roots
16
15. Cypselae shallowly quadrate, tubers long, cylindrical
1330. Phoebanthus
– Cypselae biconvex, never shallowly quadrate, tubers
round or elliptic; North America
1324. Helianthus
16. Annual or perennial herbs, apices of paleae trilobed;
anther thecae brown or deep purple 1324. Helianthus
– Annual or perennial herbs, shrubs; apices of paleae
round to acute; if perennial herbs, then anther thecae
yellow, otherwise brown or black; Colombia, Ecuador
and Peru
1329. Pappobolus
17. Capitula in congested corymbiform cymes arranged
in paniculiform cymes; phyllaries shortly aristate, narrow tips spreading and with long, white trichomes
1335. Stuessya
– Capitula in open corymbiform or paniculiform cymes;
phyllaries acute to acuminate, never shortly aristate,
trichomes otherwise
18
18. Phyllaries with ovate, indurate bases and linear,
spreading, herbaceous apices; shrubs; Sonoran and
Mojave deserts, Baja California; x = 18
1322. Bahiopsis
– Phyllaries various, rarely with ovate, indurate bases
and spreading, herbaceous apices; annual or perennial
herbs, shrubs; America, rarely in Sonoran or Mojave
deserts or in Baja California; x = 17 1338. Viguiera
19. Cypselae with a narrow corky rim; accessory branches
of the inflorescences mostly arising at nearly 90° from
main axis; south-western Mexico
1331. Rhysolepis
465
– Cypselae without a narrow corky rim; accessory
branches of the inflorescences mostly arising at
40–70° from main axis; America
20
20. Involucres cylindrical to narrowly campanulate,
phyllaries strongly indurate with narrow herbaceous
apices; cypselae epappose; Sonora, Mexico
1321. Alvordia
– Involucres various, if cylindrical to narrowly campanulate and phyllaries strongly indurate, then phyllaries
with a herbaceous rim, cypselae pappose and plants
from coastal ranges of Sonora (Viguiera in part) 21
21. Plants with solitary capitula, shrubs with bicoloured,
small, ovate to trullate, rarely round leaves, abaxial
side densely pubescent
1323. Calanticaria
– Plants with capitula in paniculiform cymes, rarely
solitary, if solitary, then leaves not conspicuously bicoloured
22
22. Leaves mostly auriculate; cypselae strongly flattened,
nearly linear in cross-section, mostly with embryo
as an oval protuberance in the centre of cypselae,
if cypsela plump throughout and biconvex in crosssection, then ray corollas pale lemon-yellow with conspicuous greenish or reddish corolla veins; apices of
lobes of disc corollas mostly with sclerified cells
1334. Simsia
– Leaves not auriculate; cypselae not strongly flattened,
biconvex in cross-section, corollas mostly yellow or
golden yellow, rarely with pale lemon-yellow ray corollas and veins conspicuously contrasting against corolla
background; apices of lobes of disc corollas without
sclerified cells
23
23. Cypselae with a pappus of two awns and some intervening squamellae
1338. Viguiera
– Cypsela with a pappus of several squamellae or absent
24
24. Leaves mostly lanceolate with glandular trichomes on
abaxial surface; temperate North America
1325. Heliomeris
– Leaves mostly ovate, rarely lanceolate, without glandular trichomes on abaxial surface; tropical America
1326. Hymenostephium
Genera of Helianthinae
1320. Aldama La Llave
Aldama La Llave, Nov. Veg. Descr. 1: 14 (1824); Feddema,
Ph.D. Thesis, University of Michigan, Ann Arbor: 1–132
(1966); Feddema, Phytologia 21: 308–314 (1971), rev.
Erect annuals. Leaves opposite, alternate towards
the inflorescences, blades lanceolate to narrowly
ovate. Capitula in open corymbiform cymes, radiate or discoid. Involucres campanulate to subhemispheric, phyllaries in 2–3 series, subequal. Receptacles convex to hemispheric, paleae indurate and
wrapping tightly around cypselae (perigynia), appearing tubular, deciduous with the cypsela. Ray
floret corollas yellow to orange. Disc floret corollas
yellow; anthers black. Cypselae biconvex to slightly
trigonous, black, adaxial side straight to slightly
466
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
concave, glabrous. Pappus a minute corona of lacerate squamellae or absent. x = 17. Two species,
Mexico, Central America, northern South America.
1321. Alvordia Brandegee
Alvordia Brandegee, Proc. Calif. Acad. Sci. ser. 2, 2: 174
(1889); Carter, Proc. Calif. Acad. Sci. 30: 157–174 (1964),
rev.
Agiabampoa Rose ex Hoffm. (1894).
Shrubs. Leaves opposite, blades lanceolate to ovate.
Capitula in congested corymbiform cymes or open
paniculiform cymes of congested corymbiform
cymes, discoid or radiate. Involucres cylindrical
to narrowly campanulate, phyllaries in 2–5 series,
gradate. Receptacles flat. Ray floret corollas yellow.
Disc florets 1 to few per head, corollas bright
yellow; anthers yellow. Cypselae obovoid, slightly
flattened to quadrangular, black, glabrous. Pappus
of multiple, lanceolate, lacerate squamellae or
absent. x = 17. Four species, Mexico.
1322. Bahiopsis Kellogg
Bahiopsis Kellogg, Proc. Calif. Acad. 2: 35 (1863); Schilling,
Madroño 37: 149–170 (1990), rev. as Viguiera; Schilling &
Panero, Bot. J. Linn. Soc. 140: 65–76 (2002), phylog.
Viguiera Kunth sect. Bahiopsis (Kellogg) E.E. Schill. (1990).
Shrubs. Leaves opposite, blades ovate to deltate,
unlobed or rarely laciniate. Capitula solitary and
scapose or mostly in open paniculiform cymes,
radiate. Involucres campanulate to hemispherical,
phyllaries in 2–4 series, subequal, distally tapered
and herbaceous. Receptacles convex. Ray floret
corollas golden yellow. Disc floret corollas yellow;
anthers black to reddish brown. Cypselae biconvex,
obovate, black or brown, glabrous to moderately
pubescent. Pappus of 2 awns and few lacerate
squamellae. x = 18. Twelve species, north-western
Mexico, south-western USA.
1323. Calanticaria (B.L. Rob. & Greenm.) E.E.
Schill. & Panero
Calanticaria (B.L. Rob. & Greenm.) E.E. Schill. & Panero,
Bot. J. Linn. Soc. 140: 73 (2002); González Elizondo et al.,
Acta Bot. Mexicana 53: 35–48 (2000), new spp., as member
of Viguiera; Schilling & Panero, Bot. J. Linn. Soc. 140: 65–76
(2002), phylog.
Shrubs. Leaves opposite or alternate, blades ovate,
trullate to round. Capitula solitary or few in tight
corymbiform cymes, radiate, rarely discoid. In-
Fig. 95. Compositae-Heliantheae. Helianthus laciniatus.
A. Habit, terminal and lower portions of plant. B Capitulum. C Ray corolla. D Disc corolla. E Palea. F Cypsela with
caducous awns. G Anthers. H Style arms. (Reproduced
from Flora Novo-Galiciana, vol. 12, with permission of the
University of Michigan Herbarium; see McVaugh 1984)
volucres campanulate to hemispheric, phyllaries in
2–3 series, subequal. Receptacles convex. Ray floret
corollas yellow to golden yellow. Disc floret corollas
yellow; anthers brown or yellow. Cypselae biconvex, obovate to cuneate, black or mottled black and
brown, glabrous to moderately pubescent. Pappus
of 2 awns and few, lacerate squamellae or absent.
Five species, northern Mexico.
1324. Helianthus L.
Fig. 95
Helianthus L., Sp. Pl. 2: 904 (1753); Heiser, Mem. Torrey
Bot. Club 22: 1–218 (1969), rev.; Heiser & Schilling, Taxon
30: 393–403 (1981), class.; Schilling et al., Syst. Bot. 23: 177–
187 (1998), phylog.
Erect annual or perennial herbs, occasionally with
tuberous roots. Leaves alternate, sometimes opposite, blades unlobed. Capitula solitary or in open
paniculiform cymes, discoid or radiate. Involucres
Compositae
campanulate to hemispherical, phyllaries in 2–5 series, subequal. Receptacles flat to shallowly convex,
paleae with trilobed apices. Ray corollas yellow to
golden yellow. Disc floret corollas yellow to greenish yellow, golden yellow, reddish or burgundy;
anthers black or deep purple; style arms sometimes with cylindric appendages. Cypselae biconvex, obovate to oblong, black or greyish brown and
mottled, glabrous or sparsely pubescent. Pappus of
two awns sometimes with a few squamellae in between, caducous. x = 17. Approximately 50 species,
North America.
1325. Heliomeris Nutt.
Heliomeris Nutt., Proc. Acad. Nat. Sci. Philadelphia 4: 19
(1848); Yates & Heiser, Proc. Indiana Acad. Sci. 88: 364–372
(1979), rev.; Schilling & Panero, Bot. J. Linn. Soc. 140: 65–76
(2002), phylog.
Erect annual or perennial herbs. Leaves opposite
or alternate, blades linear to ovate, sometimes revolute. Capitula solitary or in open paniculiform
cymes, radiate. Involucres hemispherical, phyllaries in 2 series, subequal. Receptacles convex to
conical. Ray corollas yellow to orange. Disc floret
corollas yellow to yellow-orange, tube narrow and
opening into a broad cylindrical throat, densely
pubescent at base of throat on abaxial surface; anthers black or reddish brown. Cypselae biconvex,
obovate, black, glabrous. Pappus absent. x = 8, 13,
14. Five species, Mexico, USA.
1326. Hymenostephium Benth.
Hymenostephium Benth. in Benth. & Hook. f., Gen. Pl. 2:
382 (1873); Schilling & Panero, Bot. J. Linn. Soc. 140: 65–76
(2002), phylog.
Erect annual or perennial herbs, sometimes
scandent shrubs. Leaves opposite, blades ovate
to deltate, rarely narrowly lanceolate. Capitula
solitary but mostly in open paniculiform cymes,
radiate or discoid. Involucres cylindric to campanulate, rarely hemispherical, phyllaries in 2–3 series,
subequal. Receptacles flat to shallowly convex.
Ray floret corollas pale yellow to golden yellow
or orange. Disc floret corollas yellow, orange or
red, rarely greenish white; anthers black, brown
or yellow. Cypselae biconvex, black or brown,
glabrous to densely pubescent, sometimes with
a prominent elaiosome. Pappus of several lacerate
round to apiculate squamellae or absent. x = 17?,
20. Approximately 26 species, Mexico to Argentina,
467
with a distinctive group of species in the Andes of
Venezuela.
1327. Iostephane Benth.
Iostephane Benth. in Benth. & Hook. f., Gen. Pl. 2: 368
(1873); Sharp, Ann. Missouri Bot. Gard. 22: 51–153 (1935);
Strother, Madroño 30: 34–38 (1983), rev.
Perennial herbs. Leaves alternate in a lax to congested rosette, leaves erect or flat, blades lanceolate
to ovate, deltate, unlobed to deeply 3-lobed.
Capitula solitary on scapes or in open paniculiform cymes, radiate. Involucres campanulate to
hemispheric, phyllaries in 2–3 series, subequal.
Receptacles convex. Ray florets sometimes styliferous but neuter, corollas pale purple to pink, white,
ochroleucous, yellow to orange. Disc floret corollas
yellow to whitish yellow, greenish yellow sometimes with purplish tips; anthers black. Cypselae
weakly compressed, mostly quadrate or round in
cross-section, oblong in outline, brownish black,
glabrous to sparsely puberulent. Pappus of 1–2
reduced awns and few smaller squamellae or most
commonly absent. x = 17. About four species,
Mexico.
1328. Lagascea Cav.
Lagascea Cav., Anales Ci. Nat. 6: 331 (1803), nom. cons.;
Stuessy, Fieldiana Bot. 38: 75–133 (1978), rev.
Erect annual or perennial herbs, erect or scandent shrubs. Leaves opposite, blades lanceolate
to ovate to oblanceolate or elliptic. Capitula of
1 or 2 (rarely 8) florets, aggregated into tight
glomerule-like clusters and subtended by leafy
bracts producing a compound capitulum, discoid.
Involucres cylindrical, phyllaries in 1 series, fused,
subequal. Receptacles epaleate. Corollas white,
pink or purple-pink, yellow or orange-yellow;
anthers black, pink or yellow. Cypselae biconvex,
oblanceolate to obovate, black, glabrous to sparsely
pubescent. Pappus absent or a small crown of
fused, lacerate, minute squamellae, sometimes
those at angles of cypsela longer. x = 17. Nine
species, neotropical, most species in southern
Mexico, introduced elsewhere.
1329. Pappobolus S.F. Blake
Pappobolus S.F. Blake, Hooker’s Icon. Pl. pl. 3057 (1916);
Panero, Syst. Bot. Monogr. 36: 1–195 (1992), rev.
Helianthopsis H. Rob. (1979).
Annual or perennial herbs, shrubs, small trees.
Leaves opposite or alternate, blades linear to
468
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
broadly ovate. Capitula solitary or in simple
dichasial or monochasial to paniculiform or
corymbiform cymes, sometimes subaxillary,
radiate. Involucres campanulate to hemispherical,
phyllaries in 3–6 series, subequal to gradate,
sometimes outermost spreading and leaf-like.
Receptacles flat to convex. Ray floret corollas
yellow to yellow-orange. Disc florets sometimes
functionally pistillate, corollas yellow to golden
yellow sometimes with blackish lobes; anthers
yellow or black. Cypselae biconvex, cuneate to
oblong, brown to black, glabrous to densely
pubescent. Pappus of 2 or many caducous awns,
sometimes with few minute lacerate squamellae
between the two awns. x = 17. About 38 species,
Colombia, Ecuador, Peru.
Shrubs, trees. Leaves alternate, blades lanceolate
to ovate, rarely cordate, unlobed to deeply lobed
or 2-pinnate. Capitula solitary or in corymbiform
cymes, discoid, disciform or radiate. Involucres
campanulate to hemispherical, phyllaries in 2–4
series, subequal to gradate, sometimes outermost
spreading and leaf-like. Receptacles flat to shallowly convex, paleae sometimes with trilobed
apices. Ray florets sometimes styliferous but
neuter or staminiferous with some pollen, corollas
white. Disc floret corollas white, sometimes
greenish white to dull yellow with age, peripheral
corollas sometimes bent outwards; anthers black.
Cypselae biconvex, cuneate to oblong, brown to
black, glabrous. Pappus of 1 or 2 awns or of a few
lacerate squamellae or absent. x = 34. About
eleven species, Galápagos Islands.
1330. Phoebanthus S.F. Blake
Phoebanthus S.F. Blake, Proc. Amer. Acad. Arts 51: 520
(1916).
Perennial herbs with tuberous roots. Leaves alternate, blades filiform to narrowly lanceolate. Capitula solitary or in open paniculiform cymes, radiate.
Involucres campanulate to hemispherical, phyllaries in 2–4 series. Receptacles convex. Ray floret
corollas yellow. Disc floret corollas yellow; anthers
black. Cypselae slightly quadrate, black, glabrous.
Pappus of 2 awns. x = 17. Two species, Florida,
USA.
1331. Rhysolepis S.F. Blake
Rhysolepis S.F. Blake, Contr. Gray Herb. 52: 36 (1917);
Blake, Contr. Gray Herb. 54: 1–205 (1918), rev.; Robinson &
Moore, Proc. Biol. Soc. Wash. 117: 323–446 (2004), rev.
Perennial herbs or weak scandent shrubs. Leaves
opposite or alternate, blades lanceolate, ovate
to oval. Capitula in open paniculiform cymes,
paracladia branching at nearly 90° from main axis,
radiate. Involucres campanulate to hemispherical,
phyllaries in 3–4 series. Receptacles convex. Ray
floret corollas golden yellow to yellow-orange.
Disc floret corollas yellow; anthers black. Cypselae
biconvex, cuneate, black, glabrous. Pappus of 2
awns and several minute squamellae, or of a few
squamellae, sometimes easily caducous. Three
species, Mexico.
1332. Scalesia Arn.
Scalesia Arn., Nat. Syst. ed. 2: 443 (1836); Eliasson, Opera
Bot. 36: 1–120 (1974), rev.
Zemisne Degener & Sherff (1935).
1333. Sclerocarpus Jacq.
Sclerocarpus Jacq., Icon. Pl. Rar. 1: 17, pl. 176 (1781); Feddema, Ph.D. Thesis, University of Michigan, Ann Arbor:
1–132 (1966); Schilling & Panero, Bot. J. Linn. Soc. 140: 65–
76 (2002), phylog.
Erect annual or perennial herbs. Leaves alternate,
blades lanceolate to ovate, unlobed to deeply lobed.
Capitula solitary or in open paniculiform cymes,
radiate. Involucres campanulate to hemispherical,
phyllaries 0 to few in 1–2 series, subequal. Receptacles conical, paleae indurate and tightly wrapping
cypselae (perigynia), appearing tubular, shed with
the cypsela. Ray floret corollas bright yellow to orange. Disc floret corollas yellow to orange, mostly
with black papillae on adaxial surface of throat; anthers black or yellow. Cypselae broadly biconvex,
black or brown, adaxial side straight to slightly concave, glabrous. Pappus a crown of lacerate squamellae or absent. x = 11, 12, 13, 18. Eight species, southwestern USA, Mexico, Central America, Venezuela,
Africa.
1334. Simsia Pers.
Simsia Pers., Syn. Pl. 2: 478 (1807); Robinson & Brettell,
Phytologia 24: 361–377 (1972), reg. rev.; Spooner, Syst. Bot.
Monogr. 30: 1–90 (1990), rev.
Erect annual or perennial herbs, shrubs. Leaves
opposite, auriculate, blades lanceolate, ovate to
subcordate, sometimes deltate, trilobate, unlobed
to shallowly lobed. Capitula solitary or in open
paniculiform cymes, radiate or discoid. Involucres
broadly cylindrical, turbinate, rarely hemispherical, phyllaries in 2–4 series, gradate. Receptacles
Compositae
convex. Ray floret corollas lemon to golden yellow,
white or cherry red. Disc floret corollas yellow
to golden yellow, white or cherry red, corollas
mostly with sclerified cells on tips of lobes; anthers
black, brown or yellow, sometimes bicoloured;
style arms long and tapered, sometimes curled.
Cypselae narrowly biconvex to biconvex, mostly
with embryo confined to centre of cypselae with
a flat rim around it, black, brown or mottled,
glabrous. Pappus of 2 awns or absent. x = 17.
About 22 species, America.
1335. Stuessya B.L. Turner
Stuessya B.L. Turner, Brittonia 32: 209 (1980).
Erect annuals. Leaves alternate, blades narrowly
lanceolate to ovate. Capitula in corymbiform
cymes arranged in open paniculiform cymes,
radiate. Involucres broadly cylindrical to campanulate or ovoid, phyllaries in 3–4 series, gradate,
shortly aristate. Receptacles shallowly convex. Ray
floret corollas bright yellow. Disc floret corollas
yellow; anthers black or yellow; style arms tapered.
Cypselae biconvex, black, glabrous. Pappus of two
awns and several minute, lacerate squamellae.
x = 17. Three species, Mexico.
1336. Syncretocarpus S.F. Blake
Syncretocarpus S.F. Blake, Bot. Jahrb. Syst. 54: 49 (1916);
Panero & Granda Paucar, Phytologia 87: 109–112 (2005),
new sp., tax.
Erect annual or perennial herbs, shrubs. Leaves alternate, blades narrowly lanceolate to ovate. Capitula in open paniculiform cymes, radiate. Involucres campanulate to hemispherical, phyllaries in
2–3 series, subequal. Receptacles shallowly convex,
paleae deciduous. Ray floret corollas golden yellow
to lemon yellow. Disc floret corollas yellow; anthers
black. Cypselae biconvex, with long sericeous trichomes (encelioid) and a distinctive corky margin,
with or without a prominent elaiosome. Pappus of
2 awns and 4 lacerate squamellae. x = 17. Two or
three species, south-central Peru.
1337. Tithonia Desf.
Tithonia Desf., Gen. Pl. 189 (1789); La Duke, Rhodora 84:
453–522 (1982), rev.
Erect annual or perennial herbs, shrubs, rarely
trees. Leaves alternate, blades narrowly ovate to
ovate, sometimes auriculate, bases attenuate. Capitula solitary or in open paniculiform cymes, in
469
most species on fistulose peduncles, radiate. Involucres hemispherical, phyllaries in 2–5 series,
subequal, sometimes outermost foliaceous. Receptacles shallowly convex. Ray floret corollas yellow
to orange-red. Disc floret corollas yellow to orange;
anthers black, brown or yellow. Cypselae biconvex to weakly quadrate, black or greyish brown or
mottled, glabrous to moderately pubescent. Pappus
a crown of fused squamellae or 2 awns with fused
squamellae or absent. x = 17. About 11 species,
south-western USA to northern South America,
one or two species introduced in most other tropical regions of the world.
1338. Viguiera Kunth
Viguiera Kunth in Humb., Bonpl. & Kunth, Nov. Gen. Sp.
folio ed. 4: 176 (1818); Blake, Contr. Gray Herb. 54: 1–
205 (1918), rev.; Panero & Schilling, Syst. Bot. 13: 371–406
(1988), sect. rev.
Erect rarely decumbent annual or perennial herbs,
shrubs, rarely trees. Leaves opposite or alternate,
linear to ovate, sometimes cordate or deltate. Capitula solitary or in congested to open paniculiform
cymes, radiate, rarely discoid. Involucres cylindrical to hemispherical, phyllaries in 2–7 series, subequal to prominently gradate. Receptacles flat to
convex, rarely conical. Ray florets sometimes styliferous but neuter, corollas yellow to yellow-orange,
rarely dull red or white. Disc floret corollas yellow
to orange, rarely dull red; anthers yellow, brown
or black, rarely reddish. Cypselae biconvex, obovate to oblong, cuneate, black, brown or greyish
brown, sometimes mottled with age, glabrous to
densely pubescent. Pappus of 2 persistent or somewhat easily caducous awns, with or rarely without
squamellae, sometimes only squamellae present, or
absent. x = 17. Approximately 140 species, America.
XXVI.8. Subtribe Montanoinae H. Rob.
(1978).
Erect perennial herbs (?), shrubs, trees. Leaves
opposite, blades mostly ovate, entire or pinnatifid,
triplinerved, rarely 5–7-plinerved. Capitula terminal, mostly in paniculiform to corymbiform or
thyrsoid cymes, radiate, rarely discoid. Involucres
mostly hemispherical, phyllaries 3–7 in 1–2 series,
subequal. Receptacles convex, paleae accrescent
after anthesis and enfolding cypselae. Ray florets
neuter, corollas creamy white to white. Disc florets
bisexual, rarely functionally staminate, corollas
470
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
pentamerous, yellow, green-yellow, greyish white,
creamy white, without fibres embedding vascular
strands; stamens 5, anthers black or yellow,
appendages ovate with glandular trichomes; styles
with two vascular strands, style arms with divided
stigmatic surfaces, apices deltoid with a linear
appendage. Cypselae weakly compressed to obconical and shallowly quadrangular in cross-section,
black to brownish black or reddish brown, striate,
glabrous to sparsely pubescent. Pappus absent.
Only one genus:
1339. Montanoa Cerv. in La Llave & Lex.
Montanoa Cerv. in La Llave & Lex., Nov. Veg. Descr. 2: 11
(1825); Funk, Mem. New York Bot. Gard. 36: 1–133 (1982),
rev.
Characters of the subtribe. x = 19. About 25 species,
most species in Mexico, Central America, northern
South America south to northern Peru.
XXVI.9. Subtribe Rojasianthinae Panero
(2002).
Shrubs or small trees. Leaves opposite, blades
palmate, suborbicular in outline, palmately lobed.
Capitula in terminal, open paniculiform dichasial
cymes, radiate. Involucres hemispherical, phyllaries in 2–3 series, subequal, herbaceous. Receptacles
shallowly convex, paleae with acicular projections,
accrescent after anthesis and enfolding cypselae.
Ray florets neuter, corollas white, sometimes
suffused with pink on abaxial surfaces. Disc florets
bisexual, corollas pentamerous, white, distal half
black, without fibres embedding the vascular
strands; stamens 5, anthers black, filaments
densely papillose, papillae black, appendages
with glandular trichomes; styles with two vascular
strands, style arms with divided stigmatic surfaces,
apices acute to acuminate. Cypselae compressed,
obovate, raspberry colour when immature, brownblack, striate, glabrous to sparsely pubescent.
Pappus of several caducous, golden, barbellate
awns.
Only one genus:
1340. Rojasianthe Standl. & Steyerm.
Rojasianthe Standl. & Steyerm., Publ. Field Mus. Nat. Hist.,
Bot. Ser. 22: 311 (1940).
Characters of the subtribe. x = 19. One species,
R. superba Standl. & Steyerm., Mexico, Guatemala.
XXVI.10. Subtribe Rudbeckiinae H. Rob.
(1978); Urbatsch et al., Syst. Bot. 25:
539–565 (2000), phylog.
Erect annual or perennial herbs. Leaves alternate,
blades ovate, unlobed to pinnatifid. Capitula in terminal, paniculiform cymes, radiate, rarely discoid.
Involucres flat to reflexed, rarely hemispherical or
hemispherical at the beginning of anthesis, phyllaries in 2–3 series, sometimes dimorphic. Receptacles conical to columnar, paleae sometimes shed
with cypsela. Ray florets neuter. Disc florets bisexual, corollas pentamerous, without fibres embedding the vascular strands; stamens 5, anthers black,
sometimes shallowly calcarate, endothecium with
mostly radial thickenings; styles with two vascular strands, style arms with divided stigmatic surfaces, these broad and touching each other, sometimes with long papillose appendages. Cypselae
compressed, quadrate, striate, oblong to obpyramidal, glabrous to sparsely pubescent, sometimes
minutely winged. Pappus a small crown of minute
scales or few squamellae or absent.
Key to the Genera
1. Leaves pinnatifid, phyllaries in 2 dimorphic series, paleae deciduous with the cypselae, cypselae
sometimes minutely winged and with multiseriate
trichomes
1341. Ratibida
– Leaves mostly entire to trilobed, rarely plants with pinnatifid leaves associated with inflorescences, phyllaries in 2 or more series, subequal, paleae not deciduous
with the cypselae, cypselae never winged, rarely with
multiseriate trichomes
1342. Rudbeckia
1341. Ratibida Raf.
Ratibida Raf., Fl. Ludov. 73 (1817); Richards, Rhodora 70:
348–393 (1968), rev.; Correll & Johnston, Man. Vasc. Pl.
Texas: 1523–1736 (1970), reg. rev.; Urbatsch & Cox, Syst.
Bot. 15: 394–402 (1990), phylog.; Urbatsch & Jansen, Syst.
Bot. 20: 28–39 (1995), phylog.; Urbatsch et al., Syst. Bot. 25:
539–565 (2000), phylog.
Leaf-blades ovate, unlobed or pinnatifid. Capitula
radiate. Involucres flat to reflexed, phyllaries in
2 series, dimorphic. Receptacles columnar, paleae
deeply keeled and densely pubescent distally, shed
with cypsela. Ray floret corollas yellow, reddish
brown or bicoloured. Disc floret corollas yellow,
Compositae
purple or greenish. Cypselae oblong to oblanceolate, tangential angles weakly defined, glabrous to
sparsely pubescent, sometimes with a minute wing.
Pappus of 1–2 small awns or projections, rarely
a minute crown or 1 awn or absent. x = 16. Seven
species, North America.
1342. Rudbeckia L.
Rudbeckia L., Sp. Pl. 2: 906 (1753); Perdue, Rhodora 59:
293–299 (1957), subg. rev.; Correll & Johnston, Man. Vasc.
Pl. Texas: 1523–1736 (1970), reg. rev.; Urbatsch & Cox, Syst.
Bot. 15: 394–402 (1990), phylog.; Urbatsch & Jansen, Syst.
Bot. 20: 28–39 (1995), phylog.; Urbatsch et al., Syst. Bot. 25:
539–565 (2000), phylog.
Dracopis Cass. (1825).
Leaf-blades ovate to broadly ovate, unlobed or pinnatifid, rarely sessile and cordate, sometimes glaucous. Capitula in terminal, paniculiform cymes,
radiate, rarely discoid. Involucres flat to reflexed,
rarely hemispheric or hemispherical at the beginning of anthesis. Receptacles conic to columnar.
Ray floret corollas golden to lemon yellow, orange
or reddish, or bicoloured. Disc floret corollas yellow to purplish, often bicoloured. Cypselae oblong
to obpyramidal, glabrous or with some thick multicellular trichomes on radial angles. Pappus a small
crown of minute scales or 2–4 small scales or absent.
x = 16, 18, 19. Seventeen species, North America.
XXVI.11. Subtribe Spilanthinae Panero
(2005).
Erect or decumbent annual or perennial herbs,
sometimes rooting at the nodes, rarely scandent
shrubs. Leaves opposite, blades linear to ovate,
sometimes reniform, unlobed, triplinerved. Capitula axillary or terminal, solitary or in simple to
congested cymes, on sometimes fistulose peduncles, discoid or radiate. Involucres campanulate to
hemispherical, phyllaries in 1–5 series, subequal,
rarely dimorphic, mostly herbaceous. Receptacles
convex to conical especially with age, mostly
paleate, paleae chartaceous, rarely coriaceous,
bases slightly decurrent. Ray florets pistillate.
Disc florets bisexual, corollas pentamerous,
rarely tetramerous, without fibres embedding
the vascular strands; stamens 5, rarely 4; styles
with two vascular strands, style arms with fused
stigmatic surfaces, apices acute and papillose,
without appendages. Ray cypselae obcompressed,
triquetrous, obovoid, sparsely to densely ciliate.
Disc cypselae compressed, sometimes peripheral
471
ones triquetrous, terete, sometimes quadrate
and narrowly rhombic, rarely square in outline,
black or brown, rarely striate, mostly shallowly or
sometimes conspicuously winged, corky, ciliate,
glabrous to sparsely pubescent. Pappus a minute
crown, a single awn fused to a broad ring around
the neck of the cypsela or more commonly of
2–3 slender awns as a continuation of the wings,
sometimes with squamellae in between.
Key to the Genera
1. Capitula with 4 florets
1347. Tetranthus
– Capitula with more than 4 florets
2
2. Cypselae strongly striate or shallowly ribbed; pappus
a whitish crown, sometimes with a single awn
1344. Oxycarpha
– Cypselae and pappus otherwise
3
3. Leaves subsessile, rarely petiolate; cypselae winged or
with broad corky edges, sometimes lacerate and ending in a stout shoulder or awn
1346. Spilanthes
– Leaves petiolate; cypselae narrowly winged or with
narrow corky edges
4
4. Plants herbaceous; leaves thin; capitula discoid or radiate; corollas yellow or white
1343. Acmella
– Plants shrubby, sometimes scandent; leaves coriaceous; capitula discoid; corollas white, rarely
purplish-white
1345. Salmea
Genera of Spilanthinae
1343. Acmella Rich. ex Pers.
Acmella Rich. ex Pers., Syn. Pl. 2: 472 (1807); Jansen, Syst.
Bot. Monogr. 8: 1–115 (1985), rev.
Erect or decumbent annual or perennial herbs,
sometimes rooting at the nodes. Leaf-blades
filiform to mostly ovate, triplinerved. Capitula terminal or axillary, solitary or in open paniculiform
cymes, discoid or radiate. Involucres hemispherical, sometimes patent with age, phyllaries in 1–3
series, subequal. Receptacles conic, paleate. Ray
florets pistillate, purplish to yellow-orange or
white. Disc floret corollas pentamerous or sometimes tetramerous, creamy white to yellow-orange;
anthers black, appendages with or without glands.
Ray cypselae triquetrous, obovoid to ellipsoid,
black, glabrous to densely pubescent, sometimes
densely ciliate. Disc cypselae compressed, obovoid
to ellipsoid, black, rarely brownish, sometimes
with a light brown corky margin, pubescence as
ray cypselae. Pappus absent or of few soft bristles,
or more commonly of two persistent, slender awns
at the angles of the cypsela. x = 12, 13. About
472
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
30 species, pantropical, most species in the New
World.
corky, ciliate margin. Pappus of 1–3 subequal
awns. x = 16. Six species, pantropical.
1344. Oxycarpha S.F. Blake
1347. Tetranthus Sw.
Oxycarpha S.F. Blake, Contr. Gray Herb. ser. 2, 53: 52 (1918).
Tetranthus Sw., Prodr. 7: 115 (1788).
Stoloniferous, succulent, perennial herbs or weak
low shrubs. Leaf-blades filiform, succulent. Capitula terminal, solitary, discoid. Involucres campanulate, phyllaries in 3–5 series, gradate, coriaceous.
Receptacles conic, paleae coriaceous, apices aristate to acicular. Corollas white, yellow (?); anthers
brown. Cypselae oblanceolate, weakly biconvex but
mostly terete with conspicuous ribs, one side with
a minute wing wider at base, black, glabrous. Pappus a crown sometimes with a single awn. One
species, O. suaedifolia S.F. Blake, Venezuela.
Prostrate perennial herbs. Leaf-blades oval to
reniform, triplinerved, unlobed. Capitula axillary,
scapose, solitary, discoid. Involucres campanulate,
phyllaries dimorphic, outer approximately 1/10
the length of inner, inner enclosing the cypselae
(perigynia). Receptacles epaleate. Disc florets 4,
corollas sometimes tetramerous, white; anthers
black, appendages with minute glandular trichomes; Cypselae quadrate, obpyramidal, black,
glabrous. Pappus a minute crown. Between two
and four species, Caribbean.
1345. Salmea DC.
XXVI.12. Subtribe Verbesininae Benth.
(1873).
Salmea DC., Cat. Pl. Horti Monsp.: 140 (1813), nom. cons.;
Blake, J. Bot. 53: 193–201 (1915), rev.; Bolick, Syst. Bot. 16:
462–477 (1991), rev.
Erect or scandent shrubs. Leaf-blades lanceolate
to ovate, coriaceous, triplinerved. Capitula in
open paniculiform cymes of variously congested
corymbiform cymes, discoid. Involucres campanulate to hemispherical, sometimes patent with
age, phyllaries in 2–5 series, subequal or gradate.
Receptacles conical, paleate. Corollas white or
creamy white, sometimes purplish; anthers brown
or black, appendages with or without glandular
trichomes. Cypselae obovoid to ellipsoid, compressed, black, rarely brownish sometimes with
a light brown corky margin, sparsely pubescent.
Pappus of 2(–3) slender awns, sometimes with
a few squamellae in between, rarely a crown.
x = 18. Ten species, neotropics.
1346. Spilanthes Jacq.
Spilanthes Jacq., Enum. Syst. Pl. 8, 28 (1760); Jansen, Syst.
Bot. 6: 231–257 (1981), rev.
Perennial herbs, some prostrate and becoming
woody at base. Leaf-blades linear to elliptic, weakly
triplinerved. Capitula terminal, rarely axillary,
solitary or in simple cymes, discoid. Involucres
hemispherical, phyllaries in 2–3 series, subequal.
Receptacles convex to conic, paleate. Corollas
white or purplish white; anthers black, appendages
without glands. Cypselae obovate, strongly compressed at maturity, shallowly quadrate, rhombic
in outline, in most species with a conspicuous
Erect, rarely decumbent annuals or perennial
herbs, shrubs, rarely vines, sometimes trees.
Leaves alternate or opposite, rarely in whorls,
rarely forming rosettes, blades various, 3–5nerved, rarely pectinate, rarely perfoliate, unlobed
to deeply lobed, pinnately veined or triplinerved.
Capitula terminal, simple, sometimes scapose, or
in paniculiform or corymbiform cymes, rarely
thyrses, discoid or radiate. Involucres cylindrical
to hemispherical, phyllaries in 2–4 series, subequal
or gradate. Receptacles flat to convex, rarely
globose, paleate. Ray florets pistillate, sometimes
pistillate but sterile or neuter, corollas rarely with
biseriate trichomes on tube. Disc florets bisexual,
corollas pentamerous, rarely tetramerous, without
fibres embedding the vascular strands; stamens
5, rarely 4; styles with two vascular strands, style
arms with divided stigmatic surfaces, appendages
lacking. Cypselae compressed, rarely obpyramidal
and quadrate, sometimes stipitate, ray cypselae
sometimes triquetrous and obpyramidal, winged
or wingless, black or brown, not striate, rarely
striate, glabrous to densely pubescent. Pappus of
awns or squamellae at angles of cypsela.
Key to the Genera
1. Cypsela quadrate, obpyramidal with a pappus of 4
awns or squamellae at angles of cypsela, sometimes
these longer and alternating with 4 squamellae
1350. Tetrachyron
– Cypsela and pappus otherwise
2
2. Cypselae winged, not stipitate
1351. Verbesina
Compositae
– Cypselae not winged, stipitate
3
3. Ray corollas white
1348. Podachaenium
– Ray corollas yellow
1349. Squamopappus
Genera of Verbesininae
1348. Podachaenium Benth.
Podachaenium Benth., Vidensk. Meddel. Dansk Naturhist.
Foren. Kjobenhavn 1852 (5/7): 98–99 (1853); Jansen et al.,
Syst. Bot. 7: 476–483 (1982), circumscr.; Turner & Panero,
Phytologia 73: 143–148 (1992), reg. rev.
Shrubs or trees. Leaves opposite, blades ovate to
broadly ovate, palmate, shallowly serrate, venation
palmate or triplinerved, rarely pectinate. Capitula
in terminal, corymbiform cymes or in thyrses, radiate. Involucres hemispherical, sometimes becoming reflexed. Receptacles convex to globose. Ray florets pistillate, corollas white to creamy white. Disc
corollas yellow to yellow-orange; anthers black or
yellow, appendages with glandular trichomes; style
arms with broadly acute to round apices. Cypselae
compressed, obovate, those of the ray obpyramidal and triquetrous, sometimes conspicuously tapered at the base, sparsely pubescent. Pappus of
several erose squamellae, those on the angles of
the cypsela longer. x = 18, 19. Six species, Mexico,
Central America, Colombia.
1349. Squamopappus R.K. Jansen, N.A. Harriman
& Urbatsch
Squamopappus R.K. Jansen, N.A. Harriman & Urbatsch,
Syst. Bot. 7: 480 (1982).
Shrubs or small trees. Leaves opposite below,
then alternate, blades ovate, shallowly serrate,
triplinerved. Capitula in terminal, compact
corymbiform cymes, radiate. Involucres campanulate to subhemispherical. Receptacles convex.
Ray florets pistillate, corollas golden yellow. Disc
corollas yellow; anthers black, appendages without
glandular trichomes; style arms with acute apices.
Cypselae compressed, obovate, those of the ray
obpyramidal and triquetrous, somewhat tapered
at the base, sparsely pubescent. Pappus of several
erose squamellae, those on the angles of the
cypsela longer, awn-like. x = 19. One species,
S. skutchii (S.F. Blake) R.K. Jansen, N.A. Harriman
& Urbatsch, Mexico, Guatemala.
1350. Tetrachyron Schltr
Tetrachyron Schltr, Linnaea 19: 744 (1847); Wussow &
Urbatsch, Syst. Bot. 4: 297–318 (1979), rev.; Jansen et al.,
Syst. Bot. 7: 476–483 (1982), circumscr.
473
Erect, open to densely branched shrubs. Leaves
opposite, blades linear to lanceolate, ovate, trullate, or rarely suborbicular to round, entire to
shallowly serrate, pinnately veined or triplinerved,
sometimes densely pubescent on abaxial surfaces.
Capitula in terminal, simple or paniculiform
cymes or thyrses, radiate. Involucres turbinate to
hemispherical. Receptacles shallowly convex to
convex. Ray florets pistillate, corollas yellow to
yellow-orange, sometimes with biseriate trichomes
on tube. Disc corollas yellow or yellow-orange;
anthers yellow, appendages with or without glandular trichomes; style arms with acute apices and
conspicuous resin and crystal deposits. Cypselae
obpyramidal, quadrate, conspicuously tapered at
the base, sometimes striate, with a conspicuous
carpopodium, sparsely pubescent. Pappus of
four awns or squamellae, sometimes squamellae
alternating with awns. x = 15, 16, 17. About eight
species, Mexico, Guatemala.
1351. Verbesina L.
Fig. 96
Verbesina L., Sp. Pl. 2: 901 (1753), nom. cons.; Robinson
& Greenman, Proc. Amer. Acad. Arts 34: 534–566 (1899),
rev.; Correll & Johnston, Man. Vasc. Pl. Texas: 1523–1736
(1970), reg. rev.; Olsen, Pl. Syst. Evol. 149: 47–63 (1985),
sect. rev.; Turner, Pl. Syst. Evol. 150: 237–262 (1985), sect.
rev.; Olsen, Brittonia 38: 362–368 (1986), sect. rev.; Olsen,
Sida 13: 45–56 (1988), sect. rev.; Panero & Jansen, Amer.
J. Bot. 84: 382–392 (1997), phylog.
Annual or perennial herbs, erect, rarely vine-like
or straggly shrubs, or trees sometimes up to
25 m tall. Leaves alternate or opposite, sometimes with petiolar wings decurrent, sometimes
forming dainty rosettes, blades linear, lanceolate,
ovate, suborbicular, sagittate, deltate, unlobed or
deeply dissected, rarely perfoliate, triplinerved,
rarely pinnately veined. Capitula terminal, solitary, sometimes scapose, or in paniculiform or
corymbiform cymes, discoid or radiate. Involucres
cylindrical, turbinate, campanulate, hemispheric.
Receptacles flat to convex, rarely globose, paleae
tapered, sometimes with rounded flat tips. Ray
florets pistillate, sometimes pistillate but sterile or
neuter, corollas white, orange, red, yellow, greenish
white, white-pink, purplish green. Disc corollas
rarely tetramerous, yellow, orange, red, green,
white, purple or a combination of these colours
on throat and lobes; anthers brown or black,
rarely red, pink or yellow, anther appendages
with or without glandular trichomes; style arms
long and tapered or with broadly acute or deltoid
474
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
hemispherical, phyllaries in 2–3 series, subequal.
Receptacles convex to conical, paleate. Ray florets
pistillate. Disc florets bisexual, corollas pentamerous, without fibres embedding the vascular
strands; stamens 5, anther appendages with glandular trichomes; styles with two vascular strands,
style arms with fused stigmatic surfaces. Cypselae
compressed and weakly quadrate, obovate, sometimes asymmetrical with adaxial side straight
to concave and abaxial side convex, glabrous.
Pappus absent or rarely of a few squamellae on ray
cypselae.
Key to the Genera
1. Leaves deeply dissected, pinnatifid; paleae linear; anthers black
1352. Hybridella
– Leaves unlobed to trilobed, sometimes lobes dissected;
paleae oval, never linear; anthers yellow
1353. Zaluzania
1352. Hybridella Cass.
Hybridella Cass., Bull. Sci. Soc. Philom. Paris 1817: 12
(1817); Olsen, Madroño 24: 29–36 (1977), re-establ.
Fig. 96. Compositae-Heliantheae. Verbesina pietatis. A Portions of flowering plant. B Flowering capitulum with facing
ray floret removed. C Ray floret. D Disc floret. E Abaxial
view of style tip. F Palea. G Slightly immature disc cypsela
(wings starting to develop). (Reproduced from Flora NovoGaliciana, vol. 12, with permission of the University of
Michigan Herbarium; see McVaugh 1984)
apices. Cypselae strongly compressed, sometimes ray cypselae triquetrous, broadly obovate,
oblanceolate to narrowly cuneate, symmetrically
to asymmetrically winged, stramineous to dark
brown or blackish, sparsely pubescent distally,
the wings thin to corky. Pappus of two awns,
erect, rarely uncinate, sometimes triquetrous ray
cypselae with 3 awns, or rarely absent. x = 16, 17,
18. About 300 species, America, highest number of
species in Mexico and tropical Andes.
XXVI.13. Subtribe Zaluzaniinae H. Rob.
(1978).
Erect annual or perennial herbs, shrubs. Leaves
alternate, petiolate, blades mostly ovate, unlobed,
rarely trilobed or pinnatifid, triplinerved. Capitula
in terminal, paniculiform or corymbiform cymes,
discoid or radiate. Involucres campanulate to
Perennial herbs. Leaf-blades lanceolate in outline,
twice pinnate. Capitula in terminal, paniculiform
cymes, radiate. Involucres hemispheric, phyllaries
reflexed. Receptacles conical, paleae linear. Ray
floret corollas yellow. Disc floret corollas yellow;
anthers black; style arms with somewhat canaliculate stigmatic surfaces, apices round to truncate.
Cypselae compressed and weakly quadrate, obovate, asymmetrical, glabrous. Pappus absent.
x = 16. One species, H. globosa Cass., Mexico.
1353. Zaluzania Pers.
Zaluzania Pers., Syn. Pl. 2: 473 (1807); Olsen, Rhodora 81:
449–500 (1979), rev.
Erect annual or perennial herbs, shrubs. Leafblades cordate to ovate, serrate to deeply laciniate.
Capitula in terminal, paniculiform cymes, discoid
or radiate. Involucres campanulate to hemispheric.
Receptacles convex to shallowly conical. Ray floret
corollas yellow, rarely white. Disc floret corollas
yellow, rarely white; anthers yellow; style arms
with acute to acuminate apices. Cypselae compressed and weakly quadrate, obovate, sometimes
asymmetrical, glabrous. Pappus absent or rarely of
a few squamellae on ray cypselae. x = 17, 18. Ten
species, Mexico, south-western USA.
Compositae
XXVI.14. Subtribe Zinniinae Benth. (1873).
Erect annual or perennial herbs, shrubs. Leaves alternate or opposite, petiolate or sessile, blades linear to ovate, unlobed, pinnately veined to triplinerved. Capitula terminal, rarely axillary, solitary
and scapose or in open paniculiform cymes on
sometimes fistulose peduncles, radiate, rarely discoid. Involucres turbinate to hemispherical, rarely
globose and phyllaries reflexed, phyllaries in 2–5
series, subequal to strongly gradate. Receptacles
convex to conic, globose, paleate, rarely epaleate.
Ray florets pistillate, rarely neuter, corollas marcescent and fused to the cypselae, rarely deciduous,
often without a tube. Disc florets bisexual, sometimes functionally staminate, corollas pentamerous, tubes and throat bases densely thickened with
additional layers of cells, without fibres embedding
the vascular strands; stamens 5, anther appendages
with glandular trichomes; styles with two vascular
strands, style arms with fused stigmatic surfaces,
rarely divided, apices acute to deltate with small
papillose appendages. Ray cypselae obpyramidal
to terete or obcompressed, black to stramineous,
sometimes with striations, sometimes with uncinate trichomes. Pappus absent or of mutiple awns
or rarely a crown, ray cypselae with either 1 or 3
awns at the angles of the cypselae.
475
– Ray corollas with a tube; cypselae terete to quadrate;
pappus of cypselae of a few teeth or a crenulate, minute
rim
1355. Heliopsis
Genera of Zinniinae
1354. Echinacea Moench
Echinacea Moench, Methodus 591 (1794); McGregor, Univ.
Kansas Sci. Bull. 48: 113–142 (1968), rev.; Correll & Johnston,
Man. Vasc. Pl. Texas: 1523–1736 (1970), reg. rev.; Urbatsch
& Cox, Syst. Bot. 15: 394–402 (1990), phylog.; Urbatsch &
Jansen, Syst. Bot. 20: 28–39 (1995), phylog.; Urbatsch et al.,
Syst. Bot. 25: 539–565 (2000), phylog.; Binns et al., Syst. Bot.
27: 610–632 (2002), rev.
Annual or perennial herbs. Leaves alternate, blades
linear to ovate, pinnately veined or triplinerved,
entire to shallowly serrate. Capitula terminal, solitary and scapose or in open paniculiform cymes,
radiate. Involucres patent or reflexed, phyllaries in
2–4 series, reflexed. Receptacles convex to strongly
conical or globose, paleate, paleae with pointed or
rounded, stiff tips. Ray florets neuter, corollas pink
or purple, rarely white or yellow. Disc florets bisexual, corollas green, pink, purple, rarely yellow;
anthers mostly black; style arms with fused stigmatic surfaces, apices acute. Cypselae compressed
to quadrate, cuneate, black, glabrous to sparsely
pubescent. Pappus a small crown. x = 11. Between
four and 11 species, Canada, USA.
Key to the Genera
1.
–
2.
–
3.
–
4.
–
5.
–
6.
–
7.
Disc florets functionally staminate
1360. Zinnia
Disc florets bisexual
2
Paleae at anthesis 1.5–2 times as long as disc florets 3
Paleae at anthesis as long as or shorter than disc florets
4
Ray florets sterile
1354. Echinacea
Ray florets fertile
1358. Tehuana
Tubes of ray and disc corollas and abaxial surfaces
of lobes of disc corollas densely covered with black,
moniliform trichomes with an expanded terminal cell;
plants aquatic; northern Mexico (western Durango
state)
1359. Trichocoryne
Tubes and lobes of ray and disc corollas glabrous or
variously pubescent but never with black moniliform
hairs; plants terrestrial, rarely aquatic; American 5
Pappus of ray cypselae with at least one broad awn as
a continuation of adaxial angle of cypsela and opposite
the ray corolla sinus
6
Pappus of ray cypselae various or absent but always
without a broad awn opposite the ray corolla sinus 7
Ray cypselae with 1 awn
1356. Philactis
Ray cypselae with 3 awns
1357. Sanvitalia
Ray corollas often without a tube; cypselae either
compressed or obcompressed, sometimes shallowly
quadrate to terete; pappus of cypselae absent or of 1
or 2 awns
1360. Zinnia
1355. Heliopsis Pers.
Heliopsis Pers., Syn. Pl. 2: 473 (1807), nom. cons.; Fisher,
Ohio J. Sci. 57: 171–191 (1957), rev.
Erect to decumbent annual or perennial herbs,
weak shrubs. Leaves opposite, sometimes alternate
distally, blades filiform to suborbicular, mostly
ovate, pinnately veined or triplinerved, entire
to deeply serrate or lobed. Capitula terminal,
solitary, sometimes on long peduncles or in
open paniculiform cymes on sometimes fistulose peduncles, radiate or discoid. Involucres
turbinate to hemispherical, phyllaries subequal,
herbaceous. Receptacles convex to conical to
columnar with age, paleae with rounded flat tips.
Ray florets pistillate, corollas yellow to orange,
purple, sometimes greenish with age. Disc florets
bisexual, corollas yellow to orange, purple; anthers
mostly black, rarely yellow; style arms with fused
stigmatic surfaces, apices acute. Cypselae ovoid
to terete, weakly 3–4-angled, greenish brown to
black, smooth to tuberculate, glabrous to sparsely
476
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
puberulent. Pappus a minute crenulate rim or of
2–3 small teeth. x = 14. Fifteen species, Central
and northern South America.
1356. Philactis Schrad.
Philactis Schrad., Index Sem. (Göttingen) (1832); Torres,
Brittonia 21: 322–332 (1969), rev.
Shrubs. Leaves opposite, petiolate, blades lanceolate to ovate, triplinerved, serrate. Capitula
terminal, solitary or in simple cymes, radiate.
Involucres hemispherical, phyllaries in 2–3 series,
subequal. Receptacles convex, paleate, paleae with
pointed, stiff tips. Ray florets pistillate, corollas
yellow to yellow orange, sometimes dull greenish
to blue-green with age, tube very short. Disc florets
bisexual, corollas yellow-orange; anthers black,
appendages ovate; style arms with fused stigmatic
surfaces, apices acute. Ray cypselae obpyramidal,
triquetrous, brown with lengthwise striations.
Disc cypselae compressed, obpyramidal, biconvex
to quadrate, brown, without phytomelanin (?)
Pappus of mostly 2, rarely many awns and of 1 awn
on adaxial side of ray cypsela facing corolla sinus.
x = 14. One polymorphic or 3 or 4 species, Mexico,
Guatemala.
1357. Sanvitalia Lam.
Sanvitalia Lam., J. Hist. Nat. 2: 176 (1792); Torres, Brittonia
16: 417–433 (1964), rev.
Erect or prostrate annual or perennial herbs, rarely
shrubs. Leaves opposite, blades linear to ovate,
mostly triplinerved, entire to shallowly serrate.
Capitula terminal, solitary or in open paniculiform cymes, radiate. Involucres hemispherical,
phyllaries in 2–4 series, subequal. Receptacles
convex, paleate, paleae with pointed, stiff tips.
Ray florets pistillate, corollas yellow, greenish
yellow or yellow-orange, without tube. Disc florets
bisexual, corollas yellow, greenish yellow or
deep purple; anthers black, brown or purplish;
style arms with fused stigmatic surfaces, apices
acute. Ray cypselae obpyramidal, triquetrous to
terete, stramineous with uncinate trichomes. Disc
cypselae compressed, obpyramidal, triquetrous to
quadrate, winged, peripheral cypselae sometimes
wingless. Pappus absent or of many awns or of
three awns one each on adaxial and tangential
sides of ray cypselae. x = 8, 9, 11. Seven species,
Mexico, Central America, Bolivia, Argentina.
1358. Tehuana Panero & Villaseñor
Tehuana Panero & Villaseñor, Syst. Bot. 21: 555 (1996)
[1997].
Erect annuals. Leaves opposite, blades ovate,
triplinerved, shallowly serrate. Capitula terminal,
solitary or in open paniculiform cymes, radiate.
Involucres patent or reflexed, phyllaries in 2–3
series, subequal. Receptacles conical to columnar
with age, paleate, paleae with acicular tips. Ray
florets pistillate, corollas orange-yellow, tube
very short. Disc florets bisexual, corollas green,
yellow-orange distally; anthers black, appendages
with glandular trichomes; style arms with fused
stigmatic surfaces, apices acute. Ray cypselae
obpyramidal, triquetrous, brown-black, minutely
strigose with lengthwise striations. Disc cypselae
obpyramidal, rounded-quadrate in cross-section,
the 4 ribs with minute twin trichomes, areas
between ribs with minute uniseriate trichomes.
Pappus absent or of one awn on adaxial side of ray
cypselae. x = 14. One species, T. calzadae Panero
& Villaseñor, Mexico.
1359. Trichocoryne S.F. Blake
Trichocoryne S.F. Blake, Contr. U.S. Natl Herb. 22: 648–649
(1924).
Aquatic annual or perennial (?) herbs. Leaves opposite, sessile, linear to narrowly ovate, uninerved, entire. Capitula axillary, solitary on short shoots, radiate. Involucres hemispherical, phyllaries in 2 series,
subequal. Receptacles convex to conical, epaleate.
Ray florets 5, pistillate, sterile, corollas white with
two green-black veins on abaxial side, not marcescent. Disc florets bisexual, corollas white to creamy
white with conspicuous black trichomes on tube
and abaxial side of lobes; anthers black; style arms
with divided stigmatic surfaces, apices acute to
deltate. Cypselae compressed to quadrate, obovate
to cuneate, black, glabrous. Pappus absent. One
species, T. connata S.F. Blake, north-western Mexico.
1360. Zinnia L.
Zinnia L., Syst. Nat., ed. 10: 1189, 1221, 1377 (1759), nom.
cons.; Torres, Brittonia 15: 1–25, 290–302 (1963), part. rev.
Crassina Scepin (1758), nom. rej.
Lepia Hill (1759), nom. rej.
Tragoceros Kunth (1818).
Erect annual or perennial herbs or shrubs. Leaves
opposite, linear to ovate, entire, rarely serrate. Ca-
Compositae
477
limbs sometimes extremely short or lacking,
apices conspicuously trilobed, rarely shallowly
trilobed. Disc florets bisexual or functionally
staminate, actinomorphic or sometimes the
peripheral zygomorphic with 3 longer abaxial
lobes, pentamerous, rarely tetramerous, throats
gradually tapered or abruptly expanded above
the tube, normally pubescent with glandular
trichomes; anthers sometimes free, appendages
lanceolate to ovate, sometimes cochleate, with or
without glandular trichomes, appendages weakly,
rarely strongly sclerified, endothecium of quadrate
to oblong or isodiametric cells with 1–5 polar
thickenings, sometimes with radial thickenings,
rarely thickenings surrounding the whole cell;
style arms of disc florets wholly or partially fused
or spreading with parallel, rarely fused stigmatic
surfaces, appendages wanting to prominent. Ray
cypselae essentially similar to disc cypselae,
except that sometimes variously enclosed by inner
phyllaries or paleae, obcompressed, broadly ovate
to suborbicular, sometimes incurved, black, rarely
with a basal caruncle or elaiosome (Guardiola),
otherwise obconic to obpyramidal, sometimes
quadrate, striate, glabrous or densely pubescent.
XXVII. Tribe Millerieae Lindl. (1829). Disc cypselae obconic to obpyramidal, sometimes incurved, quadrate to round, rarely oval in
cross-section, subterete to shallowly angled, black,
J.L. Panero
glabrous to densely pubescent. Pappus absent or
Annual or perennial herbs, shrubs or trees, variously of minute scales, acuminate scales, or
sometimes rosulate or caulirosulate; herbage barbellate or plumose bristles, bristles of equal or
sometimes conspicuously and densely glandular, unequal length, sometimes caducous.
The tribe contains 34 genera and approximately
sometimes viscous. Leaves usually opposite,
petiolate or sessile, sometimes leaf bases ampliated 400 species found mostly in central Mexico and the
or forming a cupule (invaginated) around the stem northern Andes with a few species in tropical re(Espeletiinae), blades linear to ovate, sometimes gions of the Old World, especially Africa. Some
suborbicular, entire to variously serrate to lacerate members of the Espeletiinae are the defining eleor pectinate, triplinerved, sometimes pinnately ment of the paramos of northern South America.
Millerieae have recently been resurrected
veined or pentanerved. Capitula in terminal
or axillary, open to congested paniculiform or (Panero et al. 2001c; Panero and Funk 2002)
corymbiform cymes, sometimes scapose, radiate to accommodate the genera variously placed
or discoid, rarely disciform. Involucres turbinate to by Robinson (1981) and other authors (Stuessy
hemispheric, phyllaries in 1–5+ series, sometimes 1977; Karis and Ryding 1994a) in subtribes
dimorphic or subequal to gradate, rarely outer Desmanthodiinae, Espeletiinae, Galinsogiinae,
series broadly deltate, valvate and enclosing the Guardiolinae, Melampodiinae and Milleriinae.
developing capitulum, commonly moderately to The present circumscription of the tribe is based
densely pubescent, sometimes second series fused primarily on results of chloroplast DNA sequence
to developing ray cypselae (Melampodiinae). analyses (Panero et al. 2001c). The tribe does not
Receptacles convex to conic, rarely flat, wholly or have any single character which defines it but
partially paleate, paleae flat to navicular, some- most of its species tend to have opposite leaves,
times cucullate. Ray florets pistillate, rarely neuter, glandular herbage, scarious paleae, subterete
sometimes in multiple series or rows (especially cypselae, a pappus, when present, of scales or brisin Espeletiinae), corolla rarely shallowly bilabiate, tles arranged radially on the neck of the cypsela,
pitula terminal, solitary or in open paniculiform
cymes on sometimes fistulose peduncles, radiate,
rarely discoid. Involucres turbinate, campanulate
or hemispherical, phyllaries in 2–5 series, subequal to highly gradate. Receptacles convex to conic,
paleate, paleae with rounded, flat tips. Ray florets
pistillate, corollas yellow, green, burgundy, red, orange, pink, marcescent, often without a tube. Disc
florets bisexual, sometimes functionally staminate,
sometimes zygomorphic, corollas pink, red, yellow,
yellow-orange, burgundy, sometimes with conspicuous papillae on adaxial surface; anthers yellow,
brown or black; style arms with fused stigmatic
surfaces, sometimes with glandular trichomes on
abaxial surfaces, apices acute. Ray cypselae mostly
obcompressed, triquetrous, black to brown or greyish, smooth to tuberculate, glabrous to variously
pubescent. Disc cypselae compressed, convex to
shallowly quadrate, tuberculate or smooth, glabrous to variously pubescent, sometimes winged.
Pappus absent or of one or two awns. x = 10, 11, 12.
About 25 species, south-western USA, neotropical
region.
478
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
and trilobed ray corollas. The tribe contains
five main lineages, namely Dyscritothamninae,
the three subtribes Galinsoginae, Jaegeriinae
and Desmanthodiinae, Guardiolinae, Milleriinae
(including Espeletiinae) and Melampodiinae.
Millerieae are sister to Madieae, Perityleae and
Eupatorieae, and these collectively are sister
to Heliantheae (Panero and Funk 2002). Based
on historical considerations, Guardiolinae are
tentatively placed in Millerieae, as there is no
strong support for their inclusion here or in any
of the sister tribes. Subtribe Dyscritothamninae
was recently described (Panero 2005), based on
results from molecular studies of chloroplast
DNA (Panero et al. 2001c), to include the genera Dyscritothamnus, Tetragonotheca, Tridax,
Cymophora and Bebbia. All these genera, with
the exception of Tetragonotheca, share variously
pubescent, subterete cypselae with a pappus of
plumose or barbellate scales or bristles; the pappus
is rarely absent. Tetragonotheca is distinctive in the
group by having quadrate cypselae with a pappus
of minute scales. The inclusion of Tetragonotheca
in Dyscritothamninae, although well-supported
by the available data, is nevertheless perplexing,
as its morphology is quite divergent from that
of the other genera of the subtribe. Molecular
data support the sister-group relationship of
Dyscritothamninae and Melampodiinae.
Melampodiinae have always been recognized
as a distinctive group within the Heliantheae
alliance (Stuessy 1977; Robinson 1981; Karis
and Ryding 1994a) because of their involucres
with dimorphic phyllaries, the outer series being herbaceous and the inner fused to the ray
cypselae. Furthermore, all Melampodiinae have
functionally staminate disc florets. Molecular data
support a narrow interpretation of Melampodiinae
to include only the genera Acanthospermum,
Lecocarpus and Melampodium. The cypselae of
Melampodiinae, like those of Milleriinae, lack
a pappus. Melampodiinae and Dyscritothamninae
are sister to all the other subtribes of Millerieae,
with the exception of Guardiolinae. Results from
studies based on sequence data of chloroplast DNA
(Panero et al. 2001c) and the ITS region of nuclear
ribosomal DNA (Panero and Plovanich-Jones,
unpubl. data) provide important insights into
the evolution and phylogenetic relationships of
subtribe Galinsoginae. These studies reveal that
Galinsoga, as traditionally circumscribed (Canne
1977), is a paraphyletic assemblage and that the
genera Sabazia and Alloispermum need to be
included in its concept to maintain a monophyletic
Galinsoga. I have opted to follow the traditional
circumscription of Alloispermum, Galinsoga
and Sabazia, until more comprehensive studies
are available. The monotypic genus Freya from
western Venezuela is considered here to be part
of Sabazia because it shares with this genus
opposite leaves, trilobed white ray corollas and
scarious, striate phyllaries. Aphanactis, Oteiza,
Schistocarpha and Selloa are sister to the Galinsoga
lineage. Aphanactis was revised by Turner (1980)
and subsequently emended by Robinson (1997)
who transferred all species of Aphanactis from
Mexico and Central America to Selloa, arguing
that those species were extraneous in Aphanactis
because their peduncles did not elongate after
anthesis, a characteristic seen in all species of
South American Aphanactis. Molecular studies
show that the Mexican species of Aphanactis are
sister to the Andean species, and not to Selloa.
Based on these results, Selloa is here recognized to
contain only one species, S. plantaginea of central
Mexico. Selloa plantaginea differs from Aphanactis
by having a pappus of bristles and fused stigmatic
surfaces. The same data support, albeit weakly, the
sister relationship of Oteiza and Alepidocline with
the monotypic genus Cuchumatanea in a derived
position within Alepidocline; Cuchumatanea is
here placed in the synonymy of Alepidocline.
Monographic studies of these genera may support
the placement of Alepidocline in Oteiza; I have
refrained from merging them here, as their chromosome numbers are different and, in the case of
Oteiza, may reflect past hybridization (Strother
and Panero 1994). The same studies show that
Desmanthodium and Jaegeria are basal elements
of Galinsoginae, and they are recognized here as
distinctive subtribes. Galinsoginae, Jaegeriinae
and Desmanthodiinae are sister to the Milleriinae/Espeletiinae clade. Molecular studies of
members of the Heliantheae alliance (Panero et al.
2001c) show that Milleriinae contain two main lineages, one composed of Smallanthus, Ichthyothere
and Espeletiinae, another containing the other
members of the subtribe. These results have been
corroborated independently by Rauscher (2002)
who, by using a more extensive sampling, was
able to provide some important insights into the
relationships of the genera of the subtribe. His
findings show that several genera of Milleriinae are
not monophyletic, namely Rumfordia, Sigesbeckia
and Trigonospermum. They also demonstrate that
the traditional seven genera of the Espeletiinae,
Compositae
as outlined by Cuatrecasas (1976), appear not to
be monophyletic and that the characters used
to define them, including mostly phyllotaxy
and inflorescence morphology, have evolved in
parallel several times. These data have persuaded
me to recognize in Espeletiinae only the genera
Carramboa and Espeletia. The genus Tamananthus
is here placed in Espeletiinae, as suggested by
Badillo (1992). Examination of a fragment of the
type of Tamananthus (through the kindness of
Dr. V.M. Badillo) allowed me to conclude that the
disc florets are functionally staminate and not
perfect, as stated in the original description (see
Badillo 1992). Functionally staminate disc florets
are a characteristic of nearly all Espeletiinae. The
genus differs from all other arboreal or shrubby
espeletias in having leaves without invaginated
bases. For this reason, I have opted to recognize
Tamananthus as distinctive from Espeletia.
Delimitation of monophyletic groups at the
subtribal level in Millerieae is still tentative, and
the classification provided here is based solely
on results from molecular studies of chloroplast
DNA. Similar assessments based on the nuclear
ribosomal ITS region (Rauscher 2002; Panero and
Plovanich-Jones, unpubl. data) are sometimes
incongruent with plastid phylogenies, and are
indicative of the possibility of past hybridization
or lineage sorting in the evolution of the group.
Key to the Subtribes
1. Leaves alternate, with expanded leaf bases or more
commonly forming a cupule (invaginated) around
stem; ray florets fertile
3. Espeletiinae (p. 482)
– Leaves opposite, if alternate, then leaves not forming
a cupule nor leaf bases expanded around stems and
ray florets neuter 2
2. Anther filaments pubescent 5. Guardiolinae (p. 486)
20. Guardiola
– Anther filaments without trichomes
3
3. Second series of phyllaries fused to developing cypselae forming a variously spined, verrucose or awned
conceptacle
7. Melampodiinae (p. 487)
– Phyllaries never fused to cypselae, rarely tightly
wrapped around cypselae forming a perigynium-like
structure
4
4. Ray floret cypselae tightly wrapped or completely encased in a perigynium-like structure
5
– Ray floret cypselae never enclosed nor tightly wrapped
by innermost phyllaries
6
5. Disc florets functionally staminate; ray (tubular floret)
cypselae enclosed in a perigynium-like structure
1. Desmanthodiinae (p. 479)
– Disc florets fertile; ray cypselae tightly wrapped by
flaps or hyaline wings of innermost phyllaries
6. Jaegeriinae (p. 487)
479
6. Capitula with dimorphic phyllaries normally in 2 series, the outer series broad, herbaceous, either reflexed
or erect, 2–6, inner series cucullate, scarious, phyllaries of both series normally with conspicuous, glandular stipitate trichomes, rarely covered with long
sericeous tapered trichomes; cypselae glabrous and
epappose
8. Milleriinae (p. 488)
– Capitula with subequal or gradate phyllaries, never
dimorphic, the outermost never broad and loosely
arranged around the capitulum, nor the inner ones
cucullate, phyllaries without large stipitate glandular trichomes, if present, the glandular tip smaller
than the trichome base; cypselae glabrous to densely
pubescent, with or without a pappus
7
7. Pappus various but mostly of persistent capillary or
plumose bristles of equivalent length, if of small scales,
then cypselae quadrate in cross-section and capitula
with 4, broadly deltate, valvate phyllaries, or disc and
ray corollas white, or ray corollas shallowly bilabiate
with 2, very small lobes opposite trilobed limb
2. Dyscritothamninae (p. 480)
– Pappus various but mostly caducous, if persistent, then
either of capillary bristles or scales, if of bristles, these
of equivalent size and arising from a shallowly cyathiform cypselae neck and plants with large corymbiform
cymes and broadly ovate leaves or the ray cypselae
lacking a pappus, if of scales, then ray cypselae usually enclosed by phyllary/paleae structure consisting
of the innermost phyllary subtending the ray floret
and 2–3 adjacent paleae
4. Galinsoginae (p. 483)
XXVII.1. Subtribe Desmanthodiinae Benth.
(1873).
Perennial herbs, shrubs or treelets. Leaves opposite,
sometimes perfoliate, blades ovate to lanceolate,
triplinerved or pentanerved. Capitula disciform or
discoid, in glomerules, glomerules normally with
three smaller, capitulum-like glomerules each, each
capitulum-like glomerule with 1–3 capitula in various stages of development, each glomerule subtended by green bracts rimmed with white or pink,
arranged in terminal, congested, corymbiform or
paniculiform cymes. Involucres cylindric to campanulate. Receptacles shallowly to strongly convex, epaleate? Tubular (‘ray’) florets normally 1
per glomerule, throat very short and reduced to
a ring protruding through perigynium, corollas
white. Disc florets functionally staminate, corollas
white; anthers hyaline, appendages short, round;
styles of disc florets penicillate, undivided, those
of the tubular florets large, arms lanceolate. Cypselae fusiform to oval, black, glabrous, enclosed in
perigynium. Pappus absent.
Only one genus:
480
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
shallowly bilobed, corollas mostly yellow. Disc florets bisexual, outer in discoid capitula sometimes
zygomorphic with three abaxial lobes enlarged,
corollas yellow, pink, green or white; anther appendages lanceolate to ovate, sometimes broadly
deltate, with or without glandular trichomes; style
arms with broad, parallel stigmatic surfaces, rarely
stigmatic surfaces fused, appendages short to
prominent, subulate to cylindric, papillose. Cypselae obconic to obpyramidal, sometimes quadrate,
sparsely to densely pubescent, sometimes with
multiple ridges, black to silvery-sericeous. Pappus
of multiple barbellate or plumose bristles, rarely of
scales with or without plumose projections, rarely
absent, in some species ray cypselae with smaller
pappus or epappose. x = 9, 10, 11, 20.
Approximately 43 species.
Key to the Genera
Fig. 97. Compositae-Millerieae. Desmanthodium fruticosum. A Flowering branch. B Cluster of flowering capitula.
C Flowering capitulum. D Phyllary enclosing fertile floret,
and subtending bract. E Immature phyllary and projecting
style. F Staminate floret. G Anthers. H Cypsela. (Reproduced
from Flora Novo-Galiciana, vol. 12, with permission of the
University of Michigan Herbarium; see McVaugh 1984)
1361. Desmanthodium Benth.
Fig. 97
Desmanthodium Benth., Hooker’s Icon. Pl. 12: 14, pl. 1116
(1872).
Characters of the subtribe. x = 17, 18. Eight species,
Mexico, Central America, northern South America.
1. Phyllaries dimorphic, outer series valvate, broadly
deltate, 4
1365. Tetragonotheca
– Phyllaries imbricate, subequal or gradate, never valvate, 5 to numerous
2
2. Ray and disc floret corollas white or creamy white
1363. Cymophora
– Ray and disc floret corollas of contrasting colours, if
ray corollas white, then disc corollas yellow, yellowgreen or pink
3
3. Wiry, multi-stem shrubs, mostly without leaves; capitula discoid, corollas yellow-orange
1362. Bebbia
– Annual or perennial herbs, if shrubs, these with a few
branches and with many leaves; capitula radiate or
discoid, if discoid, then plants with glaucous and linear
leaves
4
4. Leaves alternate
1364. Dyscritothamnus
– Leaves opposite
1366. Tridax
Genera of Dyscritothamninae
1362. Bebbia Greene
XXVII.2. Subtribe Dyscritothamninae
Panero (2005).
Bebbia Greene, Bull. Calif. Acad. Sci. 1: 179 (1885); Whalen,
Madroño 24: 112–123 (1977), rev.
Annual or perennial herbs, sometimes shrubs.
Leaves usually opposite, blades linear to ovate,
rarely deltate/hastate or semisucculent, mostly
triplinerved, sometimes with a single vein or
pinnately veined. Capitula terminal, solitary or in
open paniculiform or corymbiform cymes, discoid or radiate, sometimes disciform. Involucres
turbinate, campanulate to hemispheric, phyllaries
in 1–5 series, imbricate, rarely valvate, subequal
or gradate. Receptacles convex to conic, paleate.
Ray florets pistillate, rarely neuter, sometimes
Multi-stemmed shrubs forming dense, circular
mounds. Leaves opposite or alternate, shortly
petiolate, blades linear to sagittate, sometimes
hastate, with a single vein or triplinerved. Capitula in terminal, open paniculiform cymes or
sometimes broad, compact corymbiform cymes,
discoid. Involucres campanulate to hemispheric,
phyllaries in 2–3 series, subequal. Receptacles
shallowly convex, paleae shallowly trilobed.
Florets bisexual, corollas yellow-orange; anther
appendages ovate, with glandular trichomes;
Compositae
style arms long, subulate, with parallel stigmatic
surfaces, not confluent at the apices, appendages
short, tapered, papillose. Cypselae obconic, terete
to obscurely angled, sometimes with 4–5 shallow
angles, striate, brown, densely pubescent. Pappus
of several plumose bristles of equal length. Two
species, Mexico, south-western USA.
1363. Cymophora B.L. Rob.
Cymophora B.L. Rob., Proc. Amer. Acad. Arts 43: 39 (1907);
Turner & Powell, Madroño 24: 1–6 (1977), rev.; Keil et al.,
Madroño 34: 354–358 (1987), part. rev.
Annual herbs. Leaves opposite, petiolate, blades
broadly lanceolate to ovate, sometimes trullate,
rarely deltate/hastate, acute to obtuse, triplinerved.
Capitula in terminal, large, open paniculiform
cymes, discoid to disciform or radiate. Involucres cylindric to turbinate, phyllaries in 2–3
series, subequal. Receptacles convex. Ray florets
pistillate, corollas white or creamy white. Disc
florets bisexual, those on the periphery sometimes
zygomorphic with three longer abaxial lobes,
corollas white or creamy white; anther appendages
ovate, with or without glandular trichomes; style
arms with broad, parallel stigmatic surfaces, rarely
fused, appendages subulate to cylindric, papillose,
sometimes wanting. Cypselae compressed or obcompressed, sometimes broadly convex, obovate,
sparsely to densely pubescent, silvery to rusty
silvery. Pappus absent, of multiple plumose bristles
or sometimes of ciliate/fimbriate scales, cypselae
of zygomorphic florets normally with shorter
pappi or eppapose or absent (even if pappus
present in actinomorphic disc florets). x = 9. Four
to five species, Mexico, Venezuela.
1364. Dyscritothamnus B.L. Rob.
Dyscritothamnus B.L. Rob., Contr. Gray Herb. 65: 25, f. 1
(1922).
Shrubs. Leaves alternate, petiolate to subsessile,
blades filiform to lanceolate, semisucculent,
sometimes glaucous, pinnately veined. Capitula in
terminal, corymbiform cymes, discoid or radiate.
Involucres campanulate, phyllaries in 2–3 series,
subequal to gradate, herbaceous to membranaceous. Receptacles shallowly to strongly convex.
Ray florets neuter, corollas light yellow, apices
bilobed or trilobed. Disc florets bisexual, corollas
yellow; anther appendages ovate to lanceolate,
hyaline to light yellow; style arm apices long,
subulate, appendages minute, papillose. Disc
481
cypselae obconic, densely pubescent with long
shiny trichomes. Pappus of multiple barbellate to
plumose bristles. x = 11, 20. Two species, central
Mexico.
1365. Tetragonotheca L.
Tetragonotheca L., Sp. Pl. 2: 903 (1753); Turner & Dawson,
Sida 8: 296–303 (1980), rev.
Annual or perennial herbs. Leaves opposite,
petiolate or sessile, blades lanceolate to trullate,
margins entire to dentate, lobed. Capitula terminal,
solitary or in open corymbiform or paniculiform
cymes, radiate. Involucres obconic, hemispheric
after anthesis, phyllaries in two series, dimorphic,
outer series of 4 phyllaries, valvate, herbaceous,
enclosing the developing capitulum until anthesis, inner series smaller, scarious. Receptacles
conic. Ray florets pistillate, corollas yellow or
yellow-rust, apices shallowly to conspicuously
trilobed. Disc florets bisexual, corollas yellow;
anther appendages broadly ovate, with multiple
glandular trichomes extending below appendages
along connective; style arms with broad, parallel
stigmatic surfaces, appendages prominent, conic
with cylindric apices, papillose. Cypselae obconic
to obpyramidal, commonly quadrate in outline,
sometimes with multiple ridges or striae, sparsely
to moderately pubescent, black. Pappus of a crown
of minute squamellae, rarely wanting. x = 17. Four
species, southern USA, northern Mexico.
1366. Tridax L.
Fig. 98
Tridax L., Sp. Pl. 2: 900 (1753); Powell, Brittonia 17: 47–96
(1965), rev.
Ptilostephium Kunth (1829).
Mandonia Wedd. (1864).
Annual or perennial herbs, shrubs. Leaves opposite, blades linear to ovate, simple to deeply lobed,
1-veined or triplinerved. Capitula terminal, solitary or in open paniculiform cymes, discoid, disciform or radiate. Involucres campanulate to hemispheric, phyllaries in 2–5 series, subequal or gradate, herbaceous or somewhat scarious, sometimes
apical halves translucent and yellow, pink or purple. Receptacles convex to conic, paleae trilobed
with central lobe somewhat aristate. Ray florets
fertile, corollas 3-lobed, sometimes shallowly bilabiate, with two very minute adaxial lobes, white,
pink, pink-red, purple or yellow. Disc florets bisexual, corollas yellow, pink, green, pink-white, occasionally peripheral corollas in discoid capitula zy-
482
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Fig. 98. Compositae-Millerieae. Tridax mexicana. A Habit.
B Flowering capitulum. C Ray corolla. D Disc corolla. E Anthers. F Style arms. G Outer palea. H Inner palea. I Cypsela.
(Reproduced from Flora Novo-Galiciana, vol. 12, with permission of the University of Michigan Herbarium; see McVaugh 1984)
gomorphic with three expanded lobes; anther appendages ovate to lanceolate, with or without a few,
broadly cuneate glandular trichomes; style arms
with broad, parallel stigmatic surfaces, subulate
with tapered to cylindric appendages (acuminate).
Cypselae obconic, sparsely to densely pubescent,
sometimes glabrous, black to silvery. Pappus of
multiple plumose bristles, these sometimes only
as long as floral tube and appearing as squamellae,
rarely wanting. x = 9, 10. Thirty species, neotropical, one species a pantropical weed.
XXVII.3. Subtribe Espeletiinae Cuatrec.
(1976); Rauscher, Amer. J. Bot. 89:
1074–1084 (2002), phylog.
Acauli- and caulirosulate shrubs with marcescent
leaves, sometimes few-branched or trees with
most leaves concentrated on apices of branches,
a few species monocarpic, some producing
massive inflorescences of hundreds of capitula.
Leaves alternate, with very short internodes and
ampliated bases, sometimes forming a large
rosette, mostly with leaf bases wrapping around
stem forming a cupule, blades lanceolate to
fusiform, invariably densely lanate on abaxial
surface, pinnately veined. Capitula in terminal or
axillary, dichasial or monochasial, corymbiform
or paniculiform cymes, radiate or discoid. Involucres turbinate, campanulate to hemispheric;
phyllaries in 2 to numerous series, thickened or
herbaceous, glabrescent to densely lanate, flat
to cucullate, sometimes glandular, sometimes
with a few broad herbaceous phyllaries, these
fused at the base. Receptacles convex to conic,
paleate, sometimes with receptacular trichomes.
Ray florets pistillate, sometimes in 2–5 rows,
rarely bilabiate, corolla apices shallowly bilobed or
trilobed, sometimes obtuse, mostly with a dense
tuft of moniliform trichomes on tube. Disc florets
functionally staminate, rarely bisexual; anther
appendages ovate without glandular trichomes;
style arms of disc florets fused forming a thickened, penicillate style, those of the ray thickened
with broad, parallel, stigmatic surfaces, spreading.
Cypselae obcompressed to narrowly obpyramidal,
trigonous, slightly incurved, black, glabrous or
sparsely pubescent. Pappus absent, very rarely of
a few caducous setae or 3 awns. x = 19.
Approximately 90 species, Colombia, Ecuador
and Venezuela.
Key to the Genera
1. Perennial herbs or shrubs, sometimes trees, never
forming rosettes, if leaf bases forming a cupule around
stem (invaginated), then leaves very large (25–50 cm
long), shallowly bullate on adaxial surfaces, margins
undulate, ray corollas yellow, and capitula nodding
after anthesis
2
– Rosettes, if shrubs or trees and leaf bases wrapping
around stem and forming a cupule (invaginated), then
leaves without undulate margins, smooth on adaxial
surface and ray corollas either white or greenish white
or purplish-white, never yellow
1368. Espeletia
2. Perennial herbs or shrubs; leaves with ampliated bases,
leaves never forming a cupule around stem
1369. Tamananthus
– Trees, leaves with undulate margins forming a cupule
around stem
1376. Carramboa
Genera of Espeletiinae
1367. Carramboa Cuatrec.
Carramboa Cuatrec., Phytologia 35: 54 (1976).
Small trees with most leaves concentrated on apices
of the branches. Leaves alternate, ovate to pandurate, petiolate, very large, bright green, margins
undulate, adaxial surfaces shallowly bullate, peti-
Compositae
oles and abaxial surface covered in a dense lanate,
ferrugineous golden indumentum, leaf bases ampliated and forming a cupule around stem. Capitula
in axillary, compact, large corymbiform cymes, radiate, nodding after anthesis. Involucres campanulate to hemispheric, phyllaries in 2–5 series, slightly
gradate, thickened, cucullate, herbaceous to coriaceous, moderately pubsecent, sometimes densely
glandular. Receptacles convex to conic, sometimes
with trichomes between paleae. Ray florets pistillate, corollas bright yellow, shallowly bilobed or
trilobed with a dense tuft of tapered trichomes on
tube. Disc florets functionally staminate, corollas
yellow; anther appendages ovate. Cypselae obcompressed to trigonous, obcuneate, slightly incurved,
black, glabrous to very sparsely pubescent. Pappus
absent or very rarely with a few caducous setae.
Five species, Venezuela.
1368. Espeletia Mutis ex Humb. & Bonpl.
Espeletia Mutis ex Humb. & Bonpl., Pl. Aequinoct. 2: 10
(1808) [1809]; Smith & Koch, Brittonia 1: 471–530 (1935),
rev.
Libanothamnus Ernst (1870).
Coespeletia Cuatrec. (1976).
Espeletiopsis Cuatrec. (1976).
Ruilopezia Cuatrec. (1976).
Tamania Cuatrec. (1976).
Paramiflos Cuatrec. (1995).
Acauli- or caulirosulate shrubs with marcescent
leaves, sometimes tree-like with most leaves concentrated on apices of branches, a few species
monocarpic. Leaves alternate, with very short internodes and ampliated bases, sometimes forming
a large rosette, mostly with leaf bases wrapping
around stem forming a cupule, blades lanceolate
to fusiform, densely lanate on abaxial surfaces.
Capitula in terminal or axillary, dichasial or monochasial, corymbiform or paniculiform cymes, radiate or discoid. Involucres hemispheric, phyllaries
in 2 to numerous series, thickened or herbaceous,
sometimes with a few broad herbaceous phyllaries
fused at the base. Receptacles shallowly convex to
conic, sometimes with trichomes between paleae.
Ray florets pistillate, sometimes in 2 to 5 rows
or series, rarely bilabiate, corollas yellow, white,
greenish white, purplish, shallowly bilobed or
trilobed, sometimes obtuse, mostly with a dense
tuft of moniliform trichomes on tube. Disc florets
functionally staminate, rarely bisexual, corollas
yellow, white, green or purple; anther appendages
ovate. Cypselae obcompressed, trigonous, slightly
483
incurved, black, glabrous or sparsely pubescent.
Pappus absent, rarely of 3 awns. x = 19. Approximately 85 species, Colombia, Ecuador and
Venezuela.
1369. Tamananthus V.M. Badillo
Tamananthus V.M. Badillo, Ernstia 30: 25 (1985).
Perennial herbs or shrubs. Leaves alternate, sessile to shortly petiolate, bases expanded but not
forming a cupule around stem, blades lanceolate
to oval, sometimes narrowly obovate, with a single
main vein. Capitula in terminal, open paniculiform
cymes, radiate. Involucres turbinate, phyllaries in
3–5 series, herbaceous, densely pubescent. Receptacles convex, with trichomes between paleae. Ray
florets pistillate, corollas yellow, sometimes bilabiate, apices trilobed, tubes densely pubescent.
Disc florets functionally staminate or bisexual?,
corollas yellow, tubes sparsely pubescent or glabrous, lobes with glandular trichomes, nectaries
prominent and deeply lobed; anther appendages
ovate; style arms of disc florets short, apices obtuse, those of the ray florets narrowly deltate to
fusiform, with broad, parallel stigmatic surfaces.
Cypselae narrowly obpyramidal to obcompressed,
with a prominent carpopodium, black, essentially
glabrous. Pappus absent. One species, T. crinitus
V.M. Badillo, Venezuela.
XXVII.4. Subtribe Galinsoginae Benth.
(1873).
Annual or perennial herbs, shrubs. Leaves opposite, petiolate or sessile, rarely forming rosettes,
blades linear to ovate, rarely suborbicular, normally dentate, mostly triplinerved. Capitula
terminal, solitary or in open to congested corymbiform or paniculiform cymes, radiate or discoid,
rarely disciform. Involucres campanulate to
hemispheric, phyllaries in 2–5 series, subequal to
strongly gradate, herbaceous to membranaceous,
striate. Receptacles commonly convex or conic,
paleate, paleae oval to filiform, sometimes trilobed,
sometimes caducous. Ray florets pistillate, corollas
white, yellow or white suffused with pink, limbs
sometimes inconspicuous. Disc florets bisexual
or rarely functionally staminate, corollas yellow
or yellow-green, sometimes purplish; anther
appendages narrowly lanceolate to ovate, usually
without glandular trichomes; style arms with
broad parallel, rarely fused stigmatic surfaces,
appendages wanting or short, rarely cylindric,
484
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
papillose. Cypselae obconic to obpyramidal, subterete to shallowly 4- to 5-angled, black, glabrous
to moderately pubescent, sometimes cypselae
held in pocket formed by phyllary and the fused,
adjacent 2–3 paleae. Pappus absent or of a few
tapered or truncate squamellae or bristles, of
equal or different lengths, the bristles barbellate or
sometimes plumose, persistent or caducous.
Key to the Genera
1. Perennial herbs with scapose inflorescences and leaves
with very short internodes and forming a small rosette;
plants endemic to high elevation meadows in central
Mexico
1378. Selloa
– Annual or perennial herbs, sometimes shrubs,
capitulescences various, rarely scapose (Aphanactis),
if forming rosettes, then leaves either very densely
pubescent as to appear silvery or leaves very light
green (chartreuse) in colour, and plants from the
paramos and jalcas of the northern and central Andes
of South America
2
2. Pappus of disc floret cypselae of persistent capillary or
flattened bristles
3
– Pappus of disc floret cypselae various or absent, if of
bristles, these caducous
4
3. Ray floret cypselae without pappus
1371. Alloispermum
– Ray floret cypselae with pappus 1377. Schistocarpha
4. Ray florets tubular or with a very reduced limb
1373. Faxonia
– Ray florets when present not tubular
5
5. Pappus of several, caducous bristles of various lengths
6
– Pappus of several, persistent, truncate or tapered
scales or absent
7
6. Annual herbs
1370. Alepidocline
– Erect or scandent shrubs
1375. Oteiza
7. Ray cypselae enclosed in a phyllary/palea structure,
consisting of 2–3 paleae fused to adjacent phyllary
1374. Galinsoga
– Ray cypselae never enclosed by a phyllary/palea structure
8
8. Ray corollas, when present, yellow, if white, then central disc florets sterile, if discoid, then capitula on
pedicels which grow after anthesis 1372. Aphanactis
– Ray corollas white or white suffused with pink, central disc florets fertile, pedicels never growing after
anthesis
1376. Sabazia
erved. Capitula terminal, solitary or in open
paniculiform cymes, radiate. Involucres urceolate
to hemispheric, phyllaries gradate in 3–5 series,
herbaceous to scarious, striate, sometimes apices
purple-tinged. Receptacles conic, paleae filiform to
narrowly lanceolate, sometimes paleae caducous
or a few absent. Ray floret corollas inconspicuous
to showy, white to pink or light purple, apices
conspicuously, rarely shallowly trilobed. Disc
florets bisexual, corollas yellow or yellow-green;
anther appendages ovate, rarely with glandular
trichomes; style arms with parallel stigmatic
surfaces, apices deltate to acute, appendages short,
papillose. Cypselae obovate, sometimes shallowly asymmetrical and obcompressed, slightly
incurved, otherwise terete, black, glabrous, carpopodium lateral. Pappus absent or of a few
unequal caducous barbellate setae. x = 8. Five
species, Mexico, Central America, Venezuela.
1371. Alloispermum Willd.
Alloispermum Willd., Gesell. Naturf. Freunde Berlin
Mag. Neuesten Entdeck. Gesamten Naturk. 1: 139 (1807);
Robinson, Phytologia 38: 411–412 (1978), syn.
Genera of Galinsoginae
Perennial herbs or shrubs. Leaves subsessile to
petiolate, blades linear-lanceolate to ovate, triplinerved. Capitula in terminal, simple, open to
congested paniculiform or corymbiform cymes,
radiate, rarely discoid. Involucres narrowly campanulate to hemispheric, phyllaries in 3–4 series,
gradate, herbaceous to scarious. Receptacles flat to
convex, paleae oval to filiform. Ray floret corollas
white to creamy white, sometimes suffused with
purple on abaxial surfaces, apices shallowly to
moderately trilobed. Disc florets bisexual, corollas
yellow; anther appendages ovate; style arms with
broad, parallel stigmatic surfaces, appendages
wanting. Cypselae obpyramidal to obovoid, blackish to brownish purple, ray cypselae glabrous, disc
cypselae glabrous or pubescent. Pappus absent in
ray cypselae, disc cypselae with a pappus of several, very narrowly lanceolate, tapered, barbellate
scales, resembling bristles. x = 8. Approximately
15 species, Mexico, Central America and northern
Andes.
1370. Alepidocline S.F. Blake
1372. Aphanactis Wedd.
Alepidocline S.F. Blake, J. Wash. Acad. Sci. 24: 439 (1934);
Turner, Phytologia 69: 387–392 (1990), syn.
Cuchumatanea Seid. & Beaman (1966).
Aphanactis Wedd., Chlor. Andina 1: 142 (1856); Turner,
Bol. Soc. Argent. Bot. 19: 33–44 (1980), rev.; Robinson,
Brittonia 49: 71–78 (1997), tax.
Annual herbs. Leaves petiolate, blades lanceolate
to broadly ovate, obscurely to strongly triplin-
Annual or perennial herbs, variously persisting as
caespitose, matted, cushion-like shrubs, creeping
Compositae
among forbs or small rosettes. Leaves sessile,
shallowly perfoliate, blades oval to ovate or oblong,
glabrous to densely silvery-pilose, sometimes
forming rosettes. Capitula terminal, solitary or
sometimes a few in tightly clustered corymbiform
cymes, disciform or radiate. Involucres campanulate to hemispheric, phyllaries in 2–3 series,
subequal, herbaceous to scarious. Receptacles convex to shallowly conic, paleae setiform or linear,
sometimes caducous. Ray florets sometimes in
2–3 rows, corollas yellow to whitish-yellow, apices
shallowly to conspicuously trilobed, sometimes
limbs wanting and corolla appearing tubular. Disc
florets bisexual or sometimes functionally staminate, corollas yellow to yellow-green, glabrous
to moderately pubescent, sometimes glandular;
anther appendages ovate; style arms with narrow to broad parallel stigmatic surfaces, apices
acute to obtuse, essentially without appendages.
Cypselae obovoid, obcompressed to subterete,
sometimes shallowly 4–5-angled, black, essentially glabrous. Pappus absent. x = 8. Thirteen
species, Mexico, Central America and neotropical
Andes.
1373. Faxonia Brandegee
Faxonia Brandegee, Zoë 4: 403 (1894).
Annual herbs. Leaves petiolate, blades oval to
ovate, pinnately veined. Capitula in terminal, open
umbelliform cymes, radiate or disciform. Involucres campanulate, phyllaries in 2 series, subequal,
herbaceous. Receptacles convex, paleae linear or
trilobed. Ray floret limbs very reduced. Disc florets
bisexual, corollas yellow; anthers free to partially
free, appendages ovate. Cypselae subterete, slightly
incurved, black, glabrous. Pappus absent. One
species, F. pusilla Brandegee, Mexico.
1374. Galinsoga Ruiz & Pav.
Galinsoga Ruiz & Pav., Fl. Peruv. Prodr. 110, pl. 24 (1794);
Canne, Rhodora 79: 319–389 (1977), rev.; Canne, Madroño,
25: 81–93 (1978), generic limits; Canne-Hilliker, Taxon 41:
661–666 (1992), part. rev.
Stenocarpha S.F. Blake (1915).
Annual or perennial herbs, rarely shrubs. Leaves
petiolate or sessile, blades linear-lanceolate to
ovate, triplinerved. Capitula terminal, solitary
or in open simple to paniculiform cymes, radiate, rarely discoid. Involucres campanulate to
hemispheric, phyllaries in 2–4 series, subequal to
gradate, herbaceous to membranaceous or distally
485
scarious. Receptacles convex to conic, paleae oval
to filiform, sometimes weakly trilobed. Ray floret
corollas mostly white, sometimes pink or purple.
Disc florets bisexual, corollas yellow or greenish
yellow, sometimes purplish; anther appendages
ovate; style arms with broad, parallel stigmatic
surfaces, rarely with continuous stigmatic surfaces,
appendages wanting or tapered, cylindric, papillose. Cypselae obconic to shallowly obcompressed,
blackish, glabrous to moderately pubescent, rarely
ciliate, ray cypselae enclosed in structure formed
by phyllary and the fused, adjacent 2–3 paleae.
Pappus absent or of 8–20, cuneate to lanceolate,
sometimes aristate squamellae. x = 8. Fifteen
species, neotropical, 2 species adventive in the Old
World.
1375. Oteiza La Llave
Oteiza La Llave, Reg. Trimestre 1: 41 (1832).
Erect or straggly shrubs, sometimes vine-like.
Leaves petiolate or subsessile, blades narrowly to
broadly ovate, triplinerved. Capitula in terminal,
congested corymbiform cymes, radiate. Involucres campanulate to hemispheric, phyllaries in
3–5 series, strongly gradate, membranaceous to
scarious. Receptacles conic, paleae narrow. Ray
floret corollas white or whitish green. Disc florets
bisexual, corollas yellow; anther appendages ovate;
style arms with broad, parallel stigmatic surfaces,
apices acute to obtuse, appendages wanting, papillose. Cypselae obconic, black, glabrous. Pappus of
multiple, unequal caducous bristles. x = 17. Four
species, Mexico, Guatemala.
1376. Sabazia Cass.
Sabazia Cass., Dict. Sci. Nat. 46: 480 (1827); Longpre, Publ.
Mus. Michigan State Univ., Biol. Ser. 4: 287–383 (1970), rev.;
Urbatsch & Turner, Brittonia 27: 348–35 (1975), generic
limits; Turner, Wrightia 5: 302–305 (1976), generic limits.
Tricarpha Longpre (1970).
Freya V.M. Badillo (1985).
Perennial herbs, rarely weak shrubs. Leaves
petiolate or subsessile, blades lanceolate to ovate,
sometimes margins deeply dentate, triplinerved.
Capitula terminal, solitary or in open paniculiform
cymes, radiate. Involucres hemispheric, phyllaries
in 2–4 series, subequal to gradate, herbaceous
to membranaceous. Receptacles convex to conic,
paleae oblong to filiform, sometimes conspicuously trilobed. Ray floret corollas white or creamy
white, sometimes with abaxial surfaces suffused
486
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
with pink or purple-pink, shallowly to strongly
trilobed. Disc florets bisexual, corollas yellow;
anther appendages without glandular trichomes;
style arms with broad, parallel stigmatic surfaces,
sometimes touching each other, appendages
small or wanting, papillose. Cypselae obconic,
subterete, black, glabrous to sparsely pubescent.
Pappus absent or of multiple cuneate to spathulate,
sometimes filiform scales or barbellate bristles.
x = 4. Seventeen species, Mexico, Central America,
northern South America.
1377. Schistocarpha Less.
Schistocarpha Less., Linnaea 6: 409 (1831); Robinson,
Smithsonian Contr. Bot. 42: 1–20 (1979), rev.
Perennial herbs or subshrubs. Leaves petiolate,
petioles variously winged, blades ovate to broadly
ovate, triplinerved, margins sometimes deeply
serrate. Capitula in terminal, congested paniculiform cymes, radiate or disciform. Involucres
campanulate to hemispheric, phyllaries in 3–4
series, subequal to shallowly gradate, scarious.
Receptacles shallowly convex to conic, paleae
trilobed, not protruding beyond phyllaries. Ray
floret corollas white, rarely yellow, apices acute to
shallowly trilobed. Disc florets bisexual, corollas
white, yellow or yellow-green; anther appendages
narrowly lanceolate to ovate; style arms with
continuous stigmatic surfaces, apices broadly
acute to obtuse, appendages wanting. Cypselae
narrowly obpyramidal, terete, with a constriction
or narrowing below the neck, glabrous or sparsely
pubescent, black. Pappus of multiple, barbellate,
subequal bristles, more or less persistent, disposed in a radial pattern. x = 8. Sixteen species,
neotropical, most species in Mexico and Central
America.
1378. Selloa Kunth
Selloa Kunth in Humb., Bonpl. & Kunth, Nov. Gen. Sp. folio
ed. 4: 208 (1818), nom. cons.; Longpre, Publ. Mus. Michigan
State Univ., Biol. Ser. 4: 287–383 (1970), rev.; Robinson,
Brittonia 49: 71–78 (1997), tax.
Perennial herbs with thickened caudices and
roots. Leaves sessile or petiolate, internodes very
short, leaves forming a rosette, blades lanceolate
to ovate, sometimes suborbicular, bases sheathing,
apices acute to broadly acute to obtuse, glaucous on abaxial surface, triplinerved. Capitula
terminal, scapose, solitary or in simple cymes,
radiate. Involucres campanulate, phyllaries in 3
series, gradate, outermost herbaceous to membranaceous, broadly ovate, inner progressively
narrower, lanceolate and covered by exterior
phyllaries, scarious. Receptacles convex to conic,
paleae filiform. Ray floret corollas white suffused
with pink, apices trilobed. Disc florets bisexual,
corollas yellow; anther appendages broadly ovate
without glandular trichomes; style arms with
continuous stigmatic surfaces, apices acuminate,
appendages short, papillose. Cypselae obconic
to shallowly obpyramidal, subterete, sometimes
slightly obcompressed, brown to black, essentially
glabrous. Pappus of a few bristles, caducous. One
species, S. plantaginea Kunth, Mexico.
XXVII.5. Subtribe Guardiolinae H. Rob.
(1978).
Annual or perennial herbs, sometimes forming
a xylopode, shrubs, sometimes rupicolous. Leaves
opposite, blades narrowly lanceolate to deltate,
sometimes shallowly hastate to suborbicular,
dentate, acuminate to obtuse, abaxial surfaces
sometimes glaucous, sometimes semisucculent,
membranaceous. Capitula terminal, solitary on
long peduncles or in corymbiform cymes, radiate.
Involucres cylindric to campanulate, phyllaries
dimorphic in two series, outer 3 broad, navicular,
herbaceous, inner 2–3 scarious, smaller, similar
to paleae. Receptacles flat to convex, paleate. Ray
florets pistillate, corollas white, apices shallowly to
moderately bilobed or trilobed. Disc florets functionally staminate, corollas white, with a long tube
approximately 3–6 times as long as throat/lobe
length, throat campanulate, lobes approximately
twice as long as throat, spreading, corollas essentially glabrous; anther appendages ovate with
one large, basal glandular trichome; style arms of
disc florets long, very thin, subulate, those of the
ray florets shorter and broader. Cypselae obcompressed, oval to oblong, slightly obovate, brown
to black, rarely grey-black, essentially glabrous,
sometimes conspicuously striate, with a large,
truncate, flattened caruncle or elaiosome. Pappus
absent, rarely of several, somewhat lignified scales.
x = 12.
Only one genus:
1379. Guardiola Cerv. ex Hum. & Bonpl.
Fig. 99
Guardiola Cerv. ex Hum. & Bonpl., Pl. Aequinoct. 1: 143, t. 41
(1807); Robinson, Bull. Torrey Bot. Club 26: 232–235 (1899),
Compositae
487
ual, corollas yellow to greenish yellow, sometimes
marked with purple, sometimes tetramerous; anther appendages ovate with or without glandular
trichomes; style arms with parallel stigmatic surfaces, apices acute to obtuse, appendages wanting.
Cypselae clavate, obovoid, black, glabrous. Pappus
absent. x = 9.
Only one genus:
1380. Jaegeria Kunth
Jaegeria Kunth in Humb., Bonpl. & Kunth, Nov. Gen. Sp.
folio ed. 4: 218 (1818); Torres, Brittonia 20: 52–73 (1968),
rev.; Turner, Phytologia 55: 243–251 (1984), part. rev.
Characters of the subtribe. Nine species, neotropical, most species in Mexico.
XXVII.7. Subtribe Melampodiinae Less.
(1830).
Fig. 99. Compositae-Millerieae. Guardiola tulocarpus.
A Flowering branch. B Flowering capitulum. C Floret
with palea. D Style arms. E Anthers. F Cypsela. Guardiola
mexicana var. mexicana. G Cypsela. H Leaf. (Reproduced
from Flora Novo-Galiciana, vol. 12, with permission of the
University of Michigan Herbarium; see McVaugh 1984)
rev.; Van Faasen, Ph.D. Thesis, Michigan State University
(1971), rev.
Characters of the subtribe. Ten species, Mexico,
south-western USA.
XXVII.6. Subtribe Jaegeriinae Panero (2005).
Annual or perennial herbs, some aquatic, sometimes stems fistulose and rooting at nodes. Leaves
opposite, sessile or petiolate, blades lanceolate to
ovate, 3–5-nerved. Capitula terminal or axillary,
solitary or in paniculiform cymes, radiate. Involucres campanulate to hemispheric, phyllaries subequal, herbaceous, innermost trilobed with hyaline
wings and enfolding ray cypselae or ovary. Receptacles convex to shallowly conic, paleate. Ray
florets pistillate, fertile or sterile, corollas yellow
or white suffused with pink. Disc florets bisex-
Annual or perennial herbs, shrubs. Leaves opposite,
usually petiolate, entire to deeply pinnatifid or lacerate, blades linear to ovate, sometimes rhombic or
trullate, uninerved or triplinerved, sometimes pinnately veined. Capitula terminal or axillary, solitary
or in open paniculiform or corymbiform cymes,
radiate. Involucres campanulate to hemispheric,
phyllaries dimorphic in 2 series, the outer 2–5 free
to variously connate, inner fused to ray cypselae. Receptacles flat to convex, paleate, sometimes
paleae and disc florets shed as a unit. Ray florets
pistillate, corollas yellow, orange or white, rarely
greenish yellow. Disc florets functionally staminate,
corollas yellow to orange; anther appendages ovate,
without glandular trichomes; style arms of disc florets fused, the style penicillate, those of the ray
florets spreading. Cypselae enclosed in a conceptacle derived from the fused phyllary, conceptacle
obconic to trigonal, sometimes incurved, smooth,
verrucose or spinose, sometimes phyllary extended
as an aristate and coiled appendage, rarely as discor saucer-shaped collar. Pappus absent. x = 9, 10,
11, 12.
Key to the Genera
1. Paleae and disc florets caducous as a unit after anthesis;
ray corollas attached on abaxial side of the cypselae
apices
1383. Melampodium
– Paleae and disc florets not caducous as a unit after
anthesis, ray corollas not attached to the cypselae 2
2. Shrubs; cypselae with disc- or saucer-shaped collars,
these sometimes with lacerate or spiny projections;
endemic to the Galápagos Islands 1382. Lecocarpus
488
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
– Perennial herbs; cypselae without disc- or saucershaped collars; neotropical, one species in the Galápagos Islands
1381. Acanthospermum
Genera of Melampodiinae
1381. Acanthospermum Schrank
Acanthospermum Schrank, Pl. Rar. Hort. Monac. 53 (1820)
(“1819”); Blake, Contr. U.S. Natl Herb. 20: 383–392 (1921),
rev.
Annual herbs. Leaves petiolate to subsessile, blades
elliptic, trullate or ovate, entire to lobed, triplinerved. Capitula terminal or axillary, solitary. Involucres hemispheric, phyllaries in 2 series, outer
phyllaries 5–6, free to slightly connate at base, inner
fused to ray cypselae. Receptacles flat. Ray florets
pistillate, corollas yellow. Disc floret corollas yellow
to yellow-green; anther appendages ovate. Conceptacle trigonal and strongly echinate to spinose. Pappus absent. x = 10, 11. Five to six species, neotropical, adventive in other tropical regions of the world.
1382. Lecocarpus Decne
Lecocarpus Decne, Voy. Monde Venus Bot. (1846); Eliasson,
Svensk Bot. Tidskr. 65: 245–277 (1971), rev.; Adsersen, Bot.
Tidsskr. 75: 63–76 (1980), rev.
Shrubs. Leaves petiolate, blades ovate to oblong in
outline, deeply dissected to pectinate, semisucculent. Capitula terminal, solitary. Involucres hemispheric, outer phyllaries 4, herbaceous, connate
at base, inner fused to developing ray cypselae.
Receptacles flat to convex. Ray florets pistillate,
corollas yellow, apices trilobed. Disc floret corollas yellow; anther appendages lanceolate, without
glandular trichomes. Conceptacle shallowly trigonal, sometimes spinose, sometimes spines flattened, sometimes with a pappus-like structure resembling a bowl or saucer. Pappus absent. x = 11.
Three species, Galápagos Islands.
1383. Melampodium L.
Fig. 100
Melampodium L., Sp. Pl. 2: 921 (1753); Stuessy, Rhodora
74: 1–70, 161–219 (1972), rev.
Annual or perennial herbs some developing
a woody caudex, erect or prostrate. Leaves usually petiolate, blades linear to ovate-rhombic to
shallowly trullate, entire or lobed, uninerved or
triplinerved, sometimes pinnately veined. Capitula
terminal, solitary or in simple, paniculiform or
corymbiform cymes. Involucres campanulate to
Fig. 100. Compositae-Millerieae. Melampodium nutans. A
Habit. B Ray floret. C Disc floret. D Style apex. E Palea.
F Anthers. G Flowering capitulum. H Fruit, adaxial view.
Melampodium dicoelocarpum. I Fruit, lateral view. Melampodium sericeum. J Fruit, latero-adaxial view. K Flowering branchlet. Melampodium divaricatum. L Fruit, lateral
view. M Flowering capitulum. Melampodium perfoliatum.
N Fruit, lateral view. O Fruiting capitulum. (Reproduced
from Flora Novo-Galiciana, vol. 12, with permission of the
University of Michigan Herbarium; see McVaugh 1984)
hemispheric, phyllaries dimorphic, outer phyllaries 2–5, herbaceous, sometimes connate for
2/3 of their length, inner enclosing ray cypselae.
Receptacles convex to conic, paleae and disc florets
shed as a unit. Ray florets pistillate, corollas yellow,
orange or white, rarely greenish yellow. Disc floret
corollas yellow to orange; anther appendages
ovate. Conceptacle obconic, blackish to pale
brown, smooth or conspicuously tuberculate,
sometimes with phyllaries extending as a coiled or
stout appendage. Pappus absent. x = 9, 10, 11, 12.
Forty species, neotropical.
XXVII.8. Subtribe Milleriinae Benth. (1873).
Annual or perennial herbs, shrubs, sometimes
small trees. Leaves opposite, blades linear to
broadly ovate, sometimes pinnatifid, mostly
Compositae
triplinerved, sometimes pentanerved, rarely
pinnately veined. Capitula in terminal, open to
congested corymbiform or paniculifrom cymes,
rarely solitary, radiate, rarely discoid. Involucres
cylindric to hemispheric, phyllaries in 1–3 series,
outermost series normally with approximately
5 phyllaries, broad, erect, cucullate or reflexed,
sparsely to densely stipitate-glandular, inner erect,
normally smaller than outer, membranaceous
to scarious. Receptacles flat to conic, usually
paleate, paleae cucullate, variously pubescent
and glandular. Ray florets pistillate, sometimes
corollas tubular and the limb wanting, corollas
yellow, white or white suffused with purple, rarely
red or purple, apices conspicuously trilobed. Disc
florets bisexual, fertile or functionally staminate,
corollas yellow or creamy white, sometimes purple,
pentamerous, rarely tetramerous; endothecium of
mostly polar thickenings, rarely radial; style arms
of disc florets fused or spreading, with parallel
stigmatic surfaces, those of the ray spreading,
appendages wanting to well-developed, papillose.
Cypselae compressed or obcompressed, black,
glabrous. Pappus absent. x = 10, 12, 15, 16, 17.
Key to the Genera
1. Capitula with 1–3 ray florets
2
– Capitula with more than 3 ray or peripheral, tubular
florets
4
2. Phyllaries becoming leathery with age and wrapping
tightly around cypsela
1388. Milleria
– Phyllaries herbaceous or scarious, never becoming
leathery nor wrapping tightly around cypselae
3
3. Capitula with one ray floret
1392. Stachycephalum
– Capitula with more than 1 ray floret
1394. Unxia
4. Ray florets tubular; receptacles stipitate; phyllaries
white or white suffused with pink 1386. Ichthyothere
– Ray florets with a reduced to conspicuous limb, never
tubular; receptacles never stipitate; phyllaries never
white or pink
5
5. Disc corollas tetramerous, white
1387. Micractis
– Disc corollas pentamerous, white, purple or yellow 6
6. Disc florets bisexual
7
– Disc florets functionally staminate
10
7. Outermost phyllaries broad, cochleate or cucullate,
rarely lanceolate, erect and enclosing inner phyllaries
8
– Outermost phyllaries linear to spathulate, spreading or
reflexed, inner phyllaries not covered by outer phyllaries
9
8. Phyllaries and paleae with reddish striae; plants of
tropical Africa
1385. Guizotia
– Phyllaries and paleae with dull green or no striae;
plants of Mexico and Guatemala
1389. Rumfordia
9. Paleae 1.5 or 2 times as long as ovaries, not covered by
adjacent disc corollas; leaves variously lobed and/or
glaucous underneath
1384. Axiniphyllum
489
– Paleae as long as ovaries, covered by adjacent disc
corollas; leaves never lobed, nor with glaucous abaxial
surfaces
1390. Sigesbeckia
10. Outer series of phyllaries linear to spathulate, spreading or reflexed
1390. Sigesbeckia
– Outer series of phyllaries broadly ovate, sometimes
cucullate, erect, enclosing inner phyllaries
11
11. Cypselae obpyramidal, triangular in cross-section
1393. Trigonospermum
– Cypselae obcompressed, broadly biconvex, oval in
cross-section
12
12. Outer phyllaries with long, whitish, sericeous
trichomes, paleae linear, vestigial
1394. Unxia
– Outer phyllaries without long, whitish, serieceous trichomes, paleae oval to ovate
1391. Smallanthus
Genera of Milleriinae
1384. Axiniphyllum Benth.
Axiniphyllum Benth., Hooker’s Icon. Pl. 12: 16, t. 1118
(1872); Turner, Madroño 25: 46–52 (1978), rev.
Perennial herbs decumbent or erect, arising from
a thickened root or caudex. Leaves petiolate,
shallowly connate, petioles winged, blades mostly
deltate to sagittate in outline, either unlobed
or with three prominent lobes, abaxial surfaces
sometimes whitish and contrasting with bright
green adaxial surface. Capitula in terminal, open
paniculiform cymes, radiate or discoid. Involucres hemispheric, sometimes dimorphic with
herbaceous outermost phyllaries larger, otherwise
subequal to gradate in 2–3 series, herbaceous,
densely pubescent and glandular. Receptacles
convex. Ray floret corollas yellow (?). Disc florets
bisexual, corollas purple, yellow or white, throats
abruptly expanded above tube; anther appendages
ovate, without glandular trichomes; style arms
with broad, parallel stigmatic surfaces not confluent at apices, with small papillose appendages.
Cypselae obconic, shallowly to strongly quadrate
in cross-section, slightly asymmetrical, concave,
black, glabrous. Five species, Mexico.
1385. Guizotia Cass.
Guizotia Cass., Dict. Sci. Nat. 59: 237, 247, 248 (1829), nom.
cons.; Baagøe, Bot. Tidsskr. 69: 1–39 (1974), rev.
Annual or perennial herbs, shrubs, sometimes
repent. Leaves sometimes shallowly perfoliate,
petioles variously winged, sometimes with auriculate bases, blades linear to lanceolate, ovate or
deltate. Capitula terminal, solitary or in simple
cymes or more commonly open corymbiform
cymes, radiate. Involucres campanulate to hemi-
490
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
spheric, phyllaries dimorphic, the outer broad,
herbaceous, normally 5 or 6, the inner much
shorter, scarious to membranaceous. Receptacles
convex, paleae flat to cucullate, with reddish striae.
Ray floret corollas yellow, apices trilobed. Disc
florets bisexual, corollas yellow with reddish resin
canals; anther appendages ovate, normally with
one large basal glandular trichome; style arms
with parallel stigmatic surfaces, apices subulate,
appendages papillose. Cypselae obcompressed to
shallowly quadrate, black, glabrous. x = 15. Six
species, mostly eastern Africa, one species in tropical western Africa and the Indian subcontinent.
Guizotia abyssinica (L. f.) Cass. is commercially
grown for oil (niger seed) and introduced in
various parts of the world.
1386. Ichthyothere Mart.
Ichthyothere Mart., Repert. Pharm. 35: 195 (1830).
bisexual, innermost sometimes functionally staminate, corollas yellow or yellow-white, tetramerous; anther appendages ovate without glandular trichomes; style arms with parallel stigmatic
surfaces, apices deltate, appendages short, papillose. Cypselae compressed, obovate in outline,
asymmetrical, incurved, brown to black, glabrous.
Four species, eastern Africa, Madagascar.
1388. Milleria L.
Fig. 101
Milleria L., Sp. Pl. 2: 919 (1753).
Annual herbs. Leaves perfoliate or auriculate,
blades broadly lanceolate to suborbicular, sometimes trullate, triplinerved to pentanerved.
Capitula in terminal, open paniculiform cymes
with distinctive morphology, composed of several branching dichasia which end in multiple,
diverging cincinni subtended by a small bract,
radiate. Involucres campanulate, phyllaries in 2
Perennial herbs or shrubs, aromatic. Leaves petiolate or sessile, blades lanceolate to ovate, glabrous
or pubescent, membranaceous to semisucculent,
triplinerved. Capitula in terminal, commonly
congested simple, rarely open paniculiform
cymes, disciform. Involucres campanulate to
hemispheric, phyllaries in 1(–2) series, subcoriaceous, striate, white or pinkish white, wrapping
around cypselae. Receptacles convex to conic,
stipitate. Peripheral florets tubular, pistillate,
corollas white or yellow-white, bent or incurved.
Disc florets functionally staminate, corollas white
or white-yellow, nectaries well-developed; anther
appendages ovate to oval, cucullate with large
glandular trichomes, trichomes extending down
the connective; style arms of disc florets fused,
penicillate, those of the ray florets spreading.
Cypselae shallowly obcompressed, oblong, very
broadly convex, black, essentially glabrous. x = 16.
Twenty species, Panama, tropical South America.
1387. Micractis DC.
Micractis DC., Prodr. 5: 619 (1836); Schulz, Gleditschia 18:
211–218 (1990), rev.
Annual herbs. Leaves winged-petiolate or sessile,
sometimes shallowly auriculate, blades lanceolate
to ovate or deltate, triplinerved. Capitula in terminal, open paniculiform cymes, radiate. Involucres hemispheric, phyllaries in 2 series, herbaceous. Receptacles convex to shallowly conic. Ray
florets pistillate, corollas yellow or yellow-white,
mostly bilobed, sometimes trilobed. Disc florets
Fig. 101. Compositae-Millerieae. Milleria quinqueflora.
A Habit. B Cypsela. C Flowering capitulum. D Anthers.
E Disc corolla. F Undivided style of disc corolla. G Involucre
at fruiting stage. (Reproduced from Flora Novo-Galiciana,
vol. 12, with permission of the University of Michigan
Herbarium; see McVaugh 1984)
Compositae
series, outer 2, unequal, broad, flat to cupulate,
inner around 5, scarious. Receptacles flat. Ray
florets 1–2, pistillate, 3-lobed, corollas yellow. Disc
florets 4–6, functionally staminate, corollas yellow;
anther appendages broadly ovate, deep purple or
black, with glandular trichomes; style arms of ray
florets with parallel stigmatic surfaces, styles of
disc florets undivided, penicillate, apices acute.
Ray cypsela obpyriform, black, glabrous, covered
tightly by the thickened phyllaries, resembling an
anthocarp. x = 15. Two species, Mexico, Central
America, northern South America, Peru.
1389. Rumfordia DC.
Rumfordia DC., Prodr. 5: 549 (1836); Sanders, Syst. Bot. 2:
302–316 (1977), rev.
Perennial herbs or mostly straggly or multistemmed shrubs. Leaf blades ovate to hastate,
sometimes broadly trullate, bases attenuate into
winged petiole, sometimes auriculate, triplinerved
or pectinate, margins serrate. Capitula in terminal,
open paniculiform cymes, radiate. Involucres
hemispheric, outer phyllaries normally 5, foliose
and variously pubescent, inner erect and much
smaller, coriaceous to membranaceous, densely
glandular-hairy. Receptacles convex. Ray florets
in 1 or 2 rows, pistillate, corollas yellow, apices
shallowly to conspicuously trilobed. Disc florets
bisexual, corollas yellow; anther appendages
lanceolate to ovate, glabrous or rarely with one
glandular or tapered trichome; style arms with
broad, parallel stigmatic surfaces not confluent at
apices, without appendages. Cypselae compressed,
rarely subterete, shallowly to strongly asymmetrical, sometimes with a prominent neck on abaxial
side, black or black-brown, glabrous. x = 12, 15.
Seven to eight species, Mexico, Guatemala.
1390. Sigesbeckia L.
Sigesbeckia L., Sp. Pl. 2: 900 (1753); McVaugh & Anderson,
Contr. Univ. Michigan Herb. 9: 485–493 (1972), rev.; Schulz,
Haussknechtia 4: 25–35 (1988), part. rev.; Schulz, Cat. Herb.
Lipsiensis Pl. Neotrop. 2: 52–65 (1989), part. rev.
Schkuhria Moench (1794), nom. rej., non Roth (1797),
nom. cons.
Annual or perennial herbs, sometimes forming
a woody caudex with thickened roots and/or with
fistulose stems. Leaves with winged petioles, sometimes perfoliate, blades ovate to trullate, rarely
broadly ovate, triplinerved. Capitula in terminal,
open paniculiform cymes, radiate. Involucres
491
turbinate to campanulate, hemispheric, phyllaries
dimorphic, outer lanceolate to shallowly oblanceolate, herbaceous, sometimes densely glandular,
patent or strongly reflexed, inner erect, cucullate.
Receptacles shallowly convex to conic. Ray florets
pistillate, corollas yellow, white or white suffused
with pink, apices trilobed. Disc florets bisexual,
rarely functionally staminate, corollas yellow or
creamy white; anther appendages ovate to lanceolate, without glandular trichomes, sometimes
with large druses; style arms with broad, parallel
stigmatic surfaces not confluent at apices, with
small, sometimes cylindric, papillose appendages.
Cypselae compressed, obpyramidal, asymmetrical,
shallowly quadrate, rarely triquetrous, oblong in
cross-section, black, glabrous. x = 10, 15. Eight
species, pantropical, most species in Mexico.
1391. Smallanthus Mack.
Smallanthus Mack., Man. S.E. Fl. 1406 (1933); Wells,
Brittonia 17: 144–159 (1965), rev.; Robinson, Phytologia 39:
47–53 (1978), generic limits.
Annual or perennial herbs, shrubs or small trees.
Leaves petiolate, petioles variously winged, blades
lanceolate to ovate, sometimes suborbicular or
broadly deltate, entire to strongly dentate, triplinerved. Capitula in terminal, corymbiform or open
paniculiform cymes, radiate, sometimes nodding
after anthesis. Involucres hemispheric, outer
phyllaries foliose, much longer than inner, sometimes reflexed, lanceolate or broadly ovate, inner
resembling the pales, sometimes membranaceous,
striate. Receptacles convex. Ray florets pistillate,
corollas yellow or creamy white, rarely purple,
apices shallowly trilobed. Disc florets functionally
staminate, corollas yellow, white or purplish,
glabrescent to moderately pubescent, sometimes
with glandular trichomes; anther appendages
broadly ovate to deltate, without glandular trichomes; style arms of ray florets with parallel stigmatic surfaces meeting at the apices, of disc florets
variously fused, densely papillose. Ray cypselae
shallowly compressed, obpyriform, isodiametric
to oblong in cross-section, black to dark brown,
glabrous. x = 16, 17. Twenty species, neotropical.
1392. Stachycephalum Sch. Bip. ex Benth.
Stachycephalum Sch. Bip. ex Benth., Hooker’s Icon. Pl.
t. 1102 (1872).
Perennial herbs or weak, straggly, short-lived
shrubs. Leaves winged-petiolate, blades lanceolate
492
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
to broadly ovate, triplinerved. Capitula in terminal, congested corymbiform or scorpioid cymes,
radiate, sometimes the female floret somewhat
separated from the male disc florets on a stipitate,
lateral receptacle (Stachycephalum argentinum);
ray floret subtended by cochleate phyllary, disc
florets subtended by one or two paleae/phyllaries
and collectively subtended by a bract. Involucres
cylindric, phyllaries in 1 series, 1 or 2, herbaceous.
Receptacles flat, sometimes laterally stipitate,
sometimes epaleate. Ray floret pistillate, corollas
white or yellow (?). Disc florets functionally staminate, corollas white, yellow (?); anther appendages
lanceolate to narrowly oval, without glands; style
arms of ray florets with parallel stigmatic surfaces,
these meeting at the apices, styles of disc florets,
undivided, penicillate, apices acute. Ray cypselae
obpyriform, black, glabrous, sometimes sterile
(embryo fails to develop). Three species, Ecuador,
Mexico, Argentina.
1393. Trigonospermum Less.
Trigonospermum Less., Syn. Gen. Comp. 214 (1832); McVaugh & Laskowski, Univ. Michigan Herb. 9: 495–506 (1972),
rev.; Schulz, Cat. Herb. Lipsiensis Pl. Neotrop. 1: 52–59
(1988), syn.
Annual or perennial herbs. Leaves petiolate, blades
ovate to broadly ovate, triplinerved. Capitula in terminal, open or congested paniculiform or corymbiform cymes, radiate. Involucres campanulate to
hemispheric, phyllaries in 2 series, herbaceous. Receptacles convex. Ray florets pistillate, corollas yellow, apices deeply trilobed. Disc florets functionally
staminate, corollas yellow, sometimes tetramerous;
anther appendages ovate to round, without glandular trichomes; style arms of ray florets with parallel stigmatic surfaces meeting at the apices, styles
of disc florets undivided, apices acute. Cypselae
broadly obpyramidal, triquetrous, black, glabrous.
x = 15. Four species, Mexico, Central America.
1394. Unxia L. f.
Unxia L. f., Suppl. Pl. 56, 368 (1781) [1782]; Stuessy, Brittonia 21: 314–321 (1969), rev.; Stuessy, Rhodora 73: 291–292
(1971), tax.
Annual or perennial herbs, shrubs. Leaves opposite,
blades elliptic to lanceolate, triplinerved, entire
to shallowly serrate. Capitula tightly clustered in
terminal, simple cymes, radiate. Involucres hemispheric, phyllaries in 2–3 series, gradate, outermost
2–3 broadly ovate, herbaceous, strongly sericeous,
inner scarious, striate. Receptacles convex. Ray florets pistillate, corollas yellow. Disc florets functionally staminate, corollas yellow-orange; anther
appendages ovate, without glandular trichomes;
style arms of disc florets fused, penicillate, those
of the ray florets small, spreading. Cypselae obcompressed, ovoid, black, glabrous. x = 16. Three
species, Panama, northern South America.
XXVIII. Tribe Madieae Jepson (1901).
B.G. Baldwin and J.L. Panero
Annual or perennial herbs, subshrubs, shrubs,
trees or lianas. Leaves alternate, opposite or
whorled, usually sessile, blades narrowly linear to
ovate, oblanceolate to spathulate or elliptic, rarely
deltate, entire or pinnately toothed to divided,
sometimes glandular. Capitula terminal, usually
corymbose, paniculate, racemose or solitary,
sometimes glomerulate, ± umbellate or spicate,
radiate, discoid or (rarely) disciform. Involucres
cylindric to globose, phyllaries in 1–2 (rarely
3–4+) series, subequal or rarely gradate, flat to
conduplicate (in Madiinae, each ± enveloping
a ray ovary), herbaceous, distinct or connate,
often sessile- or stipitate-glandular. Receptacles
flat to conical, glabrous or setulose to villous,
epaleate or wholly to partially paleate, the paleae
usually in 1 series between ray and disc florets or,
if capitula discoid, constituting a false involucre.
Ray florets pistillate, corollas usually yellow or
white, glabrous or pubescent. Disc florets bisexual
or functionally staminate, corollas actinomorphic,
rarely zygomorphic, (4–)5-merous, glabrous or
hairy, sometimes glandular; anthers opaque, yellow to brown or reddish to dark purple; style arms
each with two narrow to broad, parallel stigmatic
surfaces, apices truncate or obtuse to deltate or
subulate, rarely deltate-acuminate, appendages
usually much shorter than stigmatic region or
wanting. Cypselae fusiform or clavate to obovate
or obpyramidal, terete, weakly or strongly angled,
compressed, or obcompressed, ray and disc often
dimorphic, black or brown, often striate, rarely
glandular. Pappus usually of ovate to setiform,
fimbriate to plumose scales, sometimes unlike
and alternating, sometimes of ciliolate to plumose
bristles, rarely absent. x = 19.
The tribe comprises 36 genera and more
than 200 species, which occur mostly in the
conterminous western USA (especially California),
Compositae
south-western Canada and north-western Mexico
(Baja California). Geographic outliers include
three genera of Madiinae endemic to the Hawaiian
islands, some species of Arnica in northern or eastern North America or Eurasia, and three species of
Baeriinae and Madiinae that occur either disjunctly
or solely in temperate South America. Tinctures
from flowers or rhizomes of Arnica (especially A.
montana) have been used widely in traditional
cultures and folk or homeopathic medicine to
promote healing of injuries. Historically, Madia
sativa was cultivated widely for seed oil, and used
by indigenous Americans for food and medicine.
Tribe Madieae was proposed by Jepson (1901)
to apply to paleate, mostly dark-anthered composites with ray-ovary-clasping phyllaries known
commonly as tarweeds, a mostly Californian
group treated previously (and subsequently
by most workers) as a subtribe, Madiinae or
‘Madieae’ (Bentham 1873a), and usually placed
in tribe Heliantheae (e.g. Stuessy 1977; Robinson
1981), with other paleate, dark-anthered taxa.
Carlquist’s (1959) anatomical studies established
the modern limits of subtribe Madiinae. Molecular
phylogenetic studies affirmed Carlquist’s (1959)
circumscription of subtribe Madiinae, and led to
reinstatement of tribe Madieae (in an expanded
sense) for a mostly western North American
lineage including tarweeds and a subset of epaleate
taxa previously treated in Helenieae, Heliantheae
or Senecioneae (Baldwin and Wessa 2000a; Baldwin
et al. 2002; Panero and Funk 2002). Based on the
molecular findings, expression of paleae and dark
anthers in tribe Madieae is not strictly homologous
with expression of those traits in Heliantheae and
other closely related tribes. A well-supported
sister-group relationship between Madiinae and
Arnica (including Mallotopus and Whitneya), although not suggested prior to the cited molecular
studies, is consistent with the close morphological,
ecological, and chromosomal similarities between
Arnica and the tarweed genus Raillardella – the sister group to all other tarweeds (Baldwin 2003a); the
two genera were closely associated in classifications
of Senecioneae from the time of Bentham (1873a),
until Carlquist (1959) showed that Raillardella
belonged in Madiinae [Arnica was disassociated
from Senecioneae by Nordenstam (1977) and
Robinson (1981)]. Genera of tribe Madieae that are
closely related to Arnica (Arnicinae) and Madiinae
include Hulsea and Eatonella (Hulseinae) and
Venegasia (Venegasiinae), all of which share with
Arnica a base chromosome number of x = 19, and
493
also occur in montane or wet habitats somewhat
similar to sites occupied by Raillardella but unlike
habitats of most other members of Madiinae.
Based on the molecular studies cited above,
woolly sunflowers and relatives (e.g. Eriophyllum,
Lasthenia) constitute a clade corresponding to
subtribe Baeriinae, which parallels the tarweeds
as examples of extensive diversification in the
California Floristic Province within tribe Madieae.
Relationships among members of tribe Madieae
are subjects of continuing phylogenetic study.
Key to the Subtribes
1. Phyllaries each partially to wholly clasping a ray
ovary or cypsela; receptacles wholly or partially
paleate (paleae indistinguishable from phyllaries in
discoid taxa); if heads discoid, plants annuals or
rhizomatous perennials with pappus of plumose,
subulate or lanceolate scales or plants of woody
life-forms
4. Madiinae (p. 497)
– Phyllaries not clasping ray ovaries or cypselae; receptacles epaleate; if heads discoid, plants annuals or rhizomatous perennials with pappus of bristles (sometimes plumose) or non-plumose scales
2
2. Leaves opposite (at least proximally); pappus of persistent bristles or, if absent, disc florets functionally
staminate and ray corollas persistent
1. Arnicinae (p. 493)
– Leaves opposite or alternate, sometimes in basal
rosettes; pappus of awns, scales, or deciduous bristles,
or, if absent, disc florets bisexual and ray corollas
caducous
3
3. Subshrubs or shrubs; leaves distinctly petiolate, blades
entire or coarsely toothed; phyllaries 3–4+-seriate, the
outer 1(–2) series often reflexed; pappus absent
5. Venegasiinae (p. 507)
– Herbs or, less commonly, subshrubs or shrubs; leaves
usually sessile or winged-petiolate (distinctly petiolate in Constancea – with blades dissected), blades
entire, toothed, lobed, or dissected; phyllaries 1–3seriate, usually erect, sometimes the outermost series
reflexed (in Hulsea); pappus present or absent
4
4. Phyllaries 1(–2)-seriate; anther endothecium of
mostly quadrate cells with (1–)2–4 polar thickenings;
style branches flat; cypselae usually not compressed
nor ciliate
2. Baeriinae (p. 494)
– Phyllaries 2–3-seriate; anther endothecium of
fusiform cells with 1 polar thickening; style branches
cucullate to canaliculate; cypselae compressed,
densely ciliate along edges
3. Hulseinae (p. 497)
XXVIII.1. Subtribe Arnicinae B.G. Baldwin
(2002).
Perennial herbs, usually rhizomatous. Leaves all
or mostly opposite, blades lanceolate to ovate or
oblanceolate to obovate or orbiculate, entire or
shallowly lobed, glabrous or tomentose, some-
494
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
times glandular. Capitula terminal, solitary or
laxly corymbose, radiate or discoid. Involucres
narrowly obconic to campanulate or hemispheric,
phyllaries in (1–)2 series. Receptacles convex,
epaleate, tomentose. Ray floret corollas yellow
to yellowish-orange or rarely cream. Disc florets
bisexual or rarely functionally staminate, corollas
yellow to yellowish-orange, 5-lobed; anthers yellow
or rarely purple (not blackened), endothecium
of oblong cells with 1–2 polar thickenings; style
arm apices much shorter than the stigmatic
lines. Cypselae cylindrical or narrowly clavate
to obovoid or obpyramidal, weakly to strongly
angled, glabrous or hairy, the walls carbonized,
striate. Pappus usually present and persistent, of
white to tawny, barbellate to plumose bristles.
x = 19 (polyploidy/apomixis extensive).
Only one genus:
1395. Arnica L.
Fig. 102
Arnica L., Sp. Pl. 2: 884 (1753); Maguire, Brittonia 4: 386–510
(1943), rev.; Wolf & Denford, Rhodora 86: 239–309 (1984),
part. rev.; Downie & Denford, Rhodora 90: 245–275 (1988),
part. rev., Gruezo & Denford, Asia Life Sci. 3:89–212 (1994),
part. rev., Baldwin, Novon 9: 460–461 (1999), tax.
Whitneya A. Gray (1865).
Mallotopus Franch. & Sav. (1878).
Characters of the subtribe. Circa 30 species, northern hemisphere (mostly montane or boreal).
XXVIII.2. Subtribe Baeriinae Benth. & Hook.
f. (1873).
Subtribe Eriophyllinae Rydb. (1915), as ‘Eriophyllanae’
Annual herbs or shrubs, glabrous or moderately
to densely lanate. Leaves usually alternate, usually
sessile, blades linear to ovate, rarely succulent, entire or 1-, rarely 2-pinnate, faces sparsely to densely
hairy, rarely glabrous. Capitula laxly to densely
paniculate or corymbose or solitary, radiate, rarely
discoid or disciform. Involucres cylindric to hemispheric, phyllaries in 1(–2) series, herbaceous, free
or variously fused. Receptacles convex to conical,
rarely globose, epaleate, variously foveolate,
sometimes stipitate. Ray floret corollas usually
yellow, sometimes tubular and limbs very reduced,
apices entire or shallowly 2- to 3-lobed. Disc florets
bisexual, corollas golden yellow, rarely tetramerous
or zygomorphic, variously hairy with moniliform
hairs, sessile or stipitate glands normally present,
sometimes abundant; anthers opaque to yellow,
appendages lanceolate to ovate with or without
Fig. 102. Compositae-Madieae. A–C Arnica dealbata. A Top
view of a capitulum. B Side view of a capitulum. C Inflorescence. D–H A. mollis. D Top view of a capitulum. E Side
view of a capitulum. F Ray floret. G Disc floret. H Habit.
(From Baldwin 2003a; illustrations by Lesley Randall)
glands; style arm apices obtuse to deltate, rarely
subulate, appendages wanting or reduced to
a proliferation of papillae, rarely 0.5–1× length
of stigmatic surfaces (Syntrichopappus). Cypselae
obcuneate to obpyramidal, sometimes narrowly
obovate, linear to fusiform, variously compressed
or obcompressed, sometimes both conditions in
the same capitulum, rarely ray and disc cypselae
dimorphic, black, rarely brown, striate, glabrous
or sparsely to densely hairy, sometimes with
multiple, globose glands (Baeriopsis). Pappus
absent or more commonly of scales (bristles in
Syntrichopappus), sometimes with those at angles
of cypsela longer and resembling awns, persistent
or caducous. x = 19.
Key to the Genera
1. Leaves succulent; cypselae densely covered with globose glands; plants of Guadalupe Island, Mexico
1397. Baeriopsis
Compositae
– Leaves usually not succulent; cypselae without or with
relatively few and widely scattered glands; plants of
continental America or Californian Channel Islands 2
2. Shrubs; leaves petiolate (blades 1–2-pinnate, dissected); plants of San Clemente, Santa Barbara, and
Santa Catalina islands
1398. Constancea
– Annual or perennial herbs; leaves sessile to subpetiolate; plants mostly of continental America
3
3. Leaves opposite
1400. Lasthenia
– Leaves alternate, at least distally
4
4. Phyllaries connate in proximal 1/2 (leaves lobed on at
least part of plant); pappus absent or coroniform
1402. Pseudobahia
– Phyllaries free or, if connate, leaves entire; pappus of
bristles or scales or absent
5
5. Disc corolla lobes narrowly lanceolate, subulate; anther appendages narrowly lanceolate; style-branch appendages subulate, length 0.5–1× stigmatic surfaces;
pappus of bristles or absent 1403. Syntrichopappus
– Disc corolla lobes never narrowly lanceolate; anther
appendages never narrowly lanceolate; style-branch
appendages reduced to an obtuse to deltate proliferation of papillae, never subulate, length much less than
0.5× stigmatic surfaces; pappus of scales or absent 6
6. Anther appendages on an extended connective, half as
long as thecae
1306. Amblyopappus
– Anther appendages never on an extended connective,
0.2–0.5× as long as thecae
7
7. Phyllaries broadly ovate, cucullate; ray corollas ‘bilabiate’ (with small adaxial lobe); ray cypselae triquetrous,
obcompressed
1401. Monolopia
– Phyllaries oval to lanceolate; ray corollas not bilabiate;
ray cypselae never triquetrous and obcompressed
1399. Eriophyllum
Genera of Baeriinae
1396. Amblyopappus Hook. & Arn.
Amblyopappus Hook. & Arn., J. Bot. (Hooker) 3: 321 (1841).
Annual herbs. Leaves mostly alternate, blades
linear to ovate in outline, pinnatifid or the
distal entire. Capitula congested-corymbose
or congested-paniculate, radiate or disciform.
Involucres cylindric to hemispheric, phyllaries
broad, somewhat cucullate. Peripheral florets
pistillate, corollas yellow, essentially tubular,
limb reduced. Disc floret corollas yellow; anther
appendages on extended connectives usually 1/2
length of thecae, ovate, without glands; style arms
with narrow stigmatic surfaces, apices truncate to
obtuse, papillose. Cypselae obcuneate, subterete
to shallowly ridged, black, sparsely to moderately
hairy and with scattered, large glands. Pappus
of multiple, broadly spathulate to cuneate scales,
central and basal areas shallowly thickened, edges
papillose. x = 8. One species, A. pusillus Hook. &
Arn., western USA, western Mexico, Peru, Chile.
495
1397. Baeriopsis J.T. Howell
Baeriopsis J.T. Howell, Leafl. W. Bot. 3: 152–154 (1942).
Caespitose shrublets. Leaves alternate, sessile,
blades narrowly oblanceolate, succulent, entire,
glabrous. Capitula solitary, radiate. Involucres
hemispheric, phyllaries in 2 series, semi-succulent.
Receptacles conical to globose. Ray floret corollas
yellow, shallowly trilobed. Disc floret corollas
golden yellow, lobes densely hairy with biseriate
hairs; anther appendages ovate, with large glands;
style arm apices deltate, appendages acuminate,
papillose. Cypselae obpyramidal to shallowly
biconvex (ray cypselae triquetrous), black-brown,
sparsely hairy, with large glands. Pappus of multiple lanceolate to linear scales. x = 8. One species,
B. guadalupensis J.T. Howell, north-western
Mexico (Guadalupe Island, Baja California).
1398. Constancea B.G. Baldwin
Constancea B.G. Baldwin, Madroño 46: 159–160 (1999)
[2000].
Shrubs, stems densely pubescent, white. Leaves
alternate, petiolate, blades broadly ovate in outline,
1–2-pinnate, lobes linear to oblanceolate, densely
white-tomentose to (adaxially) glabrate. Capitula
congested-corymbose to congested-paniculate,
radiate. Involucres cylindric to campanulate,
phyllaries in 1–2 series, with a thickened central
area or midrib. Receptacles convex. Ray floret
corollas yellow. Disc floret corollas golden yellow;
anther appendages ovate, with 1–2 glands; style
arms with narrow stigmatic surfaces, apices obtuse
to shallowly deltate, papillose. Cypselae narrowly
obpyramidal to biconvex, oval to quadrate in
cross-section, black, glabrous. Pappus mostly of 2
awn-like scales at angles of cypselae with intervening squamellae, sometimes with 3 or 4 awn-like
scales. x = 19. One species, C. nevinii (A. Gray)
B.G. Baldwin, western USA (San Clemente, Santa
Barbara and Santa Catalina islands, California).
1399. Eriophyllum Lag.
Eriophyllum Lag., Gen. Sp. Pl. 28 (1816); Constance, Univ.
Calif. Publ. Bot. 18: 69–136 (1937), rev.; Johnson, Novon 1:
119–124 (1991), tax.
Antheropeas Rydb. (1915).
Annual or perennial herbs, shrubs. Leaves mostly
alternate, blades linear to ovate or trullate in
outline, entire or toothed to pinnatifid or deeply
lobed, lobes commonly linear to oblanceolate,
496
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
sparsely to densely lanate or glabrate. Capitula
laxly or densely paniculate or corymbose, radiate
or discoid. Involucres cylindric to hemispheric,
phyllaries in 1(–2) series (usually 1 per ray floret),
with a thickened central area or midrib, sometimes
densely lanate. Receptacles convex to conical.
Ray floret corollas yellow, rarely white, entire
or shallowly trilobed. Disc floret corollas lemon
or golden yellow; anther appendages ovate, with
multiple glands, rarely eglandular; style arm apices
obtuse, appendages reduced to a papillose rim.
Cypselae cylindric to narrowly obovoid, terete
or angled with 4 or 5 sides, sometimes biconvex,
black, glabrous or sparsely hairy. Pappus absent or
more commonly of several lacerate scales. x = 4,
5, 7, or 8 (16, 24, 32, polyploid; 15, dysploid).
Thirteen species, western USA.
1400. Lasthenia Cass.
Lasthenia Cass., Opusc. Phytol. 3: 88 (1834); Ornduff, Univ.
Calif. Publ. Bot. 40: 1–92 (1966), rev.; Chan et al., Intl J. Pl.
Sci. 162: 1347–1360 (2001), phylog.
Baeria Fisch. & C.A. Mey. (Jan.? 1836).
Crockeria Greene ex A. Gray (1884).
Annual or perennial herbs, sometimes semiaquatic. Leaves opposite, blades linear to lanceolate, entire or toothed to pinnatifid, sometimes
ciliate, glabrous to sparsely hairy. Capitula laxly
paniculate or solitary, radiate. Involucres cylindric
to hemispheric, phyllaries in 1(–2) series, free or
connate. Receptacles convex to conical. Ray floret
corollas yellow, yellow-white, sometimes greenish
yellow, entire or shallowly trilobed. Disc floret
corollas yellow, sometimes tetramerous; anther
appendages ovate, without glands; style arms
with narrow stigmatic surfaces, apices deltate,
appendages acuminate to cylindric, papillose.
Cypselae subterete to biconvex, linear to fusiform,
black, sparsely hairy, sometimes sparsely glandular. Pappus absent or of 2 fragile awns, or of
awns and scales. x = 8 (4–7, dysploid; 12, 16,
24, polyploid). Eighteen species, western USA,
south-western Canada, north-western Mexico
(Baja California), Chile.
uate to shallowly lobed or entire, sparsely to moderately lanate. Capitula laxly paniculate or solitary, radiate. Involucres hemispheric, phyllaries in
1 series, broadly ovate, free or connate, lanate,
usually with black hairs towards apices. Receptacles convex to conical. Ray floret corollas yellow, rarely cream, ‘bilabiate’ (with small adaxial
lobe), limbs sometimes inconspicuous (M. congdonii), trilobed. Disc floret corollas yellow; anther appendages ovate, usually with glands; style
arms with broad stigmatic surfaces, apices obtuse,
appendages mostly lacking or reduced to papillae. Ray cypselae obcompressed, obpyramidal, incurved, triquetrous, sometimes densely pubescent,
disc cypselae flattened-biconvex, edges densely ciliate, faces glabrous or sparsely pubescent, sometimes with a longitudinal line of trichomes coinciding with linear interstice between adjacent cypselae. Pappus absent or (in M. congdonii) of mostly 2
lacerate scales, with or without 2 fragile, diverging
awns. x = 10, 11, 12, or 13. Five species, western
USA (California).
1402. Pseudobahia (A. Gray) Rydb.
Pseudobahia (A. Gray) Rydb., N. Amer. Fl. 34: 83 (1915).
Annual herbs. Leaves alternate, blades linear
to ovate in outline, commonly pinnatifid, lobes
linear, moderately lanate. Capitula laxly paniculate
or solitary, radiate. Involucres campanulate to
hemispheric, phyllaries in 1 series, bases to
proximal half connate. Receptacles conical. Ray
floret corollas yellow, entire to shallowly bilobed
or trilobed, tubes densely pubescent distally.
Disc floret corollas yellow, lobes thickened, tubes
densely pubescent distally; anther appendages
ovate, with glands; style arm apices obtuse to
subtruncate, papillose, appendages wanting.
Cypselae obcompressed or obpyramidal, biconvex
to mostly triquetrous, sometimes with 4(–5)
ridges, narrowly triangular in cross-section, black,
sparsely pubescent. Pappus absent or a crown
of scales. x = 8 (3, 4, dysploid). Three species,
western USA (California).
1401. Monolopia DC.
1403. Syntrichopappus A. Gray
Monolopia DC., Prodr. 6: 74 (1837) [1838]; Crum, Madroño
5: 250–270 (1940), rev.; Baldwin, Novon 9: 460–461 (1999),
tax.
Lembertia Greene (1897).
Syntrichopappus A. Gray, Pac. Railr. Rep. 4, 5(4): 106 (1857).
Annual herbs. Leaves mostly alternate, blades oval
to lanceolate or oblanceolate, pinnately veined, sin-
Annual herbs. Leaves alternate to subopposite,
blades narrowly to broadly linear, entire or distally
trilobed, moderately to densely lanate. Capitula
laxly paniculate or corymbose, radiate. Involucres
cylindric to campanulate, phyllaries in 1 series,
Compositae
497
oval with a conspicuous midrib, margins scarious.
Receptacles convex, shallowly stipitate. Ray floret
corollas yellow, white or pink, shallowly bilobed or
trilobed. Disc floret corollas lemon or golden yellow, with broad vascular strands, sometimes lobes
ciliate; anther appendages narrowly lanceolate,
without glands; style arms with narrow stigmatic
surfaces, apices subulate to deltate, appendages
well developed, 0.5–1× as long as stigmatic
surfaces, densely papillose. Cypselae narrowly
clavate, 3–5-angled, sparsely to densely hairy.
Pappus absent or of multiple slender, flattened,
caducous, barbellate bristles. x = 6 or 7. Two
species, western USA.
campanulate, phyllaries in (1–)2 series, narrowly
lanceolate, densely lanate abaxially. Receptacles
convex. Ray floret corollas white-yellow or whitepink, shallowly trilobed. Disc floret corollas white
or yellow-white; anther appendages ovate, without
glands; style arms with narrow stigmatic surfaces,
canaliculate adaxially, apices somewhat expanded,
obtuse, appendages lacking. Cypselae narrow,
oblong to fusiform or linear, compressed or
obcompressed, biconvex, margins densely ciliate,
faces glabrous, black, shiny. Pappus of 2 plicate
squamellae, each with a protruding awn (at edges
of cypsela). One species, E. nivea A. Gray, western
USA.
XXVIII.3. Subtribe Hulseinae B.G. Baldwin
(2002).
1405. Hulsea Torr. & A. Gray
Biennial or perennial herbs, rarely annuals. Leaves
cauline and/or in basal rosettes, alternate, sessile or
winged-petiolate, entire or dentate or rarely lobed,
glandular and/or woolly. Capitula terminal, laxly
corymbose, racemose or paniculate or often solitary, radiate. Involucres usually hemispheric, phyllaries in 1–3 series, erect or the outermost series reflexed, narrow. Receptacles flat, epaleate, glabrous.
Ray floret corollas yellow, orange, red or purplish.
Disc florets bisexual, corollas yellow or orange,
5-lobed; style arms cucullate to canaliculate, appendages much shorter than the paired stigmatic
lines or wanting. Cypselae ± cylindrical to clavate,
± compressed, 2-edged, ± thick-margined, carbonized, edges densely ciliate, faces shiny-glabrous
or sparsely to densely hairy. Pappus persistent, of 1–
2 opposite pairs of free, acuminate to truncate, noncostate, ± uniformly thick, apically erose scales.
x = 19.
Key to the Genera
1. Phyllaries in (1–)2 series; ray corollas barely surpassing phyllaries; disc florets 4–12
1404. Eatonella
– Phyllaries in 2–3 series; ray corollas showy; disc florets
20 or more
1405. Hulsea
1404. Eatonella A. Gray
Eatonella A. Gray, Proc. Amer. Acad. Arts 19: 19 (1883).
Annual herbs, sometimes densely branched,
tufted. Leaves sessile or subpetiolate, blades
linear to spathulate, entire, densely lanate. Capitula congested-paniculate or solitary, peduncles
elongating after anthesis. Involucres cylindric to
Hulsea Torr. & A. Gray, Pac. Railr. Rep. 6: 77 (1858); Wilken,
Brittonia 27: 228–244 (1975), rev.
Annual, biennial or perennial herbs. Leaves sessile
or subpetiolate, blades broadly linear to lanceolate, rarely obovate, faces sometimes densely
white-lanate. Capitula laxly paniculate or solitary,
radiate. Involucres campanulate to hemispherical,
phyllaries in 2–3 series. Receptacles flat to slightly
convex. Ray floret corollas bright to golden
yellow, sometimes orange or rusty red, entire
to shallowly trilobed. Disc floret corollas yellow
or orange; anther appendages ovate, without
glands; style arms with broad stigmatic surfaces,
branches canaliculate to cucullate, apices obtuse
to round, papillose, appendages wanting. Cypselae
narrowly obpyramidal to biconvex, compressed,
sometimes quadrate, black, striate, moderately
to densely silvery- or rusty-hairy. Pappus mostly
of 4 scales, those at the angles of each cypsela
longer, resembling awns, sometimes 4 awns. Seven
species, western USA, north-western Mexico (Baja
California).
XXVIII.4. Subtribe Madiinae Benth. & Hook. f.
(1873) (as Madieae); Carlquist et al.
(eds) Tarweeds & silverswords: evolution of the Madiinae (Asteraceae),
Missouri Bot. Gard., St. Louis (2003),
rev.
Subtribe Raillardellinae Rydb., N. Amer. Fl. 34, 4: 318
(1927), as ‘Raillardellanae’
Annual or perennial herbs, lianas, rosette shrubs,
subshrubs, shrubs or trees. Leaves alternate, opposite or whorled, usually sessile or subpetiolate,
498
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
blades usually oblanceolate to spathulate or linear
to linear-elliptic, entire or serrulate to toothed,
sometimes pinnately lobed or pinnatifid, usually
± parallel-nerved, usually sparsely to densely
± hirsute and stipitate-glandular, sometimes
villous or sericeous or rarely glabrous. Capitula
usually corymbose, paniculate, racemose, spicate,
glomerulate or ± umbellate, sometimes solitary,
radiate or discoid. Involucres cylindric to globose,
phyllaries in 1(–2) series or 0 (‘involucre’ then
interpreted as constituting 1 series of receptacular
paleae), usually conduplicate (each partly or
wholly investing a subtended floret, apices often
attenuate), usually glandular. Receptacles flat or
convex to conic, paleate (paleae usually in 1 series
interior to phyllaries or, if capitula discoid, in 1
outer series and constituting an ‘involucre’, then
sometimes connate and persistent, sometimes 1
subtending each disc floret, then usually distinct
and often soon falling, rarely in 2–3+ series),
hispidulous or glabrous. Ray floret corollas usually
yellow or whitish, apices (2–)3-lobed. Disc florets
bisexual or functionally staminate, corollas usually
yellow, 5-lobed; anthers usually reddish to dark
purple, sometimes yellow to brownish, appendages
usually lanceolate-oblong to ovate-deltate, without
glands, endothecium of mostly quadrate to cylindrical cells, with mostly 1–2(–4) polar thickenings;
style arm apices usually lanceolate to subulate,
rarely deltate or deltate-acuminate, appendages
much shorter than stigmatic lines or wanting.
Cypselae often obpyramidal, clavate or fusiform,
compressed, obcompressed or terete, usually
ray and disc dimorphic, black or brown, often
striate or ribbed, glabrous or sparsely to densely
hairy. Pappus absent or of awns, bristles or scales
(sometimes in combination) in 1–2 series, the
elements scabrid to plumose. x = 17 or 18.
Key to the Genera
1. Ray cypselae obcompressed (each mostly or completely enveloped by lateral margins of a phyllary); if
rays 0, then plants pappose annuals
2
– Ray cypselae compressed, ± terete, or ± triangular in
cross-section (if ± obcompressed, then each ± half
enveloped by lateral margins of a phyllary); if rays 0,
then plants perennials or epappose annuals
6
2. Plants annuals (1–20 cm); disc florets 1(–2)
1418. Hemizonella
– Plants annual or perennial herbs (2–150 cm); disc florets 3–120+
3
3. Plants rhizomatous perennials; disc corollas white
1421. Holozonia
– Plants annuals; disc corollas yellow (sometimes reddish with age)
4
4. Pappus of 10, broad, apically obtuse scales
1406. Achyrachaena
– Pappus absent or of bristles or apically acute scales 5
5. Heads usually calyculate (involucre proper subtended
by 2–5 bractlets); ray florets 5; disc florets 6, functionally staminate; disc pappus 0
1424. Lagophylla
– Heads not calyculate; ray florets 0 or 3–27; disc florets
4–120+, bisexual; disc pappus of 2–32 bristles or scales,
rarely 0
1425. Layia
6. Annuals; styles of disc florets hairy proximal to minute
arms; [receptacles paleate throughout, ray corollas
white with abaxial purple lines, pappus of subulate,
fimbriate to plumose scales] 1410. Blepharipappus
– Annuals or perennials; styles of disc florets glabrous
proximal to arms
7
7. Perennial herbs, ± scapiform; disc pappus of subulate,
ciliate-plumose scales
1428. Raillardella
– Annuals, leafy-stemmed perennial herbs, perennial
rosette plants, or plants woody; disc pappus 0 or of
scales (seldom both subulate and ciliate-plumose in
herbaceous taxa)
8
8. Peduncular bracts sometimes each with an apical pitgland, tack-gland(s), or spine (or apiculus); heads
radiate; ray cypselae terete to subterete or slightly
obcompressed (cross-section nearly circular except
adaxial side ± flattened to slightly bulging, or ± 3angled with abaxial side usually ± broadly 2-faced,
angle between those faces usually 90+◦ and adaxial
side ± flattened to slightly bulging), each ± half enveloped by a phyllary; if distal leaves spine-tipped or
apiculate, then each ray cypsela ± enveloped by a phyllary and sometimes compressed
9
– Peduncular bracts each without an apical pit-gland,
tack-gland, or spine (or apiculus); heads radiate or discoid; ray cypselae usually compressed, rarely ± terete
(then cross-section usually ± 3-angled with abaxial
side relatively broad, ± rounded, adaxial side ± 2faced, angle between those faces 15–70◦ ), each usually
completely or mostly enveloped by a phyllary
15
9. Annuals; leaves filiform to linear, margins often
strongly revolute; peduncular bracts usually with
tack-glands; ray corolla lobes (lateral) often spreading
(lengths often 1/2–5/6 of total laminae)
10
– Annuals, perennial herbs, subshrubs, or shrubs; leaves
linear or broader, margins seldom strongly revolute;
peduncular bracts usually without tack-glands; ray
corolla lobes ± parallel (lengths usually 1/10–1/2 of
total laminae)
11
10. Ray cypselae beaked; tack-glands absent
1427. Osmadenia
– Ray cypselae not beaked; tack-glands present
1412. Calycadenia
11. Ray corollas usually white, sometimes yellow, laminae often with abaxial purple lines; ray cypselae not
beaked or each with an inconspicuous, straight beak
(beak lengths less than diameters)
12
– Ray corollas yellow, laminae without abaxial purple
lines; ray cypselae each with an adaxial, ascending
beak (beak lengths longer than diameters)
13
12. Receptacles with paleae restricted to bases of outermost disc florets; disc florets bisexual; cypselae hairy
1411. Blepharizonia
Compositae
– Receptacles with paleae throughout; disc florets functionally staminate; cypselae glabrous 1419. Hemizonia
13. Peduncular bracts apiculate or each with an apical
spine
1414. Centromadia
– Peduncular bracts not apiculate, without apical spines
14
14. Plants annuals; peduncular bracts each with an apical
pit-gland; receptacles paleate throughout
1420. Holocarpha
– Plants annuals or perennials; peduncular bracts
without pit-glands; receptacles with paleae usually
restricted to bases of outermost disc florets (if in 2–3+
series, then perennials)
1415. Deinandra
15. Plants woody or semi-woody, evergreen; Hawaiian islands
16
– Plants herbaceous or woody with seasonal foliage;
North American or South American
18
16. Heads radiate on at least part of plant; (thick-leaved)
rosette shrubs
1409. Argyroxiphium
– Heads discoid; lianas, (fibrous-leaved) rosette shrubs,
subshrubs, shrubs, trees
17
17. Leaves 1–4 per node, free or weakly united; lianas,
subshrubs, shrubs, trees
1416. Dubautia
– Leaves 7–15 per node, strongly connate at base; rosette
shrubs
1429. Wilkesia
18. Disc pappus absent
1426. Madia
– Disc pappus of 5–21 scales (scales sometimes subulate
to setiform, bristle-like)
19
19. Heads all discoid
1413. Carlquistia
– Heads radiate on at least part of plant
20
20. Plants shrubby; Mexico
1407. Adenothamnus
– Plants herbaceous; Californian (to north-western USA
and south-western Canada)
21
21. Plants perennials
22
– Plants annuals
23
22. Involucres campanulate to hemispheric; anthers dark
purple; ray cypselae beakless
1423. Kyhosia
– Involucres campanulate to ellipsoid or globose; anthers yellow to brownish; ray cypselae beaked
1408. Anisocarpus
23. Anthers yellow to brownish
1417. Harmonia
– Anthers ± dark purple
1422. Jensia
Genera of Madiinae
499
3–8, corollas yellow, turning reddish. Disc florets
4–35, bisexual, corollas yellow to reddish, abaxially pubescent; anthers ± dark purple, appendages
ovate; styles glabrous proximal to arms, arms free,
apices lanceolate-linear, hairy and papillose. Ray
cypselae black, obcompressed, clavate, 10-ribbed,
straight, glabrous or ± scabrous, not beaked. Disc
cypselae similar, brown to black, ± scabrous. Ray
pappus absent. Disc pappus of 10 white, oblong,
apically obtuse scales. x = 8. One species, A. mollis
Schauer, western USA, western Mexico (northern
Baja California).
1407. Adenothamnus D.D. Keck
Adenothamnus D.D. Keck, Madroño 3: 6 (1935).
Shrubs, aromatic. Leaves proximally opposite,
mostly alternate; blades linear, entire, ± evenly
stipitate-glandular. Capitula ± corymbose or
borne singly, radiate. Peduncular bract tips each
without a pit-gland, tack-gland or spine. Involucres campanulate, phyllaries 8–13 in 1 series,
linear-attenuate, each fully investing a ray ovary,
ciliolate, adaxially stipitate-glandular. Receptacles
flat or convex, setulose, paleae not persistent,
in 1 series between ray and disc florets, connate
1/2+ length. Ray florets 8–13, corollas yellow.
Disc florets 12–50+, bisexual, corollas yellow,
abaxially glabrous; anthers reddish to dark purple,
appendages oblong-lanceolate; styles glabrous
proximal to arms, arms free, apices lanceolate,
hairy and papillose. Ray cypselae black, compressed, clavate, ± arcuate, glabrous, not beaked.
Disc cypselae black, ± terete, clavate, ± straight,
faces sparsely hispidulous. Ray pappus 0. Disc
pappus of 15–20 setiform, ciliate-plumose scales.
x = 14. One species, A. validus (Brandegee) D.D.
Keck, western Mexico (northern Baja California).
1406. Achyrachaena Schauer
Achyrachaena Schauer, Del. Sem. Hort. Vratislav. 3 (1837).
Annuals. Leaves proximally opposite, distally alternate, blades linear, entire or toothed, hirsute
or villous and (distal leaves) sparsely glandularpubescent. Capitula laxly ± corymbose or borne
singly, radiate. Peduncular bract tips each without a pit-gland, tack-gland or spine. Involucres
± campanulate to cylindric or ellipsoid, phyllaries 3–8 in 1 series, each fully investing a ray ovary,
± lanceolate-linear, abaxially hirsute or villous and
sparsely glandular-pubescent or eglandular. Receptacles flat to convex, setulose, paleae not persistent,
in 1 series (between rays and disc), free. Ray florets
1408. Anisocarpus Nutt.
Anisocarpus Nutt., Trans. Amer. Philos. Soc. n.s. 7: 388
(1841); Baldwin, Novon 9: 462–471 (1999), tax.
Raillardiopsis Rydb. (1927).
Perennial herbs. Leaves proximally rosulate or opposite, distally alternate, blades oblong to linear,
lanceolate-linear or oblanceolate, entire or toothed,
hirsute to strigose or pubescent and (distal leaves)
stipitate-glandular. Capitula ± corymbose or racemose or borne singly, radiate or discoid. Peduncular bract tips each without a pit-gland, tack-gland
or spine. Involucres ± globose or broadly ellipsoid
to campanulate, phyllaries same number as ray flo-
500
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
rets, in 1 series, lanceolate to lanceolate-attenuate
or oblanceolate, each 1/2 or fully investing a ray
ovary, ciliolate, abaxially stipitate-glandular, sometimes also hirtellous. Receptacles flat to convex, glabrous or setulose, paleae not persistent, in 1 series
between ray and disc florets, usually connate. Ray
florets 0, 1–3 or 7–15, corollas yellow. Disc florets
5–30, bisexual or functionally staminate, corollas
yellow, tubes and throats often abaxially glabrous,
lobes sparsely hispidulous and minutely glandular
abaxially; anthers yellow, appendages ovate; styles
glabrous proximal to arms, arms free, apices subulate, densely hairy. Ray cypselae black or greyish,
compressed or ± obcompressed, clavate, ± arcuate,
glabrous or hairy, beaked. Disc cypselae (if present)
black or greyish, ± terete, clavate, ± straight, hairy.
Ray pappus 0 or coroniform. Disc pappus of 5–8 or
11–21 linear, lanceolate, quadrate or subulate, fimbrillate, erose or ciliate-plumose scales. x = 7. Two
species, western USA, western Canada (Vancouver
Island).
1409. Argyroxiphium DC.
Fig. 103
Argyroxiphium DC., Prodr. 5: 668 (1836); Carr, Allertonia
3: 1–123 (1985), rev.; Baldwin & Robichaux in Wagner &
Funk (eds) Hawaiian biogeography: evolution on a hot
spot archipelago: 259–287 (1995), phylog.
Rosette shrubs (monocarpic or polycarpic).
Leaves in rosettes, sessile, blades narrowly linear
to lanceolate-linear or narrowly elliptic, entire,
hispid, sericeous, silvery-floccose or glabrate. Capitula elongate-racemose or -paniculate, radiate.
Peduncular bract tips each without a pit-gland,
tack-gland or spine. Involucres campanulate to
broadly or depressed-hemispheric, phyllaries
same number as ray florets, in 1 series, linear to
lanceolate, oblanceolate or elliptic-oblong, each
partially or wholly investing a ray ovary, glandularhairy and sometimes floccose-lanate. Receptacles
convex to conic, glabrous to hirsutulous, paleae
not persistent, mostly in 1 series between ray
and disc florets, connate. Ray florets (0–)1–45;
corollas reddish, purplish, white or yellow. Disc
florets 30–600, bisexual, corollas reddish, purplish,
whitish, or yellow; anthers yellowish to brownish
or reddish to dark purple, appendages ovate;
styles glabrous proximal to arms, arms free, apices
broadly deltate-acuminate to rounded-truncate,
hairy and papillose. Cypselae ± terete or angular
(usually ribbed), narrowly clavate, arcuate or
(some disc cypselae) straight, glabrous, not
beaked. Ray pappus coroniform. Disc pappus of
Fig. 103.
Compositae-Madieae. Argyroxiphium sandwicense subsp. macrocephalum. A Habit. B Capitulum.
C Phyllary, ray floret, palea, and disc floret (right to
left). D Ray floret. E Disc floret with pappus removed.
F Disc cypsela with pappus intact. (From Baldwin 2003b;
illustrations by Lesley Randall)
2–10, lanceolate-linear to oblong, fimbriate scales
(most or all abaxial on each cypsela). x = 14. Five
species, Hawaiian islands.
1410. Blepharipappus Hook.
Blepharipappus Hook., Fl. Boreal.-Am. 1: 316. 1833 (sero)
(‘1840’).
Annuals. Leaves proximally opposite, mostly alternate, blades narrowly spathulate to linear, entire, scabrous, hirsute, strigose, sericeous or villous (distal leaves usually also stipitate-glandular).
Capitula laxly corymbose or borne singly, radiate. Peduncular bract tips each without a pit-gland,
tack-gland or spine. Involucres turbinate to campanulate or hemispheric, phyllaries (2–)3–5(–8) in
1 series, ± lanceolate or oblanceolate, each ± 1/2
investing a ray ovary, abaxially ± hirsute and/or
stipitate-glandular. Receptacles convex, glabrous,
paleae not persistent, subtending all or most disc
florets. Ray florets (2–)3–5(–8); corollas whitish
Compositae
(purple-veined abaxially). Disc florets 6–25(–60+),
bisexual, corollas whitish, glabrous abaxially except
for hispidulous lobes; anthers ± dark purple, appendages hemi-orbiculate or conical; styles hairy
proximal to minute arms, arms free, apices truncate, papillose. Cypselae ± terete, ± obconic, ± villous, not beaked. Pappus 0 or of 12–18(–26) subulate, fimbriate to ciliate or plumose scales. x = 8.
One species, B. scaber Hook., western USA.
1411. Blepharizonia (A. Gray) Greene
Blepharizonia (A. Gray) Greene, Bull. Calif. Acad. Sci. 1:
279 (1885); Baldwin et al., Syst. Bot. 26: 184–194 (2001),
phylog., tax.
Hemizonia DC. subg. Blepharizonia A. Gray (1874).
Annuals, aromatic. Leaves proximally opposite
(forming winter–spring rosettes, often not persistent), mostly alternate; blades narrowly spathulate
to linear or the distal lanceolate, entire or serrate,
hirsute to strigose, sericeous, pilose or villous
(distal leaves also ± stipitate-glandular and/or
bearing tack-glands). Capitula ± corymbose,
spicate, paniculate or borne singly, radiate. Peduncular bract tips and margins usually bearing
tack-glands. Involucres campanulate or obconic to
± globose, phyllaries 5–13 in 1 series, lanceolate
or oblanceolate, each ± 1/2 investing a ray ovary,
abaxially canescent to hirsute or hispid to glabrate,
usually also bearing tack-glands. Receptacles flat,
hirtellous, paleae not persistent, in 1 series between
ray and disc florets, free. Ray florets 5–13, corollas
whitish (often red- to purple-veined abaxially),
lobes ± parallel. Disc florets 5–35(–60+), bisexual,
corollas whitish to purplish, glabrous abaxially
except for sparsely hispidulous lobes; anthers
± dark purple, appendages ovate to ovate-deltate;
styles glabrous proximal to arms, arms free, apices
subulate, densely hairy. Cypselae ± terete or
± obcompressed, clavate, 10-ribbed, ± sericeous,
not beaked. Ray pappus 0 or coroniform. Disc
pappus 0 or of 12–20+, reddish-brown, ciliate to
plumose, ± subulate scales. x = 14. Two species,
western USA (California).
1412. Calycadenia DC.
Calycadenia DC., Prodr. 5: 695 (1836); Carr, Amer. J. Bot.
64: 694–703 (1977), rev.; Baldwin & Markos, Mol. Phylog.
Evol. 10: 449–463 (1998), phylog.
Annuals, often aromatic. Leaves proximally in
rosettes or opposite, usually mostly alternate,
blades linear, usually entire, sometimes toothed
501
(revolute), the proximal often bristly-ciliate, often
scabrous. Capitula ± corymbose, glomerulate,
racemose, paniculate or spicate, or borne singly,
radiate. Peduncular bract tips each usually bearing
a tack-gland, lateral surfaces or margins sometimes with additional tack-glands. Involucres
cylindric, obconic, ovoid or campanulate, phyllaries 1–6 in 1 series, each c. 1/2 investing a ray ovary,
often with tack-glands and/or simple, sessile or
stipitate glands. Receptacles ± flat, setulose, paleae
sometimes persistent, in 1 series between ray
and disc florets, connate. Ray florets 1–6, corollas
white to reddish or yellow, sometimes bicoloured,
lobes often spreading. Disc florets 1–25, bisexual,
corollas white to reddish or yellow, lobes abaxially
hispidulous, minutely glandular or not; anthers
usually ± dark purple, appendages lanceolateovate to deltate; styles glabrous proximal to arms,
arms free, apices subulate, hairy. Ray cypselae
slightly obcompressed (± 3-angled, abaxial side
± 2-faced, angle between those faces 90+◦ , adaxial
side ± flat), rugose or smooth, glabrous or hairy,
not beaked. Disc cypselae ± terete or angular,
clavate, glabrous or hairy. Ray pappus 0. Disc
pappus of (0)6–13 subulate to lanceolate-aristate
(or shorter, wider and often alternating) scales.
x = 7 (4–6, dysploid). Ten species, western USA
(mostly California).
1413. Carlquistia B.G. Baldwin
Carlquistia B.G. Baldwin, Novon 9: 463 (1999); Baldwin &
Kyhos, Madroño 37: 43–54 (1990), rev.
Perennial herbs, rhizomatous, often matted.
Leaves proximally opposite, distally alternate,
blades lanceolate to linear, entire, hirsute to
villous and glandular-pubescent. Capitula laxly
corymbose or borne singly, discoid. Peduncular
bract tips each without a pit-gland, tack-gland or
spine. Involucres ± campanulate, phyllaries (modified paleae) (5–)7–16 in 1 series, lanceolate to
lanceolate-linear, abaxially hirsute and glandularpubescent. Receptacles flat, setulose, epaleate
(except for bracts constituting ‘involucre’). Florets
7–29, bisexual, corollas yellow, proximal throat
and lobes abaxially hirtellous; anthers yellow
to brownish, appendages broadly ovate-deltate;
styles glabrous proximal to arms, arms free,
apices subulate, densely hairy. Cypselae ± terete,
scabrellous, not beaked. Pappus of 9–17 white
to mauve or tawny, subulate, ± plumose scales
(flattened bristles). x = 8. One species, C. muirii
(A. Gray) B.G. Baldwin, western USA (California).
502
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
1414. Centromadia Greene
Centromadia Greene, Fl. Franciscana 424 (1897); Baldwin,
Novon 9: 462–471 (1999), tax.
Annual or perennial, rhizomatous herbs. Leaves
proximally in rosettes or opposite (not persistent),
mostly alternate; blades oblanceolate to linear
or lanceolate-linear, proximal 1–2-pinnatifid to
toothed, distal entire, apices of the distal usually spine-tipped, glabrous, scabrous-hirtellous,
± hirsute or villous, often also glandular. Capitula glomerulate, ± spicate-paniculate or ±
umbellate, radiate. Peduncular bracts each without
a pit-gland or tack-gland, usually spine-tipped,
sometimes apiculate. Involucres ± obconic or
urceolate, phyllaries 5–75+ (as many as ray florets),
lanceolate to lanceolate-attenuate or oblanceolate,
each usually 1/2 investing a ray ovary, abaxially
scabrous-hirtellous, hirsute, or villous and/or
glandular, apices often spine-tipped. Receptacles
flat to convex, setulose, paleae persistent, subtending all or most disc florets, free. Ray florets
5–75+; corollas yellow. Disc florets 6–200+, usually
functionally staminate, rarely bisexual; corollas
yellow, tubes/throats abaxially glabrous, lobes
often abaxially hispidulous; anthers ± reddish to
dark purple or yellow to brownish, appendages
ovate; styles glabrous proximal to arms, arms
free, apices ovate to subulate, densely hairy.
Ray cypselae ± compressed, abaxially gibbous,
glabrous, beaked or elevated adaxially. Ray pappus
0. Disc pappus 0 or of 3–12 linear, subulate or
oblanceolate scales. x = 13 (9–12, dysploid). Four
species, western USA, western Mexico (northern
Baja California).
1415. Deinandra Greene
Deinandra Greene, Fl. Franciscana 424 (1897); Baldwin,
Novon 9: 462–471 (1999), tax.
Annual and perennial herbs, subshrubs or shrubs,
often aromatic. Leaves proximally in rosettes or
opposite (usually not persistent), mostly alternate;
blades oblanceolate to linear or lanceolate-linear,
pinnatifid to toothed, serrate or entire, variously
hairy (often also sessile- or stipitate-glandular) or
glabrous with scabrous or hispid margins. Capitula
often corymbose, ± paniculate, sometimes racemose or glomerulate, radiate. Peduncular bract tips
each without a pit-gland, tack-gland or spine. Involucres ± obconic, campanulate, hemispheric or
urceolate, phyllaries 3–35 in 1 series, ± lanceolate to lanceolate-attenuate or oblanceolate, each
usually 1/2 investing a ray ovary, abaxially ± hirsute and sessile- or stipitate-glandular. Receptacles
flat to convex, glabrous or setulose, paleae deciduous, usually in 1 series (between rays and disc),
sometimes in 2–3+ series, connate or free. Ray florets 3–35, corollas deep or pale yellow. Disc florets 3–70, usually functionally staminate, seldom
bisexual, corollas yellow, tubes/throats abaxially
glabrous or minutely glandular-hairy, lobes sometimes abaxially hispidulous or glandular; anthers
usually reddish to dark purple or yellow to brownish, rarely maroon, appendages lanceolate-ovate to
deltate; styles glabrous proximal to arms, arms free,
apices subulate, densely hairy. Ray cypselae slightly
obcompressed (adaxially), clavate, abaxially gibbous, often ± arcuate, glabrous, ± beaked, beaks
offset adaxially, ascending. Ray pappus 0. Disc pappus usually of 1–15 lanceolate to subulate, entire
or ± fringed to erose scales, sometimes absent or
a crown of linear to fimbriate scales. x = 12 or 13
(9–11, dysploid). Twenty-one species, western USA
(mostly California), Mexico (Baja California).
1416. Dubautia Gaudich.
Dubautia Gaudich., Voy. Uranie 469 (1830); Carr, Allertonia
3: 1–123 (1985), rev., tax.; Carr, Pac. Sci. 53: 144–158
(1999), rev.; Baldwin & Robichaux in Wagner & Funk
(eds) Hawaiian biogeography: evolution on a hot spot
archipelago: 259–287 (1995), phylog.; Baldwin in Givnish &
Sytsma (eds) Molecular evolution and adaptive radiation:
103–128 (1997), phylog.
Railliardia Gaudich. (1830); Raillardia auct., sphalm.;
Railliarda DC., sphalm.
Subshrubs, shrubs, trees or lianas. Leaves alternate, opposite or whorled (3–4 per node), blades
linear to lanceolate-ovate, elliptic or obovate,
entire, serrulate or toothed, glabrous or variously hairy, sometimes also glandular-glutinous.
Capitula ± corymbose, racemose or paniculate
or borne singly, discoid. Peduncular bract tips
each without a pit-gland, tack-gland or spine.
Involucres cylindrical, campanulate, obconic,
ellipsoid or ovoid, phyllaries (modified paleae)
in 1–2 series, linear-oblong to obovate, abaxially
glabrous or puberulent to villous, also often
glandular. Receptacles flat to conic, hairy, paleae
deciduous, in 1–2 ‘involucral’ series or subtending
most or all florets, often connate. Florets 2–45,
corollas white or yellow, sometimes tinged or aging
purplish, abaxially glabrous or with scattered hairs
or proximally glandular; anthers yellowish to
brownish or dark purple, appendages lanceolate-
Compositae
oblong to ovate-oblong; styles glabrous proximal
to arms, arms free, apices deltate to subulate,
moderately hairy and papillose to densely hairy.
Cypselae ± terete, ± clavate, straight, glabrous
or hairy. Pappus of 8–28 lanceolate-ovate to
setiform, laciniate-fimbriate, ciliate or plumose
scales. x = 14 (13, dysploid). Twenty-four species,
Hawaiian islands.
1417. Harmonia B.G. Baldwin
Harmonia B.G. Baldwin, Novon 9: 463 (1999); Baldwin,
Madroño 48: 293–297 (2001), phylog., tax.
Annuals. Leaves proximally opposite, distally
alternate, sessile, blades linear, entire or toothed.
Capitula laxly ± umbellate to corymbose or
borne singly, radiate. Peduncular bract tips each
without a pit-gland, tack-gland or spine. Involucres obovoid to obconic, phyllaries 3–8 (same
number as rays) in 1 series, strongly conduplicate,
lanceolate to oblanceolate, each wholly investing
a ray ovary. Receptacles flat to convex, glabrous or
setulose, paleae not persistent, in 1 series (between
rays and disc), free or weakly connate. Ray florets
usually 3–8, corollas bright yellow (laminae flabelliform to obovate). Disc florets 7–30, bisexual or
functionally staminate (sometimes in same head),
corollas bright yellow, tubes/throats abaxially glabrous or proximally hirtellous-hispidulous, lobes
abaxially hispidulous; anthers yellow to brownish,
appendages ovate-dentate to hemi-orbiculate;
styles glabrous proximal to arms, arms free, apices
subulate, densely hairy. Ray cypselae black, terete
to ± compressed, weakly arcuate, gibbous or
not, beaked or beakless, glabrous. Disc cypselae
black, ± terete, ± clavate, glabrous or hairy.
Ray pappus 0 or of 3–12, lanceolate to subulate,
fimbrillate to plumose scales. Disc pappus of 7–11,
lanceolate-attenuate to subulate, linear, oblong or
quadrate, fimbriate, fimbrillate or plumose scales.
x = 9. Five species, western USA (California).
1418. Hemizonella (A. Gray) A. Gray
Hemizonella (A. Gray) A. Gray, Proc. Amer. Acad. Arts 9:
189 (1874); Baldwin, Novon 9: 462–471 (1999), tax.
Hemizonia DC. § [unranked] Hemizonella A. Gray, Proc.
Amer. Acad. Arts 6: 548 (1865).
Annuals. Leaves proximally opposite, distally
alternate (often clustered proximal to branches),
blades linear, entire or toothed, hirsute and (distal
leaves) glandular-puberulent. Capitula corymbose
or glomerulate or borne singly, radiate (peduncles
503
filiform). Peduncular bract tips each without a pitgland, tack-gland or spine. Involucres ± obovoid,
phyllaries 3–5 in 1 series, each mostly or wholly
investing a ray ovary, ± oblanceolate, abaxially
hirsute and glandular-hirtellous. Receptacles flat
to convex, glabrous or sparsely setulose, paleae
not persistent, in 1 series (between rays and disc),
connate. Ray florets 3–5; corollas pale yellow. Disc
florets 1–2, bisexual, corollas pale yellow, abaxially
pubescent; anthers yellow to brownish, appendages
hemi-orbiculate (reduced, not exceeding thecae);
styles glabrous proximal to arms, arms free,
apices subulate, densely hairy. Ray cypselae black,
obcompressed, arcuate, sparsely hispidulous or
glabrate, apices minutely beaked. Disc cypselae
black, ± terete, clavate, ± hispidulous, apices
minutely beaked. Pappus absent. x = 21–22. One
species, H. minima (A. Gray) A. Gray, western
USA, western Canada (British Columbia).
1419. Hemizonia DC.
Hemizonia DC., Prodr. 5: 692 (1836); Baldwin, Novon 9:
462–471 (1999), tax.
Annuals, often aromatic. Leaves proximally in
rosettes or opposite (persistent or not), mostly
alternate, blades narrowly elliptic to linear or
lanceolate-linear, serrulate or entire, variously
hairy, distal leaves also sometimes stipitateglandular as well. Capitula ± glomerulate,
paniculate, racemose or spicate or borne singly,
radiate. Peduncular bract tips each without a pitgland, tack-gland or spine. Involucres hemispheric
to ± urceolate or globose, phyllaries 5–14 in
1 series, linear to lanceolate or oblanceolate,
each usually 1/2 investing a ray ovary, abaxially
pubescent to hirsute or villous, and stipitateglandular. Receptacles flat to conic, glabrous,
paleae connate, forming cells around all or most
disc florets (± deliquescent). Ray florets 5–14,
corollas white or yellow, often purple-veined
abaxially. Disc florets 5–60+, functionally staminate, corollas white or yellow, abaxially glabrous;
anthers ± dark purple, appendages broadly ovate
to ovate-deltate; styles glabrous proximal to arms,
arms free or proximally united, apices lanceolatesubulate, densely hairy. Cypselae ± obcompressed
(adaxially), abaxially gibbous, glabrous, beak
absent or inconspicuous, straight. Pappus absent.
x = 14. One species, H. congesta DC., western USA
(California, southern Oregon).
504
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
1420. Holocarpha Greene
Holocarpha Greene, Fl. Franciscana, 426 (1897); Baldwin,
Ann. Missouri Bot. Gard. 93:64–93 (2006), evol., phylog.
Annuals, aromatic. Leaves proximally in rosettes or
opposite (usually not persistent), mostly alternate,
blades linear to oblanceolate, serrate to serrulate
or entire, hirsute to strigose, sericeous or villous
(distal leaves sometimes also stipitate-glandular
and/or resin-gland-dotted), apices of distal leaves
usually with a pit-gland. Capitula ± corymbose,
glomerulate, paniculate, racemose or spicate
or borne singly; radiate. Peduncular bract tips
each with a pit-gland. Involucres ± obconic or
campanulate to ± globose, phyllaries 3–16 in
1 series, elliptic, oblanceolate or obovate, each
usually 1/2 investing a ray ovary, abaxially with
pit-gland-tipped processes, otherwise glabrous or
puberulent to hispid, and/or sessile- or stipitateglandular. Receptacles flat to convex, glabrous,
paleae not persistent, subtending all or most disc
florets. Ray florets 3–16; corollas yellow, lobes ±
parallel. Disc florets 9–90, some bisexual, most
functionally staminate; corollas yellow, proximally
and distally glandular abaxially; anthers yellow to
brownish or pinkish to dark purple, appendages
ovate to ovate-deltate; styles glabrous proximal to
arms, arms free, apices subulate to aristate, densely
hairy. Ray cypselae ± obcompressed (adaxially),
abaxially gibbous, glabrous, beaks adaxial, ascending. Disc cypselae ± clavate, glabrous, not beaked.
Pappus absent. x = 4 or 6. Four species, western
USA (California).
1421. Holozonia Greene
Holozonia Greene, Bull. Torrey Bot. Club 9: 122 (1882).
Perennials, rhizomatous. Leaves proximally opposite (basally connate), distally alternate; blades
lanceolate to linear, entire, hirsute and (distal
leaves) glandular-hirtellous (glands cup-shaped).
Capitula laxly corymbose or borne singly, radiate.
Peduncular bract tips each without a pit-gland,
tack-gland or spine. Involucres ± obconic or
turbinate, phyllaries 4–10 in 1 series, each mostly
or wholly investing a ray ovary, ± lanceolate-linear,
abaxially hirsute, sometimes glandular-hirtellous
(glands cup-shaped). Receptacles flat to convex,
glabrous or setulose, paleae not persistent, in 1
series (between rays and disc), connate. Ray florets
4–10, corollas whitish (abaxially purplish-veined).
Disc florets 9–28, functionally staminate, corollas
white, tubes/throats abaxially glabrous or with
scattered hairs, lobes abaxially hispidulous; anthers ± dark purple, appendages ovate; styles
glabrous proximal to arms; arms free or proximally
united, apices subulate, hairy. Cypselae black,
obcompressed, ± clavate, glabrous, not beaked
(areolae broadly cupulate). Ray pappus 0 or
coroniform, 0.1–0.3 mm. Disc pappus 0 or of 1–5
caducous, subulate scales. x = 14. One species,
H. filipes (Hook. & Arn.) Greene, western USA
(California).
1422. Jensia B.G. Baldwin
Jensia B.G. Baldwin, Novon 9: 464 (1999).
Annuals. Leaves proximally opposite (often
crowded), distally alternate, blades spathulate
to linear, entire or toothed, hirsute to strigose,
distal leaves sometimes also stipitate-glandular.
Capitula ± umbellate, radiate. Peduncular bract
tips each without a pit-gland, tack-gland or spine.
Involucres ± obconic or urceolate to globose,
phyllaries 2–12 in 1 series, lanceolate to lanceolateattenuate, each usually wholly investing a ray
ovary, abaxially hirsute (hair tips ± uncinate).
Receptacles flat to convex, glabrous or setulose,
paleae not persistent, in 1 series (between rays and
disc), connate. Ray florets 2–12, corollas yellow,
sometimes purple-veined abaxially. Disc florets
1–65, functionally staminate, corollas yellow,
abaxially glabrous except for hispidulous lobes;
anthers ± dark purple, appendages ovate-deltate;
styles glabrous proximal to arms, arms free,
apices narrowly lanceolate-subulate, densely hairy.
Cypselae compressed, clavate, arcuate, glabrous,
beaked. Ray pappus a crown of scales, 0.1–1 mm.
Disc pappus of 5–7 (white or purple-tipped)
subulate, crisped, ciliolate scales, 2.5–3 mm. x = 8.
Two species, western USA (California).
1423. Kyhosia B.G. Baldwin
Kyhosia B.G. Baldwin, Novon 9: 465 (1999).
Perennials, rhizomatous. Leaves proximally in
rosettes or opposite, distally alternate, blades
lanceolate-linear to linear, entire, hirsute and
(distal leaves) glandular-hirtellous. Capitula laxly
± corymbose or borne singly, radiate. Peduncular
bract tips each without a pit-gland, tack-gland or
spine. Involucres ± campanulate to hemispheric,
phyllaries 8–12 in 1 series, each mostly or wholly investing a ray ovary, lanceolate to lanceolate-linear,
abaxially hirsute and glandular-hirtellous. Receptacles flat to convex, glabrous, paleae not persistent,
Compositae
in 1 series (between rays and disc), weakly connate
or free. Ray florets 8–12, corollas bright yellow.
Disc florets 28–65, bisexual, corollas bright yellow,
proximally pubescent, lobes abaxially hispidulous
and minutely stipitate-glandular; anthers ± dark
purple, appendages lanceolate-ovate to ovate;
styles glabrous proximal to arms; arms free, apices
subulate, densely hairy. Ray cypselae black, compressed, clavate, arcuate, glabrous or hispidulous,
beakless. Disc cypselae brown to black, ± terete,
straight or arcuate, hispidulous, otherwise similar
to rays. Ray pappus 0 or a crown. Disc pappus
of 5–10 stramineous to purplish, lanceolate to
subulate, ciliate to plumose scales. x = 6. One
species, K. bolanderi (A. Gray) B.G. Baldwin,
western USA (California, southern Oregon).
1424. Lagophylla Nutt.
Lagophylla Nutt., Trans. Amer. Philos. Soc. n.s. 7: 390
(1841); Thompson, A biosystematic study of Lagophylla
(Compositae: Heliantheae) and related taxa. Ph.D. Diss.,
Univ. Calif., Davis (1983), rev.
Annuals. Leaves proximally opposite (not persistent), mostly alternate, blades narrowly elliptic
to linear or (proximal) sometimes oblanceolate to spathulate, entire or (proximal) sometimes
toothed, hirsute to strigose, sericeous or villous (all
or the distal sometimes also stipitate-glandular).
Capitula ± paniculate to glomerulate, radiate.
Peduncular bract tips each without a pit-gland,
tack-gland or spine. Involucres ± hemispheric or
obovoid to obconic, phyllaries 5 in 1 series, linear
to oblanceolate, each wholly investing a ray ovary,
abaxially piloso-hirsute to hirtellous or scabrellous. Receptacles flat to convex, densely hirtellous,
paleae not persistent, in 1 series (between rays
and disc), free or proximally connate. Ray florets
5, corollas yellow (often nerved with red to purple
abaxially). Disc florets 6, functionally staminate,
corollas yellow, glabrous or hairs scattered; anthers
± dark purple, appendages ± hemi-orbiculate
to ovate-dentate; styles glabrous proximal to
arms, arms united, apices subulate, densely to
sparsely hairy. Cypselae obcompressed, glabrous,
not beaked. Pappus absent. x = 7. Four species,
western USA.
1425. Layia Hook. & Arn. ex DC.
Layia Hook. & Arn. ex DC., Prodr. 7(1): 294 (1838), nom.
cons.; Baldwin, Ann. Missouri Bot. Gard. 93:64–93 (2006),
evol., phylog.
505
Annuals. Leaves proximally in rosettes or opposite,
mostly alternate, blades ovate, lanceolate or
oblanceolate to linear, bipinnatifid to toothed or
entire, glabrous or hirsute to strigose (distal leaves
sometimes also stipitate-glandular). Capitula
± corymbose or borne singly, usually radiate,
rarely discoid. Peduncular bract tips each without
a pit-gland, tack-gland or spine. Involucres cylindrical to ± hemispheric or urceolate, phyllaries as
many as ray florets, in 1(–2) series, lanceolate to
lanceolate-attenuate or oblanceolate, each wholly
investing a ray ovary, abaxially hirsute to strigose
or scabrous, often also glandular. Receptacles flat
to convex, setulose, paleae not persistent, in 1
series (between rays and disc) or subtending ± all
disc florets, free. Ray florets 0 or 3–27, corollas
yellow, cream, white or bicoloured. Disc florets
5–120+, bisexual, corollas yellow, hirtellous or
hispidulous and/or minutely stipitate-glandular;
anthers ± dark purple or yellow to brownish, appendages narrowly triangular, lanceolate-oblong
or lanceolate-ovate to ovate; styles glabrous proximal to arms, arms free (lengths unequal), apices
subulate, hairy and papillose. Ray cypselae obcompressed, clavate, ± arcuate to falcate, glabrous to
sparsely hairy, not beaked. Disc cypselae ± clavate,
usually ± strigose to sericeous, sometimes glabrous. Ray pappus 0. Disc pappus of (0–)2–35,
ovate, elliptic, lanceolate-attenuate, subulate or
setiform, glabrous, scabrous or plumose scales
or bristles, often basally villous and/or adaxially
woolly. x = 7 or 8 (16, polyploid). Fourteen
species, western USA, western Mexico (Baja
California).
1426. Madia Molina
Madia Molina, Sag. Stor. Nat. Chili 136: 354 (1782);
Baldwin in Hind & Beentje (eds) Compositae: systematics.
Proceedings of the International Compositae Conference,
Kew, 1994, vol. 1: 377–391 (1996), phylog.; Baldwin, Novon
9: 462–471 (1999), tax.
Annuals, often aromatic. Leaves proximally in
rosettes or opposite (often crowded), distally
alternate, blades lanceolate or oblong-linear to
linear, usually entire, hirsute to strigose, often
also glandular-pubescent. Capitula corymbose,
paniculate, racemose or spicate, usually radiate.
Peduncular bract tips each without a pit-gland,
tack-gland or spine. Involucres ellipsoid to urceolate or ± globose, phyllaries as many as ray
florets, in 1 series, usually ± lanceolate-linear to
lanceolate-attenuate or oblanceolate, each mostly
506
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
or wholly investing a ray ovary, abaxially hispid or
hirsute to ± villous, and also glandular-pubescent.
Receptacles flat to convex, glabrous or setulose,
paleae persistent or falling readily, in 1 series
(between rays and disc), ± connate or free. Ray
florets (0)1–22, corollas usually yellowish. Disc
florets 1–80+, bisexual or functionally staminate,
corollas usually yellow, glabrous to usually hirtellous or hispidulous and/or stipitate-glandular;
anthers ± dark purple or yellow to brownish,
appendages oblong to orbiculate-apiculate; styles
glabrous proximal to arms, arms free, hairy
throughout, apices narrowly triangular, densely
hairy to sparsely papillose and hairy. Ray cypselae
compressed, ± triquetrous (abaxial side rounded,
adaxial side 2-faced) or rarely terete, clavate,
often arcuate, glabrous, beaked or beakless. Disc
cypselae similar, sometimes obovoid (often ±
straight, basal attachments central, apices not
beaked). Pappus absent. x = 8 (16, 24, polyploid;
14, dysploid). Ten species, (mostly western) USA,
Canada (one species in Argentina, Chile and
Hawaiian islands).
1427. Osmadenia Nutt.
Osmadenia Nutt., Trans. Amer. Philos. Soc. n.s. 7: 391
(1841); Carr, Amer. J. Bot. 64: 694–703 (1977), rev.; Baldwin
& Markos, Mol. Phylog. Evol. 10: 449–463 (1998), phylog.
Annuals, aromatic. Leaves proximally in rosettes
or opposite, mostly alternate, blades linear, entire
(revolute), scabrous and ± hirsute. Capitula laxly
paniculate or borne singly, radiate. Peduncular
bract tips each without a pit-gland, tack-gland or
spine. Involucres ovoid, phyllaries 3–5 in 1 series,
each c. 1/2 investing a ray ovary. Receptacles flat
or convex, glabrous, paleae not persistent, in 1
series between ray and disc florets, connate. Ray
florets 3–5, corollas white, fading reddish, laminae
sometimes with central red spot, lobes spreading,
lengths ± equal to total laminae. Disc florets 3–10,
bisexual, corollas white, fading reddish, lobes
sometimes with reddish blotch, tubes/throats glabrous, lobes hispidulous and minutely glandular;
anthers ± dark purple, appendages lanceolateoblong; styles glabrous proximal to arms, arms
free, apices subulate, densely hairy. Ray cypselae
slightly obcompressed (adaxially), ± triquetrous
(abaxial side usually 2-faced, adaxial side almost
flat), rugose, glabrous, beaked. Disc cypselae
angular, ± clavate, hairy. Ray pappus 0. Disc
pappus c. 8 alternating, lanceolate-attenuate
and shorter, acute to truncate, fimbriate scales.
x = 9. One species, O. tenella Nutt., western USA
(southern California), western Mexico (northern
Baja California).
1428. Raillardella (A. Gray) Benth. in Benth. &
Hook. f.
Raillardella (A. Gray) Benth. in Benth. & Hook. f., Gen. Pl.
2: 442 (1873).
Railliardia Gaudich. § [unranked] Raillardella A. Gray
(1865).
Perennials, rhizomatous. Leaves opposite, mostly
crowded in basal rosettes, blades lanceolate or
oblanceolate to linear, usually entire, glabrous or
sericeous or sparsely hirtellous and/or stipitateglandular. Capitula usually solitary on scapiform
peduncles, rarely laxly ± corymbose, radiate
or discoid. Peduncular bracts usually absent, if
present, tips each without a pit-gland, tack-gland
or spine. Involucres hemispheric or campanulate
to cylindric, phyllaries 1–13 (as many as ray
florets) or absent (the ‘involucre’ then consisting
of 1 series of receptacular paleae), lanceolate or
oblanceolate to linear, herbaceous, each usually
1/2+ investing the subtended ray floret proximally,
abaxially hirsute and ± stipitate-glandular. Receptacles flat or convex, glabrous or setulose, paleae
± persistent, in ring at periphery of receptacle
(constituting an ‘involucre’ in discoid heads),
connate or free (sometimes overlapping). Ray
florets 0 or 1–13, corollas yellow to yellow-orange
or orange to red-orange. Disc florets 7–80+,
bisexual, corollas concolourous with rays, glabrous
or sparsely minute-hairy; anthers yellowish,
appendages lanceolate-ovate to ovate-oblong;
styles glabrous proximal to arms, arms free,
apices subulate, densely hairy. Cypselae ± terete,
± clavate, strigose, not beaked. Ray pappus 0 or
8–30 ± subulate, ciliate to plumose scales. Disc
pappus 8–30 ± subulate, ciliate to plumose scales.
x = 17 or 18 (34, 35, polyploid). Three species,
western USA.
1429. Wilkesia A. Gray
Wilkesia A. Gray, Proc. Amer. Acad. Arts 2: 160 (1852);
Carr, Allertonia 3: 1–123 (1985), rev.; Baldwin & Robichaux
in Wagner & Funk (eds) Hawaiian biogeography: evolution
on a hot spot archipelago: 259–287 (1995), phylog.
Rosette shrubs, monocarpic or polycarpic. Leaves
in rosettes, whorled (7–15 per node), sessile
(basally connate, sheathing stem), blades linear,
Compositae
entire, glabrous or ± sericeous and/or glandular.
Capitula in elongate, verticilliform arrays, discoid.
Peduncular bract tips each without a pit-gland,
tack-gland or spine. Involucres broadly campanulate, phyllaries (modified paleae) in 1 series,
oblong-oblanceolate to oblanceolate, glandular.
Receptacles convex, glabrous, paleae deciduous,
in 1 ‘involucral’ series, connate. Florets 30–225,
corollas cream, lobes setulose; anthers yellow;
styles glabrous proximal to arms, arms free, hairy
in distal half, apices cuneate-deltate, acuminate,
acumen papillose. Cypselae ± terete, ± clavate,
mostly straight, sericeous. Pappus of 6–13 lanceolate, fimbriate scales. x = 14. Two species, Hawaiian
islands (Kauai).
XXVIII.5. Subtribe Venegasiinae B.G. Baldwin (2002).
Perennial herbs, subshrubs or weak shrubs.
Leaves mostly alternate, proximally opposite,
petiolate, blades rounded-deltate to ovate or
cordate, subentire or toothed, abaxially minutely
resin-dotted, adaxially glabrous. Capitula laxly
corymbose or borne singly, radiate. Involucres
hemispheric to globose. Phyllaries in 3–4+ series,
outer 1(–2) series often reflexed, broadly ovate to
deltate, inner 2(–3) series erect, membranous or
scarious and progressively smaller. Receptacles
flat or convex to shallowly conical, epaleate,
sparsely tomentose. Ray floret corollas yellow,
tubes densely pubescent and glandular, apices
entire to obscurely bilobed or trilobed. Disc florets
bisexual, corollas yellow, tubes densely glandularpubescent; anthers yellow, appendages ovate, with
numerous glands, endothecium of quadrate to
oblong cells with 2–3 polar thickenings; style arms
with broad stigmatic surfaces, apices deltate to
obtuse, papillose, appendages wanting. Cypselae
often curved, oblong with shallow ridges, black,
striate, glabrous. Pappus absent. x = 19.
Only one genus:
1430. Venegasia
Venegasia DC., Prodr. 6: 43 (1837) [1838]; Turner & Zippin,
Sida 15: 223–229 (1992), rev.
Characters of the subtribe. One species, V. carpesioides DC., western USA (southern California),
western Mexico (northern Baja California).
507
XXIX. Tribe Perityleae B.G. Baldwin
(2002).
J.L. Panero
Annual or perennial herbs but mostly rupicolous
shrubs; herbage viscid with stipitate or sessile glandular trichomes. Leaves usually opposite, usually
petiolate, sometimes succulent, sparsely to mostly
densely glandular, blades linear to ovate, cordate,
rarely hastate or deltate, triplinerved, sometimes
5-nerved or with a single vein and pectinate. Capitula discoid or radiate, solitary or in open, usually
terminal and dichasial, paniculiform or corymbiform cymes. Involucres cylindrical, campanulate
or hemispheric, rarely ovoid, phyllaries in 1 or 2
series. Receptacles flat to shallowly convex, usually epaleate. Ray florets pistillate, fertile. Disc florets 4- or 5-lobed, bisexual, fertile or functionally
staminate, corollas covered with sessile and stipitate glandular trichomes, lobes shallowly papillose
adaxially; anthers ecalcarate and ecaudate, usually white to yellow, appendages mostly ovate and
eglandular, endothecium with 1–3 polar bridges;
style arms with paired stigmatic surfaces rarely
fusing at apices, apices tapered to acute or round,
appendages present or absent. Cypselae variously
cylindrical, biconvex or obpyramidal with 2 to 4
angles, compressed, rarely obcompressed, mostly
black, walls carbonized, without striations, sometimes with corky, ciliate margins, rarely dimorphic (Galeana). Pappus of 2 (rarely 1) bristles and
a short crown, rarely of multiple bristles or vestigial squamellae or absent, sometimes of a shallow
crown of fimbriate scales, or of 4 squamellae and 4
bristles.
The tribe contains seven genera and 84 species
distributed predominantly in northern Mexico and
the south-western USA. Lycapsus is endemic to the
Desventuradas Islands off the coast of northern
Chile. Perityle has one or two species in the Andes
of southern Peru and northern Chile. Most species
of the tribe grow on rocky cliffs or very well-drained
sandy washes in the deserts of North America.
Perityleae have traditionally been recognized
as a distinctive group (subtribe) within the classical Helenieae because of their epaleate receptacles,
tetramerous corollas, 2- or 4-angled, compressed
cypselae, the densely glandular herbage, and involucres with one or two series of subequal, herbaceous phyllaries (Powell and Turner 1974). The
phylogenetic position of Perityle and related genera
has been clarified recently by molecular phyloge-
508
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
netic studies (Baldwin et al. 2002). The data of Baldwin et al. (2002) support the inclusion of Correllia
in the synonymy of Perityle, the division of Perityle
as advanced by Powell (1969, 1972, 1973, 1974), the
position of Eutetras as sister to Amauria, Pericome
and Perityle and, more importantly, the recognition of subtribe Peritylinae, as circumscribed by
Robinson (1981), at the tribal level. The decision by
Baldwin et al. (2002) to recognize Perityleae is also
supported by molecular studies using chloroplast
DNA sequences (Panero and Funk 2002). The same
studies for the first time provide strong support
for the placement of Eupatorieae as sister to Perityleae. The concept of tribe Perityleae, as circumscribed by Baldwin et al. (2002), is amended here
to include Galeana and Villanova. The inclusion of
these genera is based on evidence from molecular
phylogenetic studies of chloroplast DNA (Panero
et al. 2001c). The tribe is now composed of three
subtribes, namely Galeaninae, Lycapsinae and Peritylinae. The description of Lycapsus is based on the
literature and especially on the excellent account of
this monotypic genus by Skottsberg (1937).
Key to the Subtribes
1.
–
2.
–
Disc corollas pentamerous
Disc corollas tetramerous
Receptacles paleate
Receptacles epaleate
1. Galeaninae (p. 508)
2
2. Lycapsinae (p. 509)
3. Peritylinae (p. 509)
XXIX.1. Subtribe Galeaninae Panero and
B.G. Baldwin, subtribus nov. type:
Galeana La Llave & Lex.
Subtribui Peritylinae similis, a qua differt corollis 5-lobis
et cypselis triquetris.
Annuals with glandular herbage. Leaves alternate or opposite. Capitula in open paniculiform
cymes, radiate. Involucres hemispherical or ovoid,
phyllaries herbaceous, subequal, navicular. Receptacles epaleate. Ray florets pistillate, disc florets
pentamerous, bisexual or functionally staminate.
Cypselae triquetrous, pappus absent.
Key to the Genera
1. Ray cypselae with corky wings
– Ray cypselae without wings
1431. Galeana
1432. Villanova
1431. Galeana La Llave & Lex.
Fig. 104
Galeana La Llave & Lex., Nov. Veg. Descr. 1: 12 (1824).
Chlamysperma Less. (1832).
Fig. 104.
Compositae-Perityleae. Galeana pratensis.
A Habit. B Flowering capitulum. C Ray corolla. D Disc
corolla. E Anthers. F Style arms. G Ray cypsela. H Disc
cypsela. (Reproduced from Flora Novo-Galiciana, vol. 12,
with permission of the University of Michigan Herbarium;
see McVaugh 1984)
Leaves opposite, viscid, petiolate, blades deltate
to ovate, triplinerved. Capitula in terminal, open
corymbiform cymes. Involucres ovoid to cylindric, phyllaries 5, biseriate. Receptacles flat to
shallowly convex. Ray florets 3, fertile, corollas
creamy white. Disc florets 3–5, 5-lobed, fertile
or functionally staminate, corollas creamy white,
sparsely glandular-pubescent; anthers whitish to
yellow, appendages acute to linear; style arms with
round to deltate papillose apices. Ray cypselae
obcompressed, with corky, concave wings, black,
glabrescent. Disc cypselae obpyramidal, wingless,
black, glabrescent, tuberculate with age. x = 9.
Compositae
One species, G. pratensis Rydb., Mexico, Central
America.
1432. Villanova Lag.
Villanova Lag., Gen. Pl. 31 (1816), nom. cons.
Vasquezia Phil. (1860).
Leaves opposite or alternate, viscid, petiolate,
blades dissected, once pinnate, ovate in outline.
Capitula in terminal, open, simple or corymbiform
cymes. Involucres campanulate to hemispheric,
phyllaries few, broad. Receptacles flat to shallowly
convex. Ray florets fertile, corollas white to yellow.
Disc florets fertile, sometimes innermost functionally staminate, sparsely glandular-pubescent;
anthers yellow, appendages ovate and deeply
carinate; style arms with round to deltate papillose
apices. Cypselae triangular-obpyramidal, with
abaxial side wider than other two, essentially
glabrous. Ten species, Mexico, Central and South
America.
509
open or congested corymbiform or paniculiform
cymes, discoid or radiate. Receptacles epaleate.
Ray florets pistillate, fertile, disc florets tetramerous, bisexual. Cypselae cylindric, 4-angled or
most commonly biconvex, compressed, rarely
obcompressed. Pappus of squamellae or bristles,
most commonly of 1 or 2 bristles, rarely absent.
Key to the Genera
1.
–
2.
–
Cypselae 4-angled
2
Cypselae narrowly biconvex
3
Pappus absent or a very reduced crown 1434. Amauria
Pappus of 4 squamellae alternating with 4 bristles
1435. Eutetras
3. Leaves hastate or triangular, pappus of fimbriate scales
1436. Pericome
– Leaves various, never triangular nor hastate, pappus
of 1 or 2 or multiple bristles, never of fimbriate scales
1437. Perityle
Genera of Peritylinae
XXIX.2. Subtribe Lycapsinae H. Rob. (1980).
1434. Amauria Benth.
Perennial herbs or weak shrubs. Leaves alternate,
semisucculent, 1–2-pinnate, lobes linear. Capitula
terminal in open paniculiform cymes, radiate. Involucres campanulate. Receptacles paleate, paleae
lanceolate, involute. Ray florets pistillate, fertile,
corollas white, disc florets tetramerous, bisexual,
corollas white; anther thecae pale, appendages
ovate; style arms with acute papillose apices.
Cypselae fusiform, 4-angled, straight to slightly
incurved. Pappus absent.
Amauria Benth., Bot. Voy. Sulphur: 31 (1844); Powell,
Madroño 21: 516–525 (1972), rev.
Only one genus:
1433. Lycapsus Phil.
Lycapsus Phil., Bot. Zeitung (Berlin) 28: 499 (1870);
Robinson, Proc. Amer. Acad. Arts 49: 438–491 (1913),
rev.; Johnston, J. Arnold Arb. 16: 440–447 (1935), rev.;
Skottsberg, Göteb. Kungl. Vetensk. Vitterh. Samh. Hand.
ser. 5, B, 5: 1–88 (1937), rev.
Characters of the subtribe. One species, L.
tenuifolius Phil., San Felix and San Ambrosio
Islands off the coast of northern Chile.
XXIX.3. Subtribe Peritylinae Rydb. (1914).
Subtribe Amauriinae Rydb. (1914).
Annual or perennial herbs or rupicolous shrubs.
Leaves opposite or alternate. Capitula solitary or in
Annuals with glandular herbage. Leaves opposite
or alternate, sometimes succulent, petiolate, blades
ovate to cordate in outline, margins shallowly to
deeply lobed and serrate. Capitula radiate, in open
paniculiform cymes. Involucres campanulate or
hemispheric. Receptacles shallowly convex. Ray
corollas white, sometimes tinged with pink. Disc
corollas yellow, sparsely glandular-pubescent;
anthers white to yellow, appendages ovate; style
arms with tapered to round apices. Cypselae
cylindric, 4-angled, black, glabrous to sparsely
pubescent with variously shaped trichomes,
sometimes uncinate. Pappus absent or a shallow
crown. x = 17, 18. Three species, Baja California,
Mexico.
1435. Eutetras A. Gray
Eutetras A. Gray, Proc. Amer. Acad. Arts 15: 39 (1879);
Turner, Southw. Naturalist 11: 118–122 (1966), rev.
Rupicolous shrubs with brittle branches. Leaves
opposite, densely glandular, viscid, somewhat succulent, petiolate, blades deltate to ovate, margins
serrate. Capitula solitary or in small corymbiform
cymes, radiate, rarely discoid. Involucres campanulate to subhemispheric. Receptacles shallowly
convex. Ray corollas white. Disc corollas white
510
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
or whitish-yellow, sparsely glandular-pubescent;
anthers white to yellow, appendages oval; style
arms with tapered apices. Cypselae cylindric,
4-angled, black, glabrous to sparsely pubescent.
Pappus of 4 scales alternating with 4 bristles
arising from the cypsela angles. x = 18. Two
species, central Mexico.
1436. Pericome A. Gray
Pericome A. Gray, Smithsonian Contr. Knowl. 5: 81 (1853);
Powell, Southw. Naturalist 18: 335–339 (1973), rev.
Perennial herbs or weak shrubs. Leaves opposite,
petiolate, triangular, hastate. Capitula in congested
corymbiform cymes, discoid. Involucres campanulate to urceolate. Receptacles flat. Corollas
deep yellow or yellow-orange, sparsely glandularpubescent; anthers yellow, appendages ovate; style
arms with tapered to acute apices, essentially
without appendages. Cypselae narrowly biconvex,
compressed, with a corky margin. Pappus of
a small crown of fimbriate scales sometimes
with 1 or 2 caducous awns. x = 18. Two species,
south-western USA, northern Mexico.
1437. Perityle Benth.
Fig. 105
Perityle Benth., Bot. Voy. Sulphur 23, pl. 15 (1844); Niles,
Mem. New York Bot. Gard. 21: 1–81 (1970), rev.; Powell,
Rhodora 71: 58–93 (1969), sect. rev.; Sida 5: 61–128 (1973),
sect. rev.; Rhodora 76: 229–306 (1974), sect. rev.
Pappothrix (A. Gray) Rydb. (1914).
Corellia A.M. Powell (1973).
Annual or perennial herbs but mostly rupicolous
shrubs, some with brittle branches. Leaves opposite or alternate, petiolate, sometimes densely
glandular and viscid, blades deltate to ovate,
cordate or rarely linear to spathulate, entire to
deeply dissected. Capitula solitary or in open
corymbiform cymes, discoid or radiate. Involucres
cylindric, campanulate to hemispheric. Receptacles flat to shallowly convex. Ray corollas white or
yellow. Disc florets 4(–5)-lobed, corollas yellow,
creamy white or white, sometimes reddish; anthers
translucent whitish to yellow, appendages ovate;
style arms with tapered apices, appendages tapered, papillose, vascularized. Cypselae narrowly
biconvex, compressed, rarely obcompressed,
sometimes with wings or most commonly with
a well-developed corky, ciliate margin. Pappus of
2 (rarely 1) bristles and a short crown, rarely of
multiple bristles or vestigial squamellae or absent,
bristles retrorsely barbed. x = 17, 19. Sixty-five
Fig. 105.
Compositae-Perityleae. Perityle feddemae.
A Habit. B Flowering capitulum. C Disc corolla.
D Anthers. E Style arms. F Cypsela. (Reproduced from
Flora Novo-Galiciana, vol. 12, with permission of the
University of Michigan Herbarium; see McVaugh 1984)
species, western USA, Mexico, Central America,
Peru and Chile.
XXX. Tribe Eupatorieae Cass. (1819).
D.J.N. Hind and H. Robinson
Herbs (rarely aquatic or semi-aquatic), subshrubs,
shrubs, climbers, small trees, sometimes epiphytic.
Leaves usually opposite, rarely strictly alternate,
sometimes rosulate or verticillate, sessile or petiolate, lamina usually simple. Inflorescence usually
a corymbose panicle, sometimes spicate. Capitula
sessile or distinctly pedicellate, homogamous,
discoid, rarely with some zygomorphic outer
florets; involucre cylindrical, campanulate or
hemispherical, rarely subtended by a subinvolucral bract; phyllaries in 1 to several series, few
or numerous, imbricate, subimbricate or distant,
equal, subequal or markedly gradate, persistent or
variously deciduous, lanceolate or ovate; receptacle
flat to convex, sometimes highly conical, usually
naked, glabrous or sometimes pubescent. Florets
Compositae
few, very rarely 1, often 4 or 5 to many, commonly
fragrant; corollas funnelform to tubular, never
truly yellow, lobes relatively short, commonly 5,
very rarely 4; anther cylinders usually included
within corolla tube; apical anther appendages
obtuse or acute, rarely emarginate or lobed, as
long as broad or shorter, sometimes absent, basal
appendages short or almost absent, obtuse or
rounded; anther collars indistinct, cylindrical
or variously pronounced; nectary rarely visible;
style base glabrous or pubescent, sometimes with
a swollen node; styles usually very conspicuous
and much exserted, glabrous or rarely pubescent;
style arms linear to clavate, obtuse, stigmatic
surfaces variously papillate. Achenes obovoid or
oblong with phytomelanin in achene walls, usually
3–5-ribbed, sometimes 10-ribbed, body rarely
flattened with 2 ribs or 5 winged ribs, sometimes
glandular, glabrous or variously setuliferous;
carpopodium often paler than achene body, rarely
indistinct or absent, of several layers of variously
enlarged, sometimes ornamented cells, usually
symmetrical, rarely eccentric, annular, cylindrical,
or stopper-shaped; pappus sometimes absent and
reduced to an apical callus, rarely a laciniate crown,
or vestigial, occasionally coroniform, usually of
setae, commonly uniseriate, rarely biseriate or very
rarely multiseriate, usually persistent, sometimes
fragile, usually numerous, sometimes few, usually
equal or subequal, rarely very short, or occasionally of flattened scales or awn-like scales, rarely of
two distinct elements, very rarely of broad laciniate
setae, or of few clavate apical appendages; setae
commonly barbellate or laciniate, rarely plumose,
apices acute or obtuse, usually gradually tapering,
sometimes dilated, very rarely conspicuously
narrowing. The chromosome base number of the
tribe has been suggested to be either x = 10 (King
et al. 1976; King and Robinson 1987) or x = 17
(Yahara et al. 1989; Watanabe et al. 1995; Ito et al.
2000a).
A tribe of 17 subtribes containing c. 2200
species in 182 genera. The tribe is mostly restricted to the western hemisphere, suggesting
a Neotropical origin. There are many pantropical
and pansubtropical weeds in the tribe. Uses of
members of the tribe have been briefly summarized by King and Robinson (1987). More recent
references on the topic include Garg and Sastry
(1996; Ageratum conyzoides, Mikania micrantha),
Huang and Ling (1996; Ageratum houstonianum,
Adenostemma, Heinrich (1996; 31 Mexican species)
and Vallès et al. (1996; Eupatorium cannabinum).
511
Several recent molecular systematic studies
(Jansen and Kim 1996; Panero et al. 2001a, b, c;
Baldwin et al. 2002) clearly position Eupatorieae
nested within Heliantheae s.l. Panero et al. (2001a,
b, c) suggested positioning Eupatorieae within
a clade with Peritylinae and Galeana-Villanova
clades. Recently, Jeffrey (2002) has, whilst discussing the current state of the systematics of the
family, proposed the creation of the new rank, the
supersubtribe, for Eupatorieae, thereby creating
the Eupatoriodinae (Cass.) C. Jeffrey; this is not
followed here. The submergence of Eupatorieae
within Heliantheae would appear to be wholly
impracticable. Baldwin et al. (2002) have suggested
a modified scheme placing Eupatorieae alongside
Bahiinae (raised to tribal rank as Bahieae) and
Madieae. With the concept of ‘taxon monophyly
within our rank-based system of botanical nomenclature’ (Baldwin et al. 2002), this has apparently
left the continuing recognition of the broad concept of Heliantheae untenable if Eupatorieae are to
be maintained – a partitioning of Heliantheae into
‘tribes coordinate with Eupatorieae’ was proposed;
the result was the recognition of three new tribes.
Opinions vary widely as to the usefulness of
the classification system provided by King and
Robinson (1987). In many recent flora accounts
and checklists for South and Central America,
a large and unwieldy Eupatorium sensu lato/amplo
has been maintained (e.g. Santa Catarina/Brasil:
Cabrera and Klein 1991; Paraguay: Cabrera 1996;
Argentina: Cabrera and Freire 1999; New Mexico:
McVaugh 1984; Mexico: Turner 1997). Other
checklists and flora accounts with input from
Robinson naturally used the scheme proposed by
King and Robinson (e.g. Ecuador: Robinson 1999c;
Guyana: Funk and Robinson 1996; Peru: Dillon and
Hensold 1993). Bremer et al. (1994) briefly summarized the situation at the subtribal level, with
comments on how some authors had treated some
of the segregates proposed by King and Robinson
(1987), and provided a tribal order in part based
on the results of a series of cladograms. Inferences
may be made from a small series of molecular
systematic studies (Schilling et al. 1999; Schmidt
and Schilling 2000; Ito et al. 2000a), together
with the results presented by Funk et al. (2005).
Apart from the latter dataset, however, none of
the analyses are based on a particularly large
sample of genera, or species within those genera,
or even comparable taxa. Even the cladograms
presented provide conflicting results, although
some trends can indeed be detected. Bearing these
512
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
uncertainties in mind, we have chosen to base this
account largely on the tribal concepts proposed
by King and Robinson (1987), with modifications
based on published cladistic, molecular systematic
and cytological studies – evidently, this is certainly
not the final picture.
Classification of the Eupatorieae
1. Subtribe Hofmeisteriinae R.M. King & H. Rob. (1980)
Genera 1438–1439.
2. Subtribe Oxylobinae R.M. King & H. Rob. (1980).
Genera 1440–1448.
3. Subtribe Mikaniinae R.M. King & H. Rob. (1980).
Genus 1449.
4. Subtribe Trichocoroninae R.M. King & H. Rob. (1980).
Genera 1450–1452.
5. Subtribe Adenostemmatinae B. L. Rob. (1913).
Genera 1453–1455.
6. Subtribe Fleischmanniinae R.M. King & H. Rob. (1980).
Genera 1456–1457.
7. Subtribe Ageratinae Less. (1832).
Genera 1458–1483.
8. Subtribe Eupatoriinae Dumort. (1827).
Genera 1484–1487.
9. Subtribe Liatrinae R.M. King & H. Rob. (1980).
Genera 1488–1493.
10. Subtribe Praxelinae R.M. King & H. Rob. (1980).
Genera 1494–1500.
11. Subtribe Gyptidinae R.M. King & H. Rob. (1980).
Genera 1501–1529.
12. Subtribe Disynaphiinae R.M. King & H. Rob. (1978).
Genera 1530–1535.
13. Subtribe Ayapaninae R.M. King & H. Rob. (1980).
Genera 1536–1548.
14. Subtribe Alomiinae Less. (1832).
Genera 1549–1571.
15. Subtribe Critoniinae R.M. King & H. Rob. (1980).
Genera 1572–1611.
16. Subtribe Hebecliinae R.M. King & H. Rob. (1980).
Genera 1612–1618.
17. Subtribe Neomirandeinae R.M. King & H. Rob. (1980).
Genus 1619.
Key to the Subtribes
1. Phyllaries distant and bases not articulated; receptacles epaleaceous unsclerified, and changing shape with
maturity
5. Adenostemmatinae (p. 518)
– Phyllaries distant to imbricate and bases sclerified or
articulated; receptacle sclerified between areolae or
paleaceous
2
2. Paleae present, bearing floret in axil when removed;
capitula solitary on erect peduncles
1. Hofmeisteriinae (p. 513)
– Paleae absent or superficial on receptacle; capitula often in complex inflorescences
3
3. Capitula solitary on erect peduncles arising from
a congested group of leaves
1. Hofmeisteriinae (p. 513)
– Capitula often in complex inflorescences and peduncles not arising from pseudowhorls of leaves
4
4. Capitula with 4 subequal phyllaries and 4 florets, usually with a fifth unequal subinvolucral bract (subtending the involucre or pedicel); style arms never with
clavate apices; pappus always present
3. Mikaniinae (p. 516)
– Capitula not with 4 subequal phyllaries and 4 florets
or with clavate style arm apices or with a defective
pappus
5
5. Phyllaries all deciduous leaving a naked receptacle,
remaining appressed until lost, not spreading with age
10. Praxelinae (p. 532)
– At least some basal phyllaries persistent, phyllaries
usually spreading with age
6
6. Aquatic or sub-aquatic plants with sessile or whorled
leaves; Mexico and USA 4. Trichocoroniinae (p. 517)
– Plants terrestrial or epiphytic and not aquatic or subaquatic, usually with petiolate leaves
7
7. Leaves alternate and plants usually rosulate; eastern
USA
9. Liatrinae (p. 530)
– Leaves opposite or plants from other than USA
8
8. Capitula becoming elongate on a highly elongated and
paleaceous receptacle
13. Ayapaninae (Isocarpha)
– Capitula not becoming elongated on a highly conical
receptacle, receptacle paleaceous or epaleaceous
9
9. Plants epiphytes or growing in humus; leaves fleshy
or coriaceous, often blackening on drying; pappus of
numerous capillary setae
17. Neomirandeinae (p. 574)
– Plants not epiphytic nor growing in humus, or pappus
defective
10
10. Corolla lobes (usually) smooth on inner surface 11
– Corolla lobes papillose on inner surface
14
11. Pappus of awns, scales, crowns, or absent, never of
wingless setae; pappus awns bristle-like in some 5flowered species; plants rarely with densely spirally
inserted leaves
7. Ageratinae (p. 520)
– Pappus of capillary setae, sometimes short, rarely absent in some plants with densely spirally inserted
leaves; capitula rarely with fewer than 6 florets
12
12. Style base usually with a distinct glabrous node; phyllaries usually distant and subequal
2. Oxylobinae (p. 513)
– Style base lacking basal node; phyllaries distant or
subimbricate, usually gradate
13
13. Receptacles convex to conical; style arms linear or with
broadened apices; anther collars usually conspicuous
and expanded; cells of inner surface of corolla lobes
short and essentially isodiametric
11. Gyptidinae (p. 535)
– Receptacles usually flat; style arms filiform; anther collars scarcely discernible and cylindrical; cells of inner
surface of corolla lobes elongate with upper ends projecting as papillae
6. Fleischmaniinae (p. 520)
14. Style arms long-clavate, thickened in both width and
thickness, with few exceptions with 10-ribbed achenes, plumose pappus or with peg-like setulae on achenes; upper corolla limb often constricted beneath
corolla lobes
14. Alomiinae (p. 552)
– Style arms not thickened or thickened only at tips, if
long-clavate, then flattened; achenes never 10-ribbed,
never with plumose pappus setae and never with peg-
Compositae
15.
–
16.
–
17.
–
18.
–
like eglandular achene setulae; corolla limb cylindrical
or funnelform beneath corolla lobes
15
Style base glabrous and basal node absent; inner phyllaries usually deciduous
16
Style base with basal node or pubescent or both 18
Capitula always with 5 florets; style arm apices papillose; pappus setae sometimes with bulbous-tipped
apical cells; plants of eastern South America
12. Disynaphiinae (p. 546)
Capitula with 1–300 florets, rarely consistently
5-flowered; style arms usually smooth at least at
apices; pappus setae never with bulbous-tipped apical
cells; plants throughout tropical and subtropical
America
17
Receptacles glabrous; anther collar usually less than 5
times as long as wide
15. Critoniinae (p. 559)
Receptacles usually pubescent; anther collar usually
more than 5 times as long as wide
16. Hebecliniinae (p. 572)
Style base lacking basal node, pubescent; capitula with
fewer than 20 florets; receptacles epaleaceous; apical
cells of pappus setae usually rounded
8. Eupatoriinae (p. 528)
Style base inflated above nectary, glabrous or
pubescent; capitula usually with more than 20 florets,
sometimes receptacle paleaceous; apical cells of
pappus setae acute
13. Ayapaninae (p. 548)
XXX.1. Subtribe Hofmeisteriinae R.M. King
& H. Rob. (1980).
Subtribe Oaxacaniinae R.M. King & H. Rob. (1980).
Key to the Genera
1. Receptacles epaleaceous; pappus of barbellate tapering setae; leaves pseudowhorled below peduncle
1438. Hofmeisteria
– Receptacles paleaceous; pappus absent or a short
laciniate crown, sometimes with one long seta; stems
leafy throughout, leaves alternate 1439. Oaxacania
1438. Hofmeisteria Walp.
Hofmeisteria Walp., Repert. Bot. Syst. 6: 106 (1846); King,
Rhodora 69, 779: 35–47 (1967), rev.
Perennial herbs or woody subshrubs. Leaves usually congested at flowering nodes, lamina broadly
ovate to filiform, entire to lobed or greatly dissected. Capitula solitary on long erect bracteolate
peduncles, broadly campanulate. Phyllaries c. 50–
100, acute to pungent or obtuse; receptacle glabrous, areolate. Florets 100–175(–250), fragrant;
corollas white, pink, or lavender, nearly tubular,
usually glabrous on inner and outer surfaces, rarely
short stipitate-glandular outside; lobes with papillae at tip, or papillose inside only; style arms slightly
513
enlarged and flattened distally. Pappus setae 3–15,
barbellate, tapering, with 6–10 intervening lacerate squamellae in species with few setae. n = 18,
19. Ten species, Mexico.
1439. Oaxacania B.L. Rob. & Greenm.
Oaxacania B.L. Rob. & Greenm., Amer. J. Sci., ser. 3, 50: 151
(1895); King, Rhodora 69, 777: 3–47 (1967), rev.
Carterothamnus R.M. King (1967).
Sprawling subshrubs or shrubs. Leaf lamina
herbaceous, orbicular to reniform, 3–7-cleft to
2|3 or more towards base. Capitula terminal on
leafy branches, laxly cymose or solitary, broadly
campanulate. Phyllaries c. 30–40, inner somewhat
deciduous, apices narrowly acuminate or obtuse,
densely fringed; receptacle paleaceous, paleae
deciduous, linear-lanceolate, with single floret
partially enclosed by base of each palea. Florets
50–100; corollas white, with basal tube long
and slender with enlarged base and apex, outer
surface of tube with numerous stipitate glands,
a few smaller stipitate glands distally on limb
or corollas glabrous; lobes c. 1.5 times as long
as wide, papillose on inner surface, smooth and
somewhat thickened on outer surface; style base
conical, truncate below against nectary, arms
narrow linear, densely long-papillose or scarcely
mamillose. Pappus absent or a short laciniate
crown or setae c. 10–12, narrow, short, with one
barbed setae with bent or contorted apex. n = 18.
Two species, Baja California, Mexico.
Turner (1987a, 1997) proposed that both
Oaxacania and Carterothamnus be incorporated
into a broadened concept of Hofmeisteria, simply
noting that the only real difference was the
presence of paleae in the Oaxacaniinae. Here, we
‘sink’ Carterothamnus into a slightly broadened
concept of Oaxacania but maintain Oaxacania
separate from Hofmeisteria.
XXX.2. Subtribe Oxylobinae R.M. King & H.
Rob. (1978).
Perennial herbs or shrubs to small trees, usually
erect. Leaves usually opposite, lamina deltoid to
narrowly elliptical. Capitula clustered on short to
moderately long pedicels in terminal, pyramidically to corymbosely paniculate inflorescences,
phyllaries distant to weakly subimbricate, usually subequal; receptacle flat to slightly convex,
paleaceous or epaleaceous. Florets 7–75; corollas
narrowly funnelform or with slender basal tube
514
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
and campanulate limb, glabrous to rarely shortpubescent or glanduliferous on outer surface; lobes
often distinctly longer than wide, usually densely
papillose on inner surface, sometimes smooth or
roughened; anther collar often more than 5 times
as long as wide; apical anther appendages as long as
wide; style base often with distinct enlarged node,
glabrous; style narrowly linear, without enlarged
tips, usually densely papillose. Achenes prismatic
to fusiform, 5-ribbed; carpopodium usually distinct; pappus of numerous deciduous or persistent
capillary setae, rarely squamellae, apical cells acute.
Key to the Genera
1. Leaves trifoliate
1446. Standleyanthus
– Leaves simple
2
2. Stems with fleshy cortex; leaves absent at anthesis
1444. Pachythamnus
– Stems not unusually thickened; plants not seasonally
defoliated
3
3. Receptacles paleaceous
1443. Jaliscoa
– Receptacles epaleaceous
4
4. Pappus of persistent laciniate squamellae
1441. Oxylobus
– Pappus setae capillary and persistent or deciduous 5
5. Pappus setae easily deciduous
6
– Pappus setae persistent
7
6. Carpopodium indistinct and undifferentiated
1442. Piptothrix
– Carpopodium well defined and sometimes cylindrical
1440. Ageratina
7. Stems and leaves with a yellowish granular appearance
(resulting from concertina’d multicellular uniseriate
hairs); Colombia
1448. Jaramilloa
– Stems and leaves lacking yellowish granular appearance (when present, hairs sometimes crisped but not
concertina’d)
8
8. Inflorescence lax-corymbose; Cuba
1445. Spaniopappus
– Inflorescence densely corymbose on branchlets; Argentina, Bolivia, Brazil, Ecuador, Peru 1447. Kaunia
Genera of Oxylobinae
1440. Ageratina Spach
Ageratina Spach, Hist. Nat. Veg. Phan. 10: 286 (1841); King
& Robinson, Phytologia 24: 79–104. (1972), reg. rev.; King
& Robinson, Phytologia 69: 61–86 (1990), reg. rev.; Turner,
Phytologia Mem. 11: i–iv, 1–272 (1997), reg. rev.
Perennial, usually erect herbs or shrubs. Leaves
usually opposite, lamina narrowly elliptical to
deltoid, mostly toothed, lobed, serrate or crenate.
Capitula laxly to densely corymbose. Phyllaries
c. 30, 2- or 3-seriate, distant to weakly subimbricate, mostly subequal; receptacle usually slightly
convex, glabrous or with minute scattered hairs.
Florets 10–60, often sweetly scented; corollas
white or lavender, usually with slender basal tube
and campanulate limb in subgenera Ageratina
and Klattiella, others narrowly funnelform; lobes
distinctly longer than wide, inner surface densely
papillose, outer surface smooth, glabrous or
glanduliferous, usually with hairs in subgenus
Ageratina; anther collar cylindrical, usually elongate; anther appendage large, ovate-oblong, longer
than wide; style base usually enlarged except in
subgenus Apoda; arms rarely slightly broadened
distally, densely papillose with projecting cells
on lateral and outer surface. Achenes prismatic
or fusiform, usually 5-ribbed, setuliferous or
glanduliferous or both; carpopodium distinct,
cylindrical in subgenus Ageratina, in others
rounded or shortly stopper-shaped; pappus setae
uniseriate, 5–40, barbellate, often easily deciduous,
capillary, often enlarged distally, often with outer
series of shorter setulae. n = c. 10, c. 16, 17, 18, 20,
c. 25, c. 40, c. 42. Circa 265 species, tropics and
subtropics of the New World.
King and Robinson (1970) first provided an
infrageneric division recognizing four subgenera,
later raising subgenus Pachythamnus to generic
status. Subsequently (King and Robinson 1978),
they recognized a further two subgenera. A complete list of species in the five subgenera was provided by King and Robinson (1987).
1441. Oxylobus (Moçino ex DC.) A. Gray
Oxylobus (Moçino ex DC.) A. Gray, Proc. Amer. Acad. Arts
15: 25 (1879); Turner & Kerr, Pl. Syst. Evol. 151: 73–87
(1985), rev.
Decumbent herbs to low shrubs. Leaves opposite,
lamina small, ovate to oblong, crenate to subentire.
Capitula laxly to densely corymbose to subcymose.
Phyllaries c. 10–15, distant, 2–3-seriate, mostly
subequal; receptacle slightly convex, epaleaceous,
glabrous. Florets 20–75, slightly scented; corollas
white or pink, with a long narrow basal tube and
a narrowly campanulate limb; lobes longer than
wide, inner surface densely papillose, outer surface
smooth with stalked glands or glabrous; anther
collar cylindrical, elongate; apical anther appendages large, ovate, longer than wide; style base
enlarged; arms densely long-papillose on lateral
and outer surfaces. Achenes fusiform, 5-ribbed,
bearing short setulae; carpopodium distinct,
often stipitate, shortly rounded; pappus of a few,
Compositae
515
short, laciniate persistent squamellae. n = 16. Five
species, Colombia, Guatemala, Mexico, Venezuela.
1444. Pachythamnus (R.M. King & H. Rob.)
R.M. King & H. Rob.
1442. Piptothrix A. Gray
Pachythamnus (R.M. King & H. Rob.) R.M. King & H. Rob.,
Phytologia 23, 1: 153 (1972).
Piptothrix A. Gray, Proc. Amer. Acad. Arts 21: 383 (1886).
Erect herbs or weak shrubs. Leaves usually opposite, lamina ovate to narrowly ovate, serrulate
to serrate, apex narrowly acute to acuminate.
Capitula in a thyrsoid panicle with rather densely
corymbose branches. Phyllaries c. 7–15, distant,
biseriate, equal to subequal; receptacle slightly
convex, epaleaceous, glabrous to spinose. Florets
7–18, fragrant; corollas white, with narrow basal
tube and funnelform to slightly campanulate limb,
outer surface glabrous; lobes c. 1.5–2 times as
long as wide, inner surface mamillose to nearly
smooth, outer surface smooth; anther collar
cylindrical, narrow; apical anther appendages
large, ovate or oblong, longer than wide; style
base enlarged; arms scarcely broadened distally,
densely papillose. Achenes prismatic, 5-ribbed,
setuliferous; carpopodium indistinct; pappus setae
c. 15–75, barbellate, often deciduous, sometimes
only half as long as corolla, with outer series of
very short, small setulae. n = c. 17. Four species,
Guatemala, Mexico.
Shrubs or small trees. Stems swollen or somewhat
succulent, wrinkled when dry. Leaves opposite,
lacking at anthesis, lamina broadly ovate to deltoid,
margins with a few blunt teeth, apiculate at tip.
Capitula terminal in rather abrupt, dense corymb.
Phyllaries c. 15, distant to scarcely subimbricate,
2–3-seriate, mostly subequal; receptacle slightly
convex, with minute scattered hairs. Florets c. 15;
corollas white or lavender, narrowly funnelform,
with narrowly cylindrical basal tubes, puberulous
on upper tube and lower throat, glabrous inside;
lobes longer than wide, densely papillose on inner
surface, outer surface and margins smooth below;
anther collar cylindrical; apical anther appendage
large, ovate, slightly longer than wide; style base
not enlarged; arms not broadened distally, densely
papillose. Achenes prismatic, usually 5-ribbed,
setuliferous on ribs and sides; carpopodium
distinct; pappus setae uniseriate, c. 25, barbellate,
slender, rather easily deciduous, not broadened
distally. One species, P. crassirameus (B.L. Rob.)
R.M. King & H. Rob., El Salvador, Guatemala,
Mexico, Nicaragua.
1443. Jaliscoa S. Watson
Jaliscoa S. Watson, Proc. Amer. Acad. Arts 25: 153 (1890);
McVaugh, Fl. Novo-Galic.: 1157 (1984), rev.; Turner,
Phytologia Mem. 11: i–iv, 1–272 (1997), reg. rev.
1445. Spaniopappus B.L. Rob.
Erect perennial herbs or shrubs. Leaves usually opposite, lamina ovate to deltoid, crenulate-serrulate
to sharply serrate, apex shortly and often narrowly
acuminate. Capitula in a lax pyramidal panicle,
with rather densely corymbose lateral branches.
Phyllaries c. 15–20, distant, biseriate, subequal
to equal; receptacle slightly convex, paleaceous.
Florets 11–25; corollas white, with narrow basal
tube and funnelform to slightly campanulate limb;
lobes c. 1.5 times as long as wide, mamillose to
short-papillose on inner surface, smooth on outer
surface; anther collar elongate, cylindrical; apical
anther appendages large, ovate or triangular,
slightly longer than wide; style base sometimes enlarged; arms densely papillose. Achenes prismatic,
4–5-ribbed, glabrous or bearing a few setulae near
top; carpopodium usually distinct, symmetrical;
pappus an obscure callus border, a laciniate crown
or of rather short deciduous setae. Three species,
Mexico.
Erect perennial herbs or shrubs. Leaves opposite, lamina oblong-ovate to elliptical, entire to
remotely serrate or lobate-crenate, apex acute
to slightly acuminate, with immediate tip often
rather blunt. Capitula in rather broad and lax
corymbs. Phyllaries c. 15, weakly subimbricate,
unequally 2–3-seriate; receptacle slightly convex,
epaleaceous, glabrous. Florets 25–60; corollas
purple, distally narrowly funnelform with basal
tube narrowly cylindrical or constricted above
nectary, inner and outer surfaces glabrous; lobes
longer than wide, inner surface densely papillose
with broad papillae, margins and outer surface
papillose with strongly projecting cells; anther
collar cylindrical, elongate; apical anther appendage large, ovate, slightly longer than wide;
style base not or scarcely enlarged; arms scarcely
broadened above base, mamillose to slightly
papillose. Achenes prismatic, usually 5-ribbed,
glabrous to sparsely setuliferous; carpopodium
Spaniopappus B.L. Rob., Contr. Gray Herb. n.s. 77: 45
(1926).
516
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
1446. Standleyanthus R.M. King & H. Rob.
apex usually densely setuliferous or glanduliferous;
carpopodium distinct, symmetrical; pappus setae
uniseriate, c. 25–30, barbellate, persistent, contiguous, slender, not broadened distally. n = c. 20,
c. 26. Fourteen species, Argentina, Bolivia, Brazil,
Ecuador, Peru.
Standleyanthus R.M. King & H. Rob., Phytologia 22: 41
(1971).
1448. Jaramilloa R.M. King & H. Rob.
indistinct or distinct; pappus of c. 40 uniseriate
setae, barbellate, persistent, or of a few short
bristles and a fringe of minute squamellae. Five
species, Cuba.
Large flaccid shrub. Stems rather fleshy, leaves
opposite, lamina divided into 3 leaflets, leaflets
oblong-ovate, petiolulate, remotely crenateundulate, narrowly acuminate, venation pinnate.
Capitula in pyramidal panicles, terminal on
branches. Phyllaries c. 12, distant, c. 2-seriate,
subequal; receptacle slightly convex, epaleaceous, glabrous. Florets c. 17; corollas whitish
(?), narrowly funnelform with long cylindrical
basal tube, outer and inner surfaces glabrous;
lobes triangular, slightly longer than wide, with
dense prominent mamillae on inner surface, outer
surface smooth below and somewhat mamillose
above; anther collar cylindrical; apical anther
appendages large, oblong-ovate, slightly longer
than wide; style base scarcely thickened, arms
scarcely broadened distally, slightly mamillose.
Achenes prismatic, 4–5-ribbed, ribs prominent
and usually pale, setuliferous, greatly expanded
at base; carpopodium indistinct; pappus setae
uniseriate, c. 20, barbellate, mostly persistent,
contiguous, slender, not broadened distally. One
species, S. triptychus (B.L. Rob.) R.M. King & H.
Rob.
Jaramilloa R.M. King & H. Rob., Phytologia 47: 117 (1980);
King & Robinson, Phytologia 47: 117–120 (1980), key.
Erect shrubs or small trees, to 3 m tall. Leaves
opposite, lamina often large, broadly oblong,
serrulate to scarcely undulate, apex short-acute.
Inflorescences terminal on branches, broadly
corymbose, capitula short-pedicellate or sessile
in glomerules. Phyllaries c. 12–23, weakly subimbricate, unequally c. 3-seriate; receptacle slightly
convex, epaleaceous, glabrous. Florets 14–20;
corollas whitish, with narrow cylindrical basal
tube and narrowly campanulate limb, glabrous on
inner and lower outside surfaces; lobes scarcely to
markedly longer than wide, slightly mamillose on
inner surface; anther collar cylindrical, elongate;
apical anther appendages large, subquadrate to
shortly oblong-ovate; style base without node,
style shaft slightly thickened; arms markedly papillose. Achenes prismatic, 5-ribbed, setuliferous
or glanduliferous; carpopodium short-cylindrical,
symmetrical; pappus setae 1–2-seriate, c. 25–50,
barbellate, rather easily deciduous, slender,
contiguous, sometimes borne on outer surface
of callus, not broadened distally. Two species,
Colombia.
1447. Kaunia R.M. King & H. Rob.
Kaunia R.M. King & H. Rob., Phytologia 47: 258 (1980).
Erect shrubs or small trees. Leaves opposite, lamina usually ovate, entire to serrate, broadly to narrowly acute. Inflorescences terminal on branches,
thyrsoid to corymbose with densely corymbose
branches. Phyllaries c. 12–23, weakly subimbricate, c. 2–3-seriate, subequal to unequal; receptacle slightly convex, epaleaceous, glabrous. Florets
(10–)16–50; corollas usually white or violet, narrowly funnelform, glabrous inside and on lower
outer surface; lobes slightly longer than wide, inner surface nearly smooth, outer surface smooth
and somewhat padded, with occasional glands; anther collar cylindrical, elongate; apical anther appendages large, ovate, slightly longer than wide,
style base not enlarged; arms smooth to slightly
mamillose. Achenes prismatic, 5-ribbed, base and
XXX.3. Subtribe Mikaniinae R.M. King & H.
Rob. (1980).
Usually woody vines, sometimes erect perennial
herbs or shrubs, moderately branched, never
rosulate. Leaves opposite or whorled, sessile to
long-petiolate, lamina linear to broadly ovate,
base narrow to cordate, membranous to coriaceous. Inflorescence terminal on stems or lateral
branches, cymose to corymbose or thyrsoid,
capitula clustered, sessile to pedicellate, with
subinvolucral bract; phyllaries distant, 4, subequal,
persistent, receptacle flat, epaleaceous. Florets
4; corollas white or pink, funnelform or with
variously campanulate limb, with or without
distinct basal tube, glabrous to pilosulous or
glanduliferous on outer surface, with or without
papillae on inside of throat or lobes; lobes broadly
Compositae
517
triangular to narrowly oblong; anther collar broad;
anther cylinder exserted from corolla throat;
apical anther appendages as long as or longer
than wide; style base thick, without distinct basal
node, glabrous or sometimes papillose; arms
narrowly linear, not broadened at apex, scarcely
to strongly papillose. Achenes prismatic, 4–10ribbed; carpopodium short-cylindrical; pappus
setae numerous, persistent, capillary, apical cells
obtuse to acute.
Only one genus:
1449. Mikania Willd.
Fig. 106
Mikania Willd., Sp. Pl. 3, 3: 1742 (1803), nom. cons.; Aristeguieta, Fl. Venez. 10: 91–243 (1964), reg. rev.; Barroso,
Arch. Jard. Bot. Rio de Janeiro 16: 239–333, pl. 1–31, foto
1–57 (1958), reg. rev. Brazil; Holmes, Sida 9: 147–158 (1981),
reg. rev. USA; Holmes, Bot. Jahrb. Syst. 103: 211–246 (1982),
reg. rev. Old World; Holmes, Sida, Bot. Misc. no. 9: 69 pp.
(1993), reg. rev. Greater Antilles; Holmes & McDaniel, Fieldiana, Bot. n.s. 9: 1–57 (1982), reg. rev. Peru; Holmes &
McDaniel, Candollea 44: 32–35 (1989), reg. rev. Paraguay;
King & Robinson, Ann. Missouri Bot. Gard. 62: 965–981
(1975), reg. rev. Panama; Smith, Fl. Trop. E. Afr. Compositae, part 3 (2005), reg. rev.
Kanimia Gardner (1847).
Characters of the subtribe. n = 16, 17, 18, 19, 20,
21, 36, 54. Circa 400 species, pantropical, although
principally neotropical with a few apparent natives
in the Old World tropics.
The report of dioecy in the genus (Holmes
1991) is apparently restricted only to species from
the Greater Antilles.
XXX.4. Subtribe Trichocoroninae
R.M. King & H. Rob. (1980).
Erect to ascending, aquatic or sub-aquatic, perennial herbs, not or sparingly branched above base;
leaves opposite, sometimes verticillate, sessile.
Capitula often solitary on long peduncles, not
or laxly clustered; phyllaries distant, persistent;
receptacle convex to conical, epaleaceous, warty,
glabrous. Florets 50–125; corolla lobes papillose
inside and at tip and margins outside; apical
anther appendages about as long as wide; style
base not enlarged, glabrous; arms narrowly linear
to broadly filiform or slightly clavate, flattened
at least at tip, densely long-papillose. Achenes
prismatic with 5 ribs; carpopodium distinct,
Fig. 106.
Compositae-Eupatorieae. Mikania grazielae
(Mikaniinae). A Flowering shoot. B Node. C Floret.
(Drawings by Margaret Tebbs)
sometimes with upper rim; pappus setae short,
a crown, or absent.
Key to the Genera
1. Pappus absent; corolla tube narrow with abruptly campanulate throat
1451. Shinnersia
– Pappus of short setulae or of 5 broad scales; corolla
tube relatively short with a long, tubular or funnelform
throat
2
2. Pappus of 5 broad scales; leaves verticillate, linear
and entire; inflorescences of solitary capitula; achenes
glandular-punctate only
1452. Sclerolepis
– Pappus of short setulae; leaves opposite, sometimes
becoming alternate above, lamina usually oblong,
margins serrate; inflorescences of solitary capitula or
laxly branched and few-headed; achenes setuliferous,
setulae with acute apices
1450. Trichocoronis
Genera of Trichocoroninae
1450. Trichocoronis A. Gray
Trichocoronis A. Gray, Mem. Amer. Acad. Arts n.s. 4: 65
(1849); King & Robinson, Phytologia 19: 497–500 (1970),
rev.
518
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Erect, sub-aquatic, annual or perennial herbs.
Leaves opposite, sometimes becoming alternate
above, lamina membranous, oblong, serrate,
acute. Inflorescences monocephalic to laxly
branched. Phyllaries c. 30, 2–3-seriate, subequal,
broadly lanceolate. Florets 75–125; corollas with
a constricted, rather elongate basal tube, limb
narrowly campanulate; lobes about as long as wide;
anther collar cylindrical; apical anther appendages
subquadrate, slightly longer than wide; style arms
broadly filiform, densely long-papillose. Achenes
prismatic to somewhat fusiform, 4–5-ribbed, ribs
acutely setuliferous with sharp-pointed setulae;
carpopodium stopper-shaped with distinct upper
rim; pappus setae 2–6, short, barbellate, persistent.
n = 15. Two species, USA, Mexico.
1451. Shinnersia R.M. King & H. Rob.
Shinnersia R.M. King & H. Rob., Phytologia 19: 297 (1970).
Ascending aquatic herbs. Leaves opposite, lamina obovate-spathulate, sinuate-incised, grossly
dentate to partly dissected. Inflorescence usually
monocephalic. Phyllaries c. 25–30, 2–3-seriate,
essentially equal, oblong; receptacle convex to
conical, warty, glabrous. Florets 90–100; corollas
white, with narrow basal tube as long as limb,
throat broadly and rather abruptly roundedcampanulate, with glands on tube, with numerous
hairs on upper throat and outer surface of
lobes; lobes broader than long; anther collar
short-cylindrical; apical anther appendages ovate
to subquadrate, about as long as wide; style
arms broadly linear, flat, densely long-papillose.
Achenes prismatic, 4–5-ribbed, with a few glands,
ribs obtusely setuliferous; carpopodium shortcylindrical; pappus absent. n = c. 30. One species,
S. rivularis (A. Gray) R.M. King & H. Rob., USA,
Mexico.
1452. Sclerolepis Cass.
Sclerolepis Cass., Bull. Soc. Philom. Paris 1816: 198 (1816).
Erect, sub-aquatic to aquatic perennial herbs.
Leaves verticillate, 4–6 per node, lamina linear,
entire. Inflorescences terminal, usually monocephalic, produced only on ‘stranded’ stems,
never on aquatic and emergent stems; capitula
pedicellate; phyllaries 22–25, biseriate, subequal,
broadly lanceolate. Florets c. 50; corollas pink,
narrowly funnelform from slightly narrowed base,
with glands on tube and base of throat; lobes
short-triangular, with short thick hairs on outer
surface; anther collar narrowly cylindrical; apical
anther appendages scarcely wider than long; style
arms narrowly linear to slightly clavate, rather
flattened distally, densely long-papillose. Achenes
prismatic, 5-ribbed, serrulate to slightly crested on
ribs, with few glands near upper end; carpopodium
subcylindrical; pappus of usually 5, thick, broadly
oblong, blunt, indurate scales, margins densely
crenulate-denticulate. n = 15, c. 30. One species,
S. uniflora (Walter) Britton, Sterns & Poggenb.,
endemic to USA.
XXX.5. Subtribe Adenostemmatinae B.L.
Rob. (1913).
Annual or perennial herbs, often creeping or with
decumbent bases. Leaves opposite. Inflorescences
terminal, unbranched or cymose. Capitula pedicellate; phyllaries distant, herbaceous without articulated or sclerified bases, receptacle glabrous, without sclerified tissue between achene scars, shallowly convex, becoming more convex with age,
epaleaceous. Florets 10–200; corolla lobes smooth
on inner surface; style base lacking basal node, glabrous; arms weakly mamillose to smooth. Achenes
3–5-angled; carpopodium distinct or scarcely distinguishable from body; pappus usually with 3 or
5 viscid-tipped knobs, or absent.
Key to the Genera
1. Pappus absent
1455. Gymnocoronis
– Pappus of 3 or 5 elongate knobs
2
2. Tips of knobs spherical and not decurrent on to surface of knob; anther appendages as long as wide; pappus always of 5 knobs; style shaft always glabrous;
carpopodium scarcely distinguishable from body and
slightly asymmetrical
1454. Sciadocephala
– Tips of knobs elliptical and decurrent on to surface of
knob; anther appendages about 1/2 as long as wide;
pappus usually of 3, rarely 5 knobs; style shaft glabrous or with long hairs; carpopodium distinct and
asymmetrical
1453. Adenostemma
Genera of Adenostemmatinae
1453. Adenostemma J.R. Forst. & J.G. Forst.
Fig. 107
Adenostemma J.R. Forst. & J.G. Forst., Char. Gen. Pl.: 89
(1776); Grierson, Ceylon J. Sci. (Biol. Sci.) 10: 42–80 (1972),
reg. rev. Sri Lanka; Koyama, Mem. Natl Sci. Mus. (Tokyo)
37: 159–168 (2001), reg. rev. E. Asia; Koyama, Bull. Natl
Sci. Mus., B (Tokyo) 28: 49–60 (2002), reg. rev. Thailand;
Compositae
519
shaft with or without long hairs; arms slightly to
strongly clavate, often forming most showy part of
head, fleshy, rounded apically, scarcely mamillose
below. Achenes slightly curved, usually 3-angled
without distinct ribs or 5-angled, often tuberculate;
carpopodium forming a prominent asymmetrical
knob; pappus of usually 3 or 5 terete clavate knobs,
knobs with tips and upper outside surface covered
with an elongated mass of viscid glands. n = 10.
Circa 26 species, pantropical.
1454. Sciadocephala Mattf.
Sciadocephala Mattf., Notizbl. Bot. Gart. Berlin-Dahlem 14:
41 (1938); King & Robinson, Phytologia 29: 1–20 (1974), reg.
rev.
Fig. 107. Compositae-Eupatorieae. Adenostemma viscosum
(Adenostemmatinae). A Basal leaf. B Inflorescence. C Floret.
(Drawings by Margaret Tebbs)
Panigrahi, Kew Bull. 30: 647–655 (1975), reg. rev. Indian
reg.; King & Robinson, Phytologia 29: 1–20 (1974), reg. rev.
Perennial herbs. Leaf lamina narrowly elliptical to
broadly ovate or hastate, crenate to strongly serrate,
acute to slightly acuminate. Inflorescence very laxly
cymose. Phyllaries 10–30, biseriate, ± overlapping,
somewhat fused at base, equal to subequal; receptacle covered with discrete oval deeply concave scars.
Florets 10–60; corollas usually white, narrowly funnelform or with narrow basal tube and broadly
campanulate limb, usually with hairs or glands
on outer surface, hairs often moniliform; lobes c.
1.5 times longer than wide, non-papillose, anther
collar usually strongly expanded below; apical anther appendages distinctly shorter than wide; style
Perennial herbs. Leaf lamina narrowly ovate to elliptical or slightly obovate, entire to serrate. Capitula solitary or laxly subcymose. Phyllaries c. 6–
14, persistent, 1–2-seriate, subequal to equal, separate to base; receptacle with discrete oval scars
separated by soft tissue. Florets c. 9–15; corollas
white, narrowly funnelform, with sparse hairs on
outer surface; lobes slightly to distinctly longer than
wide; anther collar stout, not or slightly broadened below; apical anther appendages ovate, as
long as wide or slightly longer; nectary usually glabrous, rarely pubescent; style arms long and narrow, rounded apically, scarcely mamillose below.
Achenes narrowly prismatic, nearly terete, without
distinct ribs; carpopodium only slightly asymmetrical, not enlarged; pappus of 5 terete clavate knobs,
knobs with a short globular mass of glutiniferous
glands apically. n = 10. Five species, Colombia,
Ecuador, Guyana, Panama.
1455. Gymnocoronis DC.
Gymnocoronis DC., Prodr. 5: 106 (1836); King & Robinson,
Phytologia 29: 1–20 (1974), reg. rev.
Annual to perennial erect herbs. Leaf lamina lanceolate to ovate or deltoid. Inflorescence strongly
cymose. Phyllaries c. 20–50, biseriate, equal to
subequal; receptacle with discrete oval scars and
with soft tissue between. Florets 50–200; corollas
white, narrowly funnelform, with short-stalked
glands on outer surface; lobes as wide as long to
wider than long; anther collar slightly enlarged;
apical anther appendages small, wider than long;
style arms very broadly oar-shaped, surface
mamillose below, smooth above. Achenes slightly
curved, prismatic (4–)5-ribbed, glanduliferous
between ribs, ribs sometimes corky; carpopodium
520
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
short-cylindrical; pappus absent. Five species,
Argentina, Bolivia, Brazil, Guatemala, Mexico,
Paraguay, Peru, Uruguay. Recently naturalized in
Japan (Suyama 2001).
XXX.6. Subtribe Fleischmanniinae R.M.
King & H. Rob. (1980).
Erect annual or mostly perennial herbs, lacking
rosulate basal leaves. Leaves opposite, rarely alternate. Inflorescence terminal on stems or branches,
with corymbose or cymose branches, capitula
clustered, usually pedicellate. Phyllaries usually
distinctly subimbricate, gradate, persistent; receptacle flat to slightly conical, epaleaceous. Florets
(10–)20–50; corollas with short basal tube, throat
narrowly campanulate, with veins thickened below;
lobes short-triangular; anther collar slender; apical
anther appendages about as long as wide; style base
usually not enlarged, glabrous below, or with scattered glands or hairs on shaft, arms filiform, not or
scarcely broadened distally, densely papillose. Achenes prismatic, 5-ribbed or terete; carpopodium
distinct with slight projecting upper rim or
indistinct; pappus setae uniseriate, 5 to many,
barbellate, capillary, with pointed apical cells.
Key to the Genera
1. Achenes 5-ribbed; style shaft glabrous
1456. Fleischmannia
– Achenes terete; style shaft pubescent and glandular
1457. Sartorina
1456. Fleischmannia Sch. Bip.
Fleischmannia Sch. Bip., Flora 33: 417 (1850); King &
Robinson, Ann. Missouri Bot. Gard. 62: 835–1322 (1975),
reg. rev. Panama; Robinson, Proc. Biol. Soc. Wash. 114:
529–556 (2001), new spp.; Wooten & Clewell, Rhodora 73:
566–574 (1971), reg. rev. eastern North America.
Eupatorium L. sect. Microstemon Cabrera (1991).
Erect annual or perennial herbs or subshrubs.
Leaves usually opposite, lamina elliptical to
rhomboidal or broadly cordate-ovate, upper part
serrate or crenulate, or lamina dissected into long
narrow segments. Inflorescence with laxly cymose
to densely corymbose branches. Phyllaries 20–30,
subimbricate, rarely distant, 2–4-seriate, usually
unequal; receptacle glabrous or with minute
scattered hairs. Florets (10–)20–50; corollas white,
lavender, bluish or purple, with rather short basal
tube, limb narrowly funnelform with ± cam-
panulate base, with outer surface above or on
the lobes often with short hairs or glands, veins
greatly thickened in tube or throat; lobes short;
apical anther appendages broadly ovate or oblong;
style base glabrous; arms densely long-papillose.
Achenes prismatic, 5-ribbed, usually with setulae
on ribs of upper parts; carpopodium distinct with
prominent upper rim; pappus setae 5–40, slender,
sometimes slightly fragile. n = 4 (rarely), 10, 20 or
30 (rarely). Ninety-five species, neotropical.
1457. Sartorina R.M. King & H. Rob.
Sartorina R.M. King & H. Rob., Phytologia 28: 98 (1974).
Erect to partially decumbent perennial herbs.
Leaf lamina broadly ovate to deltoid, crenulate to
obtusely serrulate. Inflorescence subthyrsoid with
subcorymbose branches. Phyllaries c. 20, subimbricate, c. 3-seriate, unequal, outer phyllaries ovate,
inner oblong; receptacle flat, glabrous. Florets c.
15–22; corollas white or lavender (?), with short,
narrowly cylindrical basal tube, limb narrowly
funnelform with ± campanulate base, glabrous
outside below lobes, veins greatly thickened in tube
and throat; lobes broadly triangular, with glands
on outer surface; apical anther appendages shortly
oblong; style base slightly enlarged, glabrous;
shaft sparsely short-pubescent and stipitateglanduliferous; arms densely long-papillose.
Achenes terete, narrowed below to a point, glabrous, with 5 narrow veins, without black deposits
in walls; carpopodium undifferentiated; pappus
setae c. 15, slender, somewhat non-contiguous,
persistent. One species, S. schultzii R.M. King &
H. Rob., Mexico (?).
XXX.7. Subtribe Ageratinae Less. (1832).
Subtribe Piqueriinae Benth. & Hook. f. (1873).
Erect to procumbent annual or perennial herbs,
shrubs or trees. Leaves opposite, sometimes alternate above, rarely throughout. Inflorescence terminal, sometimes diffuse; capitula clustered, sessile
to long-pedicellate. Phyllaries distant to subimbricate, persistent; receptacle scarcely convex to distinctly conical, paleaceous or epaleaceous. Florets
3–125; corolla lobes usually papillose; apical anther
appendages short or absent to longer than wide,
sometimes apically cleft or crenulate; style base
not or rarely enlarged, usually glabrous, arms usually linear, sometimes long-clavate, not abruptly
short-clavate at tips, usually densely papillose or
Compositae
mamillose. Achenes usually prismatic, 5-ribbed,
base sometimes contorted but not long-stipitate;
carpopodium usually distinct, sometimes with upper margin procurrent on base of achene; pappus
reduced, sometimes totally lacking, coroniform, or
of awns or scales, rarely of more than 5 primary elements. Bremer’s limited analysis (1994) suggests
that the subtribe is monophyletic ‘in some but not
all the cladograms’.
Key to the Genera
1. Marginal florets conspicuously zygomorphic and with
3 ray-like, expanded outer lobes
1482. Microspermum
– Marginal florets actinomorphic (rarely with slightly
zygomorphic marginal florets in Ferreyrella, Metastevia and Revealia but never conspicuously ray-like) 2
2. Phyllaries with conspicuous broadened, often
coloured apical appendages
1464. Scherya
– Phyllaries acute, obtuse or rounded but without broadened and coloured apical appendages
3
3. Receptacles paleaceous
4
– Receptacles epaleaceous
9
4. Receptacle conical
5
– Receptacle flat or slightly convex
6
5. Apical anther appendages conspicuous and large,
somewhat longer than wide; florets > 20
1460. Ageratum
– Apical anther appendages abortive or much reduced;
florets ≤ 14
1472. Nesomia
6. Herbaceous; paleae with densely fimbriate apices 7
– Plants shrubby or tree-like with monopodial or sympodial branching
8
7. Corollas cylindrical, sparsely glandular-punctate; apical callus and abscission layer between achene and
corolla absent; carpopodium absent or obscure; Brazil
1467. Teixeiranthus
– Corollas with constricted basal tube, tube stipitateglandular; apical callus and abscission layer between
achene and corolla present; carpopodium distinct,
symmetrical and appearing short-cylindrical; Peru
1471. Ferreyrella
8. Leaves glabrous and glandular-punctate or variously
eglandular-pubescent; pappus a short crown or a few
awns; inflorescences on pseudowhorled side branches
and plants usually with terminal vegetative lead shoot;
Brazil
1458. Acritopappus
– Leaves glandular-punctate and short stipitateglandular; pappus absent; inflorescences terminal or
axillary, but not whorled, and plants without terminal
lead vegetative shoot; Guatemala, Honduras
1463. Blakeanthus
9. Receptacle conical
1460. Ageratum
– Receptacle flat or slightly convex
10
10. Achenes compressed, 2-ribbed 1484. Macvaughiella
– Achenes prismatic usually 5-ribbed or sometimes 8–
10-ribbed
11
11. Corollas 4-lobed; 4 stamens per floret; plants minute,
< 10 cm tall
12
521
– Corollas 5-lobed; 5 stamens per floret; plants usually
much more than 10 cm tall
13
12. Florets 3–7 per capitulum; phyllaries (4–)5–6; Mexico
1474. Piqueriopsis
– Florets c. 30 per capitulum; phyllaries 8–10; Costa Rica
and Panama
1483. Iltisia
13. Plants repent with side shoots as small rosettes; inflorescence of clustered scapose capitula; pedicels shorter
than leaves; Peru
1465. Ascidiogyne
– Plants erect or ascending, or if procumbent shoots
leafy throughout; inflorescences usually exceeding
leaves
14
14. Pappus absent
15
– Pappus present
21
15. Achenes setuliferous; corolla tube constricted and
long stipitate-glandular, corolla limb markedly
expanded and only sparsely long stipitate-glandular
at base
1462. Phalacraea
– Achenes glabrous; corolla tube short or indistinguishable from funnelform limb, glabrous, short- or longpubescent, rarely stipitate-glandular and then glandular hairs throughout lower half of corolla
16
16. Corolla throat densely pubescent inside; Mexico
1477. Metastevia
– Corolla throat glabrous inside
17
17. Corolla glabrous outside; Brazil (Céara)
1473. Piqueriella
– Corolla short- or long-pubescent outside
18
18. Corollas with constricted corolla tube, hairs eglandular
19
– Corollas funnelform and stipitate-glandular in lower
half; florets 12–15; phyllaries biseriate, 10–12; Brazil
(Goías)
1468. Gardnerina
19. Anther filaments papillose or short-pubescent at base;
florets 3–5; phyllaries uniseriate, 3–5; Mexico, West
Indies
1475. Piqueria
– Anther filaments glabrous; florets 6–40
20
20. Apical anther appendages as long as wide; corollas
with conspicuously campanulate limb above constricted tube; florets 6–30; phyllaries 8–10; Brazil
(Pará)
1466. Cavalcantia
– Apical anther appendages vestigial; corollas with funnelform limb above narrowed base; florets 15–40; phyllaries 10–20; Ecuador and Peru
1470. Guevaria
21. Capitula always with 5 florets and 5 phyllaries; phyllaries uniseriate; corolla throat pubescent inside
1476. Stevia
– Capitula usually with numbers of phyllaries and florets
not equal (usually with > 10 florets and > 8 phyllaries,
sometimes florets 4–6 and then phyllaries 6); phyllaries 2–4-seriate; corolla throat glabrous inside
22
22. Florets 35–70; phyllaries 4-seriate; Brazil and Uruguay
1459. Radlkoferotoma
– Florets 4–25; phyllaries 1–2-seriate
23
23. Leaves narrowly linear; style arms linear and terete
with stigmatic lines close together on inner surface
and reaching nearly to tip; Mexico and USA
1478. Carphochaete
– Leaves broadly linear to oblong, ovate or ovateelliptical; style arms filiform or with expanded tips,
and stigmatic lines on edges of inner surface and
ending well before style arm appendage
24
24. Leaves sessile or very short-petiolate
25
– Leaves distinctly petiolate
26
522
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
25. Pappus setae of distinct long awns or squamellae; achenes 8–10-ribbed; stems erect, sparingly branched
1479. Cronquistia
– Pappus short, coroniform, of many short squamellae; achenes 5–6-ribbed; stems spreading and often
procumbent, densely branched
1480. Revealia
26. Pappus of 5 squamellae; achenes usually glabrous; leaf
lamina distinctly glandular-punctate beneath; florets
12–25; Cuba and West Indies
1461. Phania
– Pappus of c. 20 short squamellae; achenes moderately setuliferous; leaf lamina lacking distinct glandular punctae beneath; florets 10–12; Peru
1469. Ellenbergia
Genera of Ageratinae
1458. Acritopappus R.M. King & H. Rob. Fig. 108
Acritopappus R.M. King & H. Rob., Phytologia 24: 401
(1972); King & Robinson, Phytologia 45: 142–157 (1980),
key.
Shrubs or trees. Leaves opposite, lamina ovate,
ovate-elliptical, lanceolate or linear, glabrous or
pubescent, often viscid, serrate to subserrulate
or nearly entire, obtuse, short-acute to longacuminate. Inflorescence on leafy side branches,
branches densely subcymose to appearing subverticillate. Phyllaries 2–3-seriate, 5–25, distant;
receptacle flat to convex, paleaceous, paleae
linear, carinate. Florets 5–100; corollas usually
pale lavender or pink, narrowly funnelform,
glandular-punctate or glandular-stipitate on
outer surfaces; lobes slightly longer than wide,
inner surface with short cells usually with short
papillae; style base not enlarged, glabrous, arms
linear, densely short-papillose. Achenes 5-ribbed,
glabrous; carpopodium distinct, short; pappus
vestigial, a short crown, or of few awns. n = 9.
Circa 16 species, Brazil.
1459. Radlkoferotoma Kuntze
Radlkoferotoma Kuntze, Revis. Gen. Pl. 1: 358 (1891);
Cabrera, Bol. Soc. Argent. Bot. 6: 239–242 (1957), key, sub
Carelia Less.
Shrubs or small trees. Leaves opposite, lamina
ovate to lanceolate, serrate. Inflorescence corymbose. Phyllaries c. 35, markedly subimbricate, c.
4–5-seriate, gradate, inner somewhat deciduous;
receptacle flat, glabrous. Florets c. 35–70; corollas
white or rosaceous, funnelform, with only minute
glands on outer surface; lobes about twice as
long as wide, smooth on both surfaces; anther
collar broadly cylindrical; style base not enlarged,
glabrous, arms linear, scarcely or not broadened
Fig. 108. Compositae-Eupatorieae. Acritopappus connatifolius (Ageratinae). A Flowering shoot. B Floret. (Drawings
by Margaret Tebbs)
distally, densely papillose. Achenes prismatic,
4–5-ribbed, sparsely setuliferous; carpopodium
cylindrical, procurrent on to base of achene ribs;
pappus of 5 short broad scales. Three species,
Brazil, Uruguay.
1460. Ageratum L.
Ageratum L., Sp. Pl. 2: 839 (1753); Johnson, Ann. Missouri
Bot. Gard. 58: 6–88 (1971), rev.; Robinson, Phytologia 69:
93–104 (1990), reg. rev.
Annual to perennial herbs or subshrubs. Leaves opposite or sometimes alternate, lamina elliptical or
lanceolate to deltoid or ovate, entire to dentate. In-
Compositae
florescence cymose to subcymose, sometimes subumbellate. Phyllaries 30–40, distant, 2–3-seriate,
equal or subequal, lanceolate, markedly indurate,
often with scarious margins; receptacle conical,
glabrous or paleaceous. Florets 20–125; corollas
white, blue or lavender, funnelform or with distinct basal tube; lobes about as long as wide, papillose on inner surface, partially papillose and sometimes hispidulous on outer; anther collar cylindrical; style base not enlarged, glabrous, arms linear, usually strongly and densely papillose. Achenes prismatic, 4–5-ribbed, glabrous or ribs setuliferous; carpopodium distinct; pappus lacking,
or coroniform, or of 5 or 6 free, flattened, sometimes awn-like scales. n = 10, 20. Circa 40 species,
Central and South America.
One species (A. houstonianum Mill.) widely
cultivated, and another (A. conyzoides L.), although sometimes cultivated, is a widespread
weed throughout the tropics in both the Old and
New Worlds.
1461. Phania DC.
Phania DC., Prodr. 5: 114 (1836).
Annual or perennial herbs or small shrubs. Leaves
opposite, lamina ovate or deltoid to palmate,
crenate-dentate to shallowly lobed. Inflorescence
of small cymes or solitary capitula terminal
on leafy branches. Phyllaries c. 10–20, distant,
biseriate, equal or subequal, broadly lanceolate to
narrowly oblanceolate; receptacle conical, epaleaceous, glabrous. Florets c. 12–25; corollas pale
lavender to white, with short constricted basal tube
abruptly expanding into short-campanulate limb,
with scattered glandular punctae on outer surface;
lobes about as long as wide or slightly wider, with
large strongly mamillose cells on inner surface;
anther collar rather long; style base not enlarged,
glabrous, arms somewhat strongly clavate distally,
short-papillose or mamillose. Achenes prismatic,
4–5-ribbed, glabrous or with few glands or setulae; carpopodia distinct, cylindrical, somewhat
procurrent on to lower ribs of achene; pappus
squamellae 5. Five species, Cuba, West Indies.
1462. Phalacraea DC.
Phalacraea DC., Prodr. 5: 105 (1836); King & Robinson,
Phytologia 29: 251–256 (1974), rev.
Perennial herbs, procumbent or erect from decumbent bases. Leaves opposite, lamina ovate to
broadly triangular, crenate to serrate. Inflores-
523
cence rather laxly cymose, with denser ultimate
branching. Phyllaries 10–18, distant, 2–3-seriate,
subequal, broad with short-acute non-scarious
tips; receptacle flat to slightly convex, glabrous.
Florets 10–18; corollas white, with constricted
basal tube and abruptly expanding campanulate
limb, many scattered minutely gland-tipped hairs
on tube, more sparse on limb; lobes slightly longer
than wide, papillose on inner surface; anther collar
often rather stout; style base not enlarged, glabrous,
arms broad, becoming slightly broader towards the
broadly rounded tips, densely papillose. Achenes
prismatic, 4–5-ribbed, moderately to densely
setuliferous; carpopodium small; pappus absent.
n = 20. Four species, Colombia, Ecuador, Peru.
1463. Blakeanthus R.M. King & H. Rob.
Blakeanthus R.M. King & H. Rob., Phytologia 24: 118 (1972).
Erect shrubs. Leaves opposite, lamina ovate, crenate to serrulate. Inflorescence a compact corymbose panicle, branches with ± glomerulous capitula. Phyllaries c. 20, distant, 2–3-seriate, subequal,
lanceolate; receptacle flat to slightly convex, paleaceous. Florets c. 25; corollas white, narrowly funnelform; lobes slightly longer than wide, glandularpunctate on outer surface, smooth on inner; anther collar slender; style base not enlarged, glabrous, arms linear, not or scarcely broader at tips,
mamillose to subpapillose. Achenes prismatic, 5ribbed, mostly glabrous, with few minute spicules
on ribs and some glands on upper callus; carpopodium symmetrical, short; pappus absent. One
species, B. cordatus (S.F. Blake) R.M. King & H.
Rob., Guatemala, Honduras.
1464. Scherya R.M. King & H. Rob.
Scherya R.M. King & H. Rob., Phytologia 38: 101 (1977).
Erect perennial herbs. Leaves opposite, lamina
linear, entire. Inflorescence terminal, subscapose,
cymose to subcymose. Phyllaries c. 20, distant,
± biseriate, subequal, with broad, rounded, chartaceous, coloured tips; receptacle flat to slightly
convex, paleaceous. Florets c. 25; corolla pale,
funnelform, with glands on outer surface; lobes
slightly longer than wide, nearly smooth and
glanduliferous on outer surface, densely papillose
on inner; anther collar rather short; style base
not enlarged, glabrous, arms filiform, densely
papillose. Achenes prismatic, 5-ribbed, glabrous,
bands of carbonization very narrow; carpopodium
symmetrical, procurrent along ribs of achene;
524
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
pappus a laciniate crown with 5 long primary
teeth, rather indurate. One species, S. bahiensis
R.M. King & H. Rob., Brazil.
1465. Ascidiogyne Cuatrec.
Ascidiogyne Cuatrec., Ann. Missouri Bot. Gard. 52: 310
(1965).
Prostrate, somewhat fleshy, stoloniferous herbs
with erect rosulate branches. Leaves of prostrate
stems opposite, congested on erect stems, lamina
ovate or obovate to narrowly elliptical, entire.
Inflorescence of clustered short 1-headed scapes.
Phyllaries 4–7, distant, biseriate, equal, broadly
oblong to ovate; receptacle flat, glabrous, foveolate. Florets 5–7; corollas white, tube strongly
constricted and pilose in upper part, limb broadly
campanulate and glabrous; lobes up to twice as
long as wide, smooth on most of inner surface;
anther collar rather short, enlarged; style base not
enlarged, glabrous, arms short, clavate, with short
papillae denser towards tip. Achenes prismatic, 5ribbed, glabrous; carpopodium slightly asymmetrical; pappus lacking or of a rim of callus forming
a high 5-lobed crown. n = 10. Two species, Peru.
The peculiar fluid-filled sac, formed from the
outer wall of the achene, is present only in the type,
and evident only in fresh material.
1466. Cavalcantia R.M. King & H. Rob.
Cavalcantia R.M. King & H. Rob., Phytologia 47: 113
(1980).
Erect annual or short-lived perennial herbs. Lower
leaves opposite, alternate above, lamina ovate
to deltoid, distinctly shallowly lobed. Capitula
strongly divaricately cymose or aggregated in
glomerules. Phyllaries 8–10, distant, biseriate,
subequal to equal, persistent, narrowly obovate to
ovate; receptacle flat or conical, glabrous. Florets
c. 6–30; corollas white; basal tube short, broad
below and constricted above, densely pubescent;
limb funnelform; lobes as long as wide, papillose
on inner surface; anther collar slightly enlarged
below; style base not enlarged, glabrous, arms
filiform, densely papillose. Achenes prismatic,
5-ribbed, glabrous; carpopodium small; pappus
absent. Two species, Brazil.
1467. Teixeiranthus R.M. King & H. Rob.
Teixeiranthus R.M. King & H. Rob., Phytologia 47: 108
(1980); King & Robinson, Phytologia 50: 379–384 (1982),
rev.
Erect or decumbent, annual or short-lived perennial herbs. Leaves opposite, lamina elliptical to
linear. Inflorescence a corymbose cyme. Phyllaries
c. 10, distant, ± biseriate, equal, persistent,
elliptical to narrowly obovate, with expanded
apices; receptacle conical, paleaceous; paleae
oblanceolate with expanded apices. Florets c. 30;
corollas pale reddish, cylindrical with extreme base
campanulate, base fused directly to top of achene,
sometimes with poorly developed abscission zone;
lobes slightly longer than wide, densely papillose
on inner surface, smooth on outer; anther collar
scarcely discernible; style base not enlarged,
glabrous, arms filiform, slightly wider distally,
densely and strongly papillose. Achenes prismatic
to subfusiform, usually 5-ribbed; carpopodium
apparently absent; pappus absent. Two species,
Brazil.
1468. Gardnerina R.M. King & H. Rob.
Gardnerina R.M. King & H. Rob., Phytologia 49: 2 (1981).
Annual or short-lived perennial herbs, erect from
decumbent bases. Leaves opposite below, alternate
above, lamina ovate to rhomboid, repand-dentate
to pinnatifid. Inflorescence a few-headed cyme.
Phyllaries c. 10–12, distant, biseriate, equal to
subequal, persistent, lanceolate, bicostate; receptacle flat, glabrous. Florets 12–15; corollas white,
funnelform, with stipitate glands outside in lower
part, throat with short non-glandular hairs inside
on and near bases of filaments; lobes c. 1.25–1.5
times as long as wide, inner surface densely
papillose, outer smooth; anther collar cylindrical;
style base not or scarcely enlarged, glabrous,
arms broadly clavate or strap-shaped, flattened,
rather fleshy, densely short-papillose, mamillose
distally. Achenes subprismatic, 5-ribbed, glabrous;
carpopodium distinct, ± inflated; pappus absent.
One species, G. angustata (Gardner) R.M. King &
H. Rob., Brazil.
1469. Ellenbergia Cuatrec.
Ellenbergia Cuatrec., Proc. Biol. Soc. Wash. 77: 142 (1964).
Erect annual herb. Leaves opposite, a few alternate above, lamina ovate to ovate-elliptical,
crenate-dentate, acute. Inflorescence a lax panicle
with subcymose branches. Phyllaries 8, distant,
biseriate, subequal, persistent, oblong-elliptical
to obovate-elliptical; receptacle flat, foveolate,
glabrous. Florets 10–12; corollas white (?), with
distinct constricted basal tube, glandular-punctate
Compositae
outside; throat broadly campanulate, papillate on
inner surface; lobes about as long as wide, densely
papillose on inner surface and margins; lower
part of filament short, densely pubescent; anther
collar slender; style base not enlarged, hirtellous,
arms shortly clavate, densely papillose. Achenes
prismatic, 5-ribbed, ribs setuliferous, narrowed
and densely setuliferous above carpopodium, carpopodium distinct, shortly cylindrical; pappus of
c. 20 short, lanceolate squamellae, densely scabrid
on margins, nearly smooth on outer surface. One
species, E. glandulata Cuatrec., Peru.
1470. Guevaria R.M. King & H. Rob.
Guevaria R.M. King & H. Rob., Phytologia 29: 259 (1974).
Small perennial herbs, decumbent or erect with
decumbent bases. Leaves opposite or alternate,
lamina ovate, crenulate to serrulate. Inflorescence
laxly paniculate with cymose branches. Phyllaries
10–20, distant, 2–3-seriate, subequal, persistent,
oblong; receptacle conical, glabrous. Florets 15–40;
corollas white, with distinct constricted basal
tube bearing many hairs; throat shortly and
broadly campanulate; lobes 1–2 times as long
as wide, densely papillose on inner surface and
margins, with short hairs and glands on outer
surface; anther collar only slightly expanded; style
base not enlarged, glabrous, arms broadly linear,
densely long-papillose. Achenes prismatic, obovate, 5-ribbed, glabrous; carpopodium cylindrical;
pappus absent. n = 10. Five species, Ecuador, Peru.
1471. Ferreyrella S.F. Blake
Ferreyrella S.F. Blake, J. Wash. Acad. Sci. 47: 407 (1958).
Small, erect, annual herbs. Leaves opposite below,
alternate above, lamina ovate to broadly elliptical,
coarsely to finely serrate. Inflorescence a diffuse
corymbose cyme. Phyllaries c. 15–25, distant,
biseriate, subequal to equal, persistent, ovate
to broadly oblanceolate, with rounded apices,
pubescent; receptacle conical, paleaceous. Florets
c. 30; corollas white, with short constricted basal
tube bearing glandular or eglandular hairs; throat
short and broad-campanulate; lobes as long as
wide or outer lobes of peripheral florets longer,
densely short-papillose on inner surface and
margins; lower part of filament glabrous; anther
collars rather short; style base not enlarged,
glabrous; arms rather short-clavate, densely shortpapillose; papillae larger and less dense below
clavate tip. Achenes prismatic, 5-ribbed, glabrous;
525
carpopodium distinct, short-cylindrical; pappus
absent. Two species, Peru.
1472. Nesomia B.L. Turner
Nesomia B.L. Turner, Phytologia 71: 208 (1991).
Perennial herb. Leaves opposite, lamina ovate to
deltoid, coarsely serrate, acuminate. Inflorescences
congested paniculate corymbs. Involucres campanulate; phyllaries distant, oblanceolate, subequal, 1–
2(–3)-ribbed, glabrous, margins ciliate; receptacles
conical, paleaceous. Florets 12–14; corollas light
purple or lavender, tube pubescent with glandular and eglandular hairs, throat campanulate, lobes
broadly triangular, glabrous; anther collars linear;
style arms linear, short-papillose. Achenes 4–5ribbed, glabrous; carpopodium conspicuous; pappus absent. One species, N. chiapensis B.L. Turner,
Mexico.
1473. Piqueriella R.M. King & H. Rob.
Piqueriella R.M. King & H. Rob., Phytologia 29: 264 (1974).
Small, annual or short-lived perennial herbs.
Leaves usually opposite, lamina ovate, with many
large teeth, shortly and narrowly acuminate.
Inflorescence a lax cyme. Phyllaries c. 6, distant, biseriate, equal, persistent, broadly oblong,
subtruncate, 3–5-denticulate; receptacle slightly
convex, glabrous. Florets c. 8; corollas whitish, with
distinct constricted basal tube, glabrous on outer
surface; throat broadly and shortly campanulate;
lobes longer than wide, densely papillose on inner
surface and margins; anther collars rather short,
stout; style base not enlarged, glabrous, arms short
and subclavate, with dense long papillae. Achenes
prismatic, 5-ribbed, glabrous; carpopodium
strongly asymmetric; pappus absent. One species,
P. brasiliensis R.M. King & H. Rob., Brazil.
1474. Piqueriopsis R.M. King
Piqueriopsis R.M. King, Brittonia 17: 352 (1965).
Minute, ephemeral, erect herbs. Leaves opposite,
lamina ovate to elliptic-rhomboid, crenulate to
undulate. Inflorescence a small panicle with
few-headed cymose branches. Phyllaries (4–)5–6,
distant, ± biseriate, equal to subequal, persistent,
obovate to oblanceolate; receptacle convex, glabrous. Florets 3–7; corollas white, with distinct
constricted short basal tube, bearing numerous
stout non-glandular hairs near base; throat short
and widely funnelform from a subcampanu-
526
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
late base; corolla lobes 4, c. 1.5 times as long
as wide, strongly mamillose on inner surface,
glabrous on outer; anther collar stout; style base
slightly enlarged, glabrous, arms short, not or
slightly clavate, mamillose. Achenes prismatic,
8–10-ribbed, glabrous; carpopodium forming
a somewhat flaring annulus; pappus absent. One
species, P. michoacana R.M. King, Mexico.
Turner (1987b, 1997) suggested that Piqueriopsis should be included within an expanded Microspermum, along with Iltisia.
1475. Piqueria Cav.
Piqueria Cav., Icon. 3: 18 (1794); Turner, Phytologia Mem.
11: i–iv, 1–272 (1997), key.
Erect, annual to perennial herbs or subshrubs.
Leaves mostly opposite, lamina ovate to lanceolate,
serrulate to serrate. Inflorescence laxly paniculate
with laxly to densely subcymose branches. Phyllaries uniseriate, 3–5, distant, equal, persistent,
obtuse to apiculate; receptacle flat, glabrous.
Florets 3–5; corollas white to slightly lavender,
with short, narrow, densely pubescent basal tube;
throat short-campanulate; lobes 1.5–3 times as
long as wide, densely papillose on inner surface,
slightly papillose on margins and tip outside;
anther collar broad; style base not enlarged,
glabrous, arms with slightly to strongly clavate
appendages, densely long-papillose on narrower
parts, becoming nearly smooth on tips. Achenes
prismatic, 5-ribbed, glabrous; carpopodium
usually asymmetrical and large; pappus absent.
n = 11, c. 22, c. 24. Six species, Central America,
Mexico, West Indies.
1476. Stevia Cav.
Fig. 109
Fig. 109. Compositae-Eupatorieae. Stevia morii (Ageratinae). A Leafy basal portion of stem. B Part of inflorescence.
C Floret. (Drawings by Margaret Tebbs)
Stevia Cav., Icon. 4: 32 (1797).
Mostly erect, annual or perennial herbs or shrubs.
Leaves opposite or in some species alternate, lamina linear to orbicular, entire to serrate or dentate,
rarely deeply lobed. Inflorescence diffuse or dense
corymbose clusters on tips of branches. Involucres cylindrical, rarely funnelform, narrow at base;
phyllaries uniseriate, 5, distant, equal to subequal,
linear to elliptical; receptacle flat to slightly convex, glabrous. Florets 5; corollas white or lavender to purple, basal and distal parts sometimes of
different colour, narrowly funnelform below lobes
or with somewhat expanded throat, usually with
hairs or glands on outer surface, with erect hairs
on inner surface of throat; corolla lobes triangular
or oblong-ovate, densely papillose on inner surface, smooth on outer surface, usually less than
2 mm long, unequal and zygomorphic in series
Podocephalae, equal in others, longest lobes shorter
than throat and tube; anther collar cylindrical or
broadened below; style base often slightly enlarged,
glabrous or in a few species papillose, arms filiform,
densely long-papillose. Achenes narrowly fusiform
to narrowly prismatic, strongly 5-ribbed, with few
to many glands or setulae; carpopodium distinct,
short; pappus at least a crown of free or united
scales, often with 1–30 bristle-like awns, 1 or 2 achenes in each head often with more reduced pappus.
n = 11, 12, 17. Circa 175–230 species, from USA
Compositae
and Mexico south to Central (excluding the West
Indies) and South America.
A difficult genus with an inadequate infrageneric division. Robinson’s original treatments
(Robinson 1930a, b, c, d, 1931a, b, c, 1932a, b)
cover much of the distribution of the genus, with
the exception of the Brazilian species. A number of
more recent regional treatments (Argentina: Ariza
Espinar and Cerana 1986; Cabrera and Freire 1997;
Mexico: McVaugh 1984; Turner 1997; North America: Grashoff 1974) have been supplemented by the
addition of many new species. A modern revision is
certainly needed to adequately document the problems of hybridization and apomixis in the genus.
1477. Metastevia Grashoff
Metastevia Grashoff, Brittonia 27: 69 (1975).
Erect perennial herbs. Leaves opposite, basal pairs
very small, lamina ovate, crenate to serrate, acute
to obtuse. Inflorescence a lax panicle. Involucre
narrowly campanulate, broad at base; phyllaries
± biseriate, 4–6, distant, equal to subequal,
ovate-oblong; receptacle slightly convex, alveolate.
Florets 4–6; corollas white, with narrow basal tube,
expanding into narrowly campanulate throat,
sparsely pubescent on inner surface of throat;
corolla lobes more than 2.5 times longer than
wide, 2–2.5 mm long, longest equal to throat and
tube, sometimes zygomorphic with outer lobes
longer, inner surface densely papillose, outer
mostly glabrous, with a few papillae at tips; anther
collar rather short; style base enlarged, glabrous,
arms filiform, densely long-papillose. Achenes
obconical, 5-ribbed, glabrous; carpopodium
distinct, short; pappus absent. One species, M.
hintonii Grashoff, Mexico.
1478. Carphochaete A. Gray
Carphochaete A. Gray, Mem. Amer. Acad. n.s. 4: 65 (1849).
Erect perennial herbs or small shrubs. Leaves
opposite, lamina linear to elliptical-spathulate.
Inflorescence single-headed or a loose corymbose
cyme. Phyllaries c. 6, 2–4-seriate, distant to
subimbricate, unequal to subequal, persistent,
elliptical to oblong-lanceolate; receptacle flat to
slightly convex, glabrous. Florets 4–6; corollas
white, pink or purple, tubular below with narrowly
funnelform throat, glabrous; lobes c. 2–3 times as
long as wide, densely papillose on inner surface,
smooth or nearly smooth on outer surface; anther
collar expanded below; style base bulbous, gla-
527
brous, arms filiform, densely papillose, terete to
slightly compressed. Achenes narrowly prismatic,
5–10-ribbed, ribs setuliferous; carpopodium
distinct; pappus of 5–10 awns, longer than corolla,
sometimes with a few shorter squamellae. n = 11.
Circa 7 or 8 species, Mexico, USA.
Turner (1987b) treated Carphochaete in a broad
sense and included the monotypic Cronquistia and
Revealia.
1479. Cronquistia R.M. King
Cronquistia R.M. King, Brittonia 20: 11 (1968).
Erect perennial herbs. Leaves mostly opposite,
often alternate above, lamina linear to oblong,
entire. Inflorescence laxly corymbose. Phyllaries
10–15, distant, 1–2-seriate, equal or subequal,
persistent, elliptical; receptacles flat to slightly
convex, glabrous. Florets (5–)7–8(–12); corollas
purplish, narrowly funnelform, with few glandular
punctae outside, inner surface glabrous; lobes c.
3 times as long as wide, inner surface densely
papillose, outer rather smooth; anther collar short;
style base not enlarged, glabrous, arms distinctly
flattened and somewhat long-clavate, densely
short-papillose. Achene prismatic, 8–l0-ribbed,
ribs setuliferous; carpopodium a narrow basal rim;
pappus of a few winged awns or squamellae, or
absent. n = 12. One species, C. pringlei (S. Watson)
R.M. King, Mexico.
1480. Revealia R.M. King & H. Rob.
Revealia R.M. King & H. Rob., Phytologia 33: 277 (1976).
Spreading and often procumbent shrubs. Leaves
opposite, lamina narrowly oblong, entire to
obscurely crenulate-serrate, rather fleshy. Inflorescence usually of single capitula on short
leafy branches. Phyllaries c. 10, distant, biseriate,
equal to subequal, persistent; receptacle flat to
slightly convex, glabrous. Florets c. 10; corollas
purple, narrowly funnelform, with numerous
small short-stalked glands on outer surface,
upper throat short-pilose with numerous often
septate hairs; lobes oblong, outer three lobes
longer, densely papillose on inner surface, nearly
smooth on outer surface; anther collar very short;
style base with distinct enlargement, glabrous,
arms filiform, terete, densely papillose. Achenes
prismatic, 5-ribbed, ribs with minute setulae;
carpopodium distinct, forming a narrow basal
rim; pappus short, coroniform with many short
squamellae. One species, R. macrocephala (Paray)
R.M. King & H. Rob., Mexico.
528
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
1481. Macvaughiella R.M. King & H. Rob.
Macvaughiella R.M. King & H. Rob., Sida 3: 282 (1968).
Erect perennial herbs or small shrubs. Leaves opposite, lamina deltoid to rhomboid, serrate to dentate, short-acute to narrowly acuminate. Inflorescence densely corymbose. Phyllaries c. 10, distant,
± biseriate, equal to subequal, persistent; receptacle convex, glabrous. Florets 16–25; corollas white,
with a narrow basal tube expanding into a narrowly
campanulate throat, with several reddish glandular punctae on outer surface; lobes slightly longer
than wide, inner surface densely papillose, outer
only slightly roughened, with many uniseriate nonglandular hairs; anther collar elongate; style base
not enlarged, glabrous, arms filiform, densely papillose, bearing numerous reddish glandular punctations. Achenes compressed, 2-angled, setuliferous; carpopodium distinct, slightly asymmetrical;
pappus setae 2, rarely 1 or 3–4, coarsely barbellate,
apical cells acute. n = c. 13. Two species, Mexico,
Guatemala, El Salvador, Honduras.
1482. Microspermum Lag.
Microspermum Lag., Gen. Sp. Pl.: 25 (1816); Rzedowski,
Bol. Soc. Bot. Mex. 32: 77–86 (1972), key.
Annual or perennial herbs. Leaves opposite, lamina
lanceolate to rhomboid-ovate, serrate, obtuse to
rounded. Capitula solitary or in lax cymose to
corymbose panicles. Phyllaries 6–18, distant,
± biseriate, subequal, persistent, receptacle convex
to low-conical, glabrous. Florets 8–25; corollas
white, with narrow basal tube expanding rather
abruptly into campanulate throat, glandularpunctate on outer surface; lobes 5, triangular, in
peripheral zygomorphic corollas outer 3 lobes
greatly expanded and oblong, partly fused at base,
rather ray-like, densely papillose on inner surface,
nearly smooth on outer surface; anther collar
cylindrical; style base not enlarged, glabrous, arms
rather short, linear, densely papillose. Achenes
prismatic (4–)5(–7)-ribbed, setuliferous and
glanduliferous; carpopodium distinct; pappus
setae 0–4, coarsely barbellate, apical cells acute.
n = 12. Seven species, Mexico.
ovate, crenate-serrate with 1 or 2 coarse teeth
on each side, obtuse to rounded. Inflorescence
cymose, with 3–7 capitula. Phyllaries 5–10, distant,
biseriate, equal, persistent; receptacle slightly
convex, glabrous. Florets c. 30; corollas white,
with narrow basal tube, expanding into broadly
campanulate throat, with a few glands externally; lobes (3–)4(–5), slightly wider than long,
sometimes with 2 outer lobes of marginal florets
enlarged, densely papillose on inner surface,
scarcely mamillose on outer surface near apices;
anther collar cylindrical; style base not enlarged,
glabrous, arms short, appendages lanceolate,
densely short-papillose, acute. Achenes prismatic,
usually 4-ribbed, glabrous; carpopodium distinct,
forming a narrow projecting rim; pappus absent.
Two species, Costa Rica, Panama.
XXX.8. Subtribe Eupatoriinae Dumort.
(1827).
Annual or perennial herbs and subshrubs, usually
erect; leaves mostly opposite or whorled, alternate
above. Inflorescence terminal, usually corymbose
or pyramidal. Capitula short-pedicellate or sessile;
phyllaries subimbricate, often broadly scarious or
sclerified; receptacle scarcely convex, glabrous. Florets 3–23; corolla lobes smooth on inner surface;
style base with hairs, usually without a basal node;
arms usually without apical enlargement (enlarged
in some Stomatanthes), densely short-papillose except on enlargements. Achenes 5-angled, glabrous;
carpopodium obsolete to enlarged; pappus setae
numerous, capillary.
Key to the Genera
1. Achene body setuliferous
2
– Achene body only with glands
3
2. Inflorescence pyramidal or thyrsoid (if corymbose,
plants African)
1486. Stomatanthes
– Inflorescence corymbose
1487. Hatschbachiella
3. Carpopodium indistinct or absent. Widespread, but
not present in South America
1484. Eupatorium
– Carpopodium large and distinct; South America
1485. Austroeupatorium
1483. Iltisia S.F. Blake
Genera of Eupatoriinae
Iltisia S.F. Blake, J. Wash. Acad. Sci. 47: 409 (1958).
1484. Eupatorium L.
Decumbent, annual or short-lived perennial
herbs, striate, short-pilose, rooting at lower nodes.
Leaves opposite, lamina orbicular to broadly
Eupatorium L., Sp. Pl.: 836 (1753); Lamont, Mem. New York
Bot. Gard. 72: 1–68 (1995), part. rev.
Eupatoriadelphus R.M. King & H. Rob. (1970).
Compositae
Annual to perennial herbs. Leaves opposite or verticillate, upper subopposite to alternate, lamina linear to ovate, deltoid or trilobed, serrate to subentire. Capitula in a corymbose or pyramidal panicle.
Phyllaries 10–22, weakly to strongly subimbricate,
2–5-seriate, sometimes inner deciduous; receptacle
flat or weakly convex. Florets 3–23; corollas white to
purple, lavender or pink, narrowly funnelform or
with constricted basal tube and narrowly to broadly
campanulate limb, outer surface with glands often
concentrated at base of throat and on outer surfaces of lobes, rarely with a few hairs; lobes usually
slightly longer than wide; anther collar cylindrical;
apical anther appendages large, ovate-triangular,
c. 1.5 times as long as wide; style base puberulous
or rarely glabrous, with or without enlargement;
arms filiform to slightly broadened or flattened distally, papillose. Achenes prismatic, 5-ribbed; carpopodium not or slightly differentiated; pappus
setae 25–40, barbellate, persistent, apical cells with
rounded to short-acute tips. n = 10, 15, 20. Fortyfive species, Eurasia, North America.
The results of the studies by Schmidt and
Schilling (2000) and those of Ito et al. (2000b)
are in conflict as to the re-recognition of Eupatoriadelphus. Schmidt and Schilling (2000) favour
a distinct Eupatoriadelphus (= the ‘Eutrochium
group’) whereas Ito et al. (2000b) imply that
Eupatorium s.s. can be divided into three ‘morphological species groups’, one of which is the
‘Eutrochium group’.
1485. Austroeupatorium R.M. King & H. Rob.
Fig. 110
Austroeupatorium R.M. King & H. Rob., Phytologia 19: 433
(1970).
Eupatorium L. sect. Austroeupatorium (R.M. King & H.
Rob.) Cabrera (1991).
Erect herbs or subshrubs. Leaves opposite below,
often subopposite or alternate above, lamina ovate
to narrowly oblong, usually crenulate to serrulate.
Capitula in a flattened corymbose panicle. Phyllaries c. 12–18, 2–3-seriate, mostly unequal; receptacle flat or slightly convex. Florets 9–23, fragrant;
corollas white, rarely lilac, narrowly funnelform
with rather narrow tube, glands on outer surface;
lobes c. 1.5 times as long as wide, without stomata; lower part of filaments slender and flexuous;
anther collar narrowly cylindrical; apical anther
appendages ovate-oblong, longer than wide, style
base not enlarged, densely puberulous; style filiform. Achenes prismatic, 5-ribbed; carpopodium
529
Fig. 110. Compositae-Eupatorieae. Austroeupatorium inulifolium (Eupatoriinae). A Flowering shoot. B Floret. (Drawings by Margaret Tebbs)
distinct; pappus setae 30–40, slender, barbellate,
persistent, apical cells often enlarged, with rounded
tips. n = 10. Thirteen species, southern South
America, one species adventive in palaeotropics.
1486. Stomatanthes R.M. King & H. Rob.
Stomatanthes R.M. King & H. Rob., Phytologia 19: 430
(1970); Robinson, Phytologia 20: 334–337 (1970), reg. rev.
Perennial herbs or subshrubs. Leaves alternate, opposite or ternate, lamina elliptical or oblanceolate
to ovate or orbicular, entire to markedly dentate.
Inflorescence usually pyramidal to thyrsoidparticulate, sometimes corymbose. Phyllaries
4–12, distant to weakly subimbricate, 2–3-seriate,
unequal to subequal; receptacle scarcely convex.
Florets 4–11; corollas white, funnelform or nearly
tubular, glabrous or glanduliferous with few to
many hairs outside; lobes as long as wide to nearly
1.5 times as long as wide; lower parts of filaments
short, thick, straight; anther collars cylindrical;
530
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
apical anther appendages ovate or slightly shorter
than wide; style base lacking basal node, covered
with numerous hairs; arms linear to filiform or
with clavate tips, papillose at least below, tips
when greatly enlarged with smooth surfaces.
Achenes prismatic, 5–8-ribbed; carpopodium usually distinct; pappus setae uniseriate, numerous,
barbellate, persistent, apical cells with obtuse or
acute tips. n = 10. Sixteen species, eastern, central
and southern Africa, Brazil, Paraguay, Uruguay.
The results of nuclear ITS sequencing of Eupatorium s.l. by Schmidt and Schilling (2000) suggest
that ‘Stomatanthes appeared to be more closely related to Praxelinae (Chromolaena) than it is to any
member of the Liatrinae–Eupatorium clade’. This
is not supported, however, by its morphology. It is
retained here in the Eupatoriinae.
epaleaceous or with a few paleae, glabrous. Florets
4–80; corollas funnelform; lobes shortly triangular
to linear-oblong, mammillose or papillose inside;
apical anther appendages slightly wider than long
to longer than wide, often truncate or retuse at tip,
style base not enlarged, glabrous; arms filiform or
linear, not or scarcely broadened above, densely
mamillose or short-papillose. Achenes prismatic,
usually 8–10-ribbed, rarely 5-ribbed, usually
setuliferous with setulae divided nearly to base,
with only glands in Hartwrightia; carpopodium
indistinct; pappus of many, capillary, barbellate
to plumose, 1–3-seriate setae, with pointed apical
cells, or lacking.
Schilling and Cox (2001) have provided an analysis of DNA sequence data which was used as the
basis for the arrangement of genera in the present
account.
1487. Hatschbachiella R.M. King & H. Rob.
Hatschbachiella R.M. King & H. Rob., Phytologia 23: 393
(1972).
Eupatorium L. sect. Gyptis (Cass.) Cabrera (1991), p.p.
Erect herbs or subshrubs. Leaves opposite or alternate, lamina elliptical to narrowly elliptical, entire
to remotely serrulate. Inflorescence diffuse, with
corymbose panicles at tips of leafy branches. Phyllaries 12–15, weakly subimbricate, 2–3-seriate, unequal; receptacle flat. Florets 10–12; corollas white,
narrowly funnelform with a narrow basal tube,
glanduliferous on outer surface; lobes c. 1.5 times
as long as wide, with or without hairs on outer
surface; filaments in lower part slender and flexuous; anther collars cylindrical; apical anther appendages ovate, as long as or longer than wide;
style base not or slightly enlarged, with a few hairs;
arms linear to narrowly clavate, distinctly papillose. Achenes prismatic, 5-ribbed, sometimes stipitate below; carpopodium distinct; pappus setae
c. 30–40, barbellate, persistent, with slender tips,
apical cells acute. n = 10. Two species, Argentina,
Brazil, Paraguay, Uruguay.
XXX.9. Subtribe Liatrinae R.M. King &
H. Rob. (1980).
Erect perennial herbs with rosulate basal leaves at
least in early stages, or small shrubs. Leaves alternate. Inflorescence terminal on stems or branches,
thyrsoid or corymbose to pseudospicate, capitula
clustered, sessile or pedicellate; phyllaries few-(2–
5)seriate, usually distinctly subimbricate, gradate,
persistent; receptacle flat or slightly convex,
Key to the Genera
1. Pappus absent; achenes 5-ribbed, glanduliferous, setulae absent
1493. Hartwrightia
– Pappus of capillary setae; achenes 8–10-ribbed, setulae
few to many
2
2. Shrubs; leaves of mature stems not or scarcely reduced
in size in upper parts; pappus setae 60–70, barbellate,
outer series shorter and more slender 1488. Garberia
– Erect perennial herbs with basal rosettes (at least, initially) and reduced upper leaves; pappus setae 12–40,
barbellate or subplumose to plumose
3
3. Pappus setae plumose or subplumose; inflorescence spiciform or racemose; receptacles epaleaceous;
corollas often pubescent inside, corolla lobes narrowly
lanceolate to linear-oblong; plants with corm-like or
deeply penetrating root systems
1489. Liatris
– Pappus setae barbellate; inflorescence corymbose or
thyrsoid; receptacles paleaceous (paleae often few);
corollas glabrous inside, corolla lobes triangular to
oblong; plants with fibrous roots
4
4. Pappus setae uniseriate; apical anther appendages obtuse to rounded
1491. Trilisia
– Pappus setae ± biseriate; apical anther appendages
distinctly retuse
5
5. Phyllaries 15–40, distinctly imbricate, 3–4-seriate;
achenes densely setuliferous; florets 12–35
1490. Carphephorus
– Phyllaries 5–10, loosely overlapping, 2–3-seriate;
achenes sparsely short-setuliferous, moderately to
densely glandular-punctate; florets 5–10
1492. Litrisa
Genera of Liatrinae
1488. Garberia A. Gray
Garberia A. Gray, Proc. Acad. Nat. Sci. Philadelphia 1879:
379 (1879).
Compositae
Small erect shrubs, glandular-punctate. Leaves
initially rosulate at base, lamina viscid, spathulate
to spathulate-obovate or rounded-obovate, entire,
often slightly retuse. Inflorescence a corymbose
panicle. Phyllaries c. 15, subimbricate, c. 3-seriate,
markedly unequal; receptacle slightly convex,
epaleaceous. Florets 5; corollas pink to purplish,
narrowly funnelform with slightly campanulate
throat, glabrous on both surfaces; lobes 4–5
times as long as wide, longer than throat; inner
surface densely and rather antrorsely shortpapillose, outer surface smooth; anther collars
short-cylindrical; apical anther appendages large,
slightly longer than wide, somewhat retuse at tips;
style arms long-linear, densely short-papillose.
Achenes prismatic, c. 10-ribbed, densely setuliferous; carpopodium lacking; pappus setae c. 60–70,
2–3-seriate, basally barbellate and distally scabrid,
persistent, outer setae somewhat shorter and
narrower, apical cells acute. n = 10. One species,
G. heterophylla (Bartram) Merr. & F. Harper, USA.
1489. Liatris Gaertn. ex Schreber
Liatris Gaertn. ex Schreber, Gen. Pl. 7: 542 (1791), nom.
cons.; Gaiser, Rhodora 48: 165–183, 216–263, 273–326, 331–
382, 393–412 (1946), rev.; Nesom, Sida 21, 3: 1305–1321
(2005), rev.
Erect perennial herbs. Leaf lamina linear, elliptic
or oblanceolate, entire. Inflorescence cymose, usually spiciform or racemiform. Phyllaries c. 20–25,
subimbricate, 3–5-seriate, mostly persistent; receptacle nearly flat, epaleaceous. Florets 3–80; corollas usually purple, sometimes lavender or white,
broadly to narrowly funnelform, usually glandularpunctate on outer surface and rarely with uniseriate hairs, inner surface often with hairs near
insertion of filaments or on lobes; corolla lobes
linear-lanceolate to linear-oblong, c. 3–4 times as
long as wide, densely mamillose or short-papillose
on at least distal part of inner surface, smooth
on outer surface, anther collars broadly cylindrical or slightly broadened below; apical anther appendages usually oblong-ovate and as long as wide
or short and obsolete; style arms narrowly linear
to scarcely broadened distally, flattened, densely
mamillose, often with glands on inner surface or
with septate hairs on upper shaft and abaxially on
lower half of arms. Achenes prismatic, c.10-ribbed,
densely setuliferous; carpopodium lacking; pappus
setae c. 12–40, 1–2-seriate, plumose or markedly
barbellate, persistent, barbellate and fimbriate on
both lateral and outer surfaces, apical cells acute.
531
n = 10, 20, 30. Forty-one species and several hybrids, Canada, Mexico, USA. Five sections are recognized (Nesom 2005). Many species cultivated.
1490. Carphephorus Cass.
Carphephorus Cass., Bull. Soc. Philom. Paris 1816: 198
(1816); Correa & Wilbur, J. Elisha Mitchell Sci. Soc. 85:
79–91 (1969), rev.; Cronquist, Vasc. Fl. SE U.S. 1: i–xvi,
1–261 (1980), rev.
Erect, scapose, perennial herbs. Leaf lamina
linear to linear-lanceolate or oblanceolate, entire.
Inflorescence an open flat-topped corymbose
cyme. Phyllaries c. 15–40, distinctly subimbricate,
c. 3–5-seriate, markedly unequal, mostly or totally
persistent; receptacle slightly convex, usually with
a few paleae. Florets c. 12–35, fragrant; corollas
lavender to purple, funnelform or with narrowly
campanulate throat, usually with glands on outer
surface, or glabrous, glabrous on inner surface;
lobes c. 1.5–2.5 times as long as wide, densely
short-papillose on inner surface, smooth on outer
surface; anther collars short-cylindrical, slightly
constricted above; apical anther appendages
medium-sized to large, ovate-oblong, as long as
wide, distinctly retuse at tip; style narrowly linear,
rather densely high-mamillose or short-papillose.
Achenes prismatic, c. 10-ribbed, moderately
to densely setuliferous and often with glands;
carpopodium lacking; pappus setae c. 35–40,
2–3-seriate, barbellate (sometimes coarsely so),
somewhat unequal, congested, apical cells acute.
n = 10. Four species, USA.
1491. Trilisa (Cass.) Cass.
Trilisa (Cass.) Cass., Dict. Sci. Nat. 26: 228 (1827); Small, Fl.
SE U.S.: 1170–1171 (1903), key.
Erect perennial herbs. Leaves sometimes extremely
aromatic, lamina oblanceolate to obovate or rather
narrowly elliptic, somewhat succulent, entire
or coarsely few-dentate, obtuse to short-acute.
Inflorescence cymose, corymbose or thyrsoid
in form. Phyllaries c. 6–12, distant to weakly
subimbricate, c. 2-seriate, slightly unequal to
subequal, inner not deciduous; receptacle slightly
convex, usually epaleaceous, or with 1 or 2 paleae.
Florets 4–10(–15); corollas pink to purplish,
narrowly funnelform, with numerous glands on
outer surface, glabrous inside; lobes c. 1.25–1.5
times as long as wide, densely short-papillose
on inner surface; anther collar cylindrical; apical
anther appendages large, slightly longer than wide,
532
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
rounded to scarcely retuse at tip; style arms filiform, densely short-papillose. Achenes prismatic,
c. 8-ribbed; carpopodium lacking; pappus setae
usually uniseriate, c. 3–5, persistent, barbellate,
with narrower setae sometimes between and
slightly outside of larger setae, apical cells acute.
n = 10. Two species, USA.
1492. Litrisa Small
Litrisa Small, Bull. Torrey Bot. Club 51: 392 (1924).
style arms short, linear, densely short-papillose.
Achenes prismatic to obpyramidal, 5-ribbed,
glanduliferous, lacking setulae; carpopodium
lacking; pappus usually lacking, rarely a single
hair-like glanduliferous seta. n = 10. One species,
H. floridana A. Gray ex S. Watson, USA.
XXX.10. Subtribe Praxeliinae R.M. King &
H. Rob. (1980).
1493. Hartwrightia A. Gray ex S. Watson
Erect to subscandent, annual to perennial herbs or
subshrubs. Leaves mostly opposite, lamina entire
to serrate, sometimes dissected, ovate or oblong
to linear. Inflorescence laxly cymose to thyrsoidpaniculate, rarely monocephalic on long erect
pedicels, phyllaries imbricate, totally deciduous;
receptacle slightly convex to highly conical, with or
without paleae. Florets 5–65; corollas white, blue,
lavender or purple, peripheral florets sometimes
with enlarged outer lobes; lobes usually longer
than wide, usually densely papillose on inner
surface; anther collar usually 5 times as long as
wide, usually distinctly broadened in lower half;
apical anther appendages usually longer than wide,
sometimes shorter, or obsolete; style base usually
not enlarged, glabrous; arms usually narrowly
linear and densely papillose. Achenes biconvex to
trigonous or prismatic, 2–5-ribbed; carpopodium
usually distinct; pappus setae uniseriate, usually
numerous, capillary, barbellate, sometimes only
5–8, with or without intermixed smaller setae, or
pappus reduced to fringe of short unequal setae,
apical cells usually sharply acute.
Hartwrightia A. Gray ex S. Watson, Proc. Amer. Acad. Arts
23: 764 (1888).
Key to the Genera
Erect, perennial herbs. Leaf lamina somewhat
fleshy, oblanceolate, entire, narrowly acute to
acuminate. Inflorescence a flat-topped corymbose
cyme. Phyllaries 5–10, weakly subimbricate,
2–3-seriate, somewhat unequal, mostly persistent,
with hairs and glands on outer surface; receptacle
slightly convex, usually glabrous or with 1 or 2
paleae. Florets 5–10; corollas purple, narrowly
funnelform, with a few glands on outer surface,
glabrous inside; lobes c. 1.5 times as long as
wide, inner surface mamillose, outer surface with
numerous glands and some hairs; anther collars
short-cylindrical, somewhat narrowed above;
apical anther appendages large, ovate, as long as
wide, retuse at tip; style arms narrowly linear,
densely short-papillose, without glands or hairs.
Achenes prismatic, 8–10-ribbed, setuliferous and
glandular; carpopodium lacking; pappus setae c.
35, barbellate, persistent, c. 2-seriate, apical cells
acute. n = 10. One species, L. carnosa Small, USA.
Erect perennial herbs. Leaf lamina ellipticoblanceolate to elliptical, usually entire, rarely
with a large tooth or small lobe. Inflorescence
an open corymbose often flat-topped cyme.
Phyllaries c. 12–15, essentially distant, mostly
subequal, persistent; receptacle slightly convex,
often with a few phyllaries inside outermost florets.
Florets 7–10; corollas pink, blue or white, broadly
funnelform, with short and rather indistinct
basal tube, with broad and slightly campanulate
throat, glandular punctae numerous on outer
surface; lobes about as long as wide or slightly
longer, densely short-papillose on inner surface,
slightly mamillose outside near margins; anther
collar short-cylindrical; apical anther appendages
oblong, slightly wider than long, truncate or
retuse apically, with median groove adaxially;
1. Achene compressed and with 2 marginal ribs
2
– Achene usually prismatic, usually 5-ribbed
3
2. Pappus of 5 long equal setae; marginal florets zygomorpic with longer outer lobes; ribs densely longsetuliferous
1499. Eitenia
– Pappus of short unequal barbellate setae; florets all
actinomorphic; ribs sparsely short-setuliferous
1496. Eupatoriopsis
3. Pappus setae 5; corolla lobes spreading and appearing
salverform
1498. Praxeliopsis
– Pappus setae numerous, usually long and capillary,
rarely short and unequal; corolla lobes erect or ascending
4
4. Pappus setae short and unequal 1497. Lomatozona
– Pappus setae long and equal or subequal
5
5. Receptacle conical
1494. Praxelis
– Receptacle flat to slightly convex
6
6. Style arms linear; pappus setae scarcely broadened or
tapering; apical anther appendages large and usually
much longer than wide
1495. Chromolaena
Compositae
533
– Style arms broadened towards apices; pappus setae
somewhat broadened at apices; anther appendages
shorter than wide
1500. Osmiopsis
Genera of Praxeliinae
1494. Praxelis Cass.
Praxelis Cass., Dict. Sci. Nat. 43: 261 (1826).
Eupatorium sect. Praxelis (Cass.) Benth. ex Baker (1876).
Erect to decumbent annual or perennial herbs or
subshrubs. Leaves opposite or whorled, lamina
ovate to elliptical or filiform, subentire to sharply
serrate. Capitula solitary on long erect peduncles
to laxly thyrsoid or rather densely corymbose,
usually campanulate. Phyllaries 15–25, 3–4-seriate,
unequal, gradate, outer falling first; receptacle
highly conical, glabrous. Florets 25–30; corollas
white, blue or lavender, narrowly funnelform
or with cylindrical throat and slightly narrower
basal tube, outer surface mostly smooth, with
a few glands; lobes 1.5–3 times as long as wide,
densely long-papillose on inner surface, usually
with some projecting cells on outer surface at
tip; anther collars with enlarged bases, narrowed
above; apical anther appendages slightly longer
than wide to distinctly longer than wide, often
toothed at tip; style base not enlarged; arms
long, narrowly linear, more broadened in distal
half, densely long-papillose. Achenes slightly
to strongly obcompressed, 3–4-ribbed, sparsely
setuliferous; carpopodium distinct, broad, highly
asymmetrical; pappus setae c. 40, persistent, not
or scarcely broadened distally. 2n = 30, 48, 51, c.
80. Sixteen species, South America, one species
adventive in Asia and Australia.
1495. Chromolaena DC.
Fig. 111
Chromolaena DC., Prodr. 5: 133 (1836).
Eupatorium sect. Cylindrocephala DC. (1836).
Eupatorium sect. Osmia (Sch. Bip.) Benth. ex Baker (1876).
Eupatorium sect. Chromolaena (DC.) Benth. ex Baker
(1876).
Erect to somewhat scandent perennial herbs,
subshrubs or shrubs. Leaves usually opposite,
lamina mostly ovate or triangular to elliptical,
sometimes linear, subentire to lobed. Capitula
usually thyrsoid to candelabriform or on laxly to
densely corymbose branches, rarely solitary on
long erect peduncles. Phyllaries 18–65, 4–6-seriate,
markedly unequal, gradate, often with expanded
herbaceous or coloured tips; receptacle flat to
Fig. 111. Compositae-Eupatorieae. Chromolaena morii
(Praxelinae). A Flowering shoot. B Floret. (Drawings by
Margaret Tebbs)
slightly convex, glabrous, sometimes paleaceous.
Florets 6–75; corollas white, blue, lavender or
purple, rather cylindrical with scarcely narrower
base, outer surface smooth below lobes, with few
to many short-stalked glands, often with rather
stiff hairs; lobes slightly to distinctly longer than
wide; usually densely papillose on inner surface,
or smooth (subgenus Osmiella R.M. King & H.
Rob.); anther collars usually broader below, narrowed above, or not broadened below (subgenus
Osmiella); apical anther appendages large, oblong,
c. 1.5 times as long as wide, entire or crenulate at
tip; style base not enlarged; arms narrowly linear
to slightly broadened distally, slightly mamillose
to densely long-papillose. Achenes prismatic,
(3–)5-ribbed, with setulae mostly on ribs; carpopodium distinct, short-cylindrical or narrowed
below; pappus setae c. 40, slender, persistent, not
534
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
or scarcely broadened distally. n = 10, 20, 29, c.
40, 50. Circa 165 species, New World tropics and
subtropics, one species a pantropical weed.
King and Robinson (1987) noted two subgenera
and provided characters in their generic description to separate species of subgenus Osmiella. Concern is growing in many tropical countries over the
alarming spread of the weed Chromolaena odorata (e.g. Zachariades et al. 2002). The taxonomy
and distribution of this species has been well documented by Gautier (1992).
1496. Eupatoriopsis Hieron.
Eupatoriopsis Hieron., Bot. Jahrb. Syst. 18 (Beibl. 43): 46
(1893).
Erect annual herbs. Leaves opposite, lamina ovate
to rather elliptical, serrate, short-acute. Inflorescence laxly cymose. Phyllaries c. 12–14, weakly
subimbricate, c. 1–2-seriate, slightly unequal to
subequal, mostly membranaceous or scarious,
outer somewhat persistent; receptacle very highly
conical to columnar, glabrous. Florets c. 30; corollas lilac, short-funnelform, with very short basal
tube, glabrous on outer surface below lobes; corolla
lobes broadly triangular, about as long as wide,
tips and inner surface densely long-papillose,
outer surface with glands; anther collars expanded
below, narrower above; apical anther appendages
narrower than thecae, subquadrate; style base not
enlarged; arms narrowly linear, slightly broadened
in distal half, densely papillose. Achenes broadly
obcompressed, slightly obcordate, usually with 2
distinctly setuliferous ribs; carpopodium distinct,
asymmetrical; pappus setae uniseriate, c. 18–20,
very short, unequal, persistent. One species, E.
hoffmanniana Hieron., Brazil.
1497. Lomatozona Baker
Lomatozona Baker in Mart., Fl. Brasil. 6, 2: 198 (1876); King
& Robinson, Phytologia 44: 455–462 (1979), key.
Erect perennial herbs or subshrubs, covered with
minute stipitate glands. Leaves opposite, lobed to
deeply dissected, lower surface with very large sessile globular glands. Inflorescence laxly cymose.
Phyllaries c. 25, markedly imbricate, c. 3–4-seriate,
markedly unequal, gradate; receptacle slightly convex, with or without paleae. Florets 10–27; corollas
white or bluish, narrowly funnelform, with occasional glands on outer surface; corolla lobes triangular to rather oblong, up to 1.5 times longer
than wide, densely papillose on inner surface, or
outer lobes of peripheral florets longer; anther collars short, slightly broader below; apical anther
appendages large, oblong-ovate, longer than wide;
style base not enlarged; arms filiform, densely papillose. Achenes prismatic, mostly 5-ribbed, setuliferous; carpopodium lacking or poorly developed; pappus setae uniseriate, 15–22, short, unequal, persistent, barbellate. Four species, Brazil.
1498. Praxeliopsis G.M. Barroso
Praxeliopsis G.M. Barroso, Arch. Jard. Bot. Rio de Janeiro
9: 176 (1949).
Erect annual (or possibly short-lived perennial)
herbs, essentially glabrous. Leaves alternate,
sessile, linear, entire. Inflorescence laxly cymose,
pedicels very long. Phyllaries c. 20–25, imbricate,
c. 3-seriate, markedly unequal; receptacle conical,
glabrous. Florets c. 16; corollas lilac, hypocrateriform, with elongate narrowly funnelform basal
tube, glabrous on both surfaces; throat very short
and spreading; lobes unequal, 1.5–3 times as long
as wide, densely papillose on inner surface; anthers
borne slightly below bases of lobes, exserted above
the spreading lobes; anther collars short and
broad, broader below; apical anther appendages
rudimentary or lacking; style base enlarged; arms
filiform, densely papillose. Achenes prismatic, 5ribbed, with numerous long setulae mostly on ribs;
carpopodium small, distinct, symmetrical; pappus
setae 5, stout, coarsely barbellate, persistent,
narrowed distally. One species, P. mattogrossensis
G.M. Barroso, Brazil.
1499. Eitenia R.M. King & H. Rob.
Eitenia R.M. King & H. Rob., Phytologia 28: 282 (1974).
Erect annual or short-lived perennial herbs.
Leaves opposite, lamina ovate, coarsely serrate to
sublobate, acute. Inflorescence a rather lax broadly
cymose panicle. Phyllaries c. 22–35, imbricate,
3–4-seriate, markedly unequal, lanceolate, outer
falling first; receptacle highly conical, glabrous.
Florets c. 40–50; corollas violet or white, narrowly
funnelform, with long cylindrical throat; lobes
strongly unequal, with outer lobes of peripheral
florets much longer and wider, all lobes oblong,
densely long-papillose on inner surface, longpapillose and with few glands on outer surface;
anther collars distinctly broadened in lower part;
apical anther appendages narrowly oblong, c. 1.5
times as long as wide; style base not enlarged;
arms filiform, densely long-papillose. Achenes
Compositae
obcompressed, usually with 2 or rarely 3 ribs, ribs
densely long-setuliferous; carpopodium distinct,
asymmetrical; pappus uniseriate, of usually 2–8
stout scabrid persistent setae, sometimes interspersed with weaker setae, narrowed distally. Two
species, Brazil.
1500. Osmiopsis R.M. King & H. Rob.
Osmiopsis R.M. King & H. Rob., Phytologia 32: 250 (1975).
Weak scandent shrubs. Leaves opposite, lamina
ovate to lanceolate often with lobed bases, upper
margins entire. Capitula numerous, terminal on
lateral branches, in small corymbose clusters.
Phyllaries c. 20, imbricate, 4–5-seriate, markedly
unequal, gradate, mostly broadly oblong; receptacle flat to slightly convex on distal surface,
glabrous. Florets 18–26; corollas white, funnelform
with a short, broadly cylindrical base, sparsely
glandular on outer surface below lobes; lobes as
long as wide to 1.5 times as long as wide, smooth on
both surfaces, densely glandular on outer surface;
anther collars slightly broader below; apical anther
appendages semicircular, wider than long; style
base not enlarged; style arms narrowly linear with
slightly broadened tips, sharply short-papillose.
Achenes short-prismatic, slightly narrowed below,
5-ribbed, with few glands and short setulae;
carpopodium distinct, short-cylindrical; pappus
uniseriate, setae c. 25–30, stout, persistent, somewhat broadened distally. One species, O. plumerii
(Urban & Ekman) R.M. King & H. Rob., Haiti.
XXX.11. Subtribe Gyptidinae R.M. King &
H. Rob. (1980).
Erect perennial herbs, subshrubs, shrubs or small
trees; leaves opposite to spirally inserted, rarely
rosulate (Bishopiella). Inflorescence terminal
on leafy branches or rarely scapose. Capitula
clustered; phyllaries distant to weakly subimbricate, persistent; receptacle scarcely convex
to strongly conical, paleaceous or epaleaceous,
glabrous or pubescent. Florets usually numerous,
rarely fewer than 10 per capitulum; corolla lobes
often papillose on inner surface; apical anther
appendages very short to longer than wide,
sometimes deeply cleft at apex; style base often not
enlarged and glabrous, sometimes with enlarged
node or pubescent or both; arms usually linear,
usually densely mamillose or papillose. Achenes
usually 5-ribbed or rarely winged; carpopodium
distinct, indistinct or apparently absent; pappus
535
usually of many capillary setae, sometimes very
short, rarely lacking or reduced to 5 fragile setae,
sometimes with enlarged round-tipped apical
cells. The greatest concentration of genera is in
Brazil, especially in Northeast Brazil.
Gyptidinae s.l., currently containing 29 genera, constitute a somewhat problematic subtribe
and is, without doubt, polyphyletic. The only full
account of Gyptidinae s.l. is that of King and Robinson (1987). Hind (1999, 2000) believes that there are
at least three distinct groups of genera, each representing a separate subtribe – the ‘Gyptis group’
(genera 1501–1519), the ‘Agrianthus group’ (genera 1520–1527) and the ‘Litothamnus group’ (genera 1528 & 1529). Some support for such division
of the subtribe is provided by the phytochemical
studies by Bohlmann et al. (1980a, b, 1981a, b, c, d,
1982a, b, 1984a, b).
Key to the Genera
1. Plants rosulate; inflorescence scapiform; leaves fleshy
1527. Bishopiella
– Plants with leafy stems, not rosulate; inflorescence
corymbose, cymose, paniculate, or of solitary capitula,
but never scapiform; leaves coriaceous or herbaceous,
if fleshy, then plants trees
2
2. Pappus of plumose or markedly barbellate setae, or absent and corollas densely pubescent, pubescence obscuring lobes
3
– Pappus barbellate, rarely simple, if absent, then corollas lacking dense pubescence, or pubescence not obscuring lobes
4
3. Corollas densely pubescent on throat and lobes; stems
distinctly ribbed; leaves sessile or with narrow petioles; apical anther appendages rounded
1510. Trichogonia
– Corollas glandular-punctate distally; stems scarcely
striate; leaves with distinctly winged petioles; apical
anther appendages deeply cleft 1511. Trichogoniopsis
4. Plants with leaves inserted in dense spiral; leaves not
usually progressively smaller upwards
5
– Plants with often lax opposite, or occasionally alternate leaves; leaves decreasing markedly in size upwards on plants
13
5. Receptacle paleaceous; leaves scale-like, small, usually
imbricate and with broad cuneate bases
6
– Receptacle epaleaceous; leaves usually medium
to large, membranous or coriaceous, sometimes
fleshy, spreading, usually with narrowed bases or
distinct petioles, or if leaves small and scale-like, then
imbricate
7
6. Phyllaries biseriate with inner shorter than outer, all
deciduous; inflorescence overtopped by upper leaves;
corollas glabrous; leaves lacking obvious secondary
venation; stems glabrous; style arms linear-filiform;
pappus absent
1521. Catolesia
– Phyllaries (2–)3–5-seriate, gradate with inner longer
than outer, persistent; inflorescences overtopping
536
7.
–
8.
–
9.
–
10.
–
11.
–
12.
–
13.
–
14.
–
15.
–
16.
–
17.
–
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
leaves; corollas glandular-punctate or stipitateglandular (sometimes sparsely so); leaves with
obvious prominent secondary venation; stems variously pubescent; style arms clavate; pappus of setae
with pectinate to subplumose margins, rarely absent
(and if so, capitula sessile to very shortly pedicellate)
1520. Agrianthus
Leaves small and scale-like, imbricate
1520. Agrianthus
Leaves medium to large, membranous or coriaceous,
or if small, then linear and closely adpressed to
stem with long eglandular hairs or with long
stipitate-glandular hairs
8
Leaves glabrous
9
Leaves variously pubescent
10
Leaves fleshy; achenes setuliferous, at least below apical callus; phyllaries scarcely longer than achenes,
corollas well exserted above involucre
1429. Morithamnus
Leaves not fleshy; achenes completely glabrous; phyllaries considerably longer than achenes, corollas ± included within involucre
1523. Bahianthus
Leaves narrowly linear and closely adpressed to stem
1522. Arrojadocharis
Leaves narrowly elliptic, ovate to obovate or oblanceolate
11
Achenes markedly winged; leaves stipitate-glandular
and eglandular-pubescent
1524. Semiria
Achenes ribbed; leaves eglandular-pubescent
12
Style shaft pubescent; corollas stipitate-glandular; apical anther appendages shorter than wide
1526. Stylotrichium
Style shaft glabrous; corollas glandular-punctate; apical anther appendages as long as or longer than wide
1525. Lasiolaena
Leaves coriaceous, sometimes fleshy, glabrous, sessile
to subsessile
14
Leaves membranous, thin, glabrous or variously
pubescent, petiolate
15
Leaves elliptical to obovate, glabrous, distinctly fleshy,
usually stoutly petiolate; achenes 5-ribbed; plants large
shrubs or smallish trees of wooded restinga or coastal
dunes; NE Brazil (Bahia)
1528. Litothamnus
Leaves orbicular or ovate, short-pubescent and distinctly glandular-punctate, very short-petiolate; achenes 7–10-ribbed; plants shrubby but never tree-like,
of upland campos; southern and central Brazil
77. Vittetia
Achenes with stipitate or long-attenuate bases
16
Achenes of ± uniform diameter or only slightly narrowed towards base
19
Pappus of short-laciniate or subplumose flattened
setae about 1/2 to 2/3 length of corollas; style base
pubescent
1512. Platypodanthera
Pappus of long capillary barbellate setae at least as
long as corollas; style base glabrous
17
Capitula large, florets 30–100; receptacle conical or
highly convex; achene ribs pale; many species in Brazil,
1 species widespread in South America
1515. Campuloclinium
Capitula small to medium, florets < 50; receptacle flat
or slightly convex; achene ribs concolorous with body;
Bolivia, southern Brazil, Paraguay, Peru
18
18. Inner phyllaries much shorter than florets; pedicels
long; anther cylinder visible and usually partially
exserted from corolla throat; style arms linear; style
base lacking basal node; glandular punctae, when
present, clear or light-coloured; southern Brazil,
Paraguay
1516. Macropodina
– Inner phyllaries equalling or slightly shorter than florets; pedicels short; anther cylinder wholly obscured
and included well within corolla throat; style arms
clavate; style base with basal node; corolla lobes, phyllaries and underside of leaves with conspicuous, often dark, amber-coloured glandular punctae; Bolivia,
Peru
1513. Neocuatrecasia
19. Carpopodium procurrent on base of achene ribs 20
– Carpopodium annular, short-cylindrical, stoppershaped or indistinct or obsolete, but not procurrent
on ribs
22
20. Achenes completely glabrous; corolla lobes pilose; apical anther appendages bilobed to emarginate, shorter
1506. Diacranthera
than wide
– Achenes glandular-punctate or setuliferous; corolla
lobes glandular-punctate; apical anther appendages
entire, rounded to acute, as long as or longer than
wide
21
21. Achenes setuliferous; style base with swollen basal
node, pubescent
1505. Dasycondylus
– Achenes glandular-punctate; style base lacking basal
node, glabrous
1504. Barrosoa
22. Pappus setae with blunt, sometimes inflated apices 23
– Pappus setae with acute apices
25
23. Phyllaries with distinct, pubescent appendages; carpopodium stopper-shaped; Argentina, Bolivia, southern Brazil, Paraguay, Uruguay
1503. Urolepis
– Phyllary apices attenuate or acute but never
appendaged; carpopodium indistinct
24
24. Corollas markedly widening above a short tube; leaves
ascending; Argentina and southern Brazil
1502. Gyptidium
– Corollas narrowly funnelform; leaves spreading; Mexico and USA (Texas)
1517. Conoclinium
25. Achenes glabrous, lacking glandular punctae or eglandular setulae
26
– Achenes setuliferous, sometimes sparsely so, with or
without or glandular punctae
27
26. Corolla lobes glabrous outside; leaves glabrous, usually viscid; phyllaries ± biseriate and conspicuously
shorter than florets; carpopodium essentially annular with ‘crenulate’ upper margin; leaves concolorous;
plants of coastal dunes of Bahia, Brazil 1509. Prolobus
– Corolla lobes pubescent outside; leaves conspicuously
discolorous and densely tomentose beneath; phyllaries 3–4-seriate and slightly shorter than florets; carpopodium stopper-shaped with distinct upper rim;
leaves discolorous; plants of the paramos of the northern Andes (Colombia, Venezuela) 1519. Lourteigia
27. Phyllary apices densely pubescent and appearing expanded; corolla lobes with hairs on outer surface; leaf
bases acute to acuminate; northern Argentina, southern Brazil, Paraguay, Uruguay
1501. Gyptis
– Phyllary apices attenuate or acute, not expanded;
corolla lobes glabrous or scattered glandular-punctate
on outer surface; leaf bases truncate
28
28. Leaf margins crenate; corollas lacking glandular punctae; florets 20–30; carpopodium with setuliferous up-
Compositae
per margin; Brazil, Colombia, Venezuela
1507. Conocliniopsis
– Leaf margins serrate or biserrate; corollas glandularpunctate; florets ≤ 10, and pedicels uniform throughout, or 40–70, with pedicels enlarged and fistulose beneath involucre; upper margin of carpopodium lacking setulae and distinct or indistinct
29
29. Florets ≤ 10; pedicels short, but uniform throughout; carpopodium broadly stopper-shaped with distinct upper margin; Bolivia, Brazil, Paraguay
1508. Bejaranoa
– Florets 40–70; pedicels conspicuous, enlarged/swollen
and fistulose distally/beneath involucre; carpopodium
indistinct and not markedly delimited above; Mexico
and southern USA
1518. Tamaulipa
XXX.11. a. Gyptis Group
1501. Gyptis (Cass.) Cass.
Gyptis (Cass.) Cass., Dict. Sci. Nat. 16: 10 (1820); King &
Robinson, Phytologia 21: 22–25 (1971), key.
Eupatorium L. sect. Gyptis (Cass.) Cabrera (1991).
537
campanulate, smooth on both surfaces; lobes not
or only slightly longer than wide, outer surface
mostly smooth, inner surface slightly to strongly
papillose; anther collars elongate, narrow; apical
anther appendages ovate, longer than wide; style
base not enlarged, glabrous; arms linear, densely
papillose with papillae 1–2 times as long as wide.
Achenes prismatic, 4–5-ribbed, glanduliferous;
carpopodium obsolete; pappus setae uniseriate,
c. 28, persistent, apical cells with rounded and
sometimes dilated tips. Two species, Argentina,
Brazil.
1503. Urolepis (DC.) R.M. King & H. Rob.
Urolepis (DC.) R.M. King & H. Rob., Phytologia 21: 304
(1971).
Eupatorium sect. Urolepis (DC.) Benth. (1876).
Perennial herbs. Leaves opposite, often becoming
alternate above, lamina ovate to bipinnatifid,
serrulate to deeply dissected. Inflorescence subscapose, usually densely corymbose or cymose
above. Phyllaries 2–3-seriate, 16–25, weakly
subimbricate, subequal, with apical pubescence;
receptacle flat, glabrous. Florets 4–26; corollas
narrowly funnelform, white, pink or violet; lobes
about as long as wide or slightly longer, strongly
papillose on both surfaces, with hairs and often
glands on outer surface, margin often with 2-celled
teeth, anther collars short-cylindrical, only slightly
expanded below; apical anther appendages ovate,
twice as long as wide, style base not enlarged,
usually glabrous; arms narrowly linear to filiform,
densely papillose. Achenes prismatic, 5-ribbed,
surfaces densely setuliferous; carpopodium very
short or vestigial; pappus setae uniseriate, c.
20–30, persistent, apical cells subacute or acute.
n = 10. Seven species, Argentina, Brazil, Paraguay,
Uruguay.
Coarse, erect annual herbs or subshrubs. Leaves
opposite, lamina broadly deltoid, dentate or denticulate. Inflorescence a corymbose or subcymose
panicle. Phyllaries c. 50, weakly subimbricate,
3–4-seriate, equal or subequal, inner phyllaries
attenuate into long densely pubescent appendage;
receptacle subglobose, densely short-pubescent.
Florets 100–150; corollas pink, with long narrow
basal tube; throat funnelform below, becoming
cylindrical above, with outer surface of tube and
throat glabrous; lobes slightly longer than wide,
with a few short-stalked glands externally, smooth
on inner and outer surface, extreme tips of lobes
papillose; anther collars slender; apical anther
appendages ovate-triangular, longer than wide;
style base not enlarged, glabrous; arms densely
papillose with very long antrorsely imbricate
papillae. Achenes prismatic, 4–5-ribbed with
occasional short-stalked glands; carpopodium
large; pappus setae uniseriate, c. 20, barbellate, persistent, enlarged distally. n = 10. One
species, U. hecatantha (DC.) R.M. King & H. Rob.,
Argentina, Bolivia, Brazil, Paraguay, Uruguay.
1502. Gyptidium R.M. King & H. Rob.
1504. Barrosoa R.M. King & H. Rob.
Gyptidium R.M. King & H. Rob., Phytologia 23: 310 (1972).
Barrosoa R.M. King & H. Rob., Phytologia 21: 26 (1971).
Erect annual herbs. Leaves opposite, sometimes
alternate above, lamina ovate to lanceolate, crenulate. Inflorescence cymose to subcymose, capitula
sessile to long-pedicellate. Phyllaries 2–3-seriate,
c. 25, distant to weakly subimbricate, subequal,
persistent; receptacle conical, hirsute or with narrow paleae. Florets 50–80; corollas white or pale
lilac, with very narrow basal tube; throat narrowly
Erect perennial herbs. Leaves opposite, sometimes
alternate above, lamina lanceolate to broadly ovate,
serrate to crenulate. Inflorescence densely corymbose. Phyllaries c. 15–25, distant, usually biseriate,
subequal; receptacle conical, glabrous, strongly
foveolate. Florets 20–55; corollas funnelform,
white, pink, blue or purple; throat smooth; lobes
slightly longer than wide, papillose on both
538
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
surfaces, outer surface with hairs and glands;
anther collars short; apical anther appendages
subquadrate to oblong, about as long as wide;
style base not enlarged, glabrous; arms densely
short-papillose. Achenes prismatic, 5-ribbed,
sparsely glanduliferous; carpopodium distinct,
procurrent on to lower part of achene ribs; pappus
setae 25–30, persistent, slender, barbellate, apical
cells acute or subacute. n = 10, c. 30. Nine species,
South America.
enlarged, densely hirsute; style arms linear to
slightly clavate, scarcely mamillose. Achenes prismatic, 5-ribbed, glabrous; carpopodium shortly
procurrent on to ribs of achene; pappus setae
uniseriate, c. 35, persistent, not enlarged at tips,
apical cells acute. Two species, Brazil.
1505. Dasycondylus R.M. King & H. Rob.
Dasycondylus R.M. King & H. Rob., Phytologia 24: 188
(1972); King & Robinson, Phytologia 24: 187–191 (1972),
key.
Erect or spreading subshrubs or shrubs, sometimes subscandent. Leaves opposite, lamina ovate
to oblong, entire to serrate. Inflorescence a corymbose panicle. Phyllaries c. 15–25, subimbricate,
3–4-seriate, unequal, inner somewhat deciduous,
outer often broad; receptacle conical, glabrous to
sparsely pilose, strongly foveolate. Florets 20–60;
corollas white, narrowly funnelform; lobes about
as long to twice as long as wide, inner surface
smooth, outer slightly papillose, glanduliferous
and short-pubescent; anther collars short, slightly
swollen; apical anther appendages ovate, longer
than wide; style base enlarged, densely hirsute;
arms slightly clavate distally, slightly mamillose.
Achenes prismatic, 5-ribbed, sparsely setuliferous
above; carpopodium distinct, procurrent on to
lower part of achene ribs, pappus setae uniseriate, c. 30–40, persistent, barbellate, apical cells
narrowly obtuse to acute. Eight species, Bolivia,
Brazil, Peru.
1506. Diacranthera R.M. King & H. Rob.
Diacranthera R.M. King & H. Rob., Phytologia 24: 192
(1972); King & Robinson, Phytologia 24: 192–194 (1972),
key.
Perennial herbs or subshrubs. Leaves opposite,
lamina ovate to elliptical, crenulate to serrulate.
Inflorescence slightly to strongly cymose with
many branches. Phyllaries c. 25, distant, biseriate,
subequal; receptacle broadly convex, glabrous. Florets 50–65; corollas pinkish, narrowly funnelform;
lobes slightly longer than wide, inner surface
smooth to slightly mamillose, outer with a few
glands, with or without hairs; anther collars short,
narrowly cylindrical; apical anther appendages
bilobed, shorter than wide; style base slightly
Fig. 112. Compositae-Eupatorieae. Conocliniopis prasiifolia
(Gyptidinae). A Flowering shoot. B Floret. (Drawings by
Margaret Tebbs)
Compositae
1507. Conocliniopsis R.M. King & H. Rob. Fig. 112
Conocliniopsis R.M. King & H. Rob., Phytologia 23: 308
(1972).
Erect subshrubs or shrubs. Leaves opposite, few
alternate above, lamina ovate, strongly crenate. Inflorescence a dense corymbose panicle or cyme;
phyllaries 12–16, distant, subequal, c. 3-seriate, persistent; receptacle high-conical, glabrous, strongly
foveolate. Florets 20–30; corollas blue or lavender,
narrowly funnelform, outer surface glanduliferous,
inside glabrous; lobes twice as long as wide, inner
surface mamillose, outer surface somewhat papillose distally; anther collars elongate, narrow; apical
anther appendages ovate, longer than wide; style
base not enlarged, glabrous; arms filiform, scarcely
broader distally, densely papillose. Achenes prismatic, 4–5-ribbed, setuliferous; carpopodium very
prominent with projecting setuliferous upper rim;
pappus setae uniseriate, 30–35, persistent, barbellate, apical cells acute to subacute. n = 10, 30. One
species, C. prasiifolia (DC.) R.M. King & H. Rob.,
Brazil, Colombia, Venezuela.
1508. Bejaranoa R.M. King & H. Rob.
Bejaranoa R.M. King & H. Rob., Phytologia 40: 52 (1978);
King & Robinson, Phytologia 40: 51–53 (1978), key.
Erect subshrubs or shrubs. Leaves alternate
or sometimes opposite, lamina ovate or ovatelanceolate, serrate to doubly serrate. Inflorescence
terminal with densely corymbose branches. Phyllaries 8–15, subimbricate, c. 4-seriate, unequal and
somewhat gradate, persistent; receptacle flat to
minutely conical, glabrous or subglabrous. Florets
4–10; corollas white to pale lavender, narrowly
funnelform, glabrous inside, outside with glands
above; lobes scarcely longer than wide, inner
surface smooth, outer densely glanduliferous and
minutely papillose near tip, lateral margins somewhat thickened; anther collars short-cylindrical;
apical anther appendages oblong, longer than
wide; style base not enlarged, glabrous; arms linear,
slightly minutely mamillose, slightly broadened
distally. Achenes prismatic, 5-ribbed, narrowed
below, setuliferous and glandular; carpopodium
short, with distinct upper rim; pappus setae
uniseriate, c. 30–55, persistent, barbellate, apical
cells acute or sometimes truncate. n = 10. Two
species, Bolivia, Brazil, Paraguay.
1509. Prolobus R.M. King & H. Rob.
Prolobus R.M. King & H. Rob., Phytologia 50: 386 (1982).
539
Erect shrubs. Leaves mostly opposite, becoming
alternate above, lamina ovate, coarsely serrate.
Inflorescence cymose, with ascending branches,
often extra-axillary. Phyllaries c. 12–15, slightly
subimbricate, ± biseriate, subequal, receptacle
flat or slightly convex, glabrous. Florets c. 12–14;
corollas pale violet, glanduliferous above on
outer surface, tube broadly cylindrical; throat
narrowly funnelform; lobes slightly longer than
wide, inner surface mamillose below, apex of outer
surface forming prominent hump, anther collar
cylindrical; apical anther appendages oblong,
scarcely longer than wide; style base not enlarged,
glabrous; arms narrowly linear, short-mamillose.
Achenes prismatic, 5-ribbed, glabrous or with
1–2 setulae above; carpopodium with distinct
crenulate upper margin; pappus setae uniseriate,
25–30, persistent, outer surfaces not flattened
nor smooth, apical cells narrow, subacute. One
species, P. nitidulus (Baker) R.M. King & H. Rob.,
Brazil.
1510. Trichogonia (DC.) Gardner
Trichogonia (DC.) Gardner, London J. Bot. 5: 459 (1846);
Barroso, Arch. Jard. Bot. Rio de Janeiro 11: 7–18 & est. 1–13
(1951), reg. rev.
Erect perennial herbs or subshrubs. Leaves usually
alternate, opposite in some species at least below,
lamina linear to broadly cordate, usually crenulate
to crenate. Inflorescence a lax to dense cymose
or corymbose panicle. Phyllaries 10–25, distant,
± biseriate, subequal to equal; receptacle flat to
slightly convex, glabrous. Florets 10–60; corollas
pink, purple or white, narrowly funnelform, basal
tube sometimes narrow and elongate, limb with
dense pubescence on upper throat and lobes; lobes
wider than long to slightly longer than wide, inner
surface smooth, outer surface densely pubescent
and with few glands; anther collars usually rather
narrowly cylindrical; apical anther appendages
slightly shorter to distinctly longer than wide, with
rounded or retuse tip, style base not enlarged,
glabrous; style arms linear or clubbed at tip,
densely papillose or mamillose except on clubbed
tips. Achenes prismatic, 5-ribbed, setuliferous at
least on ribs; carpopodium a small rim; pappus
setae uniseriate, usually c. 14–30, plumose or
strongly barbellate, persistent, in some species
some or all achenes epappose, apical cells acute.
n = 10. Thirty species, Bolivia, Brazil, Colombia,
Paraguay, Venezuela.
540
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
1511. Trichogoniopsis R.M. King & H. Rob.
Trichogoniopsis R.M. King & H. Rob., Phytologia 24: 180
(1972); Soares Nunes, Bradea 3: 129–138 (1981), part. key.
Erect perennial herbs or subshrubs. Leaves
alternate or opposite, lamina ovate, serrate. Inflorescence a lax cymose or corymbose panicle.
Phyllaries c. 18–20, distant, biseriate, subequal;
receptacle flat or shallowly convex, glabrous. Florets 25–50; corollas white, narrowly funnelform,
outer surface with few or no hairs above; lobes c.
1.5 times as long as wide, inner surface smooth,
outer smooth and glanduliferous; anther collars
narrow; apical anther appendages ovate, shorter
than wide, strongly retuse to bilobed; style base
not enlarged, glabrous; arms long-clavate, slightly
mamillose. Achenes prismatic, 5-ribbed, with
a narrow stipitate base, setuliferous; carpopodium
small; pappus setae uniseriate, c. 30, plumose,
persistent, mostly smooth on outer surface, apical
cells narrow-acute. n = 10. Four species, Brazil.
1512. Platypodanthera R.M. King & H. Rob.
Platypodanthera R.M. King & H. Rob., Phytologia 24: 189
(1972); Hind, Kew Bull. 54: 927–932 (2000), key to subspp.
Erect or ascending, rarely procumbent, annual or
perennial herbs or subshrubs. Leaves usually alternate, lamina ovate to lanceolate, serrate. Inflorescence a lax cymose or subcorymbose panicle with
elongate basal internodes; phyllaries biseriate, c. 35,
distant, mostly subequal; receptacle low-conical,
glabrous. Florets 40–130; corollas pink or lavender,
rarely white, narrowly funnelform, glabrous; lobes
slightly longer than wide, smooth on both surfaces;
anther collars short, greatly expanded below; apical
anther appendages slightly shorter than wide, truncate; style base not enlarged, glabrous, arms longand rather broad-clavate, smooth. Achenes prismatic, 5-ribbed, setuliferous; carpopodium narrowly annuliform; pappus setae uniseriate, 15–20,
congested, rather broad, coarsely barbellate to subplumose, of various lengths, mostly half to twothirds as long as corolla, smooth on outer surface,
apical cells acute. n = 10. One species, P. melissifolia
(DC.) R.M. King & H. Rob., Brazil.
1513. Neocuatrecasia R.M. King & H. Rob.
Neocuatrecasia R.M. King & H. Rob., Phytologia 20: 332
(1970); Robinson, Novon 12: 388–392 (2002), key.
Erect to procumbent perennial herbs. Leaves usually opposite, lamina ovate to deltoid or narrowly
oblong, entire or dentate to deeply lobed. Inflorescence a lax to rather dense corymbose or cymose panicle; phyllaries ± biseriate, c. 10, distant, subequal; receptacle convex, glabrous. Florets 17–50; corollas white, with narrow basal tube,
with abruptly expanded usually elongate and campanulate throat; cells of limb oblong with sinuous lateral walls; lobes longer than wide, inner
surface densely papillose; outer surface smooth
with a few hairs; anther collars short; apical anther appendages ovate, longer than wide; style base
enlarged, hairy; arms long-clavate, densely papillose. Achenes prismatic, 5-ribbed, densely setuliferous; carpopodium short-cylindrical to annuliform; pappus setae uniseriate, c. 20–30, usually
persistent and contiguous, apical cells acute. Twelve
species, Bolivia, Peru.
1514. Vittetia R.M. King & H. Rob.
Vittetia R.M. King & H. Rob., Phytologia 29: 122 (1974);
King & Robinson, Phytologia 49: 281–283 (1981), rev.
Eupatorium L. sect. Macropodina (R.M. King & H. Rob.)
Cabrera (1991), p.p.
Erect shrubs or subshrubs. Leaves opposite or
alternate, subsessile, lamina orbicular to broadly
ovate or oblong, entire to crenate-serrate. Inflorescence an ascending multi-branched corymbose
panicle. Phyllaries ± biseriate, c. 10–15, distant,
subequal, receptacle flat, glabrous to minutely
puberulous. Florets 10–12; corollas white to pink,
narrowly funnelform or with narrow basal tube
and rather campanulate limb; lobes slightly longer
than wide, smooth on both surfaces, with glands
on outer surface; anther collar short-cylindrical;
apical anther appendages ovate, as long as or
slightly longer than wide; style base not enlarged,
glabrous; arms narrowly to broadly linear or narrowly clavate, densely sharply papillose. Achenes
prismatic, 7–10-ribbed, glanduliferous and with
non-glandular hairs; carpopodium small; pappus
setae uniseriate, 30–65, persistent, apical cells
sharply acute. n = 10. Two species, Brazil.
1515. Campuloclinium DC.
Campuloclinium DC., Prodr. 5: 136 (1836).
Eupatorium sect. Campuloclinium (DC.) Benth. (1876).
Erect coarse herbs or subshrubs. Leaves opposite
or alternate, lamina ovate to narrowly oblong.
Inflorescences corymbose, capitula few to many,
moderate-sized or often large. Phyllaries 2–3seriate, c. 15–30, distant to weakly subimbricate,
Compositae
subequal; receptacle highly rounded to conical,
with small raised scars, glabrous. Florets 30–100;
corollas pink, lavender or purple, narrowly funnelform, basal tube somewhat constricted above
nectary; lobes usually slightly wider than long,
with mamillose or papillose cells on inner surface,
slightly to strong1y papillose and with glands and
often hairs outside; anther collar short-cylindrical;
apical anther appendages oblong, about as long as
wide to c. 1.5 times wider than long; style base not
to distinctly enlarged, with few to many hairs or
glabrous; arms broadly linear, flat, slightly mamillose to papillose. Achenes elongate, prismatic,
with 5 strongly setuliferous ribs and few to many
glands; carpopodium greatly enlarged; pappus
setae uniseriate, c. 25–40, persistent, coarsely
barbellate, usually elongate or sometimes short,
apical cells subacute to acute. n = 10. Fourteen
species, Argentina, Bolivia, Brazil, Colombia,
Guatemala, Honduras, Mexico, Paraguay.
1516. Macropodina R.M. King & H. Rob.
Macropodina R.M. King & H. Rob., Phytologia 24: 173
(1972).
Eupatorium L. sect. Macropodina (R.M. King & H. Rob.)
Cabrera (1991).
541
to bipinnatifid. Inflorescence laxly cymose below, with densely cymose branches. Phyllaries
2–3-seriate, c. 25, distant, mostly subequal,
lanceolate; receptacle highly conical, glabrous or
rarely pubescent. Florets 50–70; corollas blue or
white, narrowly funnelform, with glands on outer
surface; basal tube not constricted above nectary;
lobes slightly longer than wide, inner surface with
short bulging cells, mamillose to short-papillose,
outer surface papillose in distal half, anther
collars cylindrical, often narrow; apical anther
appendages ovate to ± quadrate, about as long
as wide; style base not enlarged, glabrous; arms
narrowly linear to filiform, slightly broadened
distally, densely papillose. Achenes prismatic,
5-ribbed, glabrous or with a few scattered glands,
rarely setuliferous above; carpopodium usually
obsolete, rarely distinct and asymmetrical; pappus
setae uniseriate, c. 30, barbellate, persistent, often
with slightly to distinctly enlarged tips, apical cells
obtuse to rounded. n = 10. Four species, Mexico,
USA (south-western and eastern Texas).
1518. Tamaulipa R.M. King & H. Rob.
Tamaulipa R.M. King & H. Rob., Phytologia 22: 154 (1971).
Erect subshrubs or shrubs. Leaves usually opposite,
becoming alternate above, lamina ovate, serrulate.
Inflorescence a lax cyme. Phyllaries 3–4-seriate, c.
18–40, weakly subimbricate, narrowly elliptical to
linear, outer distinctly shorter; receptacle flat to
slightly convex, glabrous. Florets (20–)25–30(–50);
corollas pale blue, narrowly funnelform, basal
tubes elongate; lobes c. 3 times as long as wide,
smooth on both surfaces, outer surface with small
glands; anther collar cylindrical; apical anther
appendages ± quadrate, slightly longer than wide;
style base not or only very slightly enlarged,
sparsely to densely hirsute, arms linear, smooth
to slightly mamillose. Achenes prismatic, 5–6 mm
long with long basal stipe, 5-ribbed, sometimes
minutely spiculiferous on ribs; carpopodium
somewhat broader than stipe of achene; pappus
setae uniseriate, c. 25–30, barbellate, persistent,
apical cells subacute to acute. Three species, Brazil,
Paraguay.
Erect woody shrubs. Leaves opposite, lamina
deltoid, subserrate to dentate. Inflorescences
terminal, corymbose; peduncles usually short,
slightly broadened and fistulose distally. Phyllaries
2–3-seriate, 30–35, distant to slightly subimbricate,
outer unequal, inner subequal; receptacle convex
to low-conical, glabrous. Florets 40–70; corollas
pale blue, narrowly funnelform, mostly glabrous
on outer surface; lobes about as long as wide,
smooth on both surfaces, with glands on outer
surface; anther collar narrowly cylindrical; apical
anther appendages oblong to ovate, slightly longer
than wide; style base not enlarged, glabrous; arms
broadly linear, smooth to slightly mamillose.
Achenes prismatic, 5–6-ribbed, setuliferous;
carpopodium sometimes indistinct, very narrow,
somewhat confluent with broadened bases of
ribs; pappus setae uniseriate, c. 35, barbellate,
persistent, capillary, apical cells acute to subacute.
n = 10. One species, T. azurea (DC.) R.M. King &
H. Rob., Mexico, USA (Texas).
1517. Conoclinium DC.
1519. Lourteigia R.M. King & H. Rob.
Conoclinium DC., Prodr 5: 135 (1836).
Lourteigia R.M. King & H. Rob., Phytologia 21: 28 (1971).
Erect, rhizomatous, perennial herbs. Leaves
opposite, lamina ovate to deltoid-ovate, crenate
Small to medium-sized subshrubs or shrubs,
sometimes procumbent. Leaves opposite, lamina
542
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
ovate to narrowly elliptical, crenulate to serrate,
lower surface often white-tomentose. Inflorescence terminal, densely corymbose. Phyllaries
3–4-seriate, c. 20–25, weakly to moderately subimbricate, unequal; receptacle convex, rarely slightly
conical, glabrous or with minute hairs. Florets
20(–40); corollas lilac, blue, purple or greenish
white, funnelform, tube narrowed above nectary;
lobes slightly longer than wide, smooth on most
of both surfaces; papillose near margins, outer
surface densely puberulous and with few to many
glandular punctae; anther collar narrowly cylindrical; apical anther appendages ovate to oblong,
slightly longer than wide; style base not enlarged,
glabrous; arms long-subclavate, densely papillose.
Achenes prismatic, 5-ribbed, strongly constricted
above, narrowed below, sparsely setuliferous to
subglabrous; carpopodium prominent, usually
asymmetrical; pappus setae uniseriate, c. 30,
inserted on an easily detached callus, slender,
barbellate, persistent on callus, apical cells acute.
n = 10. Circa 11 species, Colombia, Venezuela.
XXX.11. b. Agrianthus Group
1520. Agrianthus Mart. ex DC.
Agrianthus Mart. ex DC., Prodr. 5: 125 (1836); Mattfeld,
Notizbl. Bot. Gart. Berlin-Dahlem 8: 428–451 (1923), rev.;
Hind, Kew Bull. 245–277 (1993), key.
Erect many-branched shrubs. Leaves usually
densely spirally inserted, lamina elliptical to
oblong-lanceolate or subulate. Inflorescence
terminal on branches, abrupt, a dense cluster
of sessile or subsessile capitula. Phyllaries c.
18–40, distant to weakly subimbricate, c. 3-seriate,
subequal, lanceolate; receptacle convex to conical, glabrous, sometimes with scattered narrow
paleae. Florets 20–45; corollas usually pink or
white, rarely purplish, narrowly funnelform,
with resin ducts along veins in throat; lobes
slightly longer than wide, mamillose to slightly
papillose on inner surface, with slightly to strongly
papillose apical cap on outer surface; anther
collars prominent, cylindrical to flattened; apical
anther appendages usually longer than wide,
apices truncate or obtuse and sometimes retuse;
style base not enlarged, glabrous, shaft glabrous,
rarely glandular-punctate; arms narrowly clavate,
strongly mamillose. Achenes prismatic, 5-ribbed,
somewhat curved, setuliferous and glandular;
carpopodium annuliform, usually procurrent on
to bases of ribs; pappus rarely absent or rapidly
caducous, usually uniseriate, with 20–40 narrow
sometimes subplumose segments, flattened on
outer surface, apical cells mostly acute. Eight
species, Brazil.
1521. Catolesia D.J.N. Hind
Catolesia D.J.N. Hind, Kew Bull. 55: 942 (2000).
Poorly branched shrubs. Leaves densely spiralled, sessile, longest surrounding inflorescences
and equalling or overtopping capitula, entire,
slightly involute, apices acute, hooked upwards.
Inflorescence terminal, corymbose; involucre
campanulate, phyllaries biseriate, distant, outer
persistent, inner deciduous or easily falling; receptacle convex, smooth, glabrous, paleaceous, paleae
scattered and often indistinguishable from inner
phyllaries. Corolla pink; tube glabrous, expanding
gradually from base to lobes; lobes narrowly
triangular, apices swollen inside and curved inwards, surfaces mammillose outside; apical anther
appendages triangular, less than twice as long as
wide, apices rounded, basal anther appendages
rounded; anther collars prominent and distinctly
wider than filaments; style base lacking node,
glabrous, shaft glabrous, arms linear-filiform to
very slightly clavate, short-papillose. Achenes
5-ribbed, sparsely setuliferous towards pale apical
callus; carpopodium annular, slightly procurrent
on to base of ribs; pappus absent, or rarely minute
abortive setae initials on apical callus. One species,
C. mentiens D.J.N. Hind, Brazil.
1522. Arrojadocharis Mattf.
Arrojadocharis Mattf., Notizbl. Bot. Gart. Berlin-Dahlem
10: 1053 (1930); King & Robinson, Phytologia 44: 463–465
(1979), rev.
Annual or short-lived perennial herbs or shrubs.
Leaves spirally inserted, sessile; lamina linear. Inflorescence of solitary capitula or lax corymbs terminating leafy branches. Phyllaries c. 20, distant,
2–3-seriate, subequal, narrowly lanceolate, receptacle highly conical, paleaceous in lower part. Florets c. 50–60; corollas pink, funnelform, with small
glands on outer surface; lobes slightly longer than
wide, mamillose on inner surface, papillose on
most of outer surface; anther collars cylindrical;
apical anther appendages mostly half to two-thirds
as long as wide, style base not enlarged, glabrous,
shaft glabrous, arms clavate, mamillose. Achenes
prismatic, 5-ribbed, somewhat curved, setuliferous
and with occasional glands; carpopodium cylin-
Compositae
drical or annuliform; pappus variable, absent or
of c. 25 short or long setae, apical cells acute. Two
species, Brazil.
1523. Bahianthus R.M. King & H. Rob.
Fig. 113
Bahianthus R.M. King & H. Rob., Phytologia 23: 312 (1972).
Erect shrubs or small trees, viscid. Leaves densely
spirally inserted, with distinct narrow petioles,
lamina obovate, remotely serrate, obtuse or truncate. Inflorescence a corymbose panicle. Capitula
hemispherical; phyllaries 18–20, ± subimbricate,
c. 3-seriate, somewhat unequal to subequal, linear
543
to lanceolate; receptacle flat to slightly convex,
glabrous. Florets 15–22; corollas pink or white,
narrowly funnelform to subcylindrical, tubes
scarcely narrower than throat, resin ducts narrow
and solitary along veins of throat; lobes as long as
wide or slightly longer, glandular outside, papillose
towards tip, mamillose on inner surface; apical
anther appendages oblong-ovate, about as long
as wide; style base not enlarged, glabrous, arms
filiform with slightly clavate tips, densely patently
papillose. Achenes prismatic, 4–5-ribbed, sparsely
glandular; carpopodium enlarged, procurrent on
lower part of achene ribs; pappus setae uniseriate,
c. 30 persistent, shortly ciliate-dentate, flattened
on outer surface, apical cells narrowly rounded.
n = 10. One species, B. viscosus (Spreng.) R.M.
King & H. Rob., Brazil.
1524. Semiria D.J.N. Hind
Fig. 114
Semiria D.J.N. Hind, Kew Bull. 54: 425 (1999).
Moderately branched small tree. Leaves sessile,
spirally inserted. Inflorescence terminal, of few
to several capitula; involucres campanulate;
phyllaries biseriate, subimbricate or eximbricate; receptacle convex, glabrous, epaleaceous.
Florets c. 40, hermaphrodite; corollas pink,
corolla tube cylindrical narrowing slightly towards base, glandular-punctate throughout and
stipitate-glandular towards base; lobes mamillose inside, glandular-punctate outside; apical
anther appendages obtuse to rotund, truncate or
emarginate, basal appendages rounded or absent;
anther collar markedly dilated; style base lacking
basal node but enlarged, glabrous; shaft glabrous,
arms linear, short-papillose. Achenes 5-winged,
sparsely glandular-punctate, rarely sparsely
stipitate-glandular, wings with lighter-coloured
margins, glabrous; carpopodium ± annuliform,
upper part procurrent on wing bases; pappus
usually of 5 uniseriate subequal ascending to
spreading setae, rarely absent, setae capillary,
fragile, sometimes caducous, usually smooth,
rarely sparsely barbellate towards apices, apices
acute. One species, S. viscosa D.J.N. Hind, Brazil.
1525. Lasiolaena R.M. King & H. Rob.
Fig. 115
Lasiolaena R.M. King & H. Rob., Phytologia 24: 185 (1972);
Hind, Kew Bull. 54: 915–925 (2000), rev.
Fig. 113. Compositae-Eupatorieae. Bahianthus viscosus
(Gyptidinae). A Flowering shoot. B Floret. (Drawings by
Margaret Tebbs)
Erect shrubs. Leaves inserted in a dense spiral,
short-petiolate, lamina narrowly to broadly obovate, serrulate above, obtuse or shortly acute.
544
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Fig. 114. Compositae-Eupatorieae. Semiria viscosa (Gyptidinae). A Flowering shoot. B Floret. C Achene. D Achene
cross-section. (Drawings by Margaret Tebbs)
Inflorescence densely corymbose on tips of leafy
branches. Phyllaries c. 20, weakly subimbricate,
2–3-seriate, mostly subequal, oblong-lanceolate,
receptacle distinctly conical, pilose with a few
hairs, epaleaceous. Florets 18–45; corollas narrowly funnelform, with scattered small glands on
outer surface; lobes slightly to distinctly papillose
on both surfaces, with or without hairs on outer
surface; anther collars cylindrical to slightly
flattened; apical anther appendages two-thirds
as long as wide to slightly longer than wide,
apices truncate to obtuse, sometimes emarginate;
style base not enlarged, glabrous; shaft glabrous;
arms linear, slightly mamillose to short-papillose.
Achenes prismatic, 5-ribbed, bearing setulae and
glands; carpopodium annuliform; pappus setae
uniseriate, 20–40, persistent, congested, setae
flattened throughout with barbellate to almost
laciniate margins, apical cells sometimes with
rounded tips. Six species, Brazil.
Fig. 115. Compositae-Eupatorieae. Lasiolaena morii (Gyptidinae). A Flowering shoot. B Floret. (Drawings by Margaret Tebbs)
1526. Stylotrichium Mattf.
Fig. 116
Stylotrichium Mattf. in Pilger, Notizbl. Bot. Gart. BerlinDahlem 8: 436 (1923); Bautista et al., Brittonia 52: 213–217
(2000), key.
Erect shrubs. Leaves densely spirally inserted,
lamina narrowly obovate to orbicular, with prominent reticulate venation beneath. Inflorescence
abruptly corymbose to subumbellate on tips
of leafy branches; phyllaries c. 25, ± biseriate,
distant, subequal, oblanceolate; receptacle distinctly conical, glabrous, epaleaceous. Florets c.
25–50; corollas white, short-funnelform, with
stalked glands below lobes, outer surfaces of lobes
glandular-punctate; lobes about as long as wide,
mamillose on inner surface, papillose on most of
outer surface; anther collars cylindrical; anther
thecae reddish in most species; apical anther
appendages about half as long as wide, truncate or
Compositae
545
5-ribbed, with setulae and occasional glands;
carpopodium cylindrical or annuliform; pappus
setae absent or uniseriate, 20–25, persistent, short,
laterally densely fringed, apical cells often blunt.
Five species, Brazil.
1527. Bishopiella R.M. King & H. Rob.
Bishopiella R.M. King & H. Rob., Phytologia 48: 418 (1981).
Acaulescent, scapose, annual or short-lived
perennial herbs or subshrubs. Leaves forming
rosette, lamina fleshy, oblanceolate, entire, narrowly obtuse. Inflorescence scapiform, a small,
few-branched cyme. Phyllaries ± biseriate, c. 20,
distant, equal or subequal, linear-lanceolate; receptacle highly conical, glabrous. Florets c. 40–50;
corollas white, shortly funnelform from a broad,
tapering, scarcely narrowed base, outer surface
with a few short-stalked glands; lobes as long as
wide, strongly mamillose on inner surface, outer
surface papillose near margins and apex, anther
collar short-cylindrical; apical anther appendages
about half as long as wide, broadly rounded to
truncate at tip; style base not enlarged, glabrous;
arms broadly linear to strap-shaped, slightly
broader distally, mamillose, essentially smooth at
tip. Achenes short-prismatic, 5-ribbed, with many
long setulae; carpopodium narrowly annuliform;
pappus setae uniseriate, c. 35, persistent, somewhat flattened and margins coarsely barbellate to
almost subplumose, apical cells sharply acute. One
species, B. elegans R.M. King & H. Rob., Brazil.
XXX.11. c. Litothamnus Group
1528. Litothamnus R.M. King & H. Rob.
Litothamnus R.M. King & H. Rob., Phytologia 44: 80 (1979);
Holmes, Phytologia 81: 385–390 (1997), rev.
Fig. 116. Compositae-Eupatorieae. Stylotrichium rotundifolium (Gyptidinae). A Flowering shoot. B Achene. (Drawings by Margaret Tebbs)
emarginate at apex; style base not enlarged, glabrous; shaft and bases of arms densely pubescent
with hairs and glands; arms with clavate tips,
densely papillose below tips. Achenes prismatic,
Erect glabrous shrubs or small trees with subfleshy stems and leaves. Leaves opposite, lamina
elliptical to slightly obovate, entire. Inflorescences
corymbose, with opposite branches. Phyllaries
12–15, distant, ± biseriate, subequal, often with
reddish tips; receptacle flat, glabrous. Florets
(5–)15–25; corollas white, narrowly funnelform,
sparsely glandular-puberulous on outer surface;
lobes slightly longer than wide, inner surface
mamillose; outer surface papillose; anther collar short-cylindrical; apical anther appendages
oblong, slightly longer than wide; style base
not enlarged, glabrous; arms linear, densely and
patently papillose. Achenes prismatic, 5-ribbed,
546
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
glabrous or with few minute gland-tipped hairs
above; carpopodium annuliform; pappus setae
± biseriate, c. 30, persistent, densely barbellate,
congested, apical cells sharply acute. n = 10. Two
species, Brazil.
1529. Morithamnus R.M. King, H. Rob. &
G.M. Barroso
Morithamnus R.M. King, H. Rob. & G.M. Barroso, Phytologia 44: 452 (1979).
Erect shrubs or trees, with candelabra-like
branching, viscid. Leaves opposite or alternate,
lamina obovate to oblanceolate, entire (very rarely
crenulate), obtuse or acute. Inflorescence abruptly
terminal on branches; phyllaries c. 35, distant, biseriate, herbaceous, linear to narrowly lanceolate,
with narrow apices, inner persistent; receptacle
flat to slightly convex, glabrous. Florets c. 25–100;
corollas pink or white, narrowly funnelform;
tubes broad, gradually broadened into cylindrical
throat; resin ducts of throat paired along veins;
lobes ovate-oblong, smooth inside, outside with
glands and sometimes minutely crested; anther
collar short-cylindrical; apical anther appendages
oblong-ovate; style base not enlarged, glabrous;
arms filiform, densely papillose. Achene prismatic,
5-ribbed, moderately to densely setuliferous on
upper part of ribs; carpopodium short-cylindrical;
pappus setae uniseriate, 20–25, capillary or irregularly shortened, persistent, outer surfaces flattened
at base, margins barbellate, apical cells sharply
acute. Two species, Brazil.
XXX.12. Subtribe Disynaphiinae R.M. King &
H. Rob. (1978).
Perennial herbs, more usually shrubs or small
trees; leaves opposite or spirally inserted, lamina
usually entire or serrate to serrulate, rarely
pinnately or bipinnately lobed. Inflorescences terminal, usually densely corymbose or paniculate.
Capitula mostly short-pedicellate; involucres narrowly cylindrical; phyllaries subimbricate, most or
all usually persistent; receptacle scarcely convex,
glabrous, epaleaceous. Florets 5; corolla lobes
smooth or nearly smooth on inner surface; style
base glabrous, lacking basal node; arms linear,
distinctly and usually densely papillose. Achenes
5-angled, body mostly glabrous, sometimes angles
sparsely setuliferous; carpopodium obsolete to
annuliform or short-cylindrical; pappus uniseriate, usually of many straight capillary setae, very
rarely few and hooked, sometimes with enlarged
round-tipped apical cells.
Key to the Genera
1. Leaves pinnately or bipinnately lobed; style arm appendages with long narrow papillae
1530. Acanthostyles
– Leaves simple; style arm appendages short-papillose
2
2. Inflorescences densely paniculate 1531. Raulinoreitzia
– Inflorescences densely corymbose
3
3. Outer phyllaries as long as inner (and often dark
coloured)
1534. Grazielia
– Phyllaries in gradate series
4
4. Phyllaries variously pubescent; leaves usually densely
spirally inserted
1532. Disynaphia
– Phyllaries glabrous
5
5. Leaf venation prominent; pedicels glabrous; plants
usually viscid/viscous; lamina ovate-elliptic, broadly
lanceolate or obovate
1535. Symphyopappus
– Leaf venation, especially midrib, insculpate; pedicels
pubescent; plants never viscid; lamina narrowlanceolate or linear
1533. Campovassouria
Genera of Disynaphiinae
1530. Acanthostyles R.M. King & H. Rob.
Acanthostyles R.M. King & H. Rob., Phytologia 22: 111
(1971).
Erect shrubs. Leaves opposite, lamina narrowly
lobed to pinnately dissected. Inflorescence a long
pyramidal panicle. Phyllaries c. 25, c. 4-seriate,
with glands on outer phyllaries, otherwise
glabrous, receptacle slightly convex, glabrous.
Corollas narrowly funnelform, purple-lilac; lobes
c. 1.5 times as long as wide, with inner surface
smooth, outer surface glanduliferous; anther collar
elongate, cylindrical; apical anther appendages
oblong to triangular, longer than wide; style arms
long-linear, with stigmatic papillae elongate and
forming a brush. Achenes prismatic, 4–5-ribbed,
setuliferous; carpopodium indistinct or minutely
annuliform; pappus setae uniseriate, c. 30–40, persistent, densely divaricately barbellate on margins
and outer surface, not enlarged at apex, apical cells
acute. n = 10. One species, A. buniifolius (Hook.
& Arn.) R.M. King & H. Rob., Argentina, Bolivia,
Brazil, Paraguay, Uruguay.
1531. Raulinoreitzia R.M. King & H. Rob.
Raulinoreitzia R.M. King & H. Rob., Phytologia 22: 113
(1971).
Eupatorium L. sect. Raulinoreitzia (R.M. King & H. Rob.)
Cabrera (1991).
Compositae
547
Erect shrubs. Leaves opposite, lamina elliptical
to linear, serrulate. Inflorescence a pyramidal,
often pendulous, panicle. Phyllaries c. 4-seriate,
c. 15–20, markedly unequal, glabrous; receptacle
slightly convex, glabrous. Corolla white, narrowly
funnelform; lobes about as long as wide to longer
than wide, smooth on inner surface; anther collar
narrowly cylindrical; apical anther appendages
triangular, slightly longer than wide; style arms
linear, short-papillose or highly mamillose.
Achenes prismatic, 4–5-ribbed, mostly glabrous;
carpopodium short-cylindrical to stopper-shaped;
pappus setae uniseriate, c. 30, persistent, scabrid
mostly on margins, apices broadened with inflated round-tipped apical cells. n = 10. Three
species, Brazil, Bolivia, Peru, Argentina, Paraguay,
Uruguay.
1532. Disynaphia Hook. & Arn. ex DC.
Fig. 117
Disynaphia Hook. & Arn. ex DC., Prodr. 7: 267 (1838).
Eupatorium sect. Dysinaphia [sic!] [= Disynaphia] (Hook.
& Arn. ex DC.) Cabrera (1991).
Erect shrubs or subshrubs. Leaves alternate, usually
densely spirally inserted, lamina linear to oblong or
oblanceolate, entire to minutely serrulate. Inflorescence corymbose-paniculate. Phyllaries c. 11–15,
2–3(–4)-seriate, unequal, pubescent on outer surface, receptacle slightly convex or flat, glabrous to
slightly pubescent. Corollas broadly tubular below,
slightly broadening above, purple, pink or white;
lobes as long as wide or longer, smooth to slightly
mammillose on inner surface; anther collar short;
anther base hastate, in a few species with bases as
long as collar; apical anther appendages large, as
long as wide, often notched at tip. Achenes prismatic, 4–5-ribbed, glabrous to slightly glanduliferous, in a few species setuliferous; carpopodium
indistinct; pappus setae uniseriate, c. 35, persistent on callus, often falling as unit with callus, apical cells acute. n = 10. Sixteen species, Argentina,
Brazil, Paraguay, Uruguay.
1533. Campovassouria R.M. King & H. Rob.
Campovassouria R.M. King & H. Rob., Phytologia 22: 121
(1971).
Eupatorium L. sect. Campovassouria (R.M. King & H. Rob.)
Cabrera (1991).
Erect shrubs or subshrubs. Leaves opposite to
alternate, usually closely spaced, lamina narrowly
lanceolate to narrowly oblong or linear, entire
to serrulate. Inflorescence densely corymbose-
Fig. 117. Compositae-Eupatorieae. Disynaphia praeficta
(Disynaphiinae). A Flowering shoot. B Floret. (Drawings
by Margaret Tebbs)
paniculate. Involucre cylindrical, phyllaries c. 12,
3–4-seriate, unequal and gradate, phyllaries eventually deciduous; receptacle flat, glabrous. Corollas
narrowly funnelform, lavender to purple; lobes as
long as wide, smooth on inner surface; anther collar cylindrical; anther thecae with short rounded
bases; apical anther appendages broadly ovate to
oblong, style arms narrowly linear, with dense,
short, acute papillae. Achenes prismatic, 5-ribbed,
slightly glanduliferous; carpopodium distinct,
cylindrical; pappus setae uniseriate, c. 30–35,
congested, persistent on callus, sometimes falling
as unit with callus, apical cells acute. n = 10. One
species, C. cruciata (Vell.) R.M. King & H. Rob.,
Argentina, Brazil, Bolivia, Paraguay, Uruguay.
548
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
1534. Grazielia R.M. King & H. Rob.
Grazielia R.M. King & H. Rob., Phytologia 23, 3: 305 (1972).
Eupatorium section Dimorpholepis G.M. Barroso (1950).
Dimorpholepis (G.M. Barroso) R.M. King & H. Rob. (1971),
non Dimorpholepis A. Gray (1852).
Coarse, mostly erect, herbs, subshrubs or shrubs.
Leaves opposite, lamina ovate to lanceolate,
sometimes pinnately to bipinnately dissected into
narrow segments, often serrate. Inflorescence
densely corymbose. Phyllaries c. 15, 3–4-seriate;
unequal, with strongly differentiated, darker, linear
basal bracts in distinct outer series; receptacle flat
or slightly convex, glabrous. Corollas with broadly
tubular bases, slightly broadening above, white,
rose, lilac or purple; lobes longer than wide, mostly
smooth on inner surface; anther collar short,
cylindrical; anther thecae with short-pointed
bases; apical anther appendages longer than wide;
style base not enlarged, glabrous: style arms
linear, not or scarcely broadened distally, densely
short-papillose. Achenes prismatic, 4–5-ribbed,
often with a few short-stalked glands and short setulae; carpopodium obsolete or short-cylindrical,
procurrent upwards on lower part of achene
ribs; pappus setae c. 20–30, congested, barbellate,
persistent, tapering to a sharp point, apical cells
acute. n = 10. Eleven species, Argentina, Brazil,
Paraguay, Uruguay.
1535. Symphyopappus Turcz.
Fig. 118
Symphyopappus Turcz., Bull. Soc. Imp. Naturalistes Moscou
21: 583 (1848).
Eupatorium L. sect. Symphyopappus (Turcz.) Cabrera
(1991).
Erect shrubs or small trees, often viscid. Leaves
opposite, sometimes becoming alternate near
inflorescence, lamina ovate-lanceolate, serrate.
Inflorescence rather densely corymbose. Phyllaries 3–4(–5)-seriate, c. 15, inner unequal and
gradate, substramineous, usually mostly persistent; receptacle flat or slightly convex, glabrous or
with numerous stiff hairs. Corollas with broadly
tubular base, slightly broadening above, rarely
narrowly funnelform throughout, white, rose
or purple-pink; lobes slightly longer than wide
to twice as long as wide, inner surface smooth,
apices papillose outside and often sparsely
glandular, outer surface otherwise glabrous;
anther collar short, cylindrical; bases of anther
thecae pointed; apical anther appendages large,
triangular, twice as long as wide; style arms
Fig. 118. Compositae-Eupatorieae. Symphyopappus decussatus (Disynaphiinae). A Flowering shoot. B Floret. (Drawings by Margaret Tebbs)
linear, covered with crowded, short, erect papillae.
Achenes short-prismatic, 4–5-ribbed, glabrous or
sparsely short stipitate-glandular; carpopodium
short-cylindrical, procurrent on lower part of ribs;
pappus setae biseriate, c. 40, congested, persistent
on callus, often falling as unit with callus, setae
tapering to sharp points, apical cells acute to
obtuse, rarely with uncinate apices. n = 10. Twelve
species, Brazil.
XXX.13. Subtribe Ayapaninae R.M. King & H.
Rob. (1980).
Erect or repent annual or perennial herbs or
subshrubs, rarely vines, leaves opposite, sometimes becoming alternate above, rarely alternate
throughout. Inflorescence terminal on leafy
branches. Capitula clustered; phyllaries usually
distinctly subimbricate, gradate, persistent; receptacle slightly convex, rarely columnar, usually
glabrous, more rarely paleaceous or with hairs.
Florets 3–300; corolla lobes usually smooth; apical
anther appendage usually as long as wide, rarely
reduced or lacking; style base with enlarged
Compositae
node, with or without hairs; style arms linear
to tapering, rarely broadened apically, smooth
to strongly papillose. Achenes prismatic with 5
ribs; carpopodium distinct, strongly demarcated;
pappus rarely absent, usually of many capillary
setae, sometimes reduced in number, rarely short,
with pointed apical cells.
549
Key to the Genera
1. Receptacle paleaceous
2
– Receptacle epaleaceous
4
2. Capitula on short pedicels and congested in dense
cymes
1547. Lepidesmia
– Capitula distinctly pedicellate and arranged in panicles
3
3. Receptacle highly conical; florets many (> 100) per capitulum; anther appendages slightly longer than wide
1548. Isocarpha
– Receptacle low-conical; florets few (c. 12) per capitulum; anther appendages lacking 1546. Parapiqueria
4. Pappus of a single seta or setae short or lacking
5
– Pappus of 5 or more regularly arranged long and persistent setae
7
5. Pappus present and of one long and persistent seta;
leaves 3-partite
1545. Monogereion
– Pappus lacking or of one or few deciduous setae; leaves
simple
6
6. Plants erect or decumbent herbs; leaves petiolate;
corolla lobes with a few short hairs; pappus lacking or
of a few short deciduous setae
1543. Alomiella
– Plants repent herbs; leaves sessile; corolla lobes glabrous and papillose inside; pappus reduced to an annular ring, sometimes with a single short seta
1544. Siapaea
7. Pappus of few setae (5–10)
1542. Gymnocondylus
– Pappus of several to many setae (18–30)
8
8. Plants epiphytic vines; style arms with markedly dilated apices
1539. Gongrostylis
– Plants terrestrial; style arms linear or filiform
9
9. Corolla with conspicuously constricted basal tube
1541. Condylidium
– Corollas funnelform or narrow and tubular
10
10. Scandent shrubs; inflorescences with distinct cymose
branching
1540. Heterocondylus
– Herbs or subshrubs; inflorescences corymbose or paniculate and often only with dense cymose terminal
branching
11
11. Anther cylinder included well within corolla tube; florets 5–40 per capitulum; style base with pubescent
node
1536. Ayapana
– Anther cylinder visible in corolla throat
12
12. Corolla tube funnelform; corollas pink to violet; florets 25–150 per capitulum; style base with glabrous or
pubescent node
1537. Ayapanopsis
– Corolla tube very narrow and tubular; corollas white;
florets 150–300 per capitulum; style base with glabrous
node
1538. Polyanthina
Genera of Ayapaninae
1536. Ayapana Spach
Fig. 119
Ayapana Spach, Hist. Nat. Vég. Phan. 10: 290 (1841); King
& Robinson, Phytologia 34: 57–66 (1976), key.
Fig. 119. Compositae-Eupatorieae. Ayapana amygdalina
(Ayapaninae). A Flowering shoot with leaf. B Floret. (Drawings by Margaret Tebbs)
Erect perennial herbs. Leaves mostly opposite,
very rarely spiralled, lamina narrowly ovate to
elliptical, entire to serrulate. Inflorescence laxly
paniculate, with laxly or densely corymbose to
550
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
subcymose branches. Phyllaries 15–35, subimbricate, 4–5-seriate, lanceolate; receptacle convex,
glabrous. Florets 5–40; corollas white or pink,
narrowly funnelform to nearly tubular, glabrous
on inner surface, with glands on outer surface of
lobes; lobes short- to long-triangular, smooth on
inner surface; anther collar slender; apical anther
appendages triangular to oblong, slightly longer
than wide; style base glabrous; arms filiform or
with short tapering recurved appendages, densely
long-papillose. Achenes prismatic, 5-ribbed, with
setulae mostly on ribs; carpopodium stoppershaped; pappus setae c. 20–40, slender, barbellate,
persistent. n = 8, 10, 17, 18. Sixteen species,
neotropics.
1537. Ayapanopsis R.M. King & H. Rob.
Ayapanopsis R.M. King & H. Rob., Phytologia 24: 382
(1972).
Erect perennial herbs or subshrubs. Leaves opposite, lamina elliptical or ovate to deltoid, acute to
acuminate, serrate to nearly entire. Inflorescence
a corymbose panicle with corymbose to weakly
cymose branches. Phyllaries 30–50, subimbricate,
3–4-seriate, narrowly oblong to lanceolate; receptacle slightly convex, glabrous to minutely pilulose.
Florets 35–150; corollas pink to violet, narrowly
funnelform; lobes as long as wide or longer,
outer surface usually glanduliferous, sometimes
with non-glandular hairs, inner smooth, anther
collar cylindrical; apical anther appendages ovate,
0.6–1.5 times as long as wide; style base glabrous to
densely hirtellous; arms linear, almost smooth to
slightly mamillose. Achenes prismatic to fusiform,
5-ribbed, setuliferous and stipitate-glandular;
carpopodium short-cylindrical; pappus setae c.
15–40, barbellate, persistent. Seventeen species,
South America.
1538. Polyanthina R.M. King & H. Rob.
Polyanthina R.M. King & H. Rob., Phytologia 20: 213
(1970).
Erect perennial herbs. Leaves opposite, sometimes
alternate above, serrate. Inflorescence a lax thyrsoid or pyramidal panicle with denser cymose
branches. Phyllaries c. 40–50, subimbricate,
2–3-seriate, lanceolate; receptacle slightly convex,
shortly puberulous. Florets c. 200–300; corollas
white, very narrowly tubular, glabrous on inner
and outer surface; lobes longer than wide; filaments inserted at staggered levels on corolla tube;
anther collar slender; apical anther appendages
triangular to ovate, slightly longer than wide; style
base glabrous; arms filiform, not tapered, shortly
mamillose. Achenes prismatic, 5-ribbed, glabrous
except for a few setulae near the top; carpopodium
stopper-shaped or cylindrical; pappus setae c.
25, barbellate, persistent. n = 10. One species,
P. nemorosa (Klatt) R.M. King & H. Rob., South
America.
1539. Gongrostylus R.M. King & H. Rob.
Gongrostylus R.M. King & H. Rob., Phytologia 24: 387
(1972).
Slender epiphytic vines, sparingly branched.
Leaves opposite, lamina ovate, remotely serrate.
Inflorescences mostly axillary, corymbose with
cymose branches. Phyllaries c. 25, subimbricate, c.
3-seriate, ovate to lanceolate or linear-lanceolate;
receptacle slightly convex, glabrous. Florets c.
20; corollas white, very narrowly funnelform,
mostly glabrous with glands on outer surface of
lobes; lobes slightly longer than wide, smooth
on both surfaces; anther collar elongate; apical
anther appendages short, only half as long as
wide; style base densely hirsute; arms narrow
and slightly mamillose below, greatly enlarged
and smooth distally in broadened and thickened
fusiform tip. Achenes prismatic, 5-ribbed, glabrous; carpopodium a distinct short cylinder with
prominent upper rim; pappus setae uniseriate,
c. 30, barbellate, persistent, scarcely narrowed
towards tips. One species, G. costaricensis (Kuntze)
R.M. King & H. Rob., Colombia, Costa Rica,
Ecuador, Panama.
1540. Heterocondylus R.M. King & H. Rob.
Heterocondylus R.M. King & H. Rob., Phytologia 24: 389
(1972).
Eupatorium L. sect. Heterocondylus (R.M. King & H. Rob.)
Cabrera (1991).
Erect to subscandent perennial herbs or subshrubs.
At least lower leaves opposite, upper alternate in
some species, lamina ovate to narrowly oblong or
panduriform, entire to serrate. Inflorescence pyramidal to distinctly cymose; capitula large, sometimes nodding. Phyllaries c. 15–30, subimbricate,
3–5-seriate, oblong to lanceolate; receptacle flat,
glabrous. Florets 20–80; corollas white to pink or
reddish-purple, narrowly funnelform; cells of limb
elongate with mostly sinuous lateral walls; lobes
usually distinctly longer than wide, smooth on both
Compositae
surfaces, glabrous to sparsely glanduliferous on
outer surface; anther collar often thickened above;
apical anther appendage ovate to oblong, slightly
longer than wide; style base glabrous or hirtellous;
arms linear to broadly linear, smooth to shortmamillose. Achenes prismatic or fusiform, 4–5ribbed, with short setulae or glands; carpopodium
distinct, stopper-shaped; pappus setae c. 20–30,
barbellate, persistent, apices not enlarged or only
gradually dilated. n = c. 20. Thirteen species, Central and South America.
1541. Condylidium R.M. King & H. Rob.
Condylidium R.M. King & H. Rob., Phytologia 24: 380
(1972).
Erect to decumbent perennial herbs or subshrubs.
Leaves opposite, lamina ovate to ovate-lanceolate,
bluntly serrate to subentire, short-acuminate.
Inflorescence thyrsoid-paniculate, with laxly
and divaricately cymose branches. Phyllaries 15,
subimbricate, in 5 ranks and 3 series, suborbicular
to narrowly lanceolate; receptacle flat to slightly
convex, glabrous. Florets 5–6; corollas white, with
a short constricted basal tube, with abruptly and
rather narrowly campanulate limb; lobes slightly
longer than wide, smooth on both surfaces; lower
filament rather short, collar almost as long, cylindrical; apical anther appendage slightly longer
than wide; style base densely short-hirsute; arms
linear, densely long-papillose. Achenes prismatic,
5-ribbed, ribs setuliferous; carpopodium asymmetrical, contorted and slightly tapering; pappus
setae c. 30–40, slender, barbellate, persistent.
n = 10. Two species, neotropics.
1542. Gymnocondylus R.M. King & H. Rob.
Gymnocondylus R.M. King & H. Rob., Phytologia 24: 393
(1972).
Erect perennial herbs. Leaves opposite, lamina
ovate, crenulate, scarcely acuminate. Inflorescence
a laxly corymbose cyme. Phyllaries c. 50, distant,
2–3-seriate, unequal, narrowly lanceolate to linear;
receptacle slightly convex, glabrous. Florets 60–80;
corollas white, narrowly funnelform, basal tube
very narrow below; lobes about twice as long as
wide, smooth on inner and outer surface, outer
surface densely hirsute; anther collar slender;
apical anther appendages triangular, 1.5 times as
long as wide; style bases glabrous, arms very narrowly clavate, densely papillose. Achenes fusiform,
5-ribbed, setuliferous in upper part; carpopodium
551
stopper-shaped; pappus setae uniseriate, c. 5–10,
barbellate, persistent, enlarged distally. One
species, G. galeopsifolius (Gardner) R.M. King &
H. Rob., Brazil.
1543. Alomiella R.M. King & H. Rob.
Alomiella R.M. King & H. Rob., Phytologia 24: 395 (1972).
Erect to decumbent perennial herbs. Leaves opposite to subopposite, lamina broadly ovate, serrate,
acute. Inflorescence laxly cymose. Phyllaries 20–30,
subimbricate, 3-seriate, unequal, elliptical to oblong, receptacle flat, glabrous. Florets c. 40; corollas white, narrowly funnelform, glabrous below on
both surfaces, veins greatly thickened in tube and
throat; lobes about as long as wide, smooth on inner and outer surfaces, outer surface with a few
small twin-hairs; anther collar cylindrical, slightly
enlarged; apical anther appendages ovate, slightly
longer than wide; style base glabrous, arms linear,
densely papillose with narrow spreading papillae.
Achenes prismatic, 5-ribbed, glabrous or setuliferous; carpopodium short stopper-shaped; pappus
absent or of short deciduous setae. Two species,
Brazil.
1544. Siapaea Pruski
Siapaea Pruski, Brittonia 48: 190 (1996).
Herbs. Stems repent, usually rooting at nodes.
Leaves opposite, simple, lamina elliptic-lanceolate,
serrate, acute. Inflorescence terminal, cymose,
few-headed; phyllaries 1–2-seriate, few, distant,
subequal; receptacle convex to conical, glabrous,
epaleaceous. Florets several, hermaphrodite;
corollas cream, corolla tube glabrous, corollalobes papillose; anther cylinder included within
throat, apical anther appendages ovate, short,
basal anther appendages rounded; style base
appressed-pubescent, shaft glabrous, arms filiform. Achenes 4–5-ribbed, glabrous; carpopodium
poorly developed; pappus reduced to a ring,
sometimes with a single short bristle. One species,
S. liesneri Pruski, Venezuela.
1545. Monogereion G.M. Barroso & R M. King
Monogereion G.M. Barroso & R.M. King, Brittonia 23: 118
(1971).
Erect, short-lived perennial herbs or subshrubs.
Leaves mostly alternate, basal leaves opposite, lamina ovate, usually deeply lobed to tripartite. Inflorescence diffuse, with capitula laxly cymosely dis-
552
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
posed on ends of leafy branches. Phyllaries c. 15,
weakly subimbricate, 2–3-seriate, somewhat unequal to subequal, lanceolate, receptacle slightly
convex, glabrous. Florets c. 25–30; corollas white,
narrowly funnelform, with hairs near base of tube,
near tips of lobes and on entire upper two-thirds
of inner surface of lobes; lobes c. 1.5 times as long
as wide, smooth on both surfaces; anther collars
slender; apical anther appendages ovate, about as
long as wide; style base glabrous; arms with linear
densely papillose appendages. Achenes somewhat
fusiform, 5-ribbed, setuliferous; carpopodium distinct, very shortly stopper-shaped; pappus of a single long scabrid persistent seta and a fringe of
numerous short narrow squamellae. One species,
M. carajensis G.M. Barroso & R.M. King, Brazil.
1546. Parapiqueria R.M. King & H. Rob.
Parapiqueria R.M. King & H. Rob., Phytologia 47: 111
(1980).
Small, erect, annual or short-lived perennial
herbs. Leaves opposite, upper subopposite to
alternate, lamina linear. Inflorescence very diffuse
with many capitula, branches rather thyrsoidpaniculate. Phyllaries c. 10, distant, ± biseriate,
subequal, persistent, oblong, narrowly bicostate;
receptacle conical, paleaceous with paleae similar to phyllaries. Florets c. 12; corollas white,
with a short glabrous basal tube, limb broadly
campanulate, with a short throat; lobes 4–5,
oblong-ovate, c. 1.5 times as long as wide, smooth
on inner surface, outer surface with 1–2 short
glands; anther collars narrowly cylindrical; apical
anther appendages lacking; style base scarcely enlarged, distinctly pilosulous; arms short, tapering,
short-papillose. Achenes subfusiform, 5-ribbed,
glabrous; carpopodium shortly stopper-shaped;
pappus absent. One species, P. cavalcantei R.M.
King & H. Rob., Brazil.
glabrous below, with a few glands on lobes; lobes c.
1.5 times as long as wide, smooth on inner surface;
anther collar slender; apical anther appendage
oblong-ovate, slightly longer than wide; style base
glabrous; arms narrowly tapering, densely longpapillose. Achenes prismatic, 4–5-ribbed, with
few to many setulae mostly on ribs; carpopodium
cylindrical; pappus a corona of short setae. n = 10.
One species, L. squarrosa Klatt, Colombia, Cuba,
Venezuela.
Aristeguieta (1964) included the genus in tribe
Heliantheae along with Isocarpha.
1548. Isocarpha R. Br.
Isocarpha R. Br., Trans. Linn. Soc. London 12: 110 (1817);
Keil & Stuessy, Syst. Bot. 6: 258–287 (1981), rev.
Erect annual or perennial herbs. Leaves opposite or
alternate, lamina narrowly elliptical, entire to serrulate, narrowly acute. Inflorescence a lax, sometimes leafy panicle. Phyllaries c. 10–15, distant,
± biseriate, subequal to equal, elliptical to lanceolate, mostly strongly ribbed; receptacle highly conical to columnar, paleaceous, paleae similar to phyllaries. Florets > 100; corollas white to pink, usually
narrowly funnelform with a distinct short basal
tube, rarely cylindrical, glands on outer surface
mostly on tube and lobes; lobes somewhat longer
than wide, mostly smooth on inner surface, rarely
papillose distally and marginally on inner surface;
filaments usually inserted well above base, anther
collar cylindrical; apical anther appendage triangular to oblong, slightly longer than wide; style base
glabrous or papillose; arms rather short, sometimes slightly tapering, often spreading or strongly
coiled, densely long-papillose. Achenes prismatic,
5-ribbed, glabrous or setuliferous; carpopodium
short, stopper-shaped; pappus absent. n = 10. Five
species, southern USA to Brazil.
1547. Lepidesmia Klatt
XXX.14. Subtribe Alomiinae Less. (1832).
Lepidesmia Klatt, Bull. Herb. Boissier 4: 479 (1896).
Subtribe Kuhniinae B.L. Rob. (1913).
Erect perennial herbs. Leaves opposite, lamina
rather fleshy, lanceolate to linear-lanceolate, entire
to subentire, blunt. Inflorescence terminal, with
elongate lower internodes, ending in dense cymes.
Phyllaries c. 15, subimbricate, c. 2–3-seriate,
unequal, lanceolate; receptacle slightly convex
to flat, paleaceous; paleae lanceolate, similar to
phyllaries but more scarious and more acute.
Florets 3–7; corollas white, narrowly funnelform,
Erect annual or perennial herbs or shrubs, sparingly to densely branched, leaves opposite or
alternate, sometimes densely spiralled, usually
petiolate. Inflorescence terminal on leafy branches,
sometimes diffuse. Capitula usually clustered,
rarely solitary, usually pedicellate, phyllaries usually distinctly subimbricate, gradate, persistent;
receptacle flat or slightly convex, epaleaceous,
usually glabrous. Florets 2–100; corollas usually
Compositae
tubular; lobes usually smooth; anther appendage
usually as long as wide; style base with or without
enlarged node, node when present usually with
hairs; style arms long-clavate, both thickened and
broadened distally, rarely narrow, usually smooth.
Achenes prismatic to obcompressed, 4–10-ribbed;
carpopodium distinct, with little or no projecting
upper rim, often asymmetrical; pappus setae
usually many, capillary, sometimes flattened on
outer surface, sometimes plumose, rarely few,
winged, or absent. The generic concept applied
here follows King and Robinson (1987, 1995),
rather than Turner (1988, 1990, 1991a, 1994, 1995)
and Turner et al. (1991).
Key to the Genera
1. Pappus lacking
2
– Pappus present, although sometimes deciduous
3
2. Florets 4–5 per capitulum; corollas glabrous or
glandular-punctate; achenes with papillate setulae;
Mexico
1558. Alomia
– Florets 40–50 per capitulum; corollas with numerous
hairs and glandular punctae; achenes glabrous; Brazil
1568. Planaltoa
3. Style base with distinctly enlarged basal node
4
– Style base lacking distinctly enlarged basal node 14
4. Pappus setae easily deciduous, often caducous; Brazil
1567. Leptoclinium
– Pappus persistent
5
5. Pappus setae of two different sizes
6
– Pappus setae equal or subequal
7
6. Florets 4 per capitulum; leaves usually alternate; longer
pappus setae alternating with shorter; Brazil (Goiás)
1566. Goyazianthus
– Florets 6–8 per capitulum; leaves opposite; longest
pappus setae above ribs of achene; Brazil (Ceará, Piauí)
1563. Dissothrix
7. Leaves narrowly linear and inserted in dense spirals
1565. Pseudobrickellia
– Leaves usually ovate or oblong and opposite or
whorled
8
8. Achenes 8–10-ribbed
1549. Brickellia
– Achenes 5-ribbed
9
9. Shrubs; leaf venation pinnate; phyllaries with rounded
or obtuse apices; style base with glabrous or pubescent
node
10
– Herbs or subshrubs; leaves usually 3-veined from or
near to base; phyllaries mostly acute; style base with
pubescent node
11
10. Leaves abruptly short-petiolate, lamina oblong-ovate,
relatively small; inflorescence with ascending branchlets; achenes densely glanduliferous; Peru
1569. Crossothamnus
– Leaves attenuate to a stout petiole, lamina elliptical,
medium sized; inflorescence with spreading branchlets; achenes with few glands or glands lacking; Colombia
1571. Condylopodium
11. Leaves with spiny lobes
1550. Barroetea
553
– Leaves without spiny lobes
12
12. Corolla throats flared; corolla lobes and style arms
densely papillate
1551. Phanerostylis
– Corolla throats tubular or narrow funnelform; corolla
lobes and style arms ± smooth
13
13. Inflorescences with laxly ascending branches; florets
10–30(–75) per capitulum; carpopodium relatively large and contorted; USA (Texas), Mexico
1553. Flyriella
– Inflorescences with spreading branches; florets 3–12
per capitulum; carpopodium relatively small and
short-cylindrical; South America
1564. Austrobrickellia
14. Pappus setae basally flattened or pappus of squamellae
or awns
15
– Pappus of capillary setae
17
15. Pappus of 5 awns
1554. Ageratella
– Pappus of alternating awns and squamellae
16
16. Leaves petiolate with distinct lamina, mostly opposite;
achenes prismatic with short ascending setulae
1557. Pleurocoronis
– Leaves sessile, linear, mostly alternate; achenes
fusiform with long spreading setulae on ribs
1556. Malperia
17. Achenes moderately to densely long stipitateglandular
1559. Dyscritogyne
– Achenes short stipitate-glandular or setuliferous 18
18. Leaf lamina bases attenuate
19
– Leaf lamina bases truncate or cordate
21
19. Corollas glandular-punctate outside; achenes
5–6-ribbed; South America
1570. Helogyne
– Corollas eglandular outside; achenes 7–10-ribbed;
USA, Mexico
20
20. Leaves sessile, opposite, linear or squamulose; phyllaries 4- or > 4-seriate
1555. Asanthus
– Leaves petiolate, opposite or whorled; phyllaries 2–3seriate
1561. Steviopsis
21. Pappus setae plumose, united at base
1562. Carminatia
– Pappus setae barbellate, free at base
22
22. Florets 9–18 per capitulum; corollas tubular; achenes
with long, spreading setulae
1560. Kyrsteniopsis
– Florets 25–35 per capitulum; corollas funnelform; achenes with short setulae
1552. Brickelliastrum
Genera of Alomiinae
1549. Brickellia Elliott
Brickellia Elliott, Sketch Bot. S. Carolina 2: 290 (1824);
Robinson, Mem. Gray Herb. 1: 1–151 (1917), rev.
Erect annual or perennial herbs, subshrubs or
shrubs. Leaves opposite or alternate, lamina linear,
lanceolate, ovate, deltoid or lobate, usually dentate. Capitula usually clustered in leafy thyrsoid
panicle, sometimes corymbose or cymose, rarely
solitary and nodding on long peduncles. Phyllaries
14–45, subimbricate, 5–6-seriate, usually gradate,
unequal; receptacle flat to slightly convex. Florets
c. 4–100; corollas usually white to cream-coloured,
554
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
sometimes purplish, tubular or rarely narrowly
funnelform; lobes as long as wide to twice as
long as wide, smooth on both surfaces; anther
collars slender to slightly thickened; apical anther
appendages slightly longer than wide; style base
with distinct enlarged node, covered with dense
contorted hairs; style arms long-clavate. Achenes
prismatic, 10-ribbed, setuliferous; carpopodium
distinct, slightly to greatly longer on outer surface; pappus setae uniseriate, 10–80, persistent,
flattened on outer surfaces, barbellate to densely
plumosely fringed on lateral margins, apical cells
acute. n = 9. Circa 100 species, New World.
1550. Barroetea A. Gray
Barroetea A. Gray, Proc. Amer. Acad. Arts 15: 29 (1880).
Annual or short-lived perennial herbs. Leaves
opposite, lamina deltoid, serrate to dentate, with
prickles on tip and teeth. Inflorescence a panicle.
Phyllaries c. 4-seriate, c. 20–25, subimbricate,
unequal; receptacle flat to convex, glabrous.
Florets c. 15–35; corollas white, tubular; lobes
more than twice as long as wide, smooth on both
surfaces; anther collars very slender, elongate;
apical anther appendages oblong, blunt, slightly
longer than wide; style base enlarged, covered
with contorted hairs; arms long-clavate, surface
smooth to slightly mamillose. Achenes slightly
to strongly obcompressed, constricted below
pappus, 4-ribbed, setuliferous; carpopodium
distinct, broad, strongly asymmetrical; pappus
setae uniseriate, c. 16–30, persistent, flattened
on outer surface, densely pectinately fringed on
lateral margins; apical cells acute. n = 9. Seven
species, Mexico.
1551. Phanerostylis (A. Gray) R.M. King &
H. Rob.
Phanerostylis (A. Gray) R.M. King & H. Rob., Phytologia
24: 70 (1972).
Erect to decumbent or rhizomatous, annual or
perennial herbs. Leaves opposite, lamina ovate to
narrowly oblong. Inflorescence sparsely branched
or capitula solitary on long erect peduncles.
Phyllaries c. 30–50, subimbricate, 3–7-seriate,
unequal, spreading with age; receptacle flat or
slightly convex, glabrous. Florets 25–50; corollas
white or pale pink, with slender basal tube, throat
funnelform; lobes scarcely longer than wide to half
again as long as wide, papillose on both surfaces;
anther collars short to rather elongate; apical
anther appendages slightly shorter to distinctly
longer than wide; style base enlarged, covered
with rather long, rather straight hairs; arms longclavate, fleshy, enlarged part densely covered with
small papillae. Achenes prismatic, 4–5-ribbed,
scabrid with minute setulae; carpopodium distinct,
nearly symmetrical to asymmetrical; pappus setae
uniseriate, c. 25, persistent, essentially smooth on
outer surface, apical cells very slender, narrowly
acute. n = 9. Five species, Mexico.
1552. Brickelliastrum R.M. King & H. Rob.
Brickelliastrum R.M. King & H. Rob., Phytologia 24: 63
(1972).
Erect to decumbent subshrubs. Lower leaves
opposite, upper often alternate, lamina ovate
to triangular, slightly pungent, crenate-serrate.
Inflorescence corymbose to pyramidally paniculate. Phyllaries c. 4-seriate, c. 25, subimbricate,
unequal; receptacle slightly convex, glabrous.
Florets 25–35; corollas white, narrowly and evenly
funnelform from base; lobes slightly longer than
wide, smooth on both surfaces; anther collars
rather short, slender; apical anther appendages
oblong, slightly longer than wide; style base
not enlarged, glabrous; arms very narrowly and
long-clavate, slightly mamillose, nearly smooth.
Achenes prismatic, mostly 5–7-ribbed, scabrous
with short setulae; carpopodium distinct, shortly
rounded; pappus setae c. 25, somewhat deciduous,
barbellate, slightly flattened on outer surface,
apical cells strongly acute. n = 10. Two species,
Mexico and USA (Texas).
1553. Flyriella R.M. King & H. Rob.
Flyriella R.M. King & H. Rob., Phytologia 24: 67 (1972);
Baker & Turner, Sida 11: 300–317 (1986), rev.
Perennial herbs to subshrubs, erect or decumbent.
Leaves usually opposite, sometimes alternate
above, lamina ovate to deltoid, serrate. Inflorescence a laxly branched panicle. Phyllaries c.
30, subimbricate, c. 3-seriate, unequal, usually
lanceolate, or c. 40, with enlarged foliaceoustipped outer phyllaries; receptacle flat, glabrous.
Florets usually 10–30 (to c. 75); corollas white,
tubular to scarcely funnelform, mostly glabrous,
cells of limb mostly elongate with sinuous lateral
walls; lobes scarcely to distinctly longer than
wide, smooth on both surfaces, outer surface
with few glands, sometimes with short hairs,
anther collars narrow; apical anther appendages
Compositae
oblong-ovate, slightly longer than wide; style base
enlarged, densely hirsute, with erect or somewhat
curved hairs; arms long-clavate, smooth. Achenes
prismatic, 5-ribbed, setuliferous; carpopodium
distinct, cylindrical or asymmetrical; pappus setae
uniseriate, 20–40, capillary, barbellate, narrowed
distally, slightly flattened and barbellate on outer
surface, persistent, apical cells obtuse to acute.
n = 10. Four species, Mexico, USA.
1554. Ageratella A. Gray ex S. Watson
Ageratella A. Gray ex S. Watson, Proc. Amer. Acad. Arts 22:
419 (1887); Turner, Phytologia 78: 204–208 (1995), rev.
Erect subshrubs or shrubs. Leaves alternate or
sometimes opposite below, lamina ovate, obovate,
linear or linear-oblanceolate, entire to lobed.
Inflorescence a loose racemose or narrowly
thyrsoid panicle. Phyllaries c. 18–20, distant, 4–
5-seriate, unequal, somewhat ranked, short-ovate
to lanceolate; receptacle slightly convex, glabrous.
Florets 15; corollas whitish, tubular, somewhat
constricted above, with glands on outer surface on
tube and lobes and especially on base of throat;
lobes more than twice as long as wide, smooth
on both surfaces; anther collars slender; apical
anther appendages about twice as wide as long;
style base not enlarged, glabrous, arms long and
narrowly clavate, with surface mamillate on lower
part of appendage, becoming smoother above.
Achenes prismatic, 5-ribbed, with short setulae;
carpopodium short stopper-shaped; pappus of
4–5 persistent, scabrid awns, winged below, apical
cells acute. One species, A. microphylla (Sch. Bip.)
A. Gray ex S. Watson, Mexico.
1555. Asanthus R.M. King & H. Rob.
Asanthus R.M. King & H. Rob., Phytologia 24: 66 (1972).
Erect subshrubs. Lower leaves opposite, upper
sometimes alternate, sometimes reduced and
scale-like, lamina narrowly lanceolate to linear.
Inflorescence thyrsoid-paniculate with densely
corymbose branches. Phyllaries c. 20–25, 4- or
> 4-seriate, subimbricate; receptacle flat, glabrous.
Florets 8–14; corollas whitish, tubular and slightly
constricted above to minimally funnelform, with
sparse minute glands externally; lobes 2–3 times
as long as wide, smooth on both surfaces; anther
collars cylindrical; apical anther appendages
oblong, slightly longer than wide or longer; style
bases not enlarged, glabrous; arms long and
narrowly clavate, slightly mamillose below, usually
555
becoming smooth above. Achenes long-prismatic,
c. 10-ribbed, ribs with short setulae; carpopodium
short stopper-shaped; pappus setae 1–3-seriate,
20–100, persistent, barbellate on margins and
outer surface, apical cells strongly acute. Three
species, Mexico, USA.
1556. Malperia S. Watson
Malperia S. Watson, Proc. Amer. Acad. Arts 24: 54 (1889);
King, Rhodora 69, 777: 35–47 [43–45] (1967), rev.
Erect annual herbs. Leaves opposite below, alternate above, lamina linear, entire. Inflorescence
a loose cymose panicle. Phyllaries 20–25, subimbricate, c. 3-seriate, strongly unequal; receptacle
flat, glabrous. Florets c. 20–30; corollas white,
narrowly tubular, with scattered small glands
on outer surface; lobes scarcely spreading, c. 1.5
times as long as wide, smooth on both surfaces;
apical anther appendages oblong-ovate, slightly
longer than wide; style base not enlarged, glabrous; arms narrow and long-clavate, scarcely
mamillose. Achenes somewhat fusiform, 5-ribbed,
with spreading setulae on ribs; carpopodium
somewhat asymmetrical; pappus uniseriate, of
long awns with winged bases and intervening short
squamellae, persistent, awns scabrid, becoming
barbellate distally, with apical cells sharply acute.
n = 10. One species, M. tenuis S. Watson, Mexico,
USA.
1557. Pleurocoronis R.M. King & H. Rob.
Pleurocoronis R.M. King & H. Rob., Phytologia 12: 468
(1966); King, Rhodora 69: 35–47 [34–43] (1967), rev.
Erect, small, usually spreading shrubs. Leaves opposite, alternate above, lamina minutely rhomboid
to broadly deltoid or cordiform in outline, slightly
to deeply toothed or incised to bipinnatifid. Capitula solitary or in lax to rather dense corymbose
or subcymose panicles. Phyllaries 30–35, subimbricate, 3–4-seriate, outer short-ovate with small
herbaceous tips, inner lanceolate; receptacle flat
or slightly convex, epaleaceous, glabrous. Florets
25–30; corollas white, tubular, with scattered
minute glands on outer surface; anther collars
cylindrical; apical anther appendages ovate to
oblong, c. 15 times as long as wide; style base
not enlarged, glabrous; arms long and sometimes
rather broadly clavate, slightly mamillose. Achenes
prismatic, 4–5-ribbed, setulose; carpopodium
slightly asymmetrical; pappus setae uniseriate,
3–6, long, barbellate, persistent, with intervening
556
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
short erosely dentate squamellae, apical cells of
setae sharply acute. n = 9. Three species, Mexico,
USA.
1560. Kyrsteniopsis R.M. King & H. Rob.
1558. Alomia Kunth
Erect subshrubs or shrubs. Leaves opposite, lamina
ovate to deltoid, entire to dentate or denticulate.
Inflorescence a lax leafy thyrsoid panicle, with laxly
to densely corymbose branches. Phyllaries 20–25,
subimbricate, c. 3–4-seriate, strongly unequal,
spreading when dry; receptacle slightly convex,
glabrous. Florets 9–18; corollas greenish white
or cream-white, narrowly tubular to minimally
funnelform, with glands on outer surface of tube
and lobes or glabrous on tube with some glands
on lobes; lobes slightly longer than wide, slightly
spreading, smooth on both surfaces; anther collars
narrowly cylindrical; apical anther appendages
oblong to ovate, slightly longer than wide, style
base not enlarged, glabrous; arms linear to scarcely
long-clavate, rounded on outer surface, scarcely or
not mamillose. Achenes prismatic, 5-ribbed, with
setulae mostly on ribs, often with stipitate glands;
carpopodium distinct; pappus setae uniseriate,
25–45, barbellate, persistent, apical cells acute.
n = 10. Five species, Guatemala, Mexico.
Alomia Kunth in Humb., Bonpl. & Kunth, Nov. Gen. Sp. (ed.
folio) 4: 119 (1818).
Erect to decumbent short-lived perennial herbs.
Leaves opposite, lamina usually ovate, upper
leaves sometimes elliptical or lanceolate, usually
serrate or dentate. Inflorescence rather lax, weakly
to strongly cymose panicles. Phyllaries 25–30,
subimbricate, 2–4-seriate, unequal to subequal; receptacle broadly convex, without paleae, glabrous.
Florets 40–50; corollas white, tubular or with narrowly campanulate throat, with minute scattered
glands on outer surface; lobes longer than wide,
smooth or nearly smooth on both surfaces; anther
collars cylindrical; apical anther appendages ovate,
about as long as wide; style base not enlarged,
glabrous; arms scarcely thickened to long and
narrowly clavate, slightly mamillose to densely
short-papillose. Achenes prismatic, 5-ribbed, with
short blunt twin-hairs; carpopodium cylindrical;
pappus absent. n = 10. Five species, Mexico.
Kyrsteniopsis R.M. King & H. Rob., Phytologia 22: 146
(1971).
Pseudokyrsteniopsis R.M. King & H. Rob. (1973).
1559. Dyscritogyne R.M. King & H. Rob.
1561. Steviopsis R.M. King & H. Rob.
Dyscritogyne R.M. King & H. Rob., Phytologia 22: 158
(1971).
Steviopsis R.M. King & H. Rob., Phytologia 22: 156 (1971).
Erect perennial herbs. Leaves opposite to alternate,
lamina broadly to narrowly ovate, subserrulate
to serrate, acute. Inflorescence a loose pyramidal or corymbose panicle. Phyllaries c. 33–40,
subimbricate, c. 4–6-seriate, strongly unequal,
mostly persistent; receptacle flat to slightly convex,
glabrous. Florets 11–16; corollas white or pink,
tubular, with scattered minute glands on outer
surface, sometimes with sparse hairs inside near
base of throat; lobes scarcely longer than wide,
smooth on both surfaces; anther collars cylindrical
or narrow below; apical anther appendages oblong
to ovate, about as long as wide; style base not
enlarged, glabrous or with a few minute papillae;
arms long-clavate or strap-shaped, fleshy, nearly
smooth. Achenes prismatic to rather fusiform, 4–
5-ribbed, densely punctate- or stipitate-glandular;
carpopodium indistinct; pappus setae 1–2-seriate,
c. 35–40, persistent, barbellate on margins and
outer surface, apex not or slightly broadened,
apical cells acute. Two species, Mexico.
Erect coarse herbs. Leaves usually opposite or
verticillate, alternate above, lamina ovate to
narrowly lanceolate, serrulate, narrowly acute to
acuminate. Inflorescence a loose often leafy pyramidal or corymbose panicle, branches ascending.
Phyllaries 20–30, distant to weakly subimbricate,
c. 2–3-seriate, with some shorter exterior bracts;
receptacle slightly convex, glabrous. Florets 15–20;
corollas greyish pink to purplish, tubular to narrowly funnelform, glabrous on outer surface; lobes
1.2–2 times longer than wide, spreading, smooth
on both surfaces; anther collars broadly cylindrical; apical anther appendages oblong, slightly
longer than wide; style base not enlarged, glabrous;
arms long and narrowly clavate, fleshy, somewhat
flattened, slightly mamillose to smooth. Achenes
prismatic, 5–10-ribbed, densely to sparsely setuliferous, with few to many short-stalked glands;
carpopodium indistinct; pappus setae uniseriate,
c. 25–30, often coarsely barbellate, persistent, not
flattened nor smooth on outer surface, apices not
broadened, apical cells acute. Four species, Mexico.
Compositae
King and Robinson’s (1971, 1972, 1987) concept of Steviopsis was expanded by Turner (1988)
to include both Asanthus and Dyscritogyne. Turner
(1994) also considered the newly described Brickelliastrum villarrealii R.M. King & H. Rob. part of
his expanded Steviopsis. All four genera are kept
separate here.
1562. Carminatia Moç. ex DC.
Carminatia Moç. ex DC., Prodr. 7: 267 (1838).
Erect annual herbs. Leaves opposite, lamina deltoid
to broadly ovate, dentate. Inflorescence spiciform,
with capitula solitary or clustered at nodes along
spike. Phyllaries c. 20, subimbricate, c. 3-seriate,
lanceolate to linear; receptacle flat, glabrous.
Florets c. 10–11; corollas white, rather tubular,
either slightly broader or slightly narrower above,
glabrous on outer surface or with a few minute
glands above, veins greatly thickened towards
base; lobes slightly shorter or slightly longer than
wide, smooth on both surfaces; anther collars
slender; apical anther appendages slightly longer
than wide; style base not enlarged, glabrous; arms
narrowly linear to scarcely long-clavate, surface
slightly mamillose. Achenes prismatic, 5-ribbed,
minutely setuliferous; carpopodium strongly differentiated; pappus setae c. 9–13, often deciduous
in groups with narrowly fused bases, flattened on
outer surface, plumose, apical cells acute. n = 10.
Three species, Mexico and USA (Arizona).
1563. Dissothrix A. Gray
Dissothrix A. Gray, Hooker’s J. Bot. Kew Gard. Misc. 3: 223
(1851).
Erect annual herbs. Leaves opposite, lamina ovatelanceolate, serrate. Inflorescence a loose leafy
thyrsoid panicle with cymose branches. Phyllaries
c. 15, subimbricate, c. 3-seriate; receptacle flat,
glabrous. Florets 6–8; corollas whitish, tubular,
somewhat constricted above, glands dense at tips
of lobes, very sparse elsewhere on outer surface,
veins distinctly thickened below; lobes smooth on
both surfaces; anther collars slender; apical anther
appendages longer than wide; style base enlarged,
with numerous thin-walled hairs; arms with long
narrow mamillose bases on appendages, with
broad elongate clavate smooth tips. Achenes prismatic, 5-ribbed, with short setulae; carpopodium
stopper-shaped; pappus setae uniseriate, persistent, barbellate, of two distinct types, c. 5 longer
thicker setae above ribs of achene, c. 18 shorter
557
narrower unequal setae above areas between
ribs, apical cells of longer setae obtuse, apical
cells of others mostly short-acute. One species,
D. imbricata (Gardner) B.L. Rob., Brazil.
Known only from the type collections.
1564. Austrobrickellia R.M. King & H. Rob.
Austrobrickellia R.M. King & H. Rob., Phytologia 24: 72
(1972).
Eupatorium sect. Austrobrickellia (R.M. King & H. Rob.)
Cabrera (1995).
Erect or spreading to arching subshrubs or shrubs.
Leaves opposite, lamina ovate, entire to sharply
dentate. Inflorescence a lax leafy thyrsoid panicle,
branches densely corymbose at tips. Phyllaries c.
6–20, subimbricate, 2–4-seriate, ovate to lanceolate; receptacle flat to slightly convex, glabrous.
Florets 3–12; corollas greenish white to purple,
tubular, sometimes with slight constrictions
above and near base, glabrous on outer surface
or with few minute glands on lobes; lobes about
twice as long as wide, erect, smooth on both
surfaces; anther collars cylindrical, sometimes
short; apical anther appendages oblong, 1.25–1.5
times as long as wide; style base enlarged, with
numerous scarcely distorted ascending hairs; arms
long-clavate, mostly smooth. Achenes prismatic,
5-ribbed, with setulae often nearly restricted to
ribs, with or without glands on sides; carpopodium
shortly stopper-shaped; pappus setae uniseriate,
30–35, persistent, barbellate, with flattened mostly
uninterrupted band along middle of outer surface,
margins rather densely scabrid, apical cells mostly
obtuse, sometimes narrowly rounded. n = 10.
Three species, Argentina, Bolivia, Brazil, Paraguay.
1565. Pseudobrickellia R.M. King & H. Rob.
Fig. 120
Pseudobrickellia R.M. King & H. Rob., Phytologia 24: 74
(1972); Hind, Fl. Pico das Almas 260–261 (1996), key.
Small trees or erect, often somewhat fasciculated
shrubs. Leaves densely spirally inserted, lamina
narrowly linear, glabrous. Inflorescence terminal
on leafy branches, densely corymbose to somewhat pyramidal. Phyllaries 12–18, subimbricate,
3–4-seriate, oblong to lanceolate; receptacle flat,
epaleaceous (with a few marginal paleae?), glabrous. Florets 2–4(–8?); corollas greenish white,
tubular or minimally funnelform, glabrous on
outer surface; lobes c. twice as long as wide, erect,
smooth on both surfaces; anther collars broadly
558
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
rescence thyrsoid-paniculate, with subcymose
branches. Phyllaries 15–16, subimbricate, c. 4seriate, mostly in 4 distinct ranks; receptacle flat,
glabrous. Florets 4; corollas creamy white, tubular,
narrower above, with numerous glands on outer
surface; lobes about twice as long as wide, smooth
on both surfaces; filaments inserted near basal
fourth of corolla, lower parts short, anther collars
slightly thickened; apical anther appendages
oblong-ovate, about as long as wide; style base
enlarged, densely pubescent with short contorted
hairs; arms clavate, fleshy, mamillose below,
becoming smooth above. Achenes prismatic, c.
7-ribbed, densely setuliferous and glanduliferous;
carpopodium short stopper-shaped; pappus setae
c. 50, persistent, dimorphic, with outer smaller
series in most gaps between larger inner setae,
densely barbellate on margins, with few short free
cell apices on outer surface, apical cells acute.
n = 20. One species, G. tetrastichus (B.L. Rob.)
R.M. King & H. Rob., Brazil.
1567. Leptoclinium Benth. & Hook. f.
Leptoclinium Benth. & Hook. f., Gen. Pl. 2: 244 (1873).
cylindrical; apical anther appendages broadly
ovate, about as long as wide; style base enlarged,
hirsute with short, contorted, blunt-tipped hairs;
arms long-clavate, flattened only on inner surface, scarcely mamillose below, smooth apically.
Achenes prismatic, 5–10-ribbed, distinctly setuliferous; carpopodium narrowly annuliform to short
stopper-shaped; pappus setae c. 35, barbellate,
persistent, ± biseriate, some outer setae shorter,
outer surfaces with median glabrous band. Two
species, Brazil.
Erect shrubs. Leaves alternate, imbricate, lamina
broadly lanceolate, entire. Inflorescence of small
terminal corymbose panicles. Phyllaries c. 12,
subimbricate, c. 3-seriate, narrowly ovate to
oblong-ovate; receptacle flat to slightly convex,
glabrous. Florets 5; corollas creamy white, tubular, glabrous; lobes over twice as long as wide;
anther collars short-cylindrical; apical anther
appendages about as long as wide; style base
enlarged, with dense cluster of short contorted
hairs; arms broad- and long-clavate, with inner surface flattened, mamillose below, smooth
on broadest parts, stigmatic lines marginal
or submarginal. Achenes prismatic, 5-ribbed,
glabrous; carpopodium stopper-shaped; pappus
setae apparently uniseriate, numerous, usually
caducous, apical cells short-acute. One species,
L. trichotomum (Gardner) Benth. ex Baker, Brazil.
1566. Goyazianthus R.M. King & H. Rob.
1568. Planaltoa Taub.
Goyazianthus R.M. King & H. Rob., Phytologia 37: 461
(1977).
Planaltoa Taub., Bot. Jahrb. Syst. 21: 454 (1896); King &
Robinson, Monogr. Syst. Bot. 22: i–xii, 1–581 (1987), key.
Erect cinereo-puberulous and glandular-punctate
subshrubs. Primary leaves usually alternate,
branch leaves and bracts of inflorescence usually
opposite, lamina narrowly oblong-oblanceolate,
entire, with rounded and apiculate tips. Inflo-
Erect, densely hirtellous subshrubs or shrubs.
Leaves alternate, imbricate, lamina lanceolate
to oblong-elliptical, minutely serrulate, margins
sometimes reflexed. Inflorescence terminal,
densely corymbose or thyrsoid; capitula sessile in
Fig. 120.
Compositae-Eupatorieae. Pseudobrickellia
brasiliensis (Alomiinae). A Flowering shoot. B Floret.
(Drawings by Margaret Tebbs)
Compositae
small bracteolate clusters. Phyllaries 5–7, weakly
subimbricate, ± biseriate, subequal; receptacle
flat, glabrous. Florets 3–5; corollas pink, tubular
or minimally funnelform, outer surface densely
pubescent with eglandular hairs, stipitate glandular hairs, or short-stalked glands; lobes nearly
twice as long as wide or longer, somewhat mamillose, with or without hairs on inner surface; anther
collar short; apical anther appendages slightly
wider than long to longer than wide, style base
scarcely enlarged, densely pubescent with long
scarcely contorted hairs; arms long and narrowly
clavate with papillae below, becoming mamillose
to smooth above, or linear and densely papillose.
Achenes prismatic, 5–6-ribbed, glabrous, abruptly
constricted at base; carpopodium indistinct;
pappus lacking. n = 10. Two species, Brazil.
1569. Crossothamnus R.M. King & H. Rob.
Crossothamnus R.M. King & H. Rob., Phytologia 24: 77
(1972); King & Robinson, Phytologia 78: 381–383 (1995),
key.
Erect shrubs. Leaves opposite to alternate, lamina
ovate, serrulate to subserrulate. Inflorescence
thyrsoid-paniculate, with branches rather densely
corymbose. Phyllaries c. 20, 3–4-seriate, strongly
subimbricate, oblong. Florets c. 10; corollas white,
slightly funnelform, slightly narrowed above,
glanduliferous on outer surface; lobes slightly
longer than wide, smooth on both surfaces;
anther collars broadly cylindrical; apical anther
appendages oblong, 1.25 times as long as wide;
style base enlarged, smooth to papillose; arms
long-clavate, slightly mamillose below, smooth
above, flattened only on inner surface. Achenes
prismatic, 5–7-ribbed, densely glanduliferous
with short-stipitate glands, rarely setuliferous;
carpopodium short-cylindrical; pappus setae
uniseriate, c. 35, persistent, barbellate, without
flattened outer surfaces, apices somewhat broadened, apical cells obtuse. n = 10. Four species,
Colombia, Ecuador, Peru.
559
Florets 5–18; corollas white, pink, purple, or
according to some descriptions, yellow, tubular
and somewhat constricted above or funnelform,
with many minute glands on outer surface, at
least on lobes; lobes smooth on both surfaces;
anther collars cylindrical; apical anther appendage
slightly longer than wide; style base not enlarged,
rarely scarcely broadened, glabrous; style arms
long-clavate, flattened on inner surface, minutely
mamillose to slightly papillose to tip, rarely smooth
above. Achenes prismatic, 5–6-ribbed, usually
with both setulae and numerous minute glands;
carpopodium stopper-shaped to short-cylindrical;
pappus setae uniseriate, c. 20–30, persistent,
barbellate to plumose on margins, free cell apices
reduced or lacking along middle of outer surface,
apical cells acute. Eight species, Argentina, Bolivia,
Chile, Peru.
1571. Condylopodium R.M. King & H. Rob.
Condylopodium R.M. King & H. Rob., Phytologia 24: 397
(1972); Diaz-Piedrahita & Mendez-Ramirez, Revista Acad.
Colomb. Ci. Exact. 25: 17–19 (2001), key.
Erect to subscandent shrubs. Leaves opposite,
lamina broadly elliptical, entire to remotely serrulate. Inflorescence broadly pyramidally paniculate.
Phyllaries 20–30, subimbricate, c. 4–5-seriate,
inner deciduous; receptacle slightly convex,
puberulous. Florets c. 10–12; corollas greenish
white, minimally narrowly funnelform, with
glands above on outer surface; lobes c. 1.5 times
as long as wide, erect, smooth on both surfaces,
anther collars cylindrical, poorly demarcated at
base; apical anther appendages oblong-ovate,
slightly longer than wide, style base enlarged,
with or without somewhat contorted hairs; arms
scarcely to distinctly long-clavate, mamillose
below, in broader forms becoming smooth at tip.
Achenes prismatic, 5-ribbed, with sparse setulae
or minute glands on sides; carpopodium shortly
stopper-shaped; pappus setae uniseriate, 30–40,
barbellate, persistent, contiguous, with broadened
apices, apical cells obtuse. Four species, Colombia.
1570. Helogyne Nutt.
Helogyne Nutt., Trans. Amer. Philos. Soc. n.s. 7: 449 (1841).
Erect subshrubs or shrubs. Leaves alternate, lamina
small, elliptical to lanceolate, entire. Inflorescence
a dense rather pyramidal or thyrsoid panicle or
sometimes diffuse and leafy. Phyllaries c. 10–30,
usually subimbricate, 2–5-seriate, rarely distant
and subequal; receptacle slightly convex, glabrous.
XXX.15. Subtribe Critoniinae R.M. King & H.
Rob. (1980).
Perennial herbs, erect or scandent shrubs, or small
trees. Leaves mostly opposite, sometimes alternate.
Inflorescence terminal, lateral or rarely axillary,
usually with corymbose branches; capitula clustered, pedicellate or sessile; phyllaries weakly
560
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
to strongly subimbricate or imbricate, usually
gradate, inner usually deciduous; receptacle flat
to slightly convex, rarely conical, paleaceous or
epaleaceous. Florets 1–300; corollas usually funnelform, sometimes tubular or with campanulate
throat, glabrous or sometimes with hairs on inner
surface; corolla lobes smooth on both surfaces;
anther collar cylindrical, less than 5 times as
long as wide, rarely very short; apical anther appendages often as long as wide or longer, less often
half as long as wide or less, rarely lacking; style
base not enlarged, glabrous; style arms filiform or
linear to distinctly clavate, mamillose or distally
smooth, rarely distinctly short-papillose. Achenes
prismatic to fusiform, 5-ribbed; carpopodium
usually distinct, sometimes procurrent along ribs;
pappus rarely squamiform or absent, setae usually
uniseriate, many, capillary, barbellate, apices not
to greatly expanded, apical cells obtuse to sharply
acute.
Key to the Genera
1. Plants rosulate or subrosulate and inflorescences
scapose or scapiform (peduncle or stipe rarely with
one or two leaf-like bracts)
2
– Plants coarse herbs to small trees with leafy stems 3
2. Capitula with 9–10 florets; pappus setae about as long
as corollas; apical anther appendages vestigial; top of
corolla tube conspicuously constricted and tube distinctly inflated; eastern Cuba
1577. Ciceronia
– Capitula with 30–50 florets; pappus a low crown of
deeply laciniate scales; apical anther appendages large,
nearly as long as wide; top of corolla tube slightly
narrowed; central Cuba
1576. Antillia
3. Individual capitula single-flowered and aggregated
into spherical clusters; leaves mostly alternate
4
– Individual capitula with 3 or 4 florets; leaves opposite
or alternate
5
4. Pappus of squamellae
1591. Mexianthus
– Pappus of capillary setae
1592. Neohintonia
5. Pappus absent; plants coastal epiphytic herbs or
shrubs
1611. Tuberostylis
– Pappus usually present; plants terrestrial herbs,
shrubs, trees or vines
6
6. Leaves distinctly alternate (except sometimes at base)
7
– Leaves mostly opposite or subopposite
8
7. Leaves densely spirally inserted, short-petiolate; phyllaries strongly subimbricate, margins markedly scarious; stems and leaves lacking stipitate glands; capitula
sessile in spiciform inflorescences, with 6–8 florets;
Peru
1585. Nothobaccharis
– Leaves remote, with slender petioles; phyllaries distant, margins herbaceous; stems and leaves with numerous stipitate glands; capitula pedicellate in corymbose inflorescences with 30–60 florets; Argentina, Bolivia
1584. Bishovia
8. Capitula densely clustered and sessile in leaf axils
1609. Uleophytum
– Capitula not sessile nor clustered in leaf axils
9
9. Receptacles with many paleae; capitula with 100–300
florets
1582. Eupatoriastrum
– Receptacle with few paleae or epaleaceous; capitula
with 4–70 florets (up to 100 in Aristeguietia)
10
10. Leaves bipinnate; Haiti, Dominican Republic
1578. Eupatorina
– Leaves not dissected, rarely pinnately lobed
11
11. Phyllaries distant or weakly subimbricate in up to 3
rather irregular series
12
– Phyllaries subimbricate or imbricate in 4 or more gradate series, inner phyllaries sometimes easily deciduous
17
12. Apical anther appendages absent, often appearing as
two minute lobes; phyllaries distant
1605. Ophryosporus
– Apical anther appendages distinct, 1/3 as long as wide
or longer; phyllaries weakly subimbricate
13
13. Corolla lobes as long as wide, glabrous or subglabrous
on outer surface; plants scandent; Brazil (Bahia)
1586. Santosia
– Corolla lobes not distinctly longer than wide, densely
glandular-punctate on outer surface; plants erect or
subscandent herbs or shrubs
14
14. Corolla tube basally constricted with throat distinctly
expanded near base; Argentina, Paraguay
1589. Chacoa
– Corolla tube broadly cylindrical, expanded gradually
into funnelform throat
15
15. Capitula sessile in dense spherical clusters
1583. Sphaereupatorium
– Capitula not sessile in dense spherical clusters
16
16. Receptacle with some paleae; shrubs with strongly
pinnately veined elliptical leaves
1590. Idiothamnus
– Receptacle epaleaceous; herbs or shrubs with pinnately or 3-veined leaves of various shapes
1581. Koanophyllon
17. Lower leaf surfaces and phyllaries both densely tomentose or villous; eastern Cuba
1587. Grisebachianthus
– Lower leaf surfaces and phyllaries glabrous,
glandular-punctate, puberulous or sparsely to
moderately pubescent or tomentose but never both
similarly so
18
18. Apical anther appendages shorter than wide, truncate
to bilobed; style arms often with abruptly enlarged
apices
19
– Apical anther appendages about as long as wide or
longer, usually with rounded apices, rarely retuse; style
arms with abruptly enlarged apices in some Critonia
and Cronquistianthus
24
19. Corollas pubescent on inner surface; inflorescence
with strongly ascending mostly subopposite to
alternate branches; apical anther appendages bilobed;
style arm apices not clavate; Argentina, Brazil
1608. Neocabreria
– Corollas usually glabrous on inner surface; inflorescence with numerous spreading opposite branches;
apical anther appendages usually truncate
20
Compositae
20. Pappus setae slender, mostly separate; anther thecae
reddish; outer phyllaries usually persistent, phyllaries
often whitish
1579. Fleischmanniopsis
– Pappus setae stout, contiguous; anther thecae not reddish; inner phyllaries often easily deciduous, phyllaries never whitish
21
21. Corollas funnelform; leaf lamina sparsely to densely
pubescent
1581. Koanophyllon
– Corollas cylindrical and not wider at lobes than at
base; leaf lamina glabrous (without evident hairs!) 22
22. Leaves with numerous glandular punctae on lower surface; phyllaries 7–8-seriate
1575. Adenocritonia
– Leaves glabrous and lacking glandular punctae, sometimes with internal cavities appearing as translucent
spots along veins or in areolae; phyllaries 5–6-seriate
23
23. Anther filaments inserted near base of corolla; achenes
and corollas glabrous; Jamaica, Cuba
1574. Urbananthus
– Anther filaments inserted well above corolla base;
achenes sparsely glandular-punctate and setuliferous,
corolla lobes glandular-punctate outside; Mexico,
Guatemala, Honduras, El Salvador
1575. Critoniadelphus
24. Leaves with translucent or lens-like internal secretory
pockets when viewed against light, lacking glandular
punctae; coarse vines and shrubs
1572. Critonia
– Leaves lacking secretory pockets, often with glandular
punctae
25
25. Style arm appendages broad and fleshy, often rather
strap-shaped and wrinkled when dry; Andean plants
26
– Style arm appendages not prominently broadened 29
26. Achenes glabrous or with eglandular setulae, lacking
glandular punctae
27
– Achenes with numerous glandular punctae, lacking
eglandular setulae
28
27. Inflorescences with mostly alternate ascending
branches; phyllary apices acute; carpopodium symmetrical with little or no upper rim
1595. Aristeguietia
– Inflorescence with many spreading opposite branches;
phyllaries apices rounded; carpopodium asymmetrical with distinct upper rim 1606. Cronquistianthus
28. Phyllaries many-ribbed; carpopodium cylindrical and
procurrent on ribs; pappus setae flattened and smooth
on outer surface, especially near base
1599. Grosvenoria
– Phyllaries 2–4-ribbed; carpopodium annuliform or
short stopper-shaped with straight upper edge; pappus setae not flattened on outer surface
1598. Badilloa
29. Pappus setae with markedly dilated spinose apices;
capitula with 50–70 florets
1610. Amboroa
– Pappus setae without or with only slightly dilated
apices; capitula with 5–35 florets
30
30. Leaves with numerous yellowish vermiform hairs
when young; corollas with large spreading lobes from
apex of narrowly cylindrical basal tube and throat
1600. Corethamnium
– Leaves lacking yellowish vermiform hairs; corolla
lobes not appearing to spread directly from tube 31
561
31. Capitula all with distinct, sometimes elongate,
pedicels
32
– Capitula at least partially sessile or subsessile in clusters
35
32. Inner phyllaries persistent; pappus setae fragile;
inflorescences with prominently spreading opposite
branches
33
– Inner phyllaries easily deciduous; pappus setae persistent; inflorescences with ascending often alternate
branches
34
33. Pappus setae slightly but distinctly dilated at apices;
capitula with 18–35 florets; corollas glabrous; Central
America
1593. Peteravenia
– Pappus setae not broadened at apices; capitula with
10–12 florets; corolla lobes glandular-punctate outside; South America
1588. Lorentzianthus
34. Leaf lamina 3-veined from base; most capitula with
large foliose bract at base; Mexico
1580. Viereckia
– Leaf lamina pinnately veined with ascending veins;
capitula without foliose bract at base; Brazil, Uruguay
1603. Malmeanthus
35. Pappus setae barbellate below and tapering to
a smooth apex in distal half
1597. Austrocritonia
– Pappus setae lacking smooth tapering distal halves,
barbellate throughout and sometimes with broadened
or dilated apices
36
36. Leaves elliptical with widely spreading pinnate venation
37
– Leaves mostly ovate with ascending or 3-veined secondary venation
38
37. Leaves stiffly coriaceous, glandular-punctate and
scabrid on both surfaces; phyllaries partly pubescent;
corolla lobes with sclerified apices on outer surface;
capitula with c. 10 florets; style arms wider than thick;
Colombia
1602. Imeria
– Leaves thinly coriaceous, glabrous above, paler
beneath; phyllaries sparsely pubescent or glabrous;
corolla lobes lacking sclerified apices; capitula with
6–7 florets; style arms narrowed above stigmatic area
becoming terete and filiform; Venezuela
1601. Castanedia
38. Corolla lobes 2–3 times longer than wide, throat
pubescent inside; apices of pappus setae dilated
1607. Steyermarkina
– Corolla lobes about as long as wide, throat glabrous
inside; apices of pappus setae not enlarged
39
39. Carpopodium clearly procurrent on to base of ribs;
petiole more than one third the length of lamina; style
arms slender and terete above stigmatic area
1594. Critoniella
– Carpopodium not markedly extending upwards on
to base of ribs; petioles usually less than a quarter
the length of lamina; style arms slightly flattened and
broadened
40
40. Inflorescences with thyrsoid-paniculate branches;
most phyllaries easily deciduous; receptacle strongly
convex, ± hemispherical; involucres campanulate;
plants scandent; Peru
1604. Hughesia
– Inflorescence branches with dense glomerate clusters
of capitula; most phyllaries usually persistent; receptacle flat; involucres cylindrical; plants erect or arching
herbs; Colombia, Ecuador, Peru 1596. Asplundianthus
562
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
Genera of Critoniinae
1572. Critonia P. Browne
Critonia P. Browne, Civ. Nat. Hist. Jamaica: 490 (1756).
Coarse subshrubs to small trees or woody vines
or climbers. Leaves opposite, lamina elliptical to
broadly ovate, entire to serrate. Inflorescences usually thyrsoid-paniculate, with branches opposite
and usually spreading at 90° angles; involucres usually cylindrical to fusiform. Phyllaries c. 20–25,
subimbricate to weakly imbricate, 4–6-seriate, unequal, gradate, all but outer easily deciduous; receptacle flat to slightly convex, glabrous. Florets 4–12;
corollas whitish, tubular or narrowly funnelform,
glabrous outside or rarely a few glands on lobes;
lobes oblong to long-triangular, erect to slightly
spreading, smooth on inner surface, sometimes
roughened at tip outside; anther collar moderately
narrow; style base not enlarged, glabrous, arms
filiform to slightly spathulate, smooth to slightly
mamillose. Achenes prismatic, 3-ribbed, glabrous
or setuliferous; carpopodium a narrow rim or short
cylinder; pappus setae uniseriate, 25–35, persistent, congested, apices slightly enlarged and more
densely barbellate, apical cells usually acute. n =
10. Forty-three species, Central and South America, Greater Antilles.
1573. Critoniadelphus R.M. King & H. Rob.
Critoniadelphus R.M. King & H. Rob., Phytologia 22: 52
(1971).
Erect shrubs to small trees. Leaves opposite,
lamina elliptical to broadly lanceolate, entire to
serrulate. Inflorescences pyramidally paniculate,
involucres cylindrical to narrowly campanulate.
Phyllaries 25–30, strongly subimbricate to imbricate, 5–6-seriate, unequal, gradate, glabrous, inner
deciduous; receptacle flat, glabrous. Florets 3–8;
corollas white, tubular to minimally funnelform,
glabrous below lobes, lobes equilaterally triangular, smooth on inner surface, glandular-punctate
on outer; anther collar rather long-cylindrical;
style base not enlarged, glabrous, arms distinctly
broadened at apex, flattened, mostly mamillose,
becoming smooth apically. Achenes prismatic,
5-ribbed, sparsely setuliferous and glanduliferous;
carpopodium symmetrical, short-cylindrical;
pappus setae uniseriate, c. 30–35, contiguous, persistent, not broadened distally, apical cells acute.
Two species, El Salvador, Guatemala, Honduras,
Mexico.
1574. Urbananthus R.M. King & H. Rob.
Urbananthus R.M. King & H. Rob., Phytologia 22: 55 (1971).
Erect shrubs or small trees. Leaves opposite, lamina elliptical, entire to subserrulate, short- to longacuminate. Inflorescence pyramidally paniculate;
capitula cylindrical to narrowly campanulate. Phyllaries c. 20–30, 5–6-seriate, unequal, gradate, inner
deciduous; receptacle usually slightly convex, glabrous. Florets 4–10; corollas white, tubular, glabrous, lobes oblong to oblong-ovate, mostly erect,
smooth on inner surface; anther collars cylindrical; style base not enlarged, glabrous, arms with
minutely spathulate tips, mamillose except at tips.
Achenes prismatic, 5-ribbed with somewhat thickened ribs, glabrous; carpopodium distinct; pappus
setae uniseriate, c. 30, persistent, contiguous, not
broadened at tips, apical cells acute. Two species,
Cuba, Jamaica.
1575. Adenocritonia R.M. King & H. Rob.
Adenocritonia R.M. King & H. Rob., Phytologia 33: 281
(1976); Robinson, Phytologia 71: 176–180 (1991), key.
Erect shrubs. Leaves opposite, lamina ovate,
apex narrowly acuminate and entire, intervening margins serrulate to serrate. Inflorescence
a broad corymbose cyme with densely corymbose branches. Phyllaries strongly subimbricate,
6–7-seriate, markedly unequal, lower persistent,
broadly ovate, inner 12–15, 3–4-seriate, somewhat deciduous, narrowly oblong; receptacle
slightly convex, glabrous. Florets 5; corollas white,
tubular, base slightly inflated, glabrous outside
below lobes; lobes about as long as wide, inner
surface smooth with short-oblong cells, outer
with numerous glands; anther collars cylindrical;
style base not enlarged, glabrous, arms distinctly
and shortly spathulate at tip. Achenes prismatic,
5-ribbed, glabrous or sparsely glandular-punctate;
carpopodium short-cylindrical; pappus setae
uniseriate, c. 30–35, persistent, contiguous, not
noticeably broadened at tips, apical cells acute.
Three species, Guatemala, Jamaica, Mexico.
1576. Antillia R.M. King & H. Rob.
Antillia R.M. King & H. Rob., Phytologia 21: 398 (1971).
Erect subrosulate perennial herbs, subscapose.
Leaves mostly opposite, lamina oblanceolate
with tapering base, crenate-lobate. Inflorescence
scapose, with few ascending mostly elongate
branches above. Capitula broadly campanulate;
Compositae
563
phyllaries c. 25, distant to very weakly subimbricate, 2–3-seriate, mostly subequal, persistent;
receptacle slightly convex, glabrous. Florets c.
40–50; corollas white (?), funnelform, with scattered minute glands and short rather pointed
hairs on outer surface of throat and lobes; lobes
slightly longer than wide, smooth on both surfaces; anther collars cylindrical; style base not
enlarged, glabrous, arms narrowly linear, slightly
broader at tip, mamillose except at tip. Achenes
prismatic, 7–8-ribbed, setuliferous mostly on ribs;
carpopodium distinct; pappus a short crown of
deeply laciniate scales, marginal cells of scales
acute. One species, A. brachychaeta (B.L. Rob.)
R.M. King & H. Rob., Cuba.
2–3-seriate, unequal, gradate, persistent or with
a few inner deciduous; receptacle flat to slightly
convex, glabrous. Florets 13–20; corollas blue,
lavender or pale purple, narrowly funnelform,
outer surface with numerous glandular punctae
mostly on lobes and at slightly constricted top of
basal tube, lobes about as long as wide, smooth
on both surfaces, anther collars narrow; style base
not enlarged, glabrous, arms only slightly broadened at tips, mamillose except at tips. Achenes
prismatic, 4–5-ribbed, setuliferous; carpopodium
distinct; pappus setae uniseriate, c. 40, persistent,
contiguous, apical cells short-acute. One species,
E. sophiifolia (L.) R.M. King & H. Rob., Dominican
Republic, Haiti.
1577. Ciceronia Urban
1579. Fleischmanniopsis R.M. King & H. Rob.
Ciceronia Urban, Repert. Spec. Nov. Reg. Veg. 21: 324
(1925).
Fleischmanniopsis R.M. King & H. Rob., Phytologia 21: 402
(1971); King & Robinson, Phytologia 36: 193–200 (1977),
key.
Perennial rosulate herbs. Leaves densely inserted, spiralled or perhaps partially opposite,
lamina oblanceolate, lobate-crenate. Inflorescence scapiform, with few slender branches
distally. Phyllaries c. 10, weakly subimbricate,
c. 3-seriate, subequal, persistent; receptacle flat,
glabrous. Florets 9–10; corollas lavender, with
short broad basal tube shortly constricted above,
throat broad and short-funnelform with slightly
campanulate base, sparsely glandular-punctate
on outer surface, densely on outer surfaces of
lobes; lobes about as long as wide, smooth on
both surfaces; anther collars cylindrical; style
base not enlarged, glabrous, arms with stigmatic
portion slightly broadened, then narrowing and
broadening distally and smooth at apex. Achenes
prismatic, c. 8 ribbed, densely setuliferous mostly
on ribs, glandular-punctate mostly between ribs;
carpopodium distinct; pappus setae uniseriate, c.
45, barbellate, persistent, with tips of teeth and
apical cells blunt. One species, C. chaptalioides
Urban, Cuba.
Erect perennial herbs. Leaves opposite, lamina
ovate-lanceolate, usually serrate. Inflorescence
a rather diffuse corymbose to thyrsoid panicle
with densely corymbose branchlets. Phyllaries
c. 15–20, markedly subimbricate, 3–5-seriate,
markedly unequal, gradate, inner usually persistent; receptacle flat or slightly convex, glabrous.
Florets 5–10; corollas white, narrowly funnelform
without distinct constriction at top of basal tube,
nearly glabrous outside, rarely with few hairs
inside near bases of filaments; lobes about as
long as wide, smooth on both surfaces, with short
papillae only at tips; anther collar slender; style
base not enlarged, glabrous, arms rather abruptly
spathulate at tips, slender bases of appendages
short-papillose, becoming mamillose near tips.
Achenes prismatic, 4- or 5-ribbed, ribs with very
few to many setulae; carpopodium distinct; pappus
setae uniseriate, c. 30–40, persistent, contiguous to
somewhat separate, apical cells acute, sometimes
distorted. n = 10. Five species, El Salvador,
Guatemala, Mexico.
1578. Eupatorina R.M. King & H. Rob.
Eupatorina R.M. King & H. Rob., Phytologia 21: 396 (1971).
Erect perennial calcicolous herbs. Leaves opposite,
larger congested at base, decrescent and increasingly remote above, lamina deeply bipinnatifid
with small rounded to oblong ultimate segments,
fern-like. Inflorescence a lax thyrsoid panicle,
with branches ending in lax cymes; pedicels
slender. Phyllaries c. 12, moderately subimbricate,
1580. Viereckia R.M. King & H. Rob.
Viereckia R. M. King & H. Rob., Phytologia 31: 118 (1975).
Erect moderately branched shrubs. Leaves opposite, lamina deltoid, serrulate, sharply acute.
Inflorescence terminal, with small rather corymbose clusters; capitula few, usually with 1 or 2
foliose bracts immediately below involucre. Phyllaries c. 20, subimbricate, 4–5-seriate, unequal,
564
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
elliptical, entire to serrate, rarely irregularly lobed.
Inflorescences pyramidally paniculate to corymbose. Phyllaries 7–16, distant to strongly subimbricate, usually weakly subimbricate, 2–4-seriate,
unequal to subequal, mostly spreading at maturity, inner sometimes deciduous; receptacle flat to
slightly convex, glabrous, epaleaceous. Florets 5–
20; corollas whitish to greenish yellow, rarely violet, funnelform with broadly cylindrical basal tube;
lobes as wide as long to 1.5 times as long as wide,
smooth on both surfaces, with numerous clustered
short-capitate glands and sometimes a few hairs
on outer surface; anther collars cylindrical; style
base not enlarged, glabrous, arms usually distinctly
broadened and becoming smooth apically, without
glands. Achenes prismatic, 5-ribbed, ribs and upper surfaces bearing setulae, with few or no glands;
carpopodium distinct; pappus setae uniseriate, c.
30–35, persistent, usually rather stout, usually long,
rarely short or lacking, apical cells acute. n = 10,
20. Circa 120 species, neotropics.
1582. Eupatoriastrum Greenm.
Eupatoriastrum Greenm., Proc. Amer. Acad. Arts 39: 93
(1903).
Fig. 121. Compositae-Eupatorieae. Koanophyllon adamantium (Critoniinae). A Flowering shoot. B Floret. (Drawings
by Margaret Tebbs)
gradate, 2–4-costate, inner easily deciduous;
receptacle slightly convex, glabrous to scarcely
puberulous. Florets 10–12; corollas whitish (?),
narrowly funnelform, glabrous; lobes about
twice as long as wide, smooth on both surfaces;
anther collar narrowly cylindrical; style base not
enlarged, glabrous, arms linear, not broadened
distally, mamillose or shortly papillose. Achenes
prismatic, 5-ribbed, setulae mostly on ribs, a few
minute glands at upper end; carpopodium distinct; pappus setae uniseriate, c. 30–35, slender,
persistent, slightly broadened distally, apical cells
acute. One species, V. tamaulipasensis R.M. King
& H. Rob., Mexico.
1581. Koanophyllon Arruda
Fig. 121
Koanophyllon Arruda, Diss. Pl. Brazil: 38? (1810).
Eupatorium L. sect. Laevia Cabrera (1991).
Shrubs or small trees, rarely vines. Leaves opposite, rarely alternate, lamina broadly lanceolate to
Erect perennial herbs or subshrubs. Leaves opposite, lamina deltoid or broadly ovate, serrate, basal
sometimes deeply lobed. Inflorescence a very lax
thyrsoid panicle. Phyllaries c. 50, weakly to moderately subimbricate, 3–5-seriate, unequal to subequal, rather persistent; receptacle strongly convex,
paleaceous. Florets 100–300; corollas pink, purple,
red or whitish, narrowly funnelform, inner and
outer surfaces of throat glabrous; lobes about as
long as wide, smooth on both surfaces, with clustered short-stipitate glands on outer surface, with
or without hairs outside; anther collar narrowly
cylindrical; style base not enlarged, glabrous, arms
filiform with scarcely to distinctly broadened tips,
weakly mamillose except at expanded tips. Achenes
prismatic, 4–5-ribbed, with numerous setulae on
sides; carpopodium distinct; pappus setae uniseriate, 15–35, slender, persistent to ± deciduous, with
non-contiguous bases, apical cells acute. n = 16.
Four species, Costa Rica, El Salvador, Guatemala,
Mexico.
1583. Sphaereupatorium (O. Hoffm.) Kuntze ex
B.L. Rob.
Sphaereupatorium (O. Hoffm.) Kuntze ex B.L. Rob., Contr.
Gray Herb. n.s. 61: 24 (1920).
Eupatorium sect. Sphaereupatorium O. Hoffm. (1897).
Compositae
Erect perennial herbs or shrubs. Leaves opposite,
lamina broadly ovate, shallowly mucronatedentate, acuminate. Inflorescence laxly thyrsoidpaniculate, with branches and branchlets at right
angles; capitula sessile in terminal globose clusters.
Phyllaries c. 12–20, distant, 3–4-seriate, slightly
unequal, ovate-lanceolate, partially deciduous;
receptacle complex, with convex lobes, paleaceous
or with interspersed phyllaries from incompletely
separated capitula. Florets c. 11; corollas white,
narrowly funnelform, with broadly cylindrical
basal tube, outer surface sparsely glandularpunctate, inner surface glabrous; lobes about as
long as wide, smooth on both surfaces, with small
cluster of glands in middle of outer surface; anther
collar narrowly cylindrical; style base not enlarged,
glabrous, arms spathulate distally, mamillose except at tips. Achenes prismatic, 5-ribbed, with
a few setulae on ribs; carpopodium symmetrical;
pappus setae uniseriate, c. 20, scarcely flexuous,
persistent, not broadened distally, apical cells
acute. One species, S. scandens (Gardner) R.M.
King & H. Rob., Bolivia, Brazil.
1584. Bishovia R.M. King & H. Rob.
Bishovia R.M. King & H. Rob., Phytologia 39: 339 (1978).
Erect perennial herbs or subshrubs. Leaves alternate above, opposite near base, lamina ovate to
broadly ovate, serrate to sublobate, acute. Inflorescence a diffuse somewhat leafy cyme. Phyllaries c. 25, distant to scarcely subimbricate, ± biseriate, subequal, distinctly bicostate on outer surface, persistent; receptacle flat to slightly convex,
glabrous. Florets 30–60, fragrant; corollas lavender, narrowly funnelform, with cylindrical basal
tube; lobes longer than wide, smooth on both surfaces, glabrous or with minute hairs or glands, anther collar narrowly cylindrical; style base not enlarged, glabrous, arms linear, only slightly enlarged
distally, densely mamillose. Achenes prismatic, 5ribbed, setuliferous and with small glands; carpopodium distinct; pappus setae uniseriate, c. 30–
40, slender, persistent, contiguous, scarcely broadened distally, apical cells acute. n = c. 10. Two
species, Argentina, Bolivia.
1585. Nothobaccharis R.M. King & H. Rob.
Nothobaccharis R.M. King & H. Rob., Phytologia 41: 397
(1979).
Erect shrubs. Leaves densely spirally inserted,
lamina small, suborbicular to elliptical, dentate
565
to crenate. Inflorescence a dense thyrsoid panicle, with branches usually spiciform; capitula
crowded. Phyllaries c. 15, distinctly subimbricate,
3–4-seriate, markedly unequal, gradate, rather
persistent; receptacle flat to slightly convex,
glabrous. Florets 6–8; corollas whitish, narrowly
funnelform, glabrous on inner and most of outer
surfaces; lobes c. 1.5 times as long as wide, smooth
on both surfaces, outer surface with cluster of
short-stipitate glands in upper half; anther collar
narrowly cylindrical; style base not enlarged, glabrous, arms slightly enlarged distally, mamillose,
becoming smooth at tips. Achenes prismatic,
5-ribbed, glanduliferous and sparsely setuliferous;
carpopodium distinct; pappus setae uniseriate, c.
30–35, contiguous, persistent, slightly broadened
and distinctly more barbellate distally, apical cells
acute. One species, N. candolleana (Steud.) R.M.
King & H. Rob., Peru.
1586. Santosia R.M. King & H. Rob.
Santosia R.M. King & H. Rob., Phytologia 45: 463 (1980).
Woody vines. Leaves opposite, lamina ovate to
ovate-elliptical, subentire. Inflorescences elongate,
terminal, thyrsoid panicles. Phyllaries c. 13,
subimbricate, c. 3-seriate, unequal, inner rather
easily deciduous; receptacle flat to slightly convex,
glabrous. Florets 1–10; corollas white, narrowly
funnelform, with broadly cylindrical base, essentially glabrous on outer surface; lobes more than
twice as long as wide, smooth on both surfaces,
with an occasional gland on outer surface; anther
collar short-cylindrical; style base not enlarged,
glabrous, arms linear to ± filiform, mamillose
to short-papillose below, essentially smooth
at slightly broadened tips. Achenes prismatic,
5-ribbed, sparsely setuliferous mostly on ribs;
carpopodium forming a basal ring; pappus setae
uniseriate, c. 20, persistent, broadened and somewhat flattened externally near base, sometimes
slightly broadened distally, apical cells acute. One
species, S. talmonii R.M. King & H. Rob., Brazil.
1587. Grisebachianthus R.M. King & H. Rob.
Grisebachianthus R.M. King & H. Rob., Phytologia 32: 268
(1975); Borhidi, Acta Bot. Acad. Sci. Hung. 29: 181–215
(1983), key.
Erect to spreading shrubs. Leaves opposite,
lamina broadly elliptical or oblong to broadly
ovate, entire to remotely subserrulate, rounded to
shortly acute. Inflorescences terminal, cymose to
566
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
somewhat pyramidal; capitula in irregular congested glomerules. Phyllaries c. 15–25, markedly
subimbricate, 4–5-seriate, congested, unequal,
gradate, inner deciduous; receptacle flat, glabrous.
Florets 12–60; corollas white to pink or purple,
narrowly funnelform, glands on outer surface
often restricted to lobes, glabrous on inner surface;
lobes distinctly longer than wide, smooth on both
surfaces; anther collar narrowly cylindrical; style
base not enlarged, glabrous, arms linear, slightly
spathulate distally, mamillose to short-papillose
except at tip. Achenes prismatic, 5-ribbed, sparsely
setuliferous and glanduliferous; carpopodium
short-cylindrical; pappus setae uniseriate, c.
20–30, rather stout, persistent, contiguous, not to
scarcely broadened distally, apical cells obtuse to
short-acute. Eight species, Cuba.
1588. Lorentzianthus R.M. King & H. Rob.
Lorentzianthus R.M. King & H. Rob., Phytologia 32: 273
(1975).
Erect shrubs. Leaves opposite, lamina ovate, often
rather large, serrate, narrowly acuminate. Inflorescence a pyramidal panicle. Phyllaries c. 20, subimbricate, c. 5-seriate, unequal, gradate, stramineous;
receptacle slightly convex, glabrous. Florets c. 10–
12; corollas whitish to purple, narrowly funnelform
with narrowly cylindrical basal tube, mostly glabrous outside below lobes, glabrous on inner surface; lobes smooth on both surfaces, with clustered
short-stipitate glands on outer surface; anther collars narrowly cylindrical; style base not enlarged,
glabrous, arms linear, not or scarcely broadened at
tips, mamillose, becoming smooth at tip. Achenes
prismatic with broad base, 5-ribbed, setuliferous
on ribs; carpopodium annuliform; pappus setae
uniseriate, 35–40, slender, rather fragile, tips not
broadened, apical cells sharply acute. n = 10. One
species, L. viscidus (Hook. & Arn.) R.M. King & H.
Rob., Argentina, Bolivia.
1589. Chacoa R.M. King & H. Rob.
Chacoa R.M. King & H. Rob., Phytologia 32: 275 (1975).
Eupatorium L. sect. Laevia Cabrera (1991), p.p.
Erect flexuous shrubs. Leaves opposite, lamina
ovate to deltoid, serrate. Inflorescences lax terminal
pyramidal panicles. Phyllaries c. 15, very weakly
subimbricate, appearing distant, c. 2-seriate,
subequal, lanceolate to linear, rather persistent;
receptacle flat to slightly convex, glabrous. Florets
c. 20; corollas white, with slender basal tubes and
narrowly campanulate limbs, glanduliferous on
outer surface; lobes not longer than wide, smooth
on both surfaces, numerous glands on outer surface; anther collar narrowly cylindrical; style base
not enlarged, glabrous, arms filiform, only slightly
broadened near tips, short-papillose. Achenes
prismatic, with somewhat narrowed bases, setuliferous and glanduliferous; carpopodium small;
pappus setae uniseriate, c. 30, persistent, slender,
contiguous, not broadened distally, apical cells
sharply acute. One species, C. pseudoprasiifolia
(Hassl.) R.M. King & H. Rob., Argentina, Brazil,
Paraguay.
1590. Idiothamnus R.M. King & H. Rob.
Idiothamnus R.M. King & H. Rob., Phytologia 32: 277 (1975).
Erect spreading shrubs or small trees. Leaves opposite, lamina elliptical to ovate, narrowly acuminate,
serrate to remotely subserrulate. Inflorescences terminal, corymbose. Phyllaries c. 14–21, subimbricate, c. 2–3-seriate, unequal to subequal, persistent;
receptacle convex, with few paleae. Florets c. 12–20;
corollas whitish to lavender, narrowly funnelform,
with broadly cylindrical basal tube, glabrous on
outer surface below lobes; lobes about 1–2 times
as long as wide, smooth on both surfaces, with few
to many short-stipitate glands on outer surface;
anther collar narrowly cylindrical; style base not
enlarged, glabrous, arms filiform, scarcely broadened distally, slightly mamillose to smooth. Achenes prismatic, 5-ribbed, sparsely setuliferous; carpopodium distinct; pappus setae uniseriate, 20–30,
slender, contiguous, mostly persistent, sometimes
slightly broadened distally, apical cells acute. Four
species, Argentina, Brazil, Peru, Venezuela.
1591. Mexianthus B.L. Rob.
Mexianthus B.L. Rob., Contr. Gray Herb. n.s. 80: 5 (1928).
Erect subshrub. Leaves alternate, lamina ovate,
acuminate, serrate. Inflorescence a laxly thyrsoid leafy panicle, capitula in dense spherical
glomerules at tips of branchlets. Phyllaries 3(5),
1–2-seriate, distant, subequal or with 1–2 shorter
outer phyllaries, inner forming thin sheath around
floret; receptacle minute, glabrous. Floret 1;
corollas white; with short cylindrical basal tube
and broadly funnelform to slightly campanulate
limb, glabrous on outer surface below lobes; lobes
about as long as wide, slightly mamillose on inner
surface, smooth with clustered short-stipitate
glands on outer surface; anther collar narrowly
Compositae
cylindrical; style base not enlarged, glabrous,
arms with minutely but distinctly spathulate tips,
slightly mamillose below tips. Achenes rather
fusiform, 5-ribbed, glabrous below, with a few
setulae above; carpopodium indistinct; pappus
of 5–7 laciniate squamellae. One species, M.
mexicanus B.L. Rob., Mexico.
567
narrowed and sometimes long-stipitate below,
setuliferous mostly on ribs; carpopodium shortly
stopper-shaped; pappus setae uniseriate, c. 30,
deciduous, slender and non-contiguous below,
broadened distally, apical cells acute. n = 10, c. 17.
Five species, Central America.
1594. Critoniella R.M. King & H. Rob.
1592. Neohintonia R.M. King & H. Rob.
Neohintonia R.M. King & H. Rob., Phytologia 22: 143
(1971).
Scandent subshrubs or shrubs. Leaves alternate to
subopposite or opposite, lamina ovate, serrulate,
short-acuminate. Inflorescence a lax narrowly
thyrsoid leafy panicle; heads sessile in spherical
glomerules. Phyllaries 4–5, weakly subimbricate,
in 2 subequal series, oblong, persistent; receptacle
minute, glabrous. Florets 1 or rarely 2; corollas
white, with broadly cylindrical base and funnelform throat, scattered glands on outer surface,
more numerous on lobes; lobes as long as wide,
smooth on both surfaces; anther collar narrowly
cylindrical; style base not enlarged, glabrous,
arms with minutely but distinctly spathulate tips,
slightly mamillose below tips, with glands along
inner surface. Achenes prismatic, 5-ribbed, with
setulae and a few glands on sides; carpopodium
shortly stopper-shaped; pappus uniseriate, of c.
25 slender scabrid slightly deciduous scarcely
contiguous bristles, not broadened at tips, apical
cells acute. One species, N. monantha (Sch. Bip.)
R.M. King & H. Rob., Mexico.
1593. Peteravenia R.M. King & H. Rob.
Peteravenia R.M. King & H. Rob., Phytologia 21: 394 (1971).
Erect coarse herbs or subshrubs. Leaves opposite,
lamina broadly ovate to deltoid or oblong-elliptical,
usually serrulate to serrate, short-acuminate. Inflorescence pyramidally paniculate, sometimes
laxly branched. Involucre often pale or brightly
coloured, broadly campanulate. Phyllaries c. 25,
strongly subimbricate, 3–4-seriate, unequal, gradate, inner mostly persistent; receptacle broadly
convex, glabrous. Florets 18–75; corollas white,
lavender or purple, narrowly funnelform, with
outer and inner surfaces glabrous; lobes as long as
wide or longer, smooth on both surfaces; anther
collar usually narrowly cylindrical; style base
not enlarged, glabrous, arms linear, sometimes
slightly spathulate at tip, short-papillose below,
mamillose distally. Achenes prismatic, 4–5-ribbed,
Critoniella R.M. King & H. Rob., Phytologia 30: 224 (1975).
Erect herbs or shrubs. Leaves opposite, lamina
ovate to broadly ovate, serrulate to serrate, acute
to acuminate. Inflorescence a broadly corymbose
to cymose panicle; capitula sessile on congested
glomerulate branchlets. Involucres narrowly cylindrical; phyllaries c. 18–32, markedly subimbricate
to imbricate, 4–6(–7)-seriate, markedly unequal,
gradate, inner usually rather persistent; receptacle flat, glabrous. Florets 6–25; corollas white,
lavender, bluish or purple, narrowly funnelform,
glabrous on inner surface and outside below lobes;
lobes as long as wide, smooth on both surfaces,
with few glands on outer surface; anther collar
narrowly cylindrical; style base not enlarged, glabrous, arm appendages terete and filiform, densely
short-papillose. Achenes ± fusiform, 5-ribbed,
sparsely to moderately setuliferous; carpopodium
stopper-shaped; pappus setae uniseriate, c. 40,
persistent, slender, narrowly tapering to tip, apical
cells acute. n = 10. Six species, Colombia, Peru,
Venezuela.
1595. Aristeguietia R.M. King & H. Rob.
Aristeguietia R.M. King & H. Rob., Phytologia 30: 218
(1975).
Erect to procumbent shrubs to small trees. Leaves
opposite, lamina broadly ovate to linear, often
oblong or elliptical, usually densely crenulate to
dentate. Capitula few to many, corymbose, pedicellate. Phyllaries c. 25–70, strongly subimbricate,
4–6-seriate, unequal, gradate, mostly persistent;
receptacle flat to slightly convex, rarely conical,
glabrous. Florets 13–100; corollas bluish, lavender,
purple or pink, narrowly funnelform, inner and
usually outer surfaces glabrous, some species
with few hairs or few to many small glands on
lobes; lobes slightly longer than wide, smooth on
both surfaces; anther collar narrowly cylindrical;
style base not enlarged, glabrous, arms broadly
strap-shaped, often longitudinally folded, mamillose. Achenes prismatic, 5-ribbed, usually with
setulae; carpopodium indistinct; pappus setae c.
568
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
30–45, 1–2-seriate, persistent, congested, scarcely
spreading at maturity, not broadened distally,
apical cells acute. n = 10. Twenty-one species,
Chile, Colombia, Ecuador, Peru.
1596. Asplundianthus R.M. King & H. Rob.
Asplundianthus R.M. King & H. Rob., Phytologia 30: 224
(1975).
Erect to scandent shrubs or trees. Leaves opposite, lamina ovate to lanceolate, subserrate to serrate. Inflorescence usually corymbose-paniculate,
capitula sessile in glomerules. Phyllaries c. 15–
20, subimbricate, c. 3–5-seriate, markedly unequal,
gradate, inner deciduous; receptacle flat, glabrous.
Florets 6–10; corollas lilac, lavender or purple, narrowly funnelform, usually with glands on outer
surface; lobes as long as wide, smooth on both
surfaces; anther collar narrowly cylindrical; style
base not enlarged, glabrous, arms narrowly linear,
mamillose. Achenes prismatic, 5-ribbed, glabrous
or sparsely setuliferous, rarely with a few glands;
carpopodium stopper-shaped; pappus setae c. 30–
40, slender, persistent, tapering distally, apical cells
acute. n = 10. Seven species, Colombia, Ecuador,
Peru.
1597. Austrocritonia R.M. King & H. Rob.
Fig. 122
Fig. 122. Compositae-Eupatorieae. Austrocritonia angulicaulis (Critoniinae). A Flowering shoot. B Floret. (Drawings
by Margaret Tebbs)
Austrocritonia R.M. King & H. Rob., Phytologia 31: 115
(1975).
Erect shrubs or small trees. Leaves opposite, lamina
ovate to elliptical, entire or remotely serrulate to
closely serrate. Inflorescence broadly corymbose;
capitula sessile in clusters. Phyllaries 12–25,
markedly subimbricate, c. 3–5, markedly unequal,
gradate, inner easily deciduous; receptacle flat to
slightly convex, glabrous. Florets 5 or c. 10; corollas
white, narrowly funnelform, glabrous on inner
and outer surfaces; lobes ± twice as long as wide,
smooth on both surfaces, glabrous; anther collar
cylindrical; style base not enlarged, glabrous, arms
linear, mamillose. Achenes prismatic, 5-ribbed,
glanduliferous or glabrous; carpopodium distinct,
long-cylindrical; pappus setae usually uniseriate,
c. 40–50, persistent, congested, narrowed and
becoming essentially smooth distally, apical cells
acute. n = 10. Four species, Brazil.
1598. Badilloa R.M. King & H. Rob.
Badilloa R.M. King & H. Rob., Phytologia 30: 230 (1975).
Erect shrubs, pubescent. Leaves opposite, lamina
oblong to lanceolate, serrate to remotely subserrulate. Inflorescence corymbose. Phyllaries c. 16–25,
markedly subimbricate, 4-seriate, markedly unequal, gradate, inner rather deciduous; receptacle
flat to slightly convex, glabrous. Florets usually
4–10(–23); corollas white, lavender, pink or violet,
narrowly funnelform, glabrous on inner surface
and on outer surface of throat; lobes c. 1.5 times
as long as wide, smooth on both surfaces, with
glands or a few small hairs on outer surface near
tips; anther collar usually broadly cylindrical; style
base not enlarged, glabrous, arms broadly linear
or strap-shaped, slightly mamillose. Achenes prismatic, 5-ribbed, body densely glandular-punctate;
carpopodium short-cylindrical to annuliform;
pappus setae uniseriate, c. 30–35, persistent,
contiguous, not distinctly broadened nor flattened
on outer surface at base, not or slightly broadened
distally, apical cells acute. Ten species, Colombia,
Ecuador, Peru, Venezuela.
Compositae
1599. Grosvenoria R.M. King & H. Rob.
Grosvenoria R.M. King & H. Rob., Phytologia 30: 221 (1975).
Erect shrubs or small trees. Leaves opposite, lamina ovate to narrowly elliptical, entire to remotely
serrate, sharply acute to short-acuminate.
Inflorescence broadly corymbose paniculate. Phyllaries 12–15, markedly subimbricate, 3–5-seriate,
markedly unequal, gradate, inner easily deciduous;
receptacle flat to slightly convex, glabrous. Florets
5–10, fragrant; corollas pink to white, narrowly
funnelform, with tube glabrous, without hairs on
inner surface; cells of throat oblong with sinuous
lateral walls; lobes c. 1.5 times as long as wide,
smooth on both surfaces; anther collar narrowly
cylindrical; style base not enlarged, glabrous,
arms very elongate, broadly linear, somewhat
broader near tip, rounded to obtusely mamillose
Achenes prismatic, 5-ribbed, glandular-punctate;
carpopodium procurrent on to bases of ribs;
pappus setae uniseriate, c. 30–40, persistent,
congested, bases broadened and flattened, longer
setae distinctly broadened distally, apical cells
acute to obtuse. Four species, Ecuador, Peru.
1600. Corethamnium R.M. King & H. Rob.
Corethamnium R.M. King & H. Rob., Phytologia 39: 55
(1978).
Erect shrubs. Leaves opposite, lamina ovate to
suborbicular, crenate-serrulate, short-obtuse to
rounded. Inflorescence of dense small corymbose
panicles on leafy branches. Phyllaries c. 16–18,
subimbricate, c. 4–5-seriate, markedly unequal,
gradate, mostly persistent; receptacle flat, glabrous. Florets c. 6; corollas white, with narrowly
cylindrical tube and throat without external
differentiation, with only lobes spreading, outer
surface very sparsely glanduliferous, inner surface
glabrous; lobes 3 times as long as wide, smooth on
both surfaces; anther collars cylindrical; style base
lacking basal node, glabrous, arms narrowly linear
to filiform, terete distally, distinctly papillose.
Achenes prismatic, 5-ribbed, glabrous or with very
few small glands; carpopodium a short cylindrical
ring without a distinct upper rim; pappus setae
1–2-seriate, c. 45, coarse, congested, persistent,
less barbellate and narrowed distally, apical cells
subacute. One species, C. chocoensis R.M. King &
H. Rob., Colombia.
1601. Castanedia R.M. King & H. Rob.
Castanedia R.M. King & H. Rob., Phytologia 39: 58 (1978)
(‘Castenedia’).
569
Erect shrubs. Leaves opposite, lamina elliptical
or elliptical-oblong to slightly obovate, remotely
serrulate, short-acute to obtuse. Inflorescence
a dense corymbose panicle. Involucre cylindrical; phyllaries c. 25, subimbricate, c. 4-seriate,
markedly unequal, gradate, inner easily deciduous;
receptacle flat, glabrous. Florets 6–7; corollas white,
narrowly funnelform, sparsely glanduliferous on
narrow tube and on throat, more glands on lobes;
lobes slightly longer than wide, smooth on both
surfaces, anther collar cylindrical; style base not
enlarged, glabrous, arms with broader stigmatic
part, in appendage becoming filiform and terete
with short papillae. Achenes prismatic, 5-ribbed,
glabrous; carpopodium shortly stopper-shaped;
pappus setae c. 60, congested, persistent, c.
2-seriate, narrowed distally, apical cells sharply
acute. One species, C. santamartensis R.M. King &
H. Rob., Colombia.
1602. Imeria R.M. King & H. Rob.
Imeria R.M. King & H. Rob., Phytologia 32: 271 (1975).
Erect slender shrubs or small trees. Leaves opposite, lamina thickly coriaceous to subcoriaceous,
ovate to elliptical, entire or serrate, shortly acute
to acuminate. Inflorescences terminal, corymbose,
capitula sessile in dense clusters. Phyllaries c. 20–
25, markedly subimbricate, 4–5-seriate, unequal,
gradate, outer persistent, inner deciduous; receptacle slightly convex, sparsely hirsute. Florets 8–
10; corollas rose-coloured or pink, narrowly funnelform, glabrous, lobes slightly longer than wide,
smooth on inner surface, densely mamillose to papillose outside on the tips forming sclerified cap;
anther collars shortly cylindrical; style base not
enlarged, glabrous, arms linear, slightly mamillose. Achenes prismatic, 7–9-ribbed, glabrous; carpopodium very shortly cylindrical; pappus setae
c. 50, 1–2-seriate, congested, subpersistent, longer
setae broadened at tips, apical cells subacute to
obtuse. One species, I. memorabilis (Maguire &
Wurd.) R.M. King & H. Rob., Brazil, Venezuela.
1603. Malmeanthus R.M. King & H. Rob.
Malmeanthus R.M. King & H. Rob., Phytologia 47: 225
(1980).
Eupatorium L. sect. Gyptis (Cass.) Cabrera (1991), p.p.
Erect shrubs. Leaves opposite, lamina ovate,
serrulate to subentire, scarcely acuminate. Inflorescences terminal, corymbose-paniculate, with
ascending mostly alternate branching. Involucres
570
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
campanulate; phyllaries c. 20, strongly subimbricate, c. 4-seriate, markedly unequal, gradate,
stramineous, inner easily deciduous; receptacle
slightly convex, glabrous. Florets 5–22; corollas
whitish, narrowly funnelform, with broad basal
tube, glabrous; lobes about as long as wide to twice
as long as wide, smooth, with or without sparse
glands on outer surface; anther collar cylindrical;
style base not enlarged, glabrous, arms narrowly
linear, slightly to distinctly mamillose. Achenes
prismatic, narrowed at base, 5-costate, setuliferous
or glandular-punctate; carpopodium obsolete;
pappus setae mostly uniseriate, c. 30–35, slender,
persistent, not or scarcely broader and not more
barbellate distally, apical cells acute. Three species,
Argentina, Brazil, Paraguay, Uruguay.
slightly convex, glabrous. Florets 3–12; corollas
white, with constricted basal tube and narrowly
funnelform or campanulate limb, usually glanduliferous outside, glabrous or rarely puberulous
inside; lobes 1–2 times as long as wide, smooth;
anther collar cylindrical; style base not enlarged,
glabrous; style arms with strongly knob-like
smooth tips, mamillose below. Achenes prismatic,
5-ribbed, with minutely gland-tipped hairs and/or
setulae, base narrowed and sometimes striate;
carpopodium slightly stopper-shaped; pappus of
setae, broad laciniate scales, vestigial squamellae
or lacking, setae barbellate to subplumose, persistent, not broadened distally, apical cells acute.
n = 10. Thirty-seven species, Argentina, Bolivia,
Brazil, Chile, Ecuador, Peru.
1604. Hughesia R.M. King & H. Rob.
1606. Cronquistianthus R.M. King & H. Rob.
Hughesia R.M. King & H. Rob., Phytologia 47: 252 (1980).
Cronquistianthus R.M. King & H. Rob., Phytologia 23: 410
(1972).
Woody vines. Leaves opposite, lamina ovate,
subentire, remotely and minutely serrulate,
shortly acute, minutely acuminate. Inflorescences
terminal on lateral branches, distinctly thyrsoidpaniculate with spreading branches; capitula
sessile or subsessile in small clusters. Phyllaries c.
18, markedly subimbricate, c. 4-seriate, markedly
unequal, gradate, inner easily deciduous; receptacle convex or hemispherical, glabrous. Florets c. 9;
corollas purple in distal half when dry, narrowly
funnelform with broadly cylindrical basal tubes,
glabrous; lobes about as long as wide, smooth;
anther collars cylindrical; style base not enlarged,
glabrous, arms linear, densely mamillose or
short-papillose. Achenes prismatic, 5-ribbed, with
short setulae above, glabrous below; carpopodium
short-cylindrical, slightly procurrent on to ribs;
pappus setae uniseriate, c. 30, slender, persistent,
not or scarcely broadened at tips, apical cells acute.
One species, H. reginae R.M. King & H. Rob., Peru.
1605. Ophryosporus Meyen
Ophryosporus Meyen, Reise Erde 1: 402 (1834); Robinson,
Proc. Amer. Acad. Arts 42: 17–27 (1906), rev.
Erect herbs or subshrubs, rarely scandent. Leaves
usually opposite, lamina broadly lanceolate to
elliptical, coarsely to scarcely serrate, acute to
short-acuminate. Inflorescence corymbose or
thyrsoid, with corymbose branches; lateral capitula sometimes from axils of lower phyllaries
of central capitula. Phyllaries 4–8, distant, 1–2seriate, mostly subequal, persistent; receptacle
Erect or flexuous shrubs. Leaves opposite, lamina ovate or lanceolate to linear, entire to serrate,
mostly acute, not acuminate. Inflorescence a terminal corymbose panicle. Phyllaries 12–25, markedly
subimbricate, 3–5-seriate, markedly unequal, gradate, inner easily deciduous; receptacle flat, glabrous. Florets 8–18, fragrant; corollas white, lavender or bluish, narrowly funnelform with broadly
cylindrical basal tube, outer surface glabrous or
with few glands or scattered hairs, inner usually
glabrous, rarely with small hairs; lobes as long
as wide or slightly longer, smooth; anther collar
narrowly cylindrical; style base not enlarged, glabrous, arms broadly linear or narrowly linear with
slightly dilated tips, short-papillose. Achenes prismatic, usually 5-ribbed, with few to many setulae predominantly on ribs, usually without glands;
carpopodium distinct; pappus setae uniseriate, c.
30–35, persistent, contiguous, usually ± as long as
corolla, more coarsely barbellate below, narrowed
and sometimes nearly smooth at tips, apical cells
acute. 20 species, Colombia, Ecuador, Peru.
Turner (1991b) did not accept Cronquistianthus other than as a group of species within
Eupatorium s. lat.
1607. Steyermarkina R.M. King & H. Rob.
Steyermarkina R.M. King & H. Rob., Phytologia 22: 43
(1971).
Eupatorium L. sect. Steyermarkina (R.M. King & H. Rob.)
Cabrera (1991).
Compositae
Vines or flexuous shrubs. Leaves opposite, lamina
ovate, entire, obtuse to acute. Inflorescence a lax
thyrsoid panicle. Phyllaries c. 15–20, markedly
subimbricate, 4–5-seriate, strongly unequal, gradate; receptacle convex to slightly conical, usually
glabrous. Florets 3–5; corollas white, narrowly
funnelform, outer surface glabrous or with minute
glands or large hairs on base of throat and lobes,
inner densely pilose on throat; lobes c. 2–4 times
as long as wide, cut to below bases of anther
sacs, smooth; anther collar narrowly cylindrical;
style base not enlarged, glabrous, arms linear,
mamillose. Achenes prismatic, 5–6-ribbed, densely
short-setuliferous; carpopodium distinct, short;
pappus setae mostly uniseriate, c. 30, slender,
congested, persistent, slightly flattened on outer
surface, longer setae distinctly broadened at tips,
apical cells shortly to sharply acute. Four species,
Brazil, Venezuela.
1608. Neocabreria R.M. King & H. Rob.
Neocabreria R.M. King & H. Rob., Phytologia 23: 151 (1972);
King & Robinson, Phytologia 38: 424–428 (1978), key.
Eupatorium L. sect. Gyptis (Cass.) Cabrera (1991), p.p.
Eupatorium L. sect. Heterocondylus (R.M. King & H. Rob.)
Cabrera (1991), p.p.
Erect subshrubs. Leaves opposite, lamina narrowly
elliptical, closely serrulate to crenate-serrulate. Inflorescence a corymbose panicle. Phyllaries 25–30,
strongly subimbricate, 3–4-seriate, markedly unequal, gradate, inner easily deciduous; receptacle
flat to slightly convex, glabrous to densely hirsute.
Florets 6–25; corollas white to rose-purple, narrowly funnelform, glabrous on outer surface, with
numerous hairs on inner; lobes as long as wide
or longer, smooth, glabrous or with sparse glands
on outer surface, anther collar cylindrical; style
base not enlarged, glabrous, arms broadly linear,
almost smooth or distinctly mamillose. Achenes
prismatic, 4–5-ribbed, with narrow base, setuliferous or glandular-punctate above; carpopodium
indistinct; pappus setae uniseriate, 30–40, slender, contiguous, persistent, not broadened distally,
apical cells acute. Five species, Argentina, Brazil,
Paraguay.
1609. Uleophytum Hieron.
Uleophytum Hieron., Verh. Bot. Vereins Prov. Brandenburg
48: 198 (1906).
Woody vines. Leaves opposite, lamina broadly
oblong-ovate, minutely denticulate, acuminate.
571
Inflorescence of numerous capitula clustered in
axils of leaves. Phyllaries c. 25, subimbricate,
3–4-seriate, unequal, gradate, mostly persistent;
receptacle flat, glabrous. Florets c. 55–60; corollas
whitish (?), narrowly funnelform, glabrous on
inner and lower outside surfaces; lobes slightly
longer than wide, smooth, with numerous glands
clustered on outer surface; anther collar cylindrical; style base not enlarged, glabrous, arms
narrowly linear, nearly filiform below, slightly
broadened distally, mamillose. Achenes prismatic,
4–5-ribbed, glabrous except for few glands near
top; carpopodium stopper-shaped; pappus setae
uniseriate, c. 30, closely contiguous, persistent, not
or slightly broader and not or slightly more barbellate distally, apical cells obtuse to short-acute.
One species, U. scandens Hieron., Peru.
1610. Amboroa Cabrera
Amboroa Cabrera, Bol. Soc. Argent. Bot. 6: 91 (1956).
Small erect subshrubs or shrubs. Leaves opposite,
lamina narrowly elliptical, remotely serrulate,
sharply acute. Inflorescences on slender peduncles,
with solitary capitula or a pair of sessile capitula.
Phyllaries c. 25–40, markedly subimbricate, c.
4–5-seriate, unequal, gradate, inner persistent;
receptacle convex, alveolate, glabrous. Florets
50–≥80; corollas white, narrowly funnelform, with
long cylindrical basal tube and rather cylindrical
throat, glabrous; lobes short compared to length of
corolla, longer than wide, smooth; anther collars
cylindrical; style base not enlarged, glabrous, arms
narrowly linear, becoming nearly filiform, with
dense short-erect papillae. Achenes prismatic,
5–6-ribbed, glabrous; carpopodium indistinct;
pappus setae uniseriate, c. 15–25, non-contiguous,
slender, with bases broad and flattened, extremely
slender and smooth for most of length, becoming greatly enlarged and subplumose distally
with many large densely projecting obtuse or
short-acute cells. Two species, Bolivia, Peru.
1611. Tuberostylis Steetz
Tuberostylis Steetz in Seemann, Bot. Voy. Herald 142
(1854).
Creeping to scandent herbs or shrubs. Leaves
opposite, lamina slightly fleshy, glabrous, obovate
to elliptical, entire to crenulate, obtuse to shortacuminate. Inflorescences terminal on lateral
branches or sessile in axils of leaves; capitula
sessile in small panicles or axillary fascicles. Phyl-
572
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
laries 25–30, subimbricate, 4–5-seriate, unequal,
gradate, spreading at maturity or inner deciduous;
receptacle flat or slightly convex, glabrous or with
narrow paleae inside marginal florets. Florets
c. 10–20; corollas white, narrowly tubular with
slightly thickened base, glabrous on inner surface
and lower outside surface; lobes 1–3 times as long
as wide, smooth, mamillose at tips, a few minute
glands on outer surface, anther collar cylindrical;
style base not enlarged, glabrous, arms narrowly
linear, slightly mamillose, slightly broadened
distally. Achenes prismatic to cylindrical, 5-veined
or 5-ribbed, whitish and strongly corticated when
mature, glabrous; carpopodium short, indistinct;
pappus absent. Two species, Colombia, Ecuador,
Panama.
XXX.16. Subtribe Hebecliniinae R.M. King &
H. Rob. (1980).
Erect perennial herbs or shrubs to small trees,
never rosulate. Leaves usually opposite, often with
long petioles; lamina often broad with rounded
to cordate bases. Inflorescence laxly cymose to
densely corymbose or thyrsoid-paniculate, with
clustered capitula; phyllaries usually strongly
subimbricate, gradate, with inner phyllaries easily
deciduous, rarely distant and persistent; receptacle
slightly convex to strongly hemispherical, usually
epaleaceous, often with hairs. Florets 4–150; corollas narrowly funnelform, rarely with hairs inside
of throat, cells of throat elongate with sinuous
lateral walls; lobes triangular, usually about as
long as wide, smooth on both surfaces; anther
collar usually elongate, 5–10 times as long as wide,
of even width; apical anther appendage usually
longer than wide, style base not enlarged, glabrous;
arms narrowly linear to filiform, mamillose to
short-papillose, rarely with enlarged tips. Achenes
prismatic, 5-ribbed; carpopodium annuliform to
turbinate, procurrent on to ribs of achene; pappus
setae usually uniseriate, rarely biseriate, usually
of numerous capillary setae, usually persistent,
apical cells sharply acute.
Key to the Genera
1. Receptacle paleaceous; florets numerous (c. 200) per
capitulum
1618. Matudina
– Receptacle epaleaceous (rarely paleaceous in Decachaeta); florets few to many (4–80) per capitulum
2
2. Pappus absent or setae easily deciduous
3
– Pappus setae present and persistent
4
3. Pappus absent; receptacle glabrous; phyllaries ± biseriate
1616. Erythradenia
– Pappus present, setae deciduous; receptacle pubescent
(rarely paleaceous); phyllaries 3–4-seriate
1615. Decachaeta
4. Phyllaries distant and subequal; achenes with a stipitate base
1613. Amolinia
– Phyllaries imbricate and gradate; achenes prismatic 5
5. Receptacle markedly convex to hemispherical; style
arm appendages filiform
1612. Hebeclinium
– Receptacle slightly convex; style arm appendages linear to slightly clavate
6
6. Carpopodium symmetrical; receptacle sparsely to
densely pubescent, rarely glabrous 1614. Bartlettina
– Carpopodium strongly asymmetrical; receptacle glabrous
1617. Guayania
Genera of Hebecliniinae
1612. Hebeclinium DC.
Hebeclinium DC., Prodr. 5: 136 (1836); Diaz Piedrahita &
Mendez Ramirez, Revista Acad. Colomb. Ci. Exact. 24:
25–44 (2000), reg. rev.
Eupatorium section Hebeclinium (DC.) Benth. ex Baker
(1876).
Large herbs or subshrubs. Leaves opposite, lamina
broadly ovate to deltoid or lanceolate, usually
crenate or serrate-pinnate. Inflorescence a lax
cyme with widely spreading branches. Involucres
broadly campanulate; phyllaries c. 25–40, 3–5seriate, markedly unequal, outer persistent, inner
deciduous; receptacle hemispherical, sclerified
throughout, glabrous to densely hirsute. Florets
(12–)20–80; corollas white or pink, outer surface
glabrous below, inner surface of throat in some
species with numerous hairs; lobes slightly longer
than wide, usually with prominent multicellular
uniseriate hairs and few glands on outer surface; anther collar usually slender; apical anther
appendages large, ovate-triangular to oblong,
slightly longer than wide; style arms narrowly
filiform, terete, mamillose, sometimes somewhat
broadened distally. Achenes narrowed below,
4–5-ribbed, sometimes setulose; carpopodium
scarcely distinct; pappus setae uniseriate, c.
30–40, barbellate, slender, persistent, capillary,
sometimes broadened distally. n = 10. Twenty
species, neotropics.
1613. Amolinia R.M. King & H. Rob.
Amolinia R.M. King & H. Rob., Phytologia 24: 265 (1972).
Erect shrubs or small trees. Leaves opposite, lamina
ovate, entire, narrowly acuminate. Inflorescence
Compositae
a corymbose panicle. Involucres narrowly campanulate; phyllaries c. 15, ± distant, 2–3-seriate,
slightly unequal, linear; receptacle slightly convex,
usually short-pubescent. Florets 20–25; corollas
white, with cylindrical basal tube, outer surface
sparsely glanduliferous, inner surface smooth
and glabrous; lobes ovate-triangular, anther collar
slender and elongate; apical anther appendage
large, oblong, slightly longer than wide; style arms
narrowly linear, mamillose. Achenes elongate,
5-ribbed, densely glanduliferous and sparsely
setuliferous; carpopodium very short; pappus
setae uniseriate, c. 30 (40–50?), slender, barbellate,
persistent, not broadened distally. n = 10. One
species, A. heydeana (B.L. Rob.) R.M. King & H.
Rob., Guatemala, Mexico.
1614. Bartlettina R.M. King & H. Rob.
Bartlettina R.M. King & H. Rob., Phytologia 22: 160 (1971).
Erect shrubs or small trees. Leaves opposite, lamina
lanceolate to broadly ovate. Inflorescence usually
corymbose-paniculate. Involucres broadly campanulate; phyllaries 3–5-seriate, c. 20–50, weakly
to strongly subimbricate, inner deciduous; receptacle broadly convex, usually sparsely to densely
pubescent. Florets 8–150; corollas white, lavender,
blue or purple, inner surface glabrous; lobes triangular, outer surface usually densely puberulous,
often glanduliferous; anther collars very elongate;
apical anther appendage large, oblong-ovate to
long-triangular, longer than wide; style arms
narrowly linear, sometimes slightly broadened
distally, nearly smooth to short-papillose. Achenes
5-ribbed, glabrous to sparsely setuliferous, rarely
with glands; carpopodium slightly to distinctly
enlarged, symmetrical; pappus setae 1–2-seriate,
c. 30–40, slender, barbellate, persistent, rarely
broadened distally. n = 10, 16. Circa 37 species,
tropical Central and South America.
573
narrowly funnelform, with cylindrical basal tube,
outer surface glabrous or sparsely glanduliferous;
lobes triangular; anther collar rather narrow, often
long; apical anther appendage short, much wider
than long, margin strongly reflexed; style arms narrowly linear, slightly broadened distally, mamillose.
Achenes 4–5-ribbed, with numerous setulae mostly
on ribs; carpopodium short-cylindrical or stoppershaped; pappus setae uniseriate, c. 10–30, slender,
barbellate, somewhat deciduous, often distinctly
dilated distally. n = 16. Seven species, Costa Rica,
Guatemala, Mexico, Panama.
1616. Erythradenia (B.L. Rob.) R.M. King &
H. Rob.
Erythradenia (B.L. Rob.) R.M. King & H. Rob., Brittonia 21:
285 (1969).
Erect or arching subshrubs, with numerous
reddish or yellowish glandular punctae. Leaves
alternate, lamina broadly ovate, often with lobed
dentate margins, shortly and often broadly acuminate. Inflorescence an elongate leafy thyrsoid
panicle with sessile or subsessile capitula in small
clusters. Involucres short, broadly campanulate;
phyllaries c. 10, weakly subimbricate, ± biseriate,
somewhat unequal, persistent; receptacle slightly
convex, glabrous. Florets c. 6; corollas white,
with broadly cylindrical basal tube, outer surface
glanduliferous, inner glabrous; lobes triangular,
about as long as wide, with numerous glands on
outer surface; anther collar narrowly cylindrical;
apical anther appendage very short, much wider
than long, reflexed; style arm appendages narrowly
linear, slightly broader distally, mamillose. Achenes 5-ribbed, setuliferous; carpopodium a narrow
rim, not procurrent on to ribs; pappus setae
obsolete, short, barbellate, persistent, scarcely
distinguishable from setulae of achene ribs. n = 16.
One species, E. pyramidalis (B.L. Rob.) R.M. King
& H. Rob., Mexico.
1615. Decachaeta DC.
Decachaeta DC., Prodr. 5: 133 (1836); King & Robinson,
Brittonia 21: 275–284 (1969), rev.
Erect or arching subshrubs or shrubs. Leaves alternate or sometimes opposite, lamina elliptical, ovate
or suborbicular, sometimes rather lobed, margins
serrate or crenate, acuminate. Inflorescence a thyrsoid panicle, usually leafy. Involucres campanulate;
phyllaries 3–4-seriate, c. 10–15, subimbricate; receptacle slightly to strongly convex, sometimes hirsute. Florets 4–30; corollas white, sometimes bluish,
1617. Guayania R.M. King & H. Rob.
Guayania R.M. King & H. Rob., Phytologia 21: 302 (1971);
Pruski, Fl. Venez. Guayana 3: 293–295 (1997), rev.
Erect perennial herbs or shrubs. Leaves opposite,
lamina elliptical to broadly ovate, serrate. Inflorescences strongly cymose, ultimate branchlets with
clusters of sessile or subsessile capitula. Involucres
campanulate; phyllaries c. 12–25, subimbricate,
± 3–4-seriate, unequal, rather persistent; receptacle convex to hemispherical, glabrous. Florets
574
A.A. Anderberg, B.G. Baldwin, R.G. Bayer et al.
5–25; corollas bluish-white, lavender or white,
with cylindrical basal tube, outer surface glabrous
below lobes, inner glabrous; lobes triangular,
outer surface densely short-pubescent; anther
collars slender; apical anther appendage large,
triangular, longer than wide; style arm appendages
filiform, essentially terete, densely short-papillose.
Achenes 5-ribbed, glabrous or with few setulae;
carpopodium strongly asymmetrical; pappus setae
uniseriate, c. 30–40, slender, barbellate, persistent,
narrowed distally. Five species, Brazil, Colombia,
Guyana, Venezuela.
1618. Matudina R.M. King & H. Rob.
Matudina R.M. King & H. Rob., Phytologia 26: 171 (1973).
Coarse sprawling to erect calciphilous subshrubs
or shrubs. Leaves opposite, lamina broadly ovate,
dentate-lobate. Inflorescences subcymose. Capitula broadly campanulate; subinvolucral bracts and
phyllaries c. 75–125, 5–6-seriate, subequal, narrowly lanceolate; receptacle broadly convex, paleaceous. Florets c. 200; corollas white, nearly cylindrical, with throat slightly constricted above, glabrous except for few glands on outer surface of
lobes; lobes small compared to length of corolla,
about as long as wide, smooth on inner and most
of outer surface: anther collar slender; apical anther appendages large, oblong, slightly longer than
wide, not retuse or grooved; style arms narrowly
linear, slightly broadened distally, ± mamillose,
apex obtuse. Achenes narrowly fusiform, 5-ribbed,
with short setulae mostly on ribs; carpopodium
symmetrical, shortly stopper-shaped; pappus setae uniseriate, c. 15–22, slender, barbellate, noncontiguous or scarcely contiguous, rather easily
deciduous, broadened and more barbellate distally.
n = 16. One species, M. corvi (McVaugh) R.M. King
& H. Rob., Mexico.
XXX.17. Subtribe Neomirandeinae R.M.
King & H. Rob. (1980).
Large herbs or shrubs to small trees, epiphytic or
humicolous. Leaves opposite or whorled, lamina
deltoid or aceriform to elliptical or oblong, often
slightly fleshy, entire to coarsely lobed and dentate. Inflorescence a broadly cymose or corymbose panicle, with clustered capitula; phyllaries c.
9–28, strongly subimbricate, 3–4-seriate, gradate,
ecostate, inner somewhat deciduous, outer persistent; receptacle flat or slightly convex, without
paleae, with or without hairs. Florets 2–28; corollas
white to reddish purple, narrowly funnelform, with
or without hairs inside throat; lobes triangular to
narrowly oblong, smooth on both surfaces; anther
collar elongate, 5–10 times as long as wide, of even
width; apical anther appendages longer than wide;
style base with or without enlargement, glabrous;
style arms narrowly linear, not strongly clavate
at tip. Achenes prismatic, 5-ribbed; carpopodium
short; pappus setae numerous, persistent, capillary,
apical cells obtuse to acute.
Bremer et al. (1994) suggested that the subtribe
be synonymized with Hebecliniinae, to date the
only author to have done so. Turner (1997) left the
‘Hebeclinium group’ and the ‘Neomirandea group’
separate. We retain the two separate in this account.
Only one genus:
1619. Neomirandea R.M. King & H. Rob.
Neomirandea R.M. King & H. Rob., Phytologia 19: 360
(1970); King & Robinson, Phytologia 19: 305–310 (1970),
rev.; King & Robinson, Ann. Missouri Bot. Gard. 62:
888–1004 (1976), reg. rev.; Scott, Systematic studies of
Neomirandea (Asteraceae–Eupatorieae). Unpubl. Ph.D.
Thesis, Univ. Austin Texas (1985), rev.
Characters of the subtribe. n = 17, 20, c. 24,
25. Twenty-eight species, Colombia, Costa Rica,
Ecuador, Mexico, Panama.
Asteroid Genus
of Uncertain Placement
C. Jeffrey
1620. Symphyllocarpus Maxim.
Symphyllocarpus Maxim., Prim. Fl. Amur.: 151, t. 8, f. 1
(1859); Smol’yaninova, Bot. Mat. Gerb. Bot. Inst. Komarova
20: 282–288 (1960), morph., syst.
Annual herb; leaves alternate, lanceolate, apically
1–3-denticulate, glabrous. Capitula congested 2–4
together in the stem bifurcations, globose, heterogamous, disciform to minutely radiate. Phyllaries
subuniseriate, equal, imbricate, membranous, margins transparent. Receptacle flat, paleate. Outer florets pistillate, numerous in several series; corolla
filiform, tubular or minutely radiate, (2–)3(–4)lobed, pale yellow. Inner florets few (6–20), perfect; corolla with campanulate 4-lobed limb, anthers ecalcarate, shortly caudate, apically rounded
to truncate, unappendaged; style arms short, ob-
Compositae
long, with sweeping hairs extending to shortly below the bifurcation; ovary adnate to 1 or 2 of the
receptacular scales. Achenes homomorphic, terete,
smooth, glandular and sparsely pilose with long,
apically divergent twin hairs, adnate to 1 or rarely
2 of the receptacular scales. Pappus absent. One
species, S. exilis Maxim., north-east Asia.
575
The lack of palynological, carpological and
molecular data precludes the assignment of this
undoubtedly Asteroid genus to a more definite
systematic position. Smol’yaninova made it the
type of its own subtribe of Inuleae which would
now be placed in Gnaphalieae, where the genus
would undoubtedly be anomalous.
Selected Bibliography to Compositae
Ahlstrand, L. 1988. Embryology of Calenduleae (Compositae). Nordic J. Bot. 5: 79–97.
Ahlstrand, L. 1992. Contributions to the embryology of Arctotideae (Compositae). The genera Dymondia Compton, Cullumia R. Br., Didelta L’Herit. and Heterolepis
Cass. Comp. Newslett. 22: 1–4.
Albach, D.C., Soltis, P.S., Olmstead, R.G. 2001. Phylogenetic
analysis of Asteridae based on sequences of four genes.
Ann. Missouri Bot. Gard. 88: 163–212.
Alvarenga, S.A.V., Ferreira, M.J.P., Rodrigues, G.V., Emerenciano, V.P. 2005. A general survey and some taxonomic implications of diterpenes in the Asteraceae.
Bot. J. Linn. Soc. 147: 291–308.
Anderberg, A.A. 1982. The genus Anvillea (Compositae).
Nordic J. Bot. 2: 297–305.
Anderberg, A.A. 1985. The genus Iphonia (CompositaeInuleae). Nordic J. Bot. 5: 169–194.
Anderberg, A.A. 1989. Phylogeny and reclassification of the
tribe Inuleae (Asteraceae). Canad. J. Bot. 67: 2277–
2296.
Anderberg, A.A. 1991a. Taxonomy and phylogeny of the
tribe Gnaphalieae (Asteraceae). Opera Bot. 104: 1–195.
Anderberg, A.A. 1991b. Taxonomy and phylogeny of the
tribe Inuleae (Asteraceae). Pl. Syst. Evol. 176: 75–123.
Anderberg, A.A. 1991c. Taxonomy and phylogeny of the
tribe Plucheeae (Asteraceae). Pl. Syst. Evol. 176: 145–
177.
Anderberg, A.A., Eldenäs, P., Bayer, R.J., Englund, M. 2005.
Evolutionary relationships in the Asteraceae tribe Inuleae (incl. Plucheeae) evidenced by DNA sequences
of ndhF; with notes on the systematic position of some
aberrant genera. Organism Diversity Evol. 5: 135–146.
Aristeguieta, L. 1964. Lepidesmia. Compositae. In: Lasser, T.
(ed.) Flora de Venezuela, vol. 10, 2. Caracas: Inst.
Botanico, pp. 529–531.
Ariza Espinar, L., Cerana, M.M. 1986. Contribucion al
conocimiento de las especies de Stevia (Asteraceae)
del centro de Argentina. Bol. Acad. Nac. Ci. Cordoba
57: 381–400.
Baagøe, J. 1980. SEM studies in ligules of Lactuceae (Compositae). Bot. Tidsskr. 75: 199–217.
Babcock, E.B., Stebbins, G.L. 1943. Systematic studies of
Cichoroideae. Univ. Calif. Publ. Bot. 18: 227–240.
Badillo, V.M. 1992. Anotaciones adicionales al género
Tamananthus Badillo (Heliantheae-Asteraceae).
Ernstia 2: 17–19.
Bain, J.F., Tyson, S., Bray, D.F. 1997. Variation in pollen wall
ultrastructure in New World Senecioneae (Asteraceae),
with special reference to Packera. Canad. J. Bot. 75:
730–735.
Baldwin, B.G. 2003a. A phylogenetic perspective on the origin and evolution of Madiinae. In: Carlquist, S., Baldwin, B.G., Carr, G.D. (eds) Tarweeds & silverswords:
evolution of the Madiinae (Asteraceae). St. Louis: Missouri Botanical Garden Press, pp. 193–228.
Baldwin, B.G. 2003b. Characteristics and diversity of
Madiinae. In: Carlquist, S., Baldwin, B.G., Carr, G.D.
(eds) Tarweeds & silverswords: evolution of the
Madiinae (Asteraceae). St. Louis: Missouri Botanical
Garden Press, pp. 17–52.
Baldwin, B.G., Wessa, B.L. 2000a. Origin and relationships
of the tarweed–silversword lineage (Compositae–
Madiinae). Amer. J. Bot. 87: 1890–1908.
Baldwin, B.G., Wessa, B.L. 2000b. Phylogenetic placement of
Pelucha and new subtribes in Helenieae sensu stricto
(Compositae). Syst. Bot. 25: 522–538.
Baldwin, B G., Wessa, B.L., Panero, J.L. 2002. Nuclear rDNA
evidence for major lineages of helenioid Heliantheae
(Compositae). Syst. Bot. 27: 161–198.
Baldwin, B.G., Wessa, B.L., Panero, J.L. 2003. Evolutionary
insights from a putative taxonomic garbage can. Tribe
Helenieae revisited and revised. Comp. Newslett. 40: 8.
Barclay, H.S., Earle, F.R. 1965. The search for new industrial
crops. V. The South African Calenduleae (Compositae)
as a source of new oil seeds. Econ. Bot. 19: 33–43.
Barkley, T.M. 1999. The segregates of Senecio, s.l., and Cacalia, s.l., in the flora of North America north of Mexico.
Sida 18: 661–672.
Batygina, T.B., Yakovlev, M.B. 1987. Comparative embryology of flowering plants. Davidiaceae-Asteraceae (in
Russian). Leningrad: Izdat. Nauka, 391 pp.
Bayer, R.J., Cross, E.W. 2002. A reassessment of tribal affinities of the enigmatic genera Printzia and Isoetopsis
(Asteraceae), based on three chloroplast sequences.
Austral. J. Bot. 50: 677–686.
Bayer, R.J., Puttock, C.F. 1999. Molecular systematics of
southern hemisphere Gnaphalieae. In: Anon. (ed.) XVI
International Botanical Congress Abstracts. St. Louis:
Missouri Botanical Garden, p. 17.
Bayer, R.J., Starr, J.R. 1998. Tribal phylogeny of the Asteraceae based on two non-coding chloroplast sequences,
the trnL intron and trnL/trnF intergenic spacer. Ann.
Missouri Bot. Gard. 85: 242–256.
Bayer, R.J., Puttock, C.F., Kelchner, S.A. 2000. Phylogeny of
South African Gnaphalieae (Asteraceae) based on two
non-coding chloroplast sequences. Amer. J. Bot. 87:
259–272.
Bayer, R.J., Greber, D.G., Bagnall, N.H. 2002. Phylogeny
of Australian Gnaphalieae (Asteraceae) based on
chloroplast and nuclear sequences, the trnL intron,
Selected Bibliography to Compositae
the trnL/trnF intergenic spacer, matK and ETS. Syst.
Bot. 27: 801–814.
Bayer, R.J., Breitwieser, I., Dillon, M., Koekemoer, M.,
Ward, J. 2003. Phylogeny of the Gnaphalieae based on
three cpDNA sequences. Comp. Newslett. 40: 8.
Beck, J.B., Nesom, G.L., Calie, P.J., Baird, G.I., Small, R.L.,
Schilling, E.E. 2004. Is subtribe Solidagininae (Asteraceae) monophyletic? Taxon 53: 691–698.
Bentham, G. 1873a. Compositae. In: Bentham, G.,
Hooker, J.D., Genera Plantarum, vol. II, 1. London: Lovell Reeve, pp. 163–533 (plus Addenda et
Corrigenda).1
Bentham, G. 1873b. Notes on the classification, history and
geographical distribution of Compositae. J. Linn. Soc.,
Bot. 13: 335–577.
Berry, P.E., Calvo, R.N. 1989. Wind pollination, selfincompatibility and shifts in pollination systems in
the high Andean genus Espeletia (Asteraceae). Amer.
J. Bot. 76. 1602–1614.
Bierner, M.W. 2001. Taxonomy of Hymenoxys subgenus Picradenia and a conspectus of the subgenera of Hymenoxys (Asteraceae: Helenieae: Tetraneurinae). Lundellia 4: 37–63.
Blackmore, S. 1981. Palynology and intergeneric relationships in subtribe Hyoseridinae (Compositae:
Lactuceae). Bot. J. Linn. Soc. 82: 1–13.
Blackmore, S. 1982. Palynology of subtribe Scorzonerinae
(Compositae: Lactuceae) and its taxonomic significance. Grana 21: 149–160.
Blackmore, S. 1986. The identification and taxonomic significance of lophate pollen in the Compositae. Canad.
J. Bot. 64: 3101–3112.
Blackmore, S., Persson, V. 1996. Palynology and systematics of the Crepidinae (Compositae: Lactuceae). In:
Hind, D.J.N., Beentje, H.J. (eds) Compositae: systematics. Proceedings of the International Compositae Conference, Kew, 1994 (D.J.N. Hind, Editor-in-Chief), vol.
1. Royal Botanic Gardens, Kew, pp. 111–122.
Blake, S.F. 1935. The genus Chionopappus of Bentham
(Asteraceae). J. Wash. Acad. Sci. 25: 488–493.
Bohlmann, F., Jakupovic, J. 1990. Progress in the chemistry
of the Vernonieae (Compositae). Pl. Syst. Evol. suppl.
4: 3–43.
Bohlmann, F., Burkhardt, T., Zdero, C. 1973. Naturally occurring acetylenes. London: Academic Press.
Bohlmann, F., Zdero, C., King, R.M., Robinson, H. 1980a.
New helangiolides from Conocliniopsis prasiifolia.
Phytochemistry 19: 1547–1549.
Bohlmann, F., Zdero, C., Robinson, H., King, R.M. 1980b.
Guaianolides from Agrianthus pungens. Phytochemistry 19: 1873–1874.
Bohlmann, F., Abraham, W.-R., Robinson, H., King, R.M.
1981a. Heliangiolides and other constituents from Bejaranoa semistriata. Phytochemistry 20: 1639–1642.
Bohlmann, F., Jakupovic, J., King, R.M., Robinson, H.
1981b. New germacranolides, guaianolides and
rearranged guaianolides from Lasiolaena santosii.
Phytochemistry 20: 1613–1622.
1
The authorship of genera described or combined in Bentham and
Hooker’s ‘Genera Plantarum’ (1873) has been variously referred to
as Benth. (1873), Benth. in Benth. & Hook. f. (1873) or Benth. &
Hook. f. (1873).
577
Bohlmann, F., Suwita, A., Robinson, H., King, R.M. 1981c.
Six guaianolides from Stylotrichium rotundifolium.
Phytochemistry 20: 1887–1890.
Bohlmann, F., Zdero, F., Pickard, J., Robinson, H.,
King, R.M. 1981d. New types of sesquiterpene lactones
and other constituents from Trichogonia species.
Phytochemistry 20: 1323–1333.
Bohlmann, F., Singh, P., King, R.M., Robinson, H. 1982a.
New guaianolides from Pseudostifftia kingii. Phytochemisty 21: 1171–1172.
Bohlmann, F., Jakupovic, J., Schuster, A., King, R.M., Robinson, H. 1982b Guaianolides and homoditerpenes from
Lasiolaena morii. Phytochemistry 21, 1: 161–165.
Bohlmann, F., Zdero, C., King, R.M., Robinson, H. 1984a
Heliangiolides and bejaranolides from Conocliniopsis
prasiifolia. Phytochemistry 23, 7: 1509–1511.
Bohlmann, F., Zdero, C., Jakupovic, J., Gerke, T.,
Wallmeyer, M., King, R.M., Robinson, H. 1984b
Neus Sesquiterpenlactone und Rosan-Derivate aus
Trichogonia-Arten. Liebigs Ann. Chem. 1984: 162–185.
Bohm, B.A., Stuessy, T.F. 1995. Flavonoid chemistry of Barnadesioideae (Asteraceae). Syst. Bot. 20: 22–27.
Bohm, B.A., Stuessy, T.F. 2001. Flavonoids of the sunflower
family (Asteraceae). Berlin Heidelberg New York:
Springer.
Bolick, M.R. 1978. Taxonomic, evolutionary and functional
considerations of Compositae pollen ultrastructure
and sculpture. Pl. Syst. Evol. 130: 209–218.
Bremer, K. 1987. Tribal interrelationships of the Asteraceae.
Cladistics 3: 210–253.
Bremer, K. 1993. New subtribes of the Lactuceae (Asteraceae). Novon 3: 328–330.
Bremer, K. 1994. Asteraceae. Cladistics & classification.
Portland, OR: Timber Press.
Bremer, K. 1996. Major clades and grades of the Asteraceae.
In: Hind, D.J.N., Beentje, H.J. (eds) Compositae: systematics. Proceedings of the International Compositae
Conference, Kew, 1994 (D.J.N. Hind, Editor-in-Chief),
vol. 1. Royal Botanic Gardens, Kew, pp. 1–7.
Bremer, K., Gustafsson, M.H.G. 1997. East Gondwana ancestry of the sunflower alliance of families. Proc. Natl
Acad. Sci. U.S.A. 94: 9188–9190.
Bremer, K., Humphries, C.J. 1993. Generic monograph of the
Asteraceae-Anthemideae. Bull. Nat. Hist. Mus. London
23: 71–177.
Bremer, K., Jansen, R.K. 1992. A new subfamily of the Asteraceae. Ann. Missouri Bot. Gard. 79: 414–415.
Bremer, K., Anderberg, A.A., Karis, P.O., Lundberg, J. 1994.
Tribe Eupatorieae. In: Bremer, K. (ed.) Asteraceae.
Cladistics & classification. Portland, OR: Timber
Press, pp. 625–680.
Brouillet, L., Allen, G., Semple, J.C., Ito, M. 2001. ITS phylogeny of North American asters (Asteraceae: Astereae). In: Proceedings Botany 2001, ASPT/BSA/IOPB
Joint Meeting, Albuquerque, New Mexico, 12–16 August 2001.
Brown, K.S. 1984. Adult-obtained pyrrolizidine alkaloids
defend ithomine butterflies against a spider predator.
Nature 309: 707–709.
Brown, G.K., Clark, W.D. 1981. Chromosome numbers in
South American Haplopappus Cass. (Compositae).
Amer. J. Bot. 68: 1218–1221.
Brown, G.K., Clark, W.D. 1982. Taxonomy of Haplopappus
sect. Gymnocoma (Compositae). Syst. Bot. 7: 199–213.
578
Selected Bibliography to Compositae
Bruhl, J.J., Quinn, C.J. 1990. Cypsela anatomy in the Cotuleae
(Asteraceae: Anthemideae). Bot. J. Linn. Soc. 102: 37–
59.
Bruhl, J.J., Quinn, C.J. 1991. Floral morphology and a reassessment of affinities in the ‘Cotuleae’ (Asteraceae).
Austral. Syst. Bot. 4: 637–654.
Cabrera, A.L. 1937. Revisión del genero Chaetanthera
(Compositae). Revista Mus. La Plata ser. 2, 1(3):
87–210.
Cabrera, A.L. 1954. Compuestas sudamericanas nuevas o
críticas, II. Notas del Museo, Universidad Nacional de
Eva Peron 17, 84: 71–80.
Cabrera, A.L. 1959. Revisión del género Dasyphyllum (Compositae). Revista Mus. La Plata, Secc. Bot. 9: 21–100.
Cabrera, A.L. 1961. Compuestas Argentinas. Clave para la
determinación de los géneros. Revista Mus. Argent. Ci.
Nat., Bot. 2: 291–362.
Cabrera, A.L. 1965. Revision del genero Mutisia (Compositae). Opera Lilloana 13: 1–227.
Cabrera, A.L. 1971. Revision del genero Gochnatia (Compositae). Revista Mus. La Plata, Secc. Bot. 12: 1–160.
Cabrera, A.L. 1977. Mutisieae – systematic review. In: Heywood, V.H., Harborne, J.B., Turner, B.L. (eds) The biology and chemistry of the Compositae, vol. 2. London:
Academic Press, pp. 1039–1066.
Cabrera, A.L. 1978. Flora de la Provincia de Jujuy, vol. 10.
Buenos Aires: INTA.
Cabrera, A.L. 1982. Revisíon del género Nassauvia (Compositae). Darwiniana 24: 283–379.
Cabrera, A.L. 1996. Compositae III, Asteroideae, Eupatorieae. In: Flora of Paraguay, vol. 25. Missouri Botanical
Garden and Conservatoire et Jardin botaniques de la
Ville de Genève, Geneva.
Cabrera, A.L., Freire, S.E. 1997. Stevia. In: Espinar, L.A.
(co-ord.) Flora Fanerogâmica Argentina, fasc. 47. 280.
Asteraceae, parte 8. Tribu II. Eupatorieae. Cordoba:
ProFlora Conicet.
Cabrera, A.L., Freire, S.E. (compilers) 1999. Eupatorieae in
Asteraceae. In: Zuloaga, O., Morrone, O. (eds) Catálogo
de las plantas vasculares de la República Argentina, II.
Monogr. Syst. Bot. 75.
Cabrera, A.L., Klein, R.M. 1989 (1991). Tribo Eupatorieae.
Compostas 4. In: Reitz, R. (ed.) Flora ilustrada Catarinense. Part 1. Monografia. Itajaí, Brazil: Herbário Barbosa Rodrigues.
Candolle, A.P. de 1836. Ordo CII: Compositae [part 1].
Prodr. 5: 4–706. Paris: Treuttel et Würtz.
Canne, J.M. 1977. A revision of the genus Galinsoga (Compositae: Heliantheae). Rhodora 79: 319–389.
Carlquist, S. 1959. Studies on Madiinae: anatomy, cytology,
and evolutionary relationships. Aliso 4: 171–236.
Carlquist, S. 1960. Wood anatomy of Cichorieae (Compositae). Trop. Woods 112: 65–91.
Carlquist, S. 1966. Wood anatomy of Compositae: a summary with comments on factors controlling wood evolution. Aliso 6: 25–44.
Carlquist, S. 1976. Tribal interrelationships and phylogeny
of the Asteraceae. Aliso 8: 465–492.
Cassini, H. 1819. Sixième mémoire sur la famille des Synanthérées, contenant les caractères des tribus. J. Phys.
Chim. Hist. Nat. Arts 88: 150–163, 189–204.
Cassini, H. 1823. Liabon. In: Cuvier, G. (ed.) Dictionnaire
des Sciences Naturelles 26: 203–211. Reprinted in
King, R.M., Dawson, H.W. (eds) 1975, Cassini on
Compositae. New York: Oriole Editions.
Cassini, H. 1825. Oligacte. In: Cuvier, G. (ed.) Dictionnaire des Sciences Naturelles 36: 16–18. Reprinted in
King, R.M., Dawson, H.W. (eds) 1975, Cassini on Compositae. New York: Oriole Editions.
Cassini, H. 1829. Tableau synoptique des Synanthérées.
Ann. Sci. Nat. Paris 17: 387–423.
Cassini, H. 1830. Zyegee. In: Cuvier, G. (ed.) Dictionnaire
des Sciences Naturelles 60: 560–619. Reprinted in
King, R.M., Dawson, H.W. (eds) 1975, Cassini on
Compositae. New York: Oriole Editions.
Cerana, M.M., Ariza Espinar, L. 1995. Sobre la presencia
de domacios en Mikania (Asteraceae). Kurtziana 24:
7–18.
Chase, M.W., Soltis, D.E., Olmstead, R.G., Morgan, D.,
Les, D., Mischler, B.D., Duvall, M.R., Price, R.A.,
Hills, H.G., Qia, Y., Kron, K.A., Rettig, J.A., Conti, E.,
Palmer, J.D., Manhart, J.R., Sytsma, K.J. Michaels, H.J.,
Kress, W.J., Donoghue, M.J., Clark, W.-D., Hendren, M., Gaut, B.S., Jansen, R.K., Kim, K.-J,
Wimpee, C.F., Smith, J.F., Furnier, G.R., Strauss, S.,
Xiang, Q., Plunkett, G.M., Soltis, P.S., Swensen, S.,
Eguiarte, L.E., Learn, G.H. Jr., Barrett, S.C.H.,
Graham, S., Dayanadan, S., Albert, V.A. 1993. Phylogenetics of seed plants: an analysis of nucleotide
sequences from the plastid gene rbc L. Ann. Missouri
Bot. Gard. 80: 528–580.
Clark, W.D. 1979. The taxonomy of Hazardia (Compositae:
Astereae). Madroño 26: 105–127.
Claßen-Bockhoff, R. 1992. Florale Differenzierung in komplex organisierten Asteraceen-Köpfen. Flora 186: 1–22.
Claßen-Bockhoff, R. 1996. Functional units beyond the level
of the capitulum and cypsela in Compositae. In: Caligari, P.D., Hind, D.J.N. (eds) Compositae: biology & utilization. Proceedings of the International Compositae
Conference, Kew, 1994 (D.J.N. Hind, Editor-in-Chief),
vol. 2. Royal Botanic Gardens, Kew, pp. 129–160.
Claßen-Bockhoff, R., Froebe, H., Langerheins, D. 1989.
Die Infloreszenzstruktur von Gundelia tournefortii L.
(Asteraceae). Flora 182: 463–479.
Cronquist, A.J. 1955. Phylogeny and taxonomy of the Compositae. Amer. Midl. Naturalist 53: 478–511.
Cross, E.W., Quinn, C.J., Wagstaff, S.J. 2002. Molecular evidence for the polyphyly of Olearia (Astereae: Asteraceae). Pl. Syst. Evol. 235: 99–120.
Cuatrecasas, J. 1970. Reinstatement of the genus Llerasia
(Compositae). Biotropica 2: 39–45.
Cuatrecasas, J. 1976. A new subtribe in the Heliantheae
(Compositae): Espeletiinae. Phytologia 35: 42–61.
D’Arcy, W. 1975. Conspectus of the Panamanian Compositae. In: Woodson, R.E., Schery, R.W. et al. (eds) Flora
of Panama, vol. Family 184, pp. 845–846. Compositae.
Ann. Missouri Bot. Gard. 62: 835–1321.
DeJong, D.C., Beaman, J.H. 1963. The genus Olivaea (Compositae, Astereae). Brittonia 15: 86–92.
DeVore, M.L., Stuessy, T.F. 1995. The place and time of the
origin of the Asteraceae, with additional comments on
the Calyceraceae and Goodeniaceae. In: Hind, D.J.N.,
Jeffrey, C., Pope, G.V. (eds) Advances in Compositae
systematics. Royal Botanic Gardens, Kew, pp. 23–40.
Dillon, M.O., Hensold, N. (compilers) 1993. Asteraceae. In:
Brako, L., Zarruchi, J.L. (eds) Catalogue of the flow-
Selected Bibliography to Compositae
ering plants and gymnosperms of Peru. Monogr. Syst.
Bot. 45: 103–189.
Dittrich, M. 1977. Cynareae – systematic review. In: Heywood, V.H., Harborne, J.B., Turner, B.L. (eds) The biology and chemistry of the Compositae, vol. 2. London:
Academic Press, pp. 999–1015.
Dittrich, M. 1996. Die Bedeutung morphologischer und
anatomischer Achänen-Merkmale für die Systematik
der Tribus Echinopseae Cass. und Carlinae Cass.
Boissiera 51: 5–102.
Eldenäs, P., Anderberg, A.A., Källersjö, M. 1998. Molecular
phylogenetics of the tribe Inuleae s.str. (Asteraceae)
based on ITS sequences of nuclear ribosomal DNA. Pl.
Syst. Evol. 210: 159–173.
Eldenäs, P., Källersjö, M., Anderberg, A.A. 1999. Phylogenetic placement and circumscription of tribes Inuleae
s.str. and Plucheeae (Asteraceae): evidence from sequences of chloroplast gene ndhF. Mol. Phylog. Evol.
13: 50–58.
Erbar, C., Leins, P. 1995. Portioned pollen release and the
syndromes of secondary pollen presentation in the
Campanulales-Asterales complex. Flora 190: 323–338.
Erbar, C., Leins, P. 2000. Some interesting features in the
capitulum and flower of Arnoldoa macbrideana Ferreyra (Asteraceae, Barnadesioideae). Bot. Jahrb. Syst.
122: 503–515.
Eriksson, T. 1991. The systematic position of the Blepharispermum group (Asteraceae–Heliantheae). Taxon 40:
33–39.
Eriksson, T. 1992. The genus Blepharispermum (Asteraceae, Heliantheae). Pl. Syst. Evol. 182: 149–227.
Eriksson, T. 1993. The genus Athroisma (Asteraceae, Heliantheae). Bot. J. Linn. Soc. 119: 101–184.
Feinbrun-Dothan, N. 1977. Flora Palaestina, vol. 3.
Jerusalem: Israel Academy of Sciences and Humanities.
Ferreyra, R. 1995. Flora of Peru. Family Asteraceae. VI.
Fieldiana, Bot. 35: 87–90.
Fioretto, A., Alfani, A. 1988. Anatomy of succulence and
CAM in 15 species of Senecio. Bot. Gaz. 149: 142–152.
Fiori, A., Paoletti, G. 1895–1899. Iconographia florae italicae. Padova: Tipografia Antoniana.
Folch, R. 1986. La vegetació dels Païssos Catalans.
Barcelona: Ketres.
Font Quer, P. 1962. Plantas medicinales. El Dioscórides renovado. Barcelona: Labor.
Francisco-Ortega, J., Santos-Guerra, A., Hines, A.,
Jansen, R.K. 1997. Molecular evidence for a Mediterranean origin of the Macaronesian endemic genus
Argyranthemum (Asteraceae). Amer. J. Bot. 84:
1595–1613.
Freire, S.E., Katinas, L. Sancho, G. 2002. Gochnatia (Asteraceae, Mutisieae) and the Gochnatia complex: taxonomic implications from morphology. Ann. Missouri
Bot. Gard. 89: 524–550.
Funk, V.A., Chan, R.A. 2003. A brief survey of the phylogeny of the Arctoteae (Compositae: Cichorioideae
s.s.). Comp. Newslett. 40: 13–14.
Funk, V.A., Robinson, H. (compilers) 1996. Asteraceae. In:
Boggan, J., Funk, V.A., Kelloff, C., Hoff, M., Cremers, G.,
Feuillet, C. (eds) Checklist of the plants of the Guianas
(Guyana, Surinam, French Guiana), 2nd edn. Washington, DC: Smithsonian Institution.
Funk, V.A., Robinson, H. 2001. A bully new genus from the
Andes (Compositae: Liabeae). Syst. Bot. 26: 216–225.
579
Funk, V.A., Chan, R., Keeley, C. 2004. Insights into the evolution of the tribe Arctoteae (Compositae: subfamily
Cichorioideae s. s. ) using trnL, ndhF, and ITS. Taxon
53: 637–655.
Funk, V.A., Bayer, R.J., Keeley, S., Chan, R., Watson, L.,
Gemeinholzer, B., Schilling, E.E., Panero, J.L., Baldwin, B.G., Garcia-Jacas, N.T., Susanna, A., Jansen, R.K.
2005. Everywhere but Antarctica: using a supertree to
understand the diversity and distribution of the Compositae. In: Friis, I., Balslev, H. (eds) Plant diversity and
complexity patterns – local, regional and global dimensions. Proceedings International Symposium, Royal
Danish Academy of Sciences and Letters, Copenhagen,
25–28 May 2003. Biol. Skr. 55: 343–373.
Gadek, P.A., Bruhl, J.J., Quinn, C.J. 1989. Exine structure
in the ‘Cotuleae’ (Anthemideae, Asteraceae). Grana 28:
163–178.
Galetto, L. 1995. Estudios sobre el nectar y los nectarios en Hyaloseris rubricunda y Barnadesia odorata
(Asteraceae–Mutisieae). Darwiniana 33: 127–133.
Gamerro, J.C. 1990. Identidad de Pseudostifftia con Moquinia (Compositae) y consideracions sobre la ubicacion tribal del taxon. Darwiniana 30: 123–136.
Garcia-Jacas, N. 1992. Estudi taxonòmic i biosistemàtic de
les espècies ibériques i nord-africanes del gènere Centaurea sect. Acrocentron subsect. Orientales. Diss., University of Barcelona.
Garcia-Jacas, N., Susanna, A., Ilarlsan, R. 1996. Aneuploidy
in the Centaureinae (Compositae): is n = 7 the end of
the series? Taxon 45: 39–42.
Garcia-Jacas, N., Susanna, A., Mozaffarian, R., Ilarslan, R.
2000. The natural delimitation of Centaurea (Asteraceae: Cardueae): ITS sequence analysis of the Centaurea jacea group. Pl. Syst. Evol. 223: 185–199.
Garcia-Jacas, N., Susanna, A., Garnatje, T., Vilatersana, R.
2001. Generic delimitation and phylogeny of the subtribe Centaureinae (Asteraceae): a combined nuclear
and chloroplast DNA analysis. Ann. Bot. London 87:
503–515.
Garcia-Jacas, N., Garnatje, T., Susanna, A., Vilatersana, R.
2002. Tribal and subtribal delimitation and phylogeny
of the Cardueae (Asteraceae): a combined nuclear and
chloroplast DNA analysis. Mol. Phylog. Evol. 22: 51–64.
Garg, S., Sastry, T.C.S. 1996. Indian Compositae in foods
and flavours – a review. In: Caligari, P.D.S., Hind, D.J.N.
(eds) Compositae: biology & utilization. Proceedings
of the International Compositae Conference, Kew, 1994
(D.J.N. Hind, Editor-in-Chief), vol. 2. Royal Botanic
Gardens, Kew, pp. 361–382.
Gautier, L. 1992. Taxonomy and distribution of a tropical
weed: Chromolaena odorata (L.) R. King & H. Robinson. Candollea 47: 645–662.
Giuliano, D.A. 2001. Clasificación infragenérica de las especies argentinas de Baccharis (Asteraceae, Astereae).
Darwiniana 39: 131–154.
Goertzen, L.R., Cannone, J.J., Gutell, R.R., Jansen, R.K.
2003. ITS secondary structure derived from comparative analysis: implications for sequence alignment
and phylogeny of Asteraceae. Mol. Phylog. Evol. 29:
216–234.
Graham, A. 1996. A contribution to the geologic history of
the Compositae. In: Hind, D.J.N., Beentje, H.J. (eds)
Compositae: systematics. Proceedings of the International Compositae Conference, Kew, 1994 (D.J.N. Hind,
580
Selected Bibliography to Compositae
Editor-in-Chief), vol. 1. Royal Botanic Gardens, Kew,
pp. 123–140.
Grashoff, J. 1974. Novelties in Stevia (Compositae: Eupatorieae). Brittonia 26: 347–384.
Grau, J. 1980. Die Testa der Mutisieae und ihre systematische
Bedeutung. Mitt. Bot. Staatssamml. München 16: 269–
332.
Grau, J., Hopf, H. 1985. Das Endosperm der Compositae.
Bot. Jahrb. Syst. 107: 251–268.
Greger, H. 1977. Anthemideae – chemical review. In: Heywood, V.H., Harborne, J.B., Turner, B.L. (eds) The biology and chemistry of the Compositae. London: Academic Press, pp. 851–898.
Gustafsson, M.H.G., Bremer, K. 1995. Morphology and
phylogenetic relationships of the Asteraceae, Calyceraceae, Campanulaceae, Goodeniaceae and related
families (Asterales). Amer. J. Bot. 82: 250–265.
Gustafsson, M.H.G., Backlund, A., Bremer, B. 1996. Phylogeny of the Asterales sensu lato based on rbcL sequences with particular reference to the Goodeniaceae.
Pl. Syst. Evol. 199: 217–242.
Gustafsson, M.H.G., Pepper, A.S.-R., Albert, V.A., Källersjö,
M. 2001. Molecular phylogeny of the Barnadesioideae
(Asteraceae). Nordic J. Bot. 21: 149–160.
Gutterman, Y., Ginott, S. 1994. Long-term protected ‘seed
bank’ in dry inflorescences of Asteriscus pygmaeus:
achene dispersal mechanism and germination. J. Arid
Environ. 26: 149–163.
Häffner, E. 2000. On the phylogeny of the subtribe Carduinae
(tribe Cardueae, Compositae). Englera 21: 1–209.
Häffner, E., Hellwig, F. 1999. Phylogeny of the tribe Cardueae
(Compositae) with emphasis on subtribe Carduinae:
an analysis based on ITS sequence data. Willdenowia
29: 27–39.
Hansen, H.V. 1985a. A taxonomic revision of the genus Gerbera (Compositae, Mutisieae) sections Gerbera, Parva,
Piloselloides (in Africa) and Lasiopus. Opera Bot. 78:
1–36.
Hansen, H.V. 1985b. Notes on Gerbera sect. Pseudoseris
(Compositae-Mutisieae). Nordic J. Bot. 5: 451–453.
Hansen, H.V. 1988. A taxonomic revision of the genera Gerbera sect. Isanthus, Leibnitzia (in Asia), and Uechtritzia
(Compositae, Mutisieae). Nordic J. Bot. 8: 61–76.
Hansen, H.V. 1990. Phylogenetic studies in the Gerberacomplex (Compositae, tribe Mutisieae, subtribe
Mutisiinae). Nordic J. Bot. 9: 469–485.
Hansen, H.V. 1991a. SEM – studies and general comments
on pollen in tribe Mutisieae (Compositae) sensu Cabrera. Nordic J. Bot. 10: 607–623.
Hansen, H.V. 1991b. Phylogenetic studies in Compositae
tribe Mutisieae. Opera Bot. 109: 1–50.
Hansen, H.V. 1997. Studies in the Goodeniaceae and the
Brunoniaceae with a discussion of their relationship
to Asteraceae and Calyceraceae. Nordic J. Bot. 17: 495–
510.
Harling, G. 1995. The genus Jungia L. fil. (Compositae-Mutisieae). Acta Regiae Soc. Sci. Litt. Gothob. Bot. 4: 1–133.
Harris, E.M. 1995. Inflorescence and floral ontogeny in
Asteraeae: a synthesis of historical and current concepts. Bot. Rev. 61: 93–275.
Harris, E.M. 1999. Capitula in the Asteridae: a widespread
and varied phenomenon. Bot. Rev. 65: 348–369.
Hartman, R.L. 1990. A conspectus of Machaeranthera
(Asteraceae: Astereae). Phytologia 68: 439–465.
Hegnauer, R. 1977. The chemistry of the Compositae. In:
Heywood, V.H., Harborne, J.B., Turner, B.L. (eds) The
biology and chemistry of the Compositae. London:
Academic Press, pp. 283–335.
Heinrich, M. 1996. Ethnobotany of Mexican Compositae:
an analysis of historical and modern sources. In: Caligari, P.D.S., Hind, D.J.N. (eds) Compositae: biology
& utilization. Proceedings of the International Compositae Conference, Kew, 1994 (D.J.N. Hind, Editor-inChief), vol. 2. Royal Botanic Gardens, Kew, pp. 475–
503.
Hellwig, F.H. 1990. Die Gattung Baccharis L. (Compositae–
Astereae) in Chile. Mitt. Bot. Staatssamml. München
29: 1–456.
Hellwig, F.H. 1993. The genera Pingraea Cass. and Neomolina Hellwig (Compositae–Astereae). Candollea 48:
203–219.
Hellwig, F.H. 1996. Taxonomy and evolution of Baccharidinae (Compositae). In: Hind, D.J.N., Beentje, H.J. (eds)
Compositae: systematics. Proceedings of the International Compositae Conference, Kew, 1994 (D.J.N. Hind,
Editor-in-Chief), vol. 1. Royal Botanic Gardens, Kew,
pp. 575–590.
Hendry, G.A.F. 1996. Fructan and the ecology and evolution
of the Compositae. In: Caligari, P.D.S., Hind, D.J.N.
(eds) Compositae: biology & utilization. Proceedings
of the International Compositae Conference, Kew, 1994
(D.J.N. Hind, Editor-in-Chief), vol. 2. Royal Botanic
Gardens, Kew, pp. 121–128.
Herz, W. 1977. Sesquiterpene lactones in the Compositae.
In: Heywood, V.H., Harborne, J.B., Turner, B.L. (eds)
The biology and chemistry of the Compositae, vol. 2.
London: Academic Press, pp. 337–357.
Heywood, V.H., Humphries, C.J. 1977. Anthemideae –
systematic review. In: Heywood, V.H., Harborne, J.B.,
Turner, B.L. (eds) The biology and chemistry of
the Compositae. London: Academic Press, pp.
851–898.
Heywood, V.H., Harborne, J.B., Turner, B.L. (eds) 1977. The
biology and chemistry of the Compositae. London:
Academic Press.
Hind, D.J.N. 1999. A new genus, Semiria (Compositae: Eupatorieae) and a discussion of its affinities within the
subtribe Gyptidinae of Bahia, Brazil. Kew Bull. 54: 425–
435.
Hind, D.J.N. 2000. A new genus, Catolesia (Compositae:
Eupatorieae), and a discussion of its affinities within
the subtribe Gyptidinae of Bahia, Brazil. Kew Bull. 55:
941–948.
Hitchcock, C.L., Cronquist, A., Ownbey, M., Thompson, J.W.
1959. Vascular plants of the Pacific Northwest, vol. 5.
Compositae. Seattle: University of Washington Press,
349 pp.
Hoffmann, O. 1894. Compositae. In: Engler, A., Prantl, K.,
Die natürlichen Pflanzenfamilien, IV. Leipzig: Engelmann, pp. 87–391.
Holmes, W.C. 1991. Dioecy in Mikania (Compositae: Eupatorieae). Pl. Syst. Evol. 175: 87–92.
Huang, Y.-P., Ling, Y.-R. 1996. Economic Compositae in
China. In: Caligari, P.D.S., Hind, D.J.N. (eds) Compositae: biology & utilization. Proceedings of the International Compositae Conference, Kew, 1994 (D.J.N. Hind,
Editor-in-Chief), vol. 2. Royal Botanic Gardens, Kew,
pp. 431–451.
Selected Bibliography to Compositae
Humbert, H. 1962. 189ème Famille – Composées, tome 2.
In: Flore de Madagascar et des Comores. Paris: Typ.
Firmin-Didot., pp. 339–622.
Iljin, M. 1960. Species novae generis Jurinea Cass. ex Asia
Media (in Russian). Bot. Mater. Gerb. Bot. Inst. Komarova Akad. Nauk S.S.S.R. 20: 344–355.
Ito, M., Yahara, T., King, R.M., Watanabe, K., Oshita, S.,
Yokoyama, J., Crawford, D.J. 2000a. Molecular phylogeny of Eupatorieae (Asteraceae) estimated from
cpDNA RFLP and its implication for the polyploid
origin hypothesis of the tribe. J. Pl. Res. 113: 91–96.
Ito, M., Watanabe, K., Kita, Y., Kawahara, T., Crawford, D.J.,
Yahara, T. 2000b. Phylogeny and phytogeography of
Eupatorium (Eupatorieae, Asteraceae): insights from
sequence data of the nrDNA ITS regions and cpDNA
RFLP. J. Pl. Res. 113: 79–89.
Jansen, R.K. 1991. Phylogeny and character evolution in the
Asteraceae based on chloroplast DNA restriction site
mapping. Syst. Bot. 16: 98–115.
Jansen, R.K., Kim, K.-J. 1996. Implications of chloroplast
DNA data for the classification and phylogeny
of the Asteraceae. In: Hind, D.J.N., Beentje, H.J.
(eds) Compositae: systematics. Proceedings of the
International Compositae Conference, Kew, 1994
(D.J.N. Hind, Editor-in-Chief), vol. 1. Royal Botanic
Gardens, Kew, pp. 317–339.
Jansen, R.K., Palmer, J.D. 1987. A chloroplast DNA inversion
marks an ancient evolutionary split in the sunflower
family (Asteraceae). Proc. Natl Acad. Sci. U.S.A. 84:
5818–5822.
Jansen, R.K., Palmer, J.D. 1988. Phylogenetic implications
of chloroplast DNA restriction site variation in the
Mutisieae (Asteraceae). Amer. J. Bot. 75: 753–766.
Jansen, R.K., Michaels, H.J., Palmer, J.D. 1991. Phylogeny
and character evolution in the Asteraceae based on
chloroplast DNA restriction site mapping. Syst. Bot.
16: 98–115.
Jeffrey, C. 1966. Notes in Compositae. I. The Cichorieae in
East Tropical Africa. Kew Bull. 18: 427–486.
Jeffrey, C. 1967. Notes on Compositae. II. The Mutisieae in
East Tropical Africa. Kew Bull. 21: 177–223.
Jeffrey, C. 1978. Compositae. In: Heywood, V.H. (ed.) Flowering plants of the world. New York: Mayflower Books,
pp. 263–268.
Jeffrey, C. 1986. The Senecioneae in East Tropical Africa.
Kew Bull. 41: 873–943.
Jeffrey, C. 1992. The tribe Senecioneae (Compositae) in the
Mascarene Islands with an annotated world check-list
of the genera of the tribe. Kew Bull. 47: 49–109.
Jeffrey, C. 2002. Systematics of Compositae at the beginning
of the 21st century (in Russian). Bot. Zhurn. (Moscow
& Leningrad) 87, 11: 1–15.
Jeffrey, C. 2004. Systema Compositarum (Asteracearum)
nova (in English). Bot. Zhurn. (Moscow & Leningrad)
89, 12: 1817–1822.
Jepson, W.L. 1901. A flora of Western Middle California.
Berkeley, CA: Encina.
Jones, S.B. 1977. Vernonieae – systematic review. In: Heywood, V.H., Harborne, J.B., Turner, B.L. (eds) The biology and chemistry of the Compositae, vol. 2. London:
Academic Press, pp. 503–521.
Kadereit, J.W. 1989. Chloroplast DNA, cladistics and the
phylogeny of the Asteraceae. Bot. Acta 102: 7–10.
581
Karis, P.O. 1992. Hoplophyllum DC., the sister group to Eremothamnus O. Hoffm. (Asteraceae)? Taxon 41: 193–
198.
Karis, P.O. 1993a. The Heliantheae sensu lato (Asteraceae),
clades and classification. Pl. Syst. Evol. 188: 139–195.
Karis, P.O. 1993b. Morphological phylogenetics of the
Asteraceae-Asteroideae, with notes on character
evolution. Pl. Syst. Evol. 186: 69–93.
Karis, P.O. 1998. Apostates Lander (Asteraceae-Astereae),
transferred to the Heliantheae sensu lato. Bot. Jahrb.
Syst. 120: 131–135.
Karis, P.O. 2006. Morphological data indicates two major
clades of the subtribe Gorteriinae (AsteraceaeArctotideae). Cladistics 22:199–221.
Karis, P.O., Ryding, O. 1994a. Tribe Heliantheae. In: Bremer, K. (ed.) Asteraceae. Cladistics & classification.
Portland, OR: Timber Press, pp. 559–624.
Karis, P.O., Ryding, O. 1994b. Tribe Helenieae. In: Bremer
K. (ed.) Asteraceae. Cladistics & classification. Portland, OR: Timber Press, pp. 521–558.
Karis, P.O., Källersjö, M., Bremer, K. 1992. Phylogenetic
analysis of the Cichorioideae (Asteraceae) with emphasis on the Mutisieae. Ann. Missouri Bot. Gard. 79:
416–427.
Karis, P.O, Eldenäs, P., Källersjö, M. 2001. New evidence
for the systematic position of Gundelia L. with notes
on delimitation of Arctoteae (Asteraceae). Taxon 50:
105–114.
Katinas, L. 2000. Implications of morphological phylogenetics for the placement of the genera Adenocaulon and
Eriachaenium (Asteraceae). Pl. Syst. Evol. 223: 229–
250.
Keeley, S.C., Chan, K. 2003. The Vernonieae: searching for
a new paradigm. Comp. Newslett. 40: 19.
Keeley, S.C., Jansen, R.K. 1991. Evidence from chloroplast
DNA for the recognition of a new tribe, the Tarchonantheae, and the tribal placement of Pluchea (Asteraceae). Syst. Bot. 16: 173–181.
Keeley, S.C., Jansen, R.K. 1995. Chloroplast DNA restriction
site variation in the Vernonieae (Asteraceae), an initial
appraisal of the relationship of New and Old World
taxa and the monophyly of Vernonia. Pl. Syst. Evol.
193: 249–265.
Kilian, N. 1997. Revision of Launaea Cass. (Compositae, Lactuceae, Sonchinae). Englera 17: 1–478.
Kim, K.-J., Jansen, R.K. 1995. ndhF sequence evolution and
the major clades in the sunflower family. Proc. Natl
Acad. Sci. U.S.A. 92: 10379–10383.
Kim, K.-J., Jansen, R.K., Wallace, R.S., Michaels, H.J.,
Palmer, J.D. 1992. Phylogenetic implications of rbcL
sequence variation in the Asteraceae. Ann. Missouri
Bot. Gard. 79: 428–445.
Kim, H.-G., Keeley, S.C., Vroom, P.S., Jansen, R.K. 1998.
Molecular evidence for an African origin of the Hawaiian endemic Hesperomannia (Asteraceae). Proc. Natl
Acad. Sci. U.S.A. 95: 15440–15445.
Kim, S.-C., Crawford, D.J., Jansen, R.K., Santos Guerra, A.
1999a. The use of a non-coding region of chloroplast
DNA in phylogenetic studies of the subtribe Sonchinae
(Asteraceae: Lactuceae). Pl. Syst. Evol. 215: 85–99.
Kim, S.-C., Crawford, D.J., Francisco-Ortega, J., Santos
Guerra, A. 1999b. Adaptive radiation and genetic
differentiation in the woody Sonchus alliance (Aste-
582
Selected Bibliography to Compositae
raceae: Sonchinae) in the Canary Islands. Pl. Syst.
Evol. 215: 101–118.
Kim, H.-G., Loockerman, D.J., Jansen, R.K. 2002. Systematic implications of ndhF sequence variation in the
Mutisieae (Asteraceae). Syst. Bot. 27: 598–609.
Kim, H.-G., Funk, V.A., Vlasek, A., Zimmer, E. 2003. A phylogeny of the Munnoziinae (Compositae, Liabeae): circumscription of Munnozia and a new placement of M.
perfoliata. Pl. Syst. Evol. 239: 171–185.
Kimball, R.T., Crawford, D.J. 2004. Phylogeny of Coreopsideae (Asteraceae) using ITS sequences suggests lability in reproductive characters. Mol. Phylog. Evol.
33: 127–139.
King, R.M., Robinson, H. 1970. Studies in the Eupatorieae
(Compositae). XIX. New combinations in Ageratina.
Phytologia 19: 208–229.
King, R.M., Robinson, H. 1971. Studies in the Eupatorieae
(Asteraceae). LIX. A new genus, Steviopsis. Phytologia
22: 156–157.
King, R.M., Robinson, H. 1972. Studies in the Eupatorieae
(Asteraceae). LXXVII. Additions to the genus Steviopsis. Phytologia 24: 60–62.
King, R.M., Robinson, H. 1978. Studies in the Eupatorieae
(Asteraceae). CLXVIII. Additions to the genus Ageratina. Phytologia 38: 323–355.
King, R.M., Robinson, H. 1987. The genera of Eupatorieae
(Asteraceae). Missouri Bot. Gard. Monogr. Syst. Bot.
22: 1–581.
King, R.M., Robinson, H. 1995. Generic limits in the Alomiinae (Eupatorieae–Asteraceae), and new combinations
in Brickelliastrum and Barroetea. Phytologia 78: 124–
126.
King, R.M., Kyhos, D.W., Powell, A.M., Raven, P.H., Robinson, H. 1976. Chromosome numbers in Compositae.
XIII. Eupatorieae. Ann. Missouri Bot. Gard. 63: 862–
888.
Koopman, W.J.M., Guetta, E., van de Wiel, C.C.M. et al.
1998. Phylogenetic relationships among Lactuca
(Asteraceae) species and related genera based on
ITS-1 DNA sequences. Amer. J. Bot. 85: 1517–1530.
Kornkven, A.B., Watson, L.E., Estes, J.R. 1998. Phylogenetic
analysis of Artemisia sect. Tridentatae (Asteraceae)
based on sequences from internal transcribed spacers (ITS) of nuclear ribosomal DNA. Amer. J. Bot. 85:
1787–1795.
Kornkven, A.B., Watson, L.E., Estes, J.R. 1999. Molecular
phylogeny of Artemisia sect. Tridentatae (Asteraceae)
based on chloroplast DNA restriction site variation.
Syst. Bot. 24: 69–84.
Lack, H.W. 1979. The subtribe Hypochoeridinae (Asteraceae, Lactuceae) in the tropics and the southern hemisphere. In: Larsen, K., Holm-Nielsen, L.B. (eds) Tropical botany. London: Academic Press, pp. 265–276.
Lammers, T.G. 1992. Circumscription and phylogeny of
the Campanulales. Ann. Missouri Bot. Gard. 79. 358–
413.
Lander, N.S. 1994. Olearia. In: Harden, G.J. (ed.) Flora of
New South Wales, vol. 3. New South Wales University
Press.
Lane, M.A. 1996. Pollination biology of Compositae. In:
Caligari, P.D.S., Hind, D.J.N. (eds) Compositae: biology
& utilization. Proceedings of the International Compositae Conference, Kew, 1994 (D.J.N. Hind, Editor-inChief), vol. 2. Royal Botanic Gardens, Kew, pp. 61–80.
Lane, M.A., Hartman, R.L. 1996. Reclassification of North
American Haplopappus (Compositae: Astereae)
completed: Rayjacksonia gen. nov. Amer. J. Bot. 83:
356–370.
Lane, M.A., Morgan, D.R., Suh, Y.-B., Simpson, B.B.,
Jansen, R.K. 1996. Relationships of North American
genera of Astereae, based on chloroplast DNA restriction site data. In: Hind, D.J.N., Beentje, H.J.
(eds) Compositae: systematics. Proceedings of the
International Compositae Conference, Kew, 1994
(D.J.N. Hind, Editor-in-Chief), vol. 1. Royal Botanic
Gardens, Kew, pp. 49–77.
Lee, J., Baldwin, B.G., Gottlieb, L.D. 2003. Phylogenetic relationships among the primarily North American genera
of Cichorieae (Compositae) based on analysis of 18S–
26S nuclear rDNA ITS and ETS sequences. Syst. Bot.
28: 616–626.
Leins, P. 1970. Die Pollenkörner und Verwandtschaftsbeziehungen der Gattung Eremothamnus (Asteraceae).
Mitt. Bot. Staatssamml. München 7: 369–376.
Leins, P. 1971. Pollensystematische Studien an Inuleen, vol.
1. Tarchonanthinae, Plucheinae, Inulinae, Buphthalminae. Bot. Jahrb. Syst. 91: 91–146.
Leins, P., Erbar, C. 1987. Studien zur Blütenentwicklung an
Compositen. Bot. Jahrb. Syst. 108: 381–401.
Leins, P., Erbar, C. 1990. On the mechanisms of secondary
pollen presentation in the Campanulales–Asterales
complex. Bot. Acta 103: 87–92.
Leins, P., Erbar, C. 2000. Die frühesten Entwicklungsstadien
der Blüten bei den Asteraceae. Bot. Jahrb. Syst. 122:
503–515.
Leins, P., Thyret, G. 1971. Pollen phylogeny and taxonomy
exemplified by an African Asteraceae group. Mitt. Bot.
Staatssamml. München 10: 280–286.
Lersten, N.R., Curtis, J.D. 1985. Distribution and anatomy
of hydathodes in Asteraceae. Bot. Gaz. 146: 106–114.
Lessing, C.F. 1832. Synopsis generum Compositarum,
earumque dispositionis novae tentamen monographus multarum capensium interjectis. Berlin:
Duncker & Humblot.
Liu, J.-Q., Gao, T.-G., Chen, Z.-D., Lu, A.-M. 2002. Molecular phylogeny and biogeography of the Qinghai-Tibet
Plateau endemic Nannoglottis (Asteraceae). Mol. Phylog. Evol. 23: 307–325.
Loockerman, D.J., Turner, B.L., Jansen, R.K. 2003. Phylogenetic relationships within the Tageteae (Asteraceae)
based on nuclear ribosomal ITS and chloroplast ndhF
gene sequences. Syst. Bot. 28: 191–207.
Lowrey, T., Urbatsch, L. 2003. Phylogenetic studies in the
Astereae: the need for generic redelimitation and a reasonable subtribal classification. Comp. Newslett. 40: 25.
Lundin, R. 2006. Nordenstamia Lundin (CompositaeSenecioneae), a new genus from the Andes of South
America. Comp. Newslett. 44: 14–18.
Lundberg, J., Bremer, K. 2003. A phylogenetic study of the
order Asterales using one morphological and three
molecular data sets. Intl. J. Pl. Sci. 164: 553–578.
Marticorena, C., Parra, O. 1975. Morfología de los granos
de polen de Hesperomannia Gray y Moquinia DC.
(Compositae–Asteraceae). Estudio comparativo con
generos afines. Gayana 29: 3–22.
McKenzie, R.J., Barker, N.P. Samuel, J., Muller, E.M., Skinner, A.K.W. 2005. Morphology of cypselae in subtribe
Arctotidinae (Compositae-Arctotideae) and its taxo-
Selected Bibliography to Compositae
nomic implications. Ann. Missouri Bot. Gard. 92: 569–
594.
McVaugh, R. 1984. Compositae. In: Anderson, W.R. (ed.)
Flora Novo-Galiciana, vol. 12. Ann Arbor: University
of Michigan Press.
Meiri, L., Dulberger, R. 1986. Stamen filament structure
in the Asteraceae: the anther collar. New Phytol. 104:
693–701.
Merxmüller, H., Leins, P., Roessler, H. 1977. Inuleae –
systematic review. In: Heywood, V.H., Harborne, J.B.,
Turner, B.L. (eds) The biology and chemistry of the
Compositae, vol. 1. London: Academic Press, pp.
577–602.
Metcalfe, C.R., Chalk, L. 1950. Anatomy of the Dicotyledons.
Oxford: Oxford University Press.
Michaels, H.J.K., Scott, K.M., Olmstead, R.G., Szaro, T.,
Jansen, R.K., Palmer, J.D. 1993. Interfamilial relationships of the Asteraceae: insights from rbcL sequence
variation. Ann. Missouri Bot. Gard. 80: 742–751.
Morgan, D.R. 2003. nrDNA external transcribed spacer
(ETS) sequence data, reticulate evolution, and the
systematics of Machaeranthera (Asteraceae). Syst.
Bot. 28: 179–190.
Morgan, D.R., Hartman, R.L. 2003. A synopsis of Machaeranthera (Asteraceae: Astereae), with recognition of
segregate genera. Sida 20: 1387–1416.
Muschler, R. 1911. Compositae africanae novae, I. Bot.
Jahrb. Syst. 46: 51–124.
Nash, D.L., Williams, L.O. 1976. Flora of Guatemala, vol. 12.
Fieldiana, Bot. 24: 1–603.
Nesom, G.L. 1989. Infrageneric taxonomy of New World
Erigeron (Compositae: Astereae). Phytologia 67: 67–
93.
Nesom, G.L. 1990. Taxonomy of the Erigeron pringlei group
(Asteraceae: Astereae). Phytologia 69: 227–235.
Nesom, G.L. 1993. Taxonomic infrastructure of Solidago and
Oligoneuron (Asteraceae: Astereae) and observations
on their phylogenetic position. Phytologia 75: 1–44.
Nesom, G.L. 1994a. Erigeron pattersonii (Asteraceae: Astereae), a new species from Nuevo Leon, Mexico. Phytologia 76: 96–100.
Nesom, G.L. 1994b. Taxonomic dispersal of Australian
Erigeron (Asteraceae: Astereae). Phytologia 76:
143–159.
Nesom, G.L. 1994c. Subtribal classification of the Astereae
(Asteraceae). Phytologia 76: 193–274.
Nesom, G.L. 1994d. Review of the taxonomy of Aster sensu
lato (Asteraceae: Astereae), emphasizing New World
species. Phytologia 77: 141–297.
Nesom, G.L. 1998 (2000). Full constitution of the Australian
genus Pappochroma (Asteraceae: Astereae). Phytologia
85: 276–279.
Nesom, G.L. 2000a. New subtribes for North American
Astereae (Asteraceae). Sida 19: 263–268.
Nesom, G.L. 2000b. Generic conspectus of the tribe Astereae
in North America, Central America, the Antilles, and
Hawaii. Sida Bot. Misc. 20: i–viii, 1–100.
Nesom, G.L. 2005. Infrageneric classification of Litaris
(Asteraceae: Eupatorieae). Sida 21, 3: 1305–1321.
Nesom, G.L., Noyes, R.D. 1999. Notes on sectional delimitations in Erigeron (Asteraceae: Astereae). Sida 18: 1161–
1165.
583
Nesom, G.L., Noyes, R.D. 2000. Batopilasia (Asteraceae:
Astereae), a new genus from Chihuahua, Mexico. Sida
19: 79–84.
Nesom, G.L., Robinson, H., Granda Paucar, A. 2001. A new
species of Chiliotrichopsis (Asteraceae: Astereae) from
Peru. Brittonia 53: 430–434.
Nijs, H.C.M. den, Menken, S.B.J. 1996. Relations between
breeding system, ploidy level and taxonomy in some
advanced sections of Taraxacum. In: Hind, D.J.N.,
Beentje, H.J. (eds) Compositae: systematics. Proceedings of the International Compositae Conference, Kew,
1994 (D.J.N. Hind, Editor-in-Chief), vol. 1. Royal
Botanic Gardens, Kew, pp. 665–677.
Nordenstam, B. 1977. Senecioneae and Liabeae – systematic
review. In: Heywood, V.H., Harborne, J.B., Turner, B.L.
(eds) The biology and chemistry of the Compositae,
vol. 2. London: Academic Press, pp. 799–830.
Nordenstam, B. 1978. Taxonomic studies in the tribe
Senecioneae (Compositae). Opera Bot. 44: 1–83.
Nordenstam, B. 1994a. Tribe Calenduleae. In: Bremer, K.
(ed.) Asteraceae. Cladistics & classification. Portland,
Timber Press, pp. 365–376.
Nordenstam, B. 1994b. New combinations in the Calenduleae. Comp. Newslett. 25: 46–49.
Nordenstam, B. 1996. Recent revision of Senecioneae and
Calenduleae systematics. In: Hind, D.J.N., Beentje, H.J.
(eds) Compositae: systematics. Proceedings of the International Compositae Conference, Kew, 1994 (D.J.N.
Hind, Editor-in-Chief), vol. 1. Kew: Royal Botanic Gardens, pp. 591–596.
Nordenstam, B. 2004. Two new species of Osteospermum
(Compositae-Calenduleae) from Southwestern Cape
Province, South Africa. Edinburgh J. Bot. 60: 259–265.
Nordenstam, B. 2006a. Additions to the genus Jacobaea Mill.
(Compositae-Senecioneae). Comp. Newslett. 44: 12–
13.
Nordenstam, B. 2006b. New combinations in Nordenstamia
(Compositae-Senecioneae) from Argentina, Bolivia, Peru and Ecuador. Comp. Newslett. 44: 19–23.
Nordenstam, B. 2006c. Canariothamnus B. Nord., a new
genus of the Compositae-Senecioneae, endemic to the
Canary Islands. Comp. Newslett. 44: 24–31.
Nordenstam, B. 2006d. Generic revisions in the tribe Calenduleae (Compositae). Comp. Newslett. 44: 38–49.
Nordenstam, B. 2006e. New genera and combinations in the
Senecioneae of the Greater Antilles. Comp. Newslett.
44: 74–93.
Nordenstam, B. 2006f. Ignurbia, a new genus of the
Asteraceae-Senecioneae from Hispaniola. In: Sipman, H., Raus, Th., Kilian, N. (eds) Festschrift Werner
Greuter. Willdenowia spec. vol. 31, 1, pp. 463–468.
Nordenstam, B., Pelser, P.B. 2005. Dauresia and Mesogramma: one new and one resurrected genus of the
Asteraceae-Senecioneae from Southern Africa. Comp.
Newslett. 42: 74–88.
Nordenstam, B., Källersjö, M., Eldenäs, P. 2006.
Nephrotheca, a new monotypic genus of the
Compositae-Calenduleae from the southwestern
Cape. Province Comp. Newslett. 44: 32–37.
Norlindh, T. 1943. Studies in the Calenduleae, I. Monograph
of the genera Dimorphotheca, Castalis, Osteospermum,
Gibbaria and Chrysanthemoides. Lund: Gleerup.
584
Selected Bibliography to Compositae
Norlindh, T. 1960. Additions to the monograph on Osteospermum. Bot. Notiser 113: 385–399.
Norlindh, T. 1962. Studies in Calendula maderensis DC.
With a discussion on the delimitation of Calendula L.
from Gibbaria Cass. and Osteospermum L. Bot. Notiser
115: 437–445.
Norlindh, T. 1977. Calenduleae – systematic review. In: Heywood, V.H., Harborne, J.B., Turner, B.L. (eds) The biology and chemistry of the Compositae. London: Academic Press, pp. 961–987.
Noyes, R.D. 2000. Biogeographical and evolutionary insights on Erigeron and allies (Asteraceae) from ITS
sequence data. Pl. Syst. Evol. 220: 93–114.
Noyes, R.D., Rieseberg, L.H. 1999. ITS sequence data
support a single origin for North American Astereae
(Asteraceae) and reflect deep geographical division in
Aster s.l. Amer. J. Bot. 86: 398–412.
Oberprieler, Ch. 2001. Phylogenetic relationships in Anthemis (Compositae, Anthemideae) based on nrDNA
ITS sequence variation. Taxon 50: 745–762.
Oberprieler, Ch. 2002. A phylogenetic analysis of
Chamaemelum Miller (Compositae, Anthemideae)
and related genera based upon nrDNA ITS and cpDNA
trnL/trnF IGS sequence variation. Bot. J. Linn. Soc.
138: 255–273.
Oberprieler, C. 2004a. On the taxonomic status and the phylogenetic relationships of some unispecific Mediterranean genera of Compositae-Anthemideae. I. Brocchia, Endopappus, and Heliocauta. Willdenowia 34:
39–57.
Oberprieler, C. 2004b. On the taxonomic status and the phylogenetic relationships of some unispecific Mediterranean genera of Compositae-Anthemideae. II. Daveaua, Leococyclus, and Nananthea. Willdenowia 34:
341–350.
Oberprieler, C. 2005. Temporal and spatial diversification
of Circum-Mediterranean Compositae-Anthemideae.
Taxon 54: 951–966.
Oberprieler, Ch., Vogt, R. 2000. The position of Castrilanthemum Vogt & Oberprieler and the phylogeny of
Mediterranean Anthemideae (Compositae) as inferred
from nrDNA ITS and cpDNA trnL/trnF IGS sequence
variation. Pl. Syst. Evol. 225: 145–170.
Oberprieler, Ch., Vogt, R. 2002. Cladanthus Cass., p. 197. In:
Greuter, W., Raus, Th. (eds) Med-Checklist Notulae,
21. Willdenowia 32: 195–208.
Olmstead, R.G., Bremer, K., Scott, R.M., Palmer, J.D. 1993.
A parsimony analysis of the Asteridae sensu lato based
on rbcL sequences. Ann. Missouri Bot. Gard. 80: 700–
722.
Ortiz, S., Paiva, J.A.R., Rodríguez-Oubiña, J. 1996. An outline of the genus Anisopappus Hook. & Arn. (Compositae). Anales Jard. Bot. Madrid 54: 378–391.
Palmer, J.H. 1996. Floral initiation and production in the
oil-seed sunflower. In: Caligari, P.D.S., Hind, D.J.N.
(eds) Compositae: biology & utilization. Proceedings
of the International Compositae Conference, Kew, 1994
(D.J.N. Hind, Editor-in-Chief), vol. 2. Royal Botanic
Gardens, Kew, pp. 161–178.
Panero, J.L. 2005. New combinations and infrafamilial taxa
in the Asteraceae. Phytologia 87: 1–14.
Panero, J.L., Funk, V.A. 2002. Toward a phylogenetic subfamilial classification for the Compositae (Asteraceae).
Proc. Biol. Soc. Wash. 115: 909–922.
Panero, J.L., Jansen, R.K., Clevinger, J.A. 1999. Phylogenetic
relationships of the subtribe Ecliptinae (Asteraceae:
Heliantheae) based on chloroplast DNA restriction site
data. Amer. J. Bot. 86: 413–427.
Panero, J.L., Baldwin, B.G., Schilling, E.E., Clevinger, J.A.
2001a. Molecular phylogenetic studies of members
of tribes Helenieae, Heliantheae and Eupatorieae
(Asteraceae). 1. Introduction. In: Osborn, J.M. (prog.
dir.) Botany 2001, Abstracts, part 3. Systematics. St.
Louis, MO: Botanical Society of America.
Panero, J.L., Baldwin, B.G., Schilling, E.E., Clevinger, J.A.
2001b. Molecular phylogenetic studies of members of
tribes Helenieae, Heliantheae and Eupatorieae (Asteraceae). 2. Tribal/generic relationships. In: Osborn, J.M.
(prog. dir.) Botany 2001, Abstracts, part 3. Systematics.
St. Louis, MO: Botanical Society of America.
Panero, J.L., Baldwin, B.G., Schilling, E.E., Clevinger, J.A.
2001c. Molecular phylogenetic studies of members
of tribes Helenieae, Heliantheae and Eupatorieae
(Asteraceae). 3. General systematics and proposed
taxonomic changes in current classification. In:
Osborn, J.M. (prog. dir.) Botany 2001, Abstracts, part
3. Systematics. St. Louis, MO: Botanical Society of
America.
Pelser, P.B., Veldkamp, J.-F., van der Meijden R. 2006.
New combinations in Jacobaea Mill. (Asteraceae:
Senecioneae). Comp. Newslett. 44: 1–11.
Petit, D.P. 1997. Generic interrelationships of the Cardueae
(Compositae): a cladistic analysis of morphological
data. Pl. Syst. Evol. 207: 173–203.
Petit, D.P. 1998. Le genre Echinops L. (Compositae, Cardueae). I. Position phylétique et interprétation de l’incapitulescence. Candollea 43: 467–481.
Petit, D.F., Mathez, J., Qaid, H. 2000. Phylogeny of the Cardueae (Asteraceae) based on analysis of morphological
and palynological characters. Bocconea 13: 41–53.
Pittman, A.B., Bates, V. 1989. A new species of Polymnia
(Compositae: Heliantheae) from the Ouchita mountain
region of Arkansas. Sida 13: 481–486.
Poljakov, P.P. 1967. Origin and classification of the Compositae (in Russian). Akad. Nauk Kazakh. S.S.R. Inst.
Bot., 335 pp.
Powell, A.M. 1969. Taxonomy of Perityle section Pappothrix
(Compositae-Peritylinae). Rhodora 71: 58–93.
Powell, A.M. 1972. Taxonomy of Amauria (CompositaePeritylinae). Madroño 21: 516–525.
Powell, A.M. 1973. Taxonomy of Perityle section Laphamia
(Compositae-Helenieae-Peritylinae). Sida 5: 61–128.
Powell, A.M. 1974. Taxonomy of Perityle section Perityle
(Compositae-Peritylinae). Rhodora 76: 229–306.
Powell, A.M., Turner, B.L. 1974. A generic conspectus of
the subtribe Peritylinae (Asteraceae-Helenieae) and
reassessment of its tribal position. Amer. J. Bot. 61:
87–93.
Praglowski, J., Grafström, E. 1980. The pollen morphology
of the tribe Calenduleae with reference to taxonomy.
Bot. Notiser 133: 177–188.
Proksch, P., Kunze, A. 1996. Chemosystematic evidence
from prenylated acetophenones – conclusions at the
tribal, inter- and intrageneric level. In: Hind, D.J.N.,
Beentje, H.J. (eds) Compositae: systematics. Proceedings of the International Compositae Conference, Kew,
1994 (D.J.N. Hind, Editor-in-Chief), vol. 1. Royal
Botanic Gardens, Kew, pp. 295–305.
Selected Bibliography to Compositae
Pullaiah, T. 1983. Studies in the embryology of Senecioneae
(Compositae). Pl. Syst. Evol. 142: 61–70.
Rauscher, J.T. 2002. Molecular phylogenetics of the Espeletia complex (Asteraceae): evidence from nrDNA ITS
sequences of the closest relatives of an Andean adaptive radiation. Amer. J. Bot. 89: 1074–1084.
Reese, H. 1989. Die Entwicklung von Perikarp und Testa
bei Calenduleae und Arctotideae (Asteraceae) – ein
Beitrag zur Systematik. Bot. Jahrb. Syst. 110: 325–419.
Reitbrecht, F. 1974. Fruchtanatomie und Systematik der
Anthemideae (Asteraceae). Diss. philosophischen
Fakultät, Universität Wien, 160 S.
Reveal, J.L. 1997. Early suprageneric names in Asteraceae.
Comp. Newslett. 30: 29–45.
Roberts, R.P., Urbatsch, L.E. 2004. Molecular phylogeny of
Chrysothamnus and related genera (Asteraceae, Astereae) based on nuclear ribosomal 3′ ETS and ITS sequence data. Syst. Bot. 29: 199–215.
Robins, D.J. 1977. Senecioneae – chemical review. In: Heywood, V.H., Harborne, J.B., Turner, B.L. (eds) The biology and chemistry of the Compositae. London: Academic Press, pp. 831–850.
Robinson, B.L. 1930a. Observations on the genus Stevia.
Contr. Gray Herb. n.s. 90: 36–58, pl. 1.
Robinson, B.L. 1930b. The stevias of the Argentine republic.
Contr. Gray Herb. n.s. 90: 58–79.
Robinson, B.L. 1930c. The stevias of Paraguay. Contr. Gray
Herb. n.s. 90: 79–90.
Robinson, B.L. 1930d. The stevias of North America. Contr.
Gray Herb. n.s. 90: 90–160.
Robinson, B.L. 1931a. The stevias of Colombia. Contr. Gray
Herb. n.s. 96: 28–36.
Robinson, B.L. 1931b. The stevias of Venezuela. Contr. Gray
Herb. n.s. 96: 37–43.
Robinson, B.L. 1931c. The stevias of Ecuador. Contr. Gray
Herb. n.s. 96: 43–49.
Robinson, B.L. 1932a. The stevias of Peru. Contr. Gray Herb.
n.s. 100: 20–36.
Robinson, B.L. 1932b. The stevias of Bolivia. Contr. Gray
Herb. n.s. 100: 36–39.
Robinson, H. 1977. An analysis of the characters and relationships of the tribes Eupatorieae and Vernonieae
(Asteraceae). Syst. Bot. 2: 199–208.
Robinson, H. 1978a. Studies in the Heliantheae (Asteraceae).
XIV. Validation of subtribes. Phytologia 41: 39–44.
Robinson, H. 1978b. Studies in the Heliantheae (Asteraceae). XII. Re-establishment of the genus Smallanthus. Phytologia 39: 47–53.
Robinson, H. 1979. Two new genera of the Vernonieae (Asteraceae) from Brazil, Heterocypsela and
Pseudostifftia. Phytologia 44: 442–450.
Robinson, H. 1981. A revision of the tribal and subtribal
limits of the Heliantheae (Asteraceae). Smithsonian
Contr. Bot. 51: 1–102.
Robinson, H. 1983a. A generic review of the tribe Liabeae
(Asteraceae). Smithsonian Contr. Bot. 54: 1–69.
Robinson, H. 1983b. Studies in the Liabeae (Asteraceae).
XVI. New taxa from Peru. Phytologia 54: 62–65.
Robinson, H. 1984. Style rotation in the Asteraceae. Taxon
33: 400–404.
Robinson, H. 1990a. Notes on Sinclairia and Liabellum in
Mesoamerica (Liabeae: Asteraceae). Phytologia 69: 57–
60.
585
Robinson, H. 1990b. A redelimitation of Microliabum Cabrera (Asteraceae: Liabeae). Syst. Bot. 15: 736–744.
Robinson, H. 1992. Observations on the unique form of
sweeping hairs on the styles of the Eremothamneae
(Asteraceae). Taxon 41: 199–200.
Robinson, H. 1994. Notes on the tribes Eremothamneae, Gundelieae and Moquinieae, with comparisons
of their pollen. Taxon 43: 33–44.
Robinson, H. 1996. The status of generic and subtribal
revisions in the Vernonieae. In: Hind, D.J.N., Beentje, H.J. (eds) Compositae: systematics. Proceedings of
the International Compositae Conference, Kew, 1994
(D.J.N. Hind, Editor-in-Chief), vol. 1. Royal Botanic
Gardens, Kew, pp. 511–529.
Robinson, H. 1997. New species of Aphanactis in
Ecuador and Bolivia and new combinations in Selloa
(Heliantheae: Asteraceae). Brittonia 49: 71–78.
Robinson, H. 1999a. Revisions of Paleotropical Vernonieae
(Asteraceae). Proc. Biol. Soc. Wash. 112: 220–247.
Robinson, H. 1999b. Generic and subtribal classification
of American Vernonieae. Smithsonian Contr. Bot. 89:
i–iv, 1–116.
Robinson, H. (compiler) 1999c. Eupatorieae in Asteraceae.
In: Jørgensen, R.M., León-Yánez, S. (eds) Catalogue
of vascular plants of Ecuador. Monogr. Syst. Bot. 75:
260–314.
Robinson, H., Brettell, R.D. 1972. Tribal revisions in the
Asteraceae. I. The relationship of Geissolepis. Phytologia 24: 299–301.
Robinson, H., Brettell, R.D. 1973a. Tribal revisions in the
Asteraceae. III. A new tribe, Liabeae. Phytologia 25:
404–407.
Robinson, H., Brettell, R.D. 1973b. Tribal revisions in the
Asteraceae. V. The relationship of Rigiopappus. Phytologia 26: 69–70.
Robinson H., Brettell, R.D. 1973c. Tribal revisions in the
Asteraceae. VIII. A new tribe, Ursinieae. Phytologia
26: 76–86.
Robinson, H., Brettell, R.D. 1973d. Tribal revisions in the
Asteraceae. X. The relationship of Plagiocheilus. Phytologia 26: 159–162.
Robinson, H., Brettell, R.D. 1974. Studies in the Liabeae
(Asteraceae). II. Preliminary survey of the genera. Phytologia 28: 43–63.
Robinson, H., Funk, V.A. 1987. A phylogenetic analysis of
Leiboldia, Lepidonia, and a new genus Stramentopappus (Vernonieae: Asteraceae). Bot. Jahrb. Syst. 108:
213–228.
Robinson, H., Kahn, B. 1986. Trinervate leaves, yellow
flowers, tailed anthers, and pollen variation in
Distephanus Cassini (Vernonieae: Asteraceae). Proc.
Biol. Soc. Wash. 99: 493–501.
Robinson, H., Marticorena, C. 1986. A palynological study
of the Liabeae (Asteraceae). Smithsonian Contr. Bot.
64: i–iv, 1–50.
Robinson, H., Moore, A.J. 2004. New species and new combinations in Rhysolepis (Asteraceae: Heliantheae). Proc.
Biol. Soc. Wash. 117: 423–446.
Robinson, H., Powell, A.M., King, R.M., Weedin, J.F. 1985.
Chromosome numbers in Compositae. XI. Liabeae.
Ann. Missouri Bot. Gard. 72: 469–479.
Robinson, H., Powell, A.M., Carr, G.D., King, R.M.,
Weedin, J.F. 1989. Chromosome numbers in Composi-
586
Selected Bibliography to Compositae
tae. XVI. Eupatorieae II. Ann. Missouri Bot. Gard. 76:
1004–1011.
Ross-Craig, S. 1962. Drawings of British plants, vol. 17. London: Bell & Hyman.
Ross-Craig, S. 1963. Drawings of British plants, vol. 18. London: Bell & Hyman.
Rydberg, P.A. 1927. Carduaceae, Liabeae, Neurolaeneae, Senecioneae (pars). N. Amer. Flora 34: 289–360.
Ryding, O., Bremer, K. 1992. Phylogeny, distribution, and
classification of the Coreopsideae (Asteraceae). Syst.
Bot. 17: 649–659.
Sáenz, A.A. 1981. Anatomía y morfología de frutos de Heliantheae. Darwiniana 23: 37–117.
Sandwith, N.Y. 1956. Contributions to the flora of tropical
America. LXI. Notes on Philoglossa. Kew Bull. 1956:
289–293.
Schilling, E.E., Cox, P.B. 2001. Abstract 555. Systematic analysis of Liatrinae (Asteraceae). In: Osborn, J.M. (prog.
dir.) Botany 2001, Abstracts, part 3. Systematics. St.
Louis, MO: Botanical Society of America.
Schilling, E.E., Panero, J.L. 2002. A revised classification
of subtribe Helianthinae (Asteraceae-Heliantheae). 1.
Basal lineages. Bot. J. Linn. Soc. 140: 65–76.
Schilling, E.E., Panero, J.L., Cox, P.B. 1999. Chloroplast DNA
restriction data support a narrowed interpretation of
Eupatorium (Asteraceae). Pl. Syst. Evol. 219: 209–233.
Schmidt, G.J., Schilling, E.E. 2000. Phylogeny and biogeography of Eupatorium (Asteraceae: Eupatorieae) based
on nuclear ITS sequence data. Amer. J. Bot. 87: 716–726.
Seaman, F.C. 1982. Sesquiterpene lactones as taxonomic
characters in the Asteraceae. Bot. Rev. 48: 121–592.
Seaman, F.C., Funk, V.A. 1983. Cladistic analysis of complex natural products: developing transformation series from sesquiterpene lactone data. Taxon 32: 1–27.
Sell, P.D. 1975. Taxonomic and nomenclatural notes on the
Compositae subfamily Cichorioideae. Bot. J. Linn. Soc.
71: 236–274.
Sennikov, A.N., Illarinova, I.D. 2001. The morphological
and anatomical structure of the achenes of species of
the genus Hieracium (Asteraceae) and related genera
(in Russian). Bot. Zhurn. (Moscow & Leningrad) 86, 3:
37–59.
Shannon, R.K., Wagner, W.L. 1997. Oparanthus revisited.
In: Lorence, D. (ed.) Botanical results of the 1988 Fatu
Hiva Expedition to the Marquesas Islands. Allertonia
7: 273–295.
Shih, C., Chen, Y.L. 1996. Faberiopsis Shih et Y.L. Chen,
genus novum familiae Compositarum sinensium. Acta
Phytotax. Sin. 34: 438–439.
Short, P.S., Wilson, K.E., Nailon, J. 1989. Notes on the fruit
anatomy of Australian members of the Inuleae (Compositae). Muelleria 7: 57–79.
Skottsberg, C. 1937. Die Flora der Desventuradas Inseln
(San Felix und San Ambrosio). Göteb. Kungl. Vetensk.
Vitterh. Samh. Handl. ser. 5 B 5: 1–88.
Skvarla, J.J., Turner, B.L., Patel, V.C., Tomb, A.S. 1977.
Pollen morphology in the Compositae and in morphologically related families. In: Heywood, V.H.,
Harborne, J.B., Turner, B.L. (eds) The biology and
chemistry of the Compositae. London: Academic
Press, pp. 141–265.
Smith, C.R., Wilson, T.L., Melvin, E.H., Wolff, I.A. 1960.
Dimorphecolic acid – a unique hydroxydienoid fatty
acid. J. Amer. Chem. Soc. 82: 1417–1421.
Solbrig, O.T. 1977. Chromosomal cytology and evolution
in the family Compositae. In: Heywood, V.H., Harborne, J.B., Turner, B.L. (eds) The biology and chemistry of the Compositae. London: Academic Press, pp.
267–281.
Stebbins, G.L. 1953. A new classification of the tribe Cichorieae, family Compositae. Madroño 12: 65–81.
Stix, E. 1960. Pollenmorphologische Untersuchungen an
Compositen. Grana Palynol. 2: 41–114.
Strother, J.L. 1977. Tageteae – systematic review. In: Heywood, V.H., Harborne, J.B., Turner, B.L. (eds) The biology and chemistry of the Compositae, vol. 2. London:
Academic Press, pp. 769–783.
Strother, J.L. 1986. Renovation of Dyssodia (Composiate:
Tageteae). Sida 11: 371–378.
Strother, J.L., Panero, J.L. 1994. Chromosome studies: Latin
American Compositae. Amer. J. Bot. 81: 770–775.
Stuessy, T.F. 1977. Heliantheae – systematic review. In: Heywood, V.H., Harborne, J.B., Turner, B.L. (eds) The biology and chemistry of the Compositae, vol. 2. London:
Academic Press, pp. 621–671.
Stuessy, T.F., Garver, D. 1996. The defensive role of pappus in
heads of Compositae. In: Caligari, P.D.S., Hind, D.J.N.
(eds) Compositae: biology & utilization. Proceedings
of the International Compositae Conference, Kew, 1994
(D.J.N. Hind, Editor-in-Chief), vol. 2. Royal Botanic
Gardens, Kew, pp. 81–91.
Stuessy, T.F, Spooner, D.M. 1988. The adaptive and phylogenetic significance of receptacular bracts in the Compositae. Taxon 37: 114–126.
Stuessy, T.F., Sang, T., DeVore, M.L. 1996. Phylogeny and
biogeography of the subfamily Barnadesioideae with
implications for early evolution of Compositae. In:
Hind, D.J.N., Beentje, H.J. (eds) Compositae: systematics. Proceedings of the International Compositae Conference, Kew, 1994 (D.J.N. Hind, Editor-in-Chief), vol.
1. Royal Botanic Gardens, Kew, pp. 463–490.
Susanna, A., Garcia-Jacas, N., Soltis, D.E., Soltis, P.S. 1995.
Phylogenetic relationships in tribe Cardueae (Asteraceae) based on ITS sequences. Amer. J. Bot. 82: 1056–
1068.
Susanna, A., Garcia-Jacas, N., Hidalgo, O., Vilatersana, R.,
Garnatje, T. 2006. The Cardueae (Compositae) revisited: insights from a ITS, trnL-trnF and matK nuclear and chloroplast DNA analysis. Ann. Missouri Bot.
Gard. 93:150–171.
Suyama, C. 2001. A new naturalized plant, Gymnocoronis
spilanthoides DC. (in Japanese). J. Phytogeogr. Taxon.
49: 183–184.
Swenson, U. 1995. Systematics of Abrotanella, an Amphipacific genus of Asteraceae (Senecioneae). Pl. Syst.
Evol. 197: 149–193.
Swenson, U., Bremer, K. 1999. On the circumscription of the
Blennospermatinae (Asteraceae, Senecioneae) based
on ndhF sequence data. Taxon 48: 7–14.
Tadesse, M., Crawford, D.J., Smith, E.B. 1995. Comparative
capitular morphology and anatomy of Coreopsis L. and
Bidens L. (Compositae) including a review of generic
boundaries. Brittonia 47: 61–91.
Tadesse, M., Crawford, D.J., Kim, S.-C. 2001. A cladistic
analyses of morphological features in Bidens L. and
Coreopsis L. (Asteraceae: Heliantheae) with notes on
generic delimitation and systematics. In: Friis, I., Ryding, O. (eds) Biodiversity research in the Horn of Africa
Selected Bibliography to Compositae
region. Biologiske Skrifter, The Royal Danish Academy
of Sciences and Letters, Copenhagen, pp. 85–102.
Tegel, F. 2002. Die Testaepidermis der Lactuceae
(Asteraceae) – ihre Diversität und systematische Bedeutung. Diss., Universität München (http://
edoc.ub.uni-muenchen.de/archive/00000104/01/Tegel_
Friedrich.pdf).
Thiele, M. 1988. Bau und Funktion des Antheren-GriffelKomplexes der Compositen. Diss. Bot. 117, 169 pp.
Stuttgart: J. Cramer.
Tomb, A.S. 1976. Pollen morphology in tribe Lactuceae
(Compositae). Grana 15: 79–89.
Tomb, A.S. 1977. Lactuceae – systematic review. In: Heywood, V.H., Harborne, J.B., Turner, B.L. (eds) The biology and chemistry of the Compositae. London: Academic Press, pp. 1067–1079.
Torrell, M., Garcia-Jacas, N., Susanna, A., Vallès, J. 1999.
Phylogeny in Artemisia (Asteraceae, Anthemideae) inferred from nuclear ribosomal DNA (ITS) sequences.
Taxon 48: 721–736.
Turner, B.L. 1980. La taxonomía del género Aphanactis
(Asteraceae-Heliantheae). Bol. Soc. Argent. Bot. 19:
33–44.
Turner, B.L. 1987a. Submergence of the genera Caterothamnus and Oaxacania into Hofmeisteria (Eupatorieae,
Asteraceae). Phytologia 63: 415–416.
Turner, B.L. 1987b. Taxonomy of Carphochaete (Asteraceae–
Eupatorieae). Phytologia 64, 2: 145–162.
Turner, B.L. 1988. Submergence of the genera Asanthus
and Discritogyne within Steviopsis (Asteraceae, Eupatorieae), including new combinations. Phytologia 64:
259–262.
Turner, B.L. 1989. Revisionary treatment of the genus
Sinclairia, including Liabellum (Asteraceae, Liabeae).
Phytologia 67: 168–206.
Turner, B.L. 1990. A new species of Steviopsis (Asteraceae:
Eupatorieae) from Nuevo Leon, Mexico. Phytologia 68:
410–412.
Turner, B.L. 1991a. Brickellia sonorana (Asteraceae), a new
species from México belonging to the subgenus
Phanerostylis. Phytologia 71: 51–56.
Turner, B.L. 1991b. Recension of the Cronquistianthus group
of Eupatorium (Asteraceae). Phytologia 70: 158–177.
Turner, B.L. 1994. Taxonomic status of Brickelliastrum villarrealii R.M. King & H. Robins. (Asteraceae, Eupatorieae). Phytologia 76: 389–390.
Turner, B.L. 1995. Synopsis of Ageratella (Asteraceae, Eupatorieae). Phytologia 78: 204–208.
Turner, B.L. 1997. Eupatorieae. In: Turner, B.L. (ed.) The
Compositae of Mexico. A systematic account of the
family Asteraceae, vol. 1. Phytologia Mem. 11: i–iv, 1–
272.
Turner, B.L., Powell, A.M. 1977. Helenieae – systematic review. In: Heywood, V.H., Harborne, J.B., Turner, B.L.
(eds) The biology and chemistry of the Compositae.
London: Academic Press, pp. 699–737.
Turner, B.L., Powell, A.M., Cuatrecasas, J. 1967. Chromosome numbers in the Compositae. XI. Peruvian
species. Ann. Missouri Bot. Gard. 54: 172–177.
Turner, B.L., Kim, K.J., Norris, J. 1991. Taxonomic status
of Barroetea glutinosa (Asteraceae, Eupatorieae) and
its allies: morphological evidence for the transfer of
Barroetea to Brickellia. Phytologia 71: 38–50.
587
Urbatsch, L.E., Roberts, R.P. 2004. New combinations in the
genus Gundlachia and four new genera of Astereae
(Asteraceae) from northern Mexico and the southern
United States. Sida 21: 243–257.
Urbatsch, L.E., Roberts, R.P., Karaman, V. 2003. Phylogenetic evaluation of Xylothamia, Gundlachia, and
related genera (Asteraceae, Astereae) based on ETS
and ITS nrDNA sequence data. Amer. J. Bot. 90:
634–649.
Urtubey, E., Stuessy, T.F. 2001. New hypotheses of phylogenetic relationships in Barnadesioideae (Asteraceae)
based on morphology. Taxon 50: 1043–1063.
Urtubey, E., Tellería, M.C. 1998. Pollen morphology of the
subfamily Barnadesioideae (Asteraceae) and its phylogenetic and taxonomic significance. Rev. Palaeobot.
Palynol. 104: 19–37.
Valadon, L.R.G. 1977. Arctoteae and Calenduleae – chemical
review. In: Heywood, V.H., Harborne, J.B., Turner, B.L.
(eds) The biology and chemistry of the Compositae,
vol. 2. London: Academic Press, pp. 989–998.
Vallès, J., Blanché, C., Bonet, M.À., Agelet, A., Muntané, J.,
Raja, D., Parada, M. 1996. A contribution to the ethnobotany of the Asteraceae in Catalonia. In: Caligari, P.D.S.,
Hind, D.J.N. (eds) Compositae: biology & utilization.
Proceedings of the International Compositae Conference, Kew, 1994 (D.J.N. Hind, Editor-in-Chief), vol. 2.
Royal Botanic Gardens, Kew, pp. 453–466.
Vallès, J., Torrell, M., Garnatje, T., Garcia-Jacas, N., Vilatersana, R., Susanna, A. 2003. The genus Artemisia and its
allies: phylogeny of the subtribe Artemisiinae (Asteraceae, Anthemideae) based on nucleotide sequences
of nuclear ribosomal DNA internal transcribed spacers (ITS). Pl. Biol. 5: 274–284.
Velez, M.C. 1981. Karpologische Untersuchungen an
amerikanischen Astereae (Compositae). Mitt. Bot.
Staatssamml. München 17: 1–170.
Vezey, E.L., Watson, L.E., Skvarla, J.J., Estes, J.R. 1994. Plesiomorphic and apomorphic pollen structure characteristics of Anthemideae (Asteroideae: Asteraceae).
Amer. J. Bot. 81: 648–657.
Viswanathan, M.V., Singh, H.B. 1996. Potential industrial
uses of some less-known Asteraceae of India. In: Caligari, P.D.S., Hind, D.J.N. (eds) Compositae: biology
& utilization. Proceedings of the International Compositae Conference, Kew, 1994 (D.J.N. Hind, Editor-inChief), vol. 2. Royal Botanic Gardens, Kew, pp. 643–
659.
Voytenko, V.F. 1989a. Heterocarpy (heterodiaspory) in angiospermous plants: analysis of the concept, classification, terminology (in Russian). Bot. Zhurn. (Moscow
& Leningrad) 74: 281–297.
Voytenko, V.F. 1989b. The typology and evolution of forms
of heterocarpy in the tribe Lactuceae (Asteraceae) (in
Russian). Bot. Zhurn. (Moscow & Leningrad) 74: 1241–
1257.
Voytenko, V.F., Oparina, S.N. 1990. Comparative analysis of
anatomical structure of fruits in heterocarpous representatives of the tribe Lactuceae (Asteraceae) (in Russian). Bot. Zhurn. (Moscow & Leningrad) 75, 3: 299–
314.
Wagenitz, G. 1997. The impact of molecular methods on
the systematics of angiosperms. Bot. Acta 110: 274–
281.
588
Selected Bibliography to Compositae
Wagenitz, G., Hellwig, F.H. 1996. Evolution of characters and
phylogeny of the Centaureinae. In: Hind, D.J.N., Beentje, H.J. (eds) Compositae: systematics. Proceedings of
the International Compositae Conference, Kew, 1994
(D.J.N. Hind, Editor-in-Chief), vol. 1. Royal Botanic
Gardens, Kew, pp. 491–510.
Wagenitz, G., Dittrich, M., Damboldt, J. 1982. Centaurothamnus, eine neue Gattung der CompositaeCardueae in Arabien. Candollea 37: 101–115.
Wagner, H. 1977. Cynareae – chemical review. In: Heywood, V.H., Harborne, J.B., Turner, B.L. (eds) The biology and chemistry of the Compositae, vol. 2. London:
Academic Press, pp. 1017–1038.
Wagstaff, S.J., Breitwieser, I. 2002. Phylogenetic relationships of New Zealand Asteraceae inferred from ITS
sequences. Pl. Syst. Evol. 231: 203–224.
Wagstaff, S.J., Breitwieser, I. 2004. Phylogeny and classification of Brachyglottis (Senecioneae, Asteraceae): an
example of rapid species radiation in New Zealand.
Syst. Bot. 29: 1003–1010.
Walsh, N.G. 1999. Centipeda. In: Walsh, N.G., Entwisle, T.J.
(eds) Flora of Victoria, vol. 4. Dicotyledons, Cornaceae
to Asteraceae. Melbourne: Inkata Press, pp. 720–723.
Walsh, N.G., Entwisle, T.J. (eds) 1999. Flora of Victoria, vol.
4. Melbourne: Inkata Press.
Watanabe, K., King, R.M., Tetsukaza, Y., Ito, M.,
Yokoyama, J., Suzuki, T., Crawford, D.J. 1995. Chromosomal cytology and evolution in Eupatorieae
(Asteraceae). Ann. Missouri Bot. Gard. 82: 581–592.
Watson, L.E., Evans, T.M. 1999. Phylogeny and biogeography of the tribe Anthemideae. In: Anon. (ed.) XVI
International Botanical Congress, Abstracts, p. 358. St.
Louis: Missouri Botanical Garden.
Watson, L.E., Evans, T.M., Boluarte, T. 2000. Molecular
phylogeny of tribe Anthemideae (Asteraceae), based
on chloroplast gene ndhF. Mol. Phylog. Evol. 15:
59–69.
Watson, L.E., Bates, P.L., Evans, T.M., Unwin, M.M.,
Estes, J.R. 2002. Molecular phylogeny of subtribe
Artemisiinae (Asteraceae), including Artemisia and
its allied and segregate genera. BMC Evol. Biol. 2:
17.
Weddell, H.A. 1855–1857. Chloris Andina, vol. 1. Paris: P.
Bertrand.
Weitz, F.M. 1989. A revision of the genus Corymbium (Asteraceae). S. African J. Bot. 55: 598–629.
Werker, E. 1997. Seed anatomy. In: Handbuch der
Pflanzenanatomie, X. 3. 1. Berlin: Bornträger, 424 pp.
Whitton, J., Wallace, R.S., Jansen, R.K. 1995. Phylogenetic
relationships and patterns of character change in the
tribe Lactuceae (Asteraceae) based on chloroplast DNA
restriction site variation. Canad. J. Bot. 73: 1058–1073.
Wilson, P.G. 1992. The Lawrencella complex (Asteraceae:
Gnaphalieae: Angianthinae) of Australia. Nuytsia 8:
361–377
Wood, A.R., Nordenstam, B. 2004. An interesting new
species of Osteospermum (Asteraceae-Calenduleae)
from the western Cape Province, South Africa,
providing a link to the genus Chrysanthemoides. S.
African J. Bot. 69: 572–578.
Wulff, A.F., Hunziker, J.H., Escobar, A. 1996. Estudios cariologicos en Compositae, VII. Darwiniana 34: 213–231.
Wussow, J.R., Urbatsch, L.E., Sullivan, G.A. 1985. Calea
(Asteraceae) in Mexico, Central America, and Jamaica.
Syst. Bot. 10: 241–267.
Yahara, T., Kawahara, T., Crawford, D.J., Ito, M., Watanabe, K. 1989. Extensive gene duplication in diploid Eupatorium (Asteraceae). Amer. J. Bot. 76: 1247–1253.
Zachariades, C., Muniappan, R., Strathie, L.W. (eds) 2002.
Proceedings 5th International Workshop Biological
Control and Management of Chromolaena odorata,
Durban, South Africa, 23–25 October 2000. Pretoria:
ARC-Plant Protection Research Institute, 186 pp.
Zardini, E.M. 1974. Sobre la presencia del genero Gerbera
en America. Bol. Soc. Argent. Bot. 16: 103–108.
Zdero, C., Bohlmann, F. 1988. Macrolide diterpenes and
other ent-labdanes from Corymbium villosum. Phytochemistry 27: 227–231.
Zdero, C., Bohlmann, F. 1990. Systematics and evolution
within the Compositae, seen with the eyes of a chemist.
Pl. Syst. Evol. 171: 1–14.
Zdero, C., Bohlmann, F., King, R.M., Robinson, H. 1986.
Further 5-methyl coumarins and other constituents
from the subtribe Mutisiinae. Phytochemistry 25: 509–
516.
Zhang, X., Bremer, K. 1993. A cladistic analysis of the tribe
Astereae (Asteraceae) with notes on their evolution and
subtribal classification. Pl. Syst. Evol. 184: 256–283.
Zhao, Z., Skvarla, J.J., Jansen, R.K., DeVore, M.L. 2000. Phylogenetic implications of pollen morphology and ultrastructure in the Barnadesioideae (Asteraceae). Lundellia 3: 26–40.
Goodeniaceae1
Goodenovieae R. Br., Prodr.: 573 (1810), nom. cons.
Brunoniaceae R. Br. (1816).
R.C. Carolin
Herbs, subshrubs, shrubs or rarely scramblers, glabrous or hairy with simple or branched or glandular hairs, sometimes viscid and varnished with age.
Leaves mostly spirally arranged, rarely opposite, often with an axillary tuft of hairs; blades simple, with
entire to dentate or rarely lobed margins; stipules
absent. Flowers in cymes, thyrses, racemes, spikes,
subumbels or heads or rarely solitary. Sepals usually 5, rarely 3, sometimes connate, sometimes reduced to a rim or obsolete. Corolla tubular, usually
slit on the adaxial side, often 2-lipped, sometimes
fan-like, sometimes with a pouch or spur; lobes 5,
often winged, often with auricles on margin of the
adaxial slit which surround the indusium. Stamens
5, opposite the sepals, epigynous or epipetalous
or apparently almost hypogynous; anthers free or
connate, 2-locular, dehiscing through longitudinal slits. Ovary superior, half-inferior or inferior,
usually 2-locular (often incompletely so), rarely 4locular or apparently 1-locular; style usually entire
with a hollow pollen-cup (indusium) usually enclosing the 2-fid or unlobed stigma, rarely 2–4-fid.
Fruit a drupe, inferior nut, or capsule dehiscing
through 2–4 valves, rarely separating transversely
into 1-seeded woody articles. Seeds with or without
endosperm, rarely with a caruncle; embryo terete
or flattened.
Eleven genera and c. 440 species, mostly in the
southern hemisphere, mainly Australia.
Vegetative Morphology. Most species in the
family are either herbs, shrubs or subshrubs, with
Scaevola enantophylla and S. oppositifolia being
the only woody scramblers. Scaevola spinescens is
thorny. Many of the herbs are relatively ephemeral,
occurring in the more arid areas of Australia and
becoming often very common after rain. Only a few
species are large shrubs. Leaves are simple and arranged spirally in almost all species, with a variety
of shapes, but opposite in Scaevola enantophylla.
1
Updated by J.W. Kadereit and M.H.G. Gustafsson.
Vegetative Anatomy. Carlquist (1969) gives
a detailed account of the wood anatomy in the
family. The species from drier habitats (“xeromorphic”) tend to have short narrow vessels
whilst those from wetter regions (“mesic”) have
long wider vessels. Abundant crystals, possibly of
calcium oxalate, are usually found in species from
drier regions. Libriform fibres have not been found
in the family and there is a relative abundance of
uniseriate rays. Krause (1912) notes the occurrence
of medullary bundles in Goodenia spp., e.g. G.
ovata, and describes them as leaf traces. Rajput
and Carolin (1984) show that the cortical bundles
in Dampiera are leaf bundles from the node(s)
above, and that the number found in any crosssection is related to the phyllotaxis of that shoot.
An illustration by Krause (1912) indicates that
there may be a similar situation in Lechenaultia.
Trichomes are widespread through the family,
and in Dampiera these may be particularly dense.
Carolin (1970) gives a detailed account of the types
to be found, including glandular, stellate, simple,
strigose and dendritic hairs. Groups of species
within Goodenia, Coopernookia and Scaevola
secrete a viscous fluid from glandular hairs on the
leaves and stems, and in some of these species
this secretion hardens to a varnish. Stomates are
anomocytic.
Inflorescence Structure. There are two
distinct types of inflorescence in the family (for
a discussion of these types, see Carolin 1967).
In Goodenia, Scaevola, Coopernookia, Selliera,
Verrauxia, Pentaptilon and Diaspasis, the primitive form of the inflorescence is a thyrse without
a terminal flower. More often, the reduction of
each constituent axillary cyme to a single flower
has resulted in a raceme or spike. In a number of
Goodenia spp., the racemes are condensed into
subumbels. In most Velleia spp., the condensation
of the main axis and the expansion of each axillary
cyme has given a very characteristic form to the
590
R.C. Carolin
plant. In Dampiera, Lechenaultia, Anthotium and
Brunonia, the inflorescence is based on a cymopanicle with a terminal flower (see also Rajput
and Carolin 1988). Tightly condensed head-like
inflorescences are found in Brunonia australis,
Dampiera wellsiana, D. eriocephala, D. dentata
and Scaevola globosa. These may elongate strongly
when fruiting.
Flower Structure. Carolin (1959; 1966) has
examined the flower structure in detail. In all
genera, the petals are connate to some degree
but, except for Brunonia, the corolla tube is slit
to the base (or nearly so) on the adaxial side.
In almost all species (Brunonia excluded), each
corolla lobe has a thin but often large, brightly
coloured wing on either side. In Scaevola, the
flower is opened out on either side of this slit,
exposing the pollen presentation apparatus and
stigma, to form a fan-shaped corolla. In other
genera, the adaxial corolla lobes are often folded
around the indusium and, in some cases, the
corolla is distinctly 2-lipped (e.g. most species of
Velleia, Dampiera and Lechenaultia, and many of
Goodenia) to such an extent that adaxial and abaxial corolla lobes may be different in colour (e.g.
Goodenia bicolor). In many species of Goodenia
and throughout Dampiera and Velleia, the lower
part of the wings of the corolla lobes on either side
of the slit are modified into auricles which enclose
the indusium. Petal venation has been described by
Gustafsson (1995), and petals were found to have
three major veins meeting at the apex. These veins
anastomose or not. The corolla shows a wide range
of colours, particularly in Lechenaultia which has
been developed as an horticultural plant. In most
species, however, the corolla is various shades of
yellow or blue. Stamens are epipetalous or free
from the petals in Velleia. The anthers are free or
connate and dehisce in the bud, and the pollen
is deposited inside a cup at the top of the style
(indusium) which grows up through the dehisced
stamens (Brough 1927; Carolin 1960; Leins and
Erbar 1989). The indusium (Fig. 123) may be
equipped with hairs (e.g. Goodenia, Velleia), or
it may be glabrous as in Dampiera. The pollen is
presented in the indusium when the flower opens.
However, pollen collection by the indusium is
rather inefficient, and much pollen is also found
around the auricles and on the outside of the
indusium. The indusium is an outgrowth from
the style (Carolin 1960) and surrounds the stigma
initially, which eventually grows out of the cup to
appear as a 2-lobed structure. In Lechenaultia,
however, the pollen is presented entirely on the
outside of the style and the stigmatic surface is
on the upper surface of the style near the top. The
homologies of the stylar parts of this genus are
not clear. The ovary is superior, half-inferior or
inferior. In Goodenia, the ovary is usually inferior
but, in G. macroplectra, the sepals are free from
the ovary and are inserted below it, whereas the
corolla is at least partially adnate to the ovary
and epigynous. Almost throughout the family
the ovary consists of 3 zones: a basal 2-locular
zone which may be very short, a middle 1-locular
zone and an upper 2-locular zone (Carolin 1959;
see also Leins and Erbar 1989). Ovules may be
inserted in either of the 2 lower zones on axile or
parietal placentas respectively. There is, however,
anatomical evidence to suggest that the ovary
is 4-carpellary (Carolin 1959, 1966). In Scaevola
porocarya, there are 4 fertile locules whereas in
Fig. 123. Goodeniaceae. Stylar cup and stigma. A Brunonia
australis. B Selliera radicans. (SEM photographs by C. Erbar
and P. Leins)
Goodeniaceae
many other species of Scaevola there are 2 fertile
loculi and 2 sterile cavities. The homologies of
the sterile cavities are unclear (Carolin 1966). The
ontogenetic studies of Leins and Erbar (1989),
however, do not appear to show any evidence for
a 4-carpellate ovary.
Embryology. Early accounts of embryology include those of Rosén (1937, 1946), and a summary
of embryological characters has been presented by
Tobe and Morin (1996). The ovule is anatropous,
unitegmic and tenuinucellar, and the integument
is 16–20 cells thick. The embryo sac is of the Polygonum type and is surrounded by an endothelium
except at the micropylar and chalazal ends. The
embryo sac invades the chalazal tissue with a multicellular extension which does not seem to have
the characteristics of a haustorium (Rosén 1937,
1946; Carolin, unpubl. data). In Brunonia, a suspensor haustorium has been claimed to exist by
Rosén (1946) but this has been doubted by Tobe and
Morin (1996). No haustorial endosperm is formed
at either the micropylar or the chalazal end. The
embryo develops according to the Solanad type
(Rosén 1937, 1946). In Goodenia, Velleia, Selliera,
Scaevola, Diaspasis and Coopernookia, the embryo
is spathulate in shape, the cotyledons being wider
than the radicle, but in Dampiera, Lechenaultia and
Anthotium the embryo is terete (Carolin, unpubl.
data). Endosperm formation is ab initio cellular or
nuclear, and in Brunonia the endosperm is eventually confined to a band of crushed cells surrounding
the embryo which has swollen cotyledons (Carolin,
unpubl. data). Anther wall development is dicotyledonous, the endothecium is fibrous, and the tapetum glandular. Mature pollen grains are 2-celled.
Fruit and Seed. The fruits are mostly capsular but in a number of species, and throughout
Scaevola, Dampiera and Diapasis, they are indehiscent. There is a fleshy mesocarp surrounding
a hard endocarp in Scaevola sect. Scaevola whilst
in Scaevola sects. Xerocarpa, Dampera, Diaspasis,
and several species of Goodenia, the mesocarp is
thin and more or less dry. In the indehiscent fruit
of Goodenia neogoodenia and Pentaptilon, the endocarp is not thickened to any degree. In Coopernookia, the seed is obloid and carunculate and,
in Scaevola, it is similar in shape but with a thin
testa, being contained within the endocarp, and no
caruncle is present. In Goodenia and related genera,
the seed is variously flattened and has a wing or rim
around the edge (Carolin 1966). In Lechenaultia,
591
each seed is surrounded by a hardened inner portion of the pericarp and each of these single-seeded
articles is dispersed separately (Morrison 1986).
Pollen Morphology. Pollen (Fig. 124) in the
family is typically tricolporate (Duigan 1961;
Gustafsson et al. 1997). Duigan (1961) distinguishes three types of pollen grain on the basis
of surface pattern and the border of the pore. In
Goodenia and Brunonia, the surface pattern is
relatively coarse but in Dampiera it is very fine.
Goodenia has a sexine which tapers gradually into
the colpi whilst Brunonia has a sexine which forms
two distinct ridges on either side of the colpi.
Pollen grains in Lechenaultia, however, are porate
and held together in one plane in rhomboidal
tetrads. There are varying numbers of aborted
grains in the tetrads, probably associated with
Fig. 124. Goodeniaceae. Pollen surface and exine structure.
A, B Dampiera. C, D Velleia. E, F Lechenaultia. (SEM photographs by M. Gustafsson)
592
R.C. Carolin
univalent formation at meiosis (Peacock 1959).
Essentially, the three groups recognized by Duigan
(1961) were also identified by Gustafsson et al.
(1997), i.e. Lechenaultia with tetrads, AnthotiumDampiera with tricolporate, rugulate grains with
a thin infratectum and rounded ora, and the
remaining genera (Scaevola-Goodenia group) with
tricolporate, microspinulose grains with distally
branched columellae (Fig. 124D) and lalongate
ora. Within the latter group, Brunonia has ridges
and depressions along the colpi.
Karyology. There are three distinct basic chromosome numbers in Goodeniaceae. Coopernookia
alone has a base number of x = 7. The other
genera can be divided into two groups: Brunonia,
Dampiera, Lechenaultia and Anthotium with a base
number of x = 9, and the remaining genera with
a base number of x = 8. There are polyploid species
within these groups, and in some species polyploid races (Peacock 1963; Peacock and SmithWhite 1978).
Dispersal. Some species are clearly animaldispersed because they have fleshy mesocarps,
particularly the drupes of many Scaevola species.
There is evidence, for instance, of emus eating
the fruits of S. tomentosa and S. spinencens, the
latter having the vernacular name of Emu Bush
in parts of Western Australia. The littoral species
S. taccada and S. calendulacea also have a fleshy
mesocarp and may be dispersed by birds but, in
S. taccada, part of the endocarp is ± corky and
it may also float some distance in sea currents.
Selliera radicans has a ± fleshy fruit containing
seeds with a very mucilaginous border, which
stick very effectively to animals (pers. obs.) and
may be dispersed this way. Dispersal by seabirds
may account for the trans-Pacific distribution of
this species, which occurs in littoral salt-marshes
as well as highland swamps. Many species of
Goodenia have seeds with a mucilaginous border
or wing, although it is doubtful if this structure
attaches the seed to animals as effectively as that
of Selliera. However, ants appear to be attracted
to this border or wing in some cases, and here
it might function as an elaiosome, e.g. the desert
species G. cycloptera (pers. obs.; A.J. Beatty, pers.
comm.). In some other Goodenia spp., the wing
on the seed is scarcely mucilaginous and seems to
function as a wind-dispersal agent, the seeds being
blown for considerable distance across the bare
soil of communities in the arid areas of Australia,
e.g. G. vilmorinae, G. triodiophila (pers. obs.).
In Brunonia, the single-seeded fruit is retained
within the hardened hairy calyx tube which has
persistent hairy lobes at the top, and this whole
structure is dispersed by the wind. Many species
of the family do not appear to have well-developed
dispersal mechanisms, although little work has
been done on most of them.
Reproductive Biology. Pollen is presented to
the vector in or around the indusium (Fig. 123).
It is deposited on the vector, usually an insect,
as it probes to the bottom of the flower where
the nectar is contained either in a single spur or
pocket, as in Goodenia, Velleia, etc. (Carolin 1959),
in several pockets as in Dampiera, or on the surface of the ovary (Carolin 1959). The hairs on and
around the indusium are agitated by the vector,
thus releasing the pollen onto its body (pers. obs.).
There are a number of structures on the corolla
which act as tactile guides for the vector. In Coopernookia these are large barbulae, which are often
hairy in Scaevola; in Goodenia they are mostly
surface hairs or large papillae, and in Goodenia
sect. Borealis they are ridges; in Dampiera, they are
sinuous ridges (Carolin et al. 1992). The vectors
observed so far are mostly native bees and Lepidoptera. Lechenaultia sect. Lechenaultia, Scaevola
coccinea and S. glabra show a bird-pollination syndrome, with well-developed corolla tubes and red,
orange or rarely bright yellow or green flowers. The
stigmatic initials later grow out of the indusium
(expect in Lechenaultia) but it appears that, in at
least some species, the pollen in the indusium is
by that time non-viable (Brough 1927). All species
examined are self-incompatible.
Phytochemistry. Inulin has been reported as
a storage carbohydrate in the family. Seco-loganin,
a simple seco-iridoid, has also been recorded. Alkaloids are also known and may be responsible for
the use of members of the family as medicines by
Australian natives.
Subdivision of and Relationships Within
the Family. Relationships within the family
have been investigated by Carolin (1959, 1978),
Gustafsson et al. (1996) and Hansen (1997). In
their analysis of rbcL sequence variation of all
genera except Pentaptilon, albeit including only
few species of the large genera, and considering
morphological variation, Gustafsson et al. (1996)
demonstrate the existence of four lineages. These
Goodeniaceae
are (1) Lechenaultia, (2) Anthotium and Dampiera,
(3) Brunonia and (4) the remaining genera
(Scaevola-Goodenia group). Brunonia, often
treated as a separate family, is well-nested within
Goodeniaceae. In the Scaevola-Goodenia group,
Diaspasis and Scaevola are successive sister genera
to a well-supported clade of Goodenia, Selliera,
Coopernookia, Velleia and Verreauxia, of which
Goodenia may not be monophyletic. The four
subgroups of the family can also be characterized
in terms of pollen morphology (Gustafsson et al.
1997). The analysis by Gustafsson et al. (1996)
confirms several but not all results of Carolin’s
(1978) earlier analysis, and some similarity can be
seen between the morphology-based conclusions
of Hansen (1997) and those of Gustafsson et al.
(1996). Howarth et al. (2003) found that Diaspasis
is deeply nested within Scaevola, while Scaevola
collaris should be excluded from this genus because
it is most closely related to species of Goodenia.
Their results also indicated that Verreauxia and
Velleia may be nested within Goodenia.
Affinities. Goodeniaceae have usually been
placed in the Campanulales with Campanulaceae,
Lobeliaceae, Compositae, Calyceraceae and Stylidiaceae. Carolin (1978) and Lammers (1992)
have examined these relationships and suggest
that this position be retained, after the removal
of Stylidiaceae. Lammers (1992) also suggests
that Menyanthaceae be included here. Indeed,
on the basis of phytochemical data, he considers
that Goodeniaceae, Compositae, Calyceraceae and
Menyanthaceae form a well-defined clade, with
Campanulaceae, Lobeliaceae (and Cyphiaceae) as
the sister-group. Both molecular and morphological evidence (Morgan and Soltis 1993; Cosner et al.
1994; Gustafsson and Bremer 1995; Olmstead et al.
2000; Soltis et al. 2000; Bremer et al. 2001; Lundberg and Bremer 2003) confirms the existence of
a well-supported Menyanthaceae, Goodeniaceae,
Calyceraceae, Compositae clade, in which Goodeniaceae are sister to Calyceraceae/Compositae,
and Menyanthaceae sister to these three families.
This clade is part of the Asterales (APG II 2003).
Distribution and Habitats. The family is almost entirely southern hemisphere in distribution
and most species are confined to Australia. They
occur in a wide variety of habitats but mostly in
more open communities, particularly in the arid
and semi-arid parts of the continent, but also
with significant numbers in the sclerophyll forests
593
and heaths of the damper climate zones. Only
a few species, such as Scaevola enantophylla and S.
glabra, occur in rain forests. Scaevola is the most
widespread genus outside Australia. Two strand
species occur almost throughout the tropics:
Scaevola taccada in the Indo-Pacific region, and
S. plumieri in America, Africa and India. Restricted
endemics are Scaevola hainanensis in southern
China, S. socotrensis on Socotra and S. wrightii on
Cuba. Many of the islands of the South Pacific have
an endemic, inland, montane Scaevola species.
New Caledonia and the Hawaiian islands each
have several species, in both cases the result of
three independent colonisations, one of which
gave rise to a small evolutionary radiation in each
of the archipelagos (Howarth et al. 2003). The
largest genus, Goodenia, also occurs northwards
from Australia, with G. purpurascens, G. pumilia
and G. armstrongiana in New Guinea, G. koningsbergeri in Java, and G. pilosa in Indonesia and
south-eastern Asia. Selliera radicans occurs in
Australia, New Zealand and Chile. Lechenaultia
filiformis is found in New Guinea and the Moluccas,
and Velleia spathulata occurs in New Guinea.
Key to the Genera
1.
–
2.
–
3.
–
4.
–
5.
–
6.
–
7.
–
8.
–
9.
–
10.
Anthers connate
2
Anthers free
6
Ovary superior
1. Brunonia
Ovary inferior
3
Ovules and seeds more than 2; fruit dehiscent or fragmenting, rarely indehiscent and then the fruit beaked
4
Ovules and seeds 1 or 2; fruit indehiscent, not beaked
5
Leaves all cauline; indusium 2-lipped
3. Lechenaultia
Leaves cauline and basal; indusium not 2-lipped
2. Antotium
Corolla without auricles; hairs simple
7. Diaspasis
Corolla with auricles; hairs branched, rarely absent
4. Dampiera
Ovules (and usually seeds) more than 2 per loculus 7
Ovules and seeds 1 or 2 per loculus
11
Seeds ovoid, obloid or irregularly compressed, without
a wing, rim or peripheral groove
8
Seeds flattened or concave-compressed, with a wing,
rim or peripheral groove
9
Corolla yellow
10. Velleia
Corolla blue to pinkish or white
5. Coopernookia
Corolla lobes without wings; fruit ± fleshy, indehiscent
9. Selliera
Corolla lobes winged; fruit a dry, dehiscent capsule 10
Flowers in thyrses, racemes, spikes or subumbels;
ovary inferior or half-inferior (or superior to the
sepals but the corolla fused to the top of the ovary in
Goodenia macroplectra)
8. Goodenia
594
R.C. Carolin
– Flowers in axillary dichasia often in the axils of basal
leaves; ovary free from the corolla for most of its length
10. Velleia
11. Fruit a capsule
5. Coopernookia
– Fruit indehiscent, nut-like or soft-shelled with several
seeds or a 1–2-seeded drupe
12
12. Ovary and fruit winged
11. Pentaptilon
– Ovary and fruit not winged
13
13. Style with stiff purplish hairs; corolla white, usually
with violet spots
8. Goodenia
– Style with whitish hairs or glabrous (purplish bristles sometimes present on back of indusium); corolla
without violet spots
14
14. Corolla blue to white
6. Scaevola
– Corolla yellow or brownish
15
15. Shrubs with alternate leaves or scramblers with opposite leaves
6. Scaevola
– Herbs with alternate or basal leaves
16
16. Plants hairy with unbranched cottony hairs or glabrous
8. Goodenia
– Plants hairy with branched and simple hairs
12. Verreauxia
Genera of Goodeniaceae
1. Brunonia Sm. ex R. Br.
Fig. 125
Brunonia Sm. ex R. Br., Prodr.: 589 (1810).
Herbs or subshrubs. Leaves basal or alternate
towards the base of the stem, ± villous. Flowers
in heads, each flower with several hyaline bracts;
heads surrounded by green involucral bracts.
Sepals connate; tubes swollen towards the top
when mature; lobes with long cilia. Petals 5, blue,
connate; tube entire; lobes spreading, not winged,
stamens epipetalous; anthers connate. Ovary superior, 1-locular; indusium scarcely compressed,
erect, with scattered hairs and almost glabrous
lips. Fruit a nut enclosed in the persistent calyx.
Seeds ellipsoid, without conspicuous endosperm.
2n = 18, 36, 72. One species, B. australis Sm. ex
R. Br., Australia south of the 17th parallel.
2. Anthotium R. Br.
Anthotium R. Br., Prodr.: 582 (1810).
Perennial glabrous herbs with 1 to many stems
and a woody rootstock. Leaves mostly basal. Flowers sessile, arranged in compact monochasia condensed into heads; bracts and bracteoles leaf-like.
Corolla 2-lipped, red to light blue or cream; tube slit
nearly to the base adaxially; lobes winged; adaxial
lobes with auricles on adjacent margins. Anthers
coherent into a tube. Ovary inferior, 2-locular, with
numerous ovules; indusium ± globular, glabrous
or hairy; stigmatic surface within the indusium.
Fig. 125. Goodeniaceae. Brunonia australis. A Habit. B Leaf
tip. C Flower, with bracts. D Flower, calyx removed. E Flower,
longitudinal section. F Fruit with persistent sepals. (Illustrations by David Mackay in Carolin 1992; reproduced with
permission from Flora of Australia)
Fruit a capsule with 4 valves. Three species, southwestern Australia.
3. Lechenaultia R. Br.
Lechenaultia R. Br., Prodr.: 581 (1810).
Latouria (Endl.) Lindley (1847).
Ericopsis C. Gardner (1923).
Perennial hairy or glabrous herbs or subshrubs
with woody rootstock, often with suckers. Leaves
sessile, linear or sometimes lanceolate to ovate.
Flowers sessile, arranged in cymes; bracts and
bracteoles usually leaf-like. Corolla 2-lipped, usually glabrous outside, variously hairy inside, red
to blue, orange, white or yellow; tube slit adaxially
or not; lobes winged, sometimes only slightly so.
Anthers cohering into a tube. Ovary inferior, 2locular, with numerous ovules; indusium 2-lipped,
usually hairy; stigmatic surface on the upper side
of the indusium. Fruit capsule-like, with 4 narrow
valves. Each seed enclosed in an article which is
Goodeniaceae
part of the hard endocarp. 2n = 18, 36. Twenty-six
to 27 species, south-western, central and northern
Australia, one species extending to New Guinea.
Three sections: sect. Lechenaultia: corolla tube
not slit; sect. Patentes D. Morrison: corolla tube
slit adaxially, adaxial corolla lobes winged; sect.
Latouria (Endl.) Benth.: corolla tube slit adaxially,
adaxial corolla lobes usually not winged.
4. Dampiera R. Br.
Dampiera R. Br., Prodr.: 587 (1810); Rajput & Carolin,
Telopea 3: 183–216 (1988), gen. class.
Subshrubs to many-stemmed perennial woody
herbs or rosette herbs, sometimes with suckers,
glabrous or with branched dendritic hairs. Flowers
in racemes or thyrses which are sometimes condensed into heads, or in irregular cymo-panicles
or apparently in clusters in the leaf axils. Sepals
usually small or sometimes obsolete. Corolla
2-lipped, blue or yellow, rarely white or pink,
usually hairy outside, often with ridges and calli
inside; tube slit to the base adaxially and often
laterally; lobes winged; adaxial lobes auriculate
on adjacent margins. Anthers connate. Ovary
inferior, 2–1-locular; indusium ± globular, usually
with a glabrous orifice; stigmatic surface within
the indusium. Ovules 1–2, sometimes bent or
horseshoe-shaped. Fruit an inferior nut. 2n = 18,
36, 54. About 66 species, Australia.Two sections:
sect. Linschotenia (Vriese) Benth.: flowers in
racemes, thyrses or heads; sect. Dampiera: flowers
in cymo-panicles or apparently clustered in leaf
axils.
5. Coopernookia Carolin
Coopernookia Carolin, Proc. Linn. Soc. New South Wales
92: 209 (1968).
Subshrubs with stellate and glandular hairs, often
viscid and becoming varnished. Flowers in leafy
thyrses or racemes, with small bracteoles; pedicels
articulate. Corolla scarcely 2-lipped, with long,
stiff, retrorse bristles inside, white to mauve or
deep pink; tube slit to the base adaxially; lobes
broadly winged, without auricles. Anthers free.
Ovary inferior, incompletely 2-locular; indusium
± flattened, with long bristles on the lips. Ovules
2–8. Fruit a capsule with 2 valves. Seeds ovoid
to obloid, glossy, carunculate, without a wing.
2n = 14. Six species, South Australia.
595
6. Scaevola L.
Scaevola L., Mant. Pl. 2: 145 (1771); Carolin, Telopea 3: 477–
515 (1990), gen. class.; Howarth et al., Amer. J. Bot. 90:
915–923 (2003), phylog.
Nigromnia Carolin (1974).
Perennial herbs, shrubs, scramblers or small trees.
Leaves alternate or rarely opposite. Flowers in terminal or axillary thyrses, racemes, spikes or apparently cymes, bracteolate. Corolla blue, white,
mauve or rarely yellow, slit on the adaxial side to the
base and fan-shaped, rarely tubular; lobes usually
winged. Anthers free. Ovary inferior, 1–4-locular,
sometimes with 2 fertile loculi and 2 sterile cavities;
ovules 1–4, indusium flattened, often pubescent
and with stiff hairs on the lips. Fruit indehiscent;
mesocarp fleshy, corky or ± dry; endocarp hard.
Seeds ovoid, without wing or caruncle. 2n = 16, 32.
Circa 130 species, 70 endemic in Australia, the others throughout the Indo-Pacific region and more
rarely in other tropical areas.
Three sections: sect. Scaevola: leaves alternate,
fruit corky or fleshy, inflorescence continuing
growth after flowering, Australia and Indo-Pacific;
sect. Enantiophyllum Miq.: leaves opposite, fruit
fleshy, inflorescence continuing growth after
flowering, Australia; sect. Xerocarpa G. Don.:
leaves alternate, fruit usually dry, inflorescence not
continuing growth after flowering.
7. Diaspasis R. Br.
Diaspasis R. Br., Prodr.: 586 (1810).
Perennial with a tough rootstock, sprinkled with
papillate hairs. Leaves sessile, linear. Flowers in terminal racemes, bracteolate. Sepals often unequal.
Corolla slightly 2-lipped, white to pink or lavender
or rarely yellowish; tube slightly slit adaxially; lobes
winged. Anthers connate. Ovary inferior, 2-locular;
indusium compressed laterally; ovules solitary in
each loculus. Fruit dry, indehiscent; endocarp hard.
2n = 16. One species, south-western Australia.
8. Goodenia Sm.
Fig. 126
Goodenia Sm., Sec. Bot. New Holl.: 15 (1793); Carolin et al.,
Fl. Austral. (1992), rev.; Sage, Nuytsia 13: 367–377 (2000),
key Australia.
Neogoodenia C. Gardner & A.S. George (1963).
Herbs or shrubs. Flowers in thyrses, racemes spikes
or subumbels, bracteolate or ebracteolate. Sepals
variously adnate to the ovary or free. Corolla adnate to the ovary, yellow, cream, pink, blue, pur-
596
R.C. Carolin
subg. Goodenia: flowers yellow, cream, blue, purplish or brownish red.
9. Selliera Cav.
Selliera Cav., Anales Hist. Nat. 1: 41 (1799).
Glabrous creeping perennial, rooting at the nodes.
Flowers in condensed axillary racemes or solitary
in the leaf axils. Corolla whitish, tinged with red,
not 2-lipped, without a pouch; tube slit to the base
adaxially but not opening out, lobes erect, wingless
or almost so. Anthers free. Ovary inferior, 2-locular;
indusium compressed, with stiff bristles on the lips;
ovules numerous. Fruit indehiscent or eventually
splitting, slightly fleshy. Seeds compressed, with
a mucilaginous wing. 2n = 16. One species, S. radicans Cav., Australia, New Zealand and Chile.
10. Velleia Sm.
Velleia Sm., Trans. Linn. Soc. London, Bot. 4: 217 (1798).
Fig. 126. Goodeniaceae. Goodenia macroplecta. A Habit.
B Flower. (Illustrations by David Mackay in Carolin et al.
1992; reproduced with permission from Flora of Australia)
plish on brownish red, usually pouched or sometimes spurred, usually bilabiate; tube slit to the base
adaxially; lobes winged; adaxial lobes sometimes
auriculate. Anthers free. Ovary usually inferior (the
sepals may be free from the ovary, although the
corolla is always at least partially adnate and the stamens are always epigynous), usually incompletely
2-locular; style simple or 2–4-fid; indusium usually with stiff bristles on the lips; ovules solitary to
numerous. Fruit usually a capsule, rarely a small 1seeded nut or a large, hard, 4-seeded drupe. Seeds
compressed, usually winged. 2n = 16, 32, 48, 64.
About 180 species, 178 in Australia, three extending
to New Guinea and southern China; one endemic
in Java.
Two subgenera: subg. Monochila (G. Don) Carolin: corolla white, usually with purplish spots near
the base of the lobes, or pale pink, rarely yellow;
Herbs or slightly shrubby plants, usually with basal
leaves. Flowers in branching cymes arising from the
axils of the leaves. Sepals sometimes reduced to 3,
usually adnate to the base of the ovary only. Corolla
usually free from the ovary except at the base, 2lipped, yellow to orange or pink to mauve or white;
tube slit to the base adaxially; lobes winged; adaxial
lobes auriculate. Anthers free. Ovary superior, incompletely 2-locular; indusium compressed, with
stiff bristles on the lips; ovules several. Fruit a capsule with 2–4 valves. Seeds compressed, winged or
with a thickened rim. 2n = 16. Twenty-one species,
all in Australia, with one extending to New Guinea.
Three sections: sect. Euthales (R. Br.) Carolin:
sepals 5, connate into a tube, south-western
Australia; sect. Menoceras R. Br.: sepals 5, free or
slightly connate at the base, southern Australia;
sect. Vellelia: sepals 3, eastern Australia.
11. Pentaptilon E. Pritz.
Pentaptilon E. Pritz., Bot. Jahrb. Syst. 35: 564 (1905).
Erect herb with a basal stock. Leaves mostly basal,
tomentose with branched hairs. Flowers in a terminal thyrse, bracteolate. Corolla 2-lipped, yellow,
throat brownish red and with dense tuft of whitish
hairs; tube slit to the base adaxially; lobes winged.
Stamens free. Ovary inferior, winged, 2-locular; indusium compressed, with stiff white bristles on the
lips; ovules several; fruit soft, indehiscent, with 3
large and 2 smaller ± bladdery wings alternating
with the sepals. Seeds compressed, with a narrow
Goodeniaceae
wing. One species, P. careyi E. Pritz., Western Australia.
12. Verreauxia Benth.
Verreauxia Benth., Fl. Austral. 4: 105 (1868).
Herbs or shrubs. Leaves basal or alternate, tomentose to villous with branched hairs. Flowers in loose
or spike-like terminal thyrses. Corolla 2-lipped, yellow, brownish in the throat; tube slit to the base
adaxially; lobes winged. Stamens free. Ovary inferior, 1-locular, with reddish or golden multicellular hairs between three of the sepaline ribs; indusium ± grooved, with short white bristles on the
lips; ovule solitary. Fruit a compressed nut. Seeds
flat, with an indistinct rim. Three species, southwestern Australia.
Selected Bibliography
APG (Angiosperm Phylogeny Group) II. 2003. An update
of the Angiosperm Phylogeny Group classification for
the orders and families of flowering plants: APG II. Bot.
J. Linn. Soc. 141: 399–436.
Bremer, K., Backlund, A., Sennblad, B., Swenson, U.,
Andreasen, K., Hjertson, M., Lundberg, J., Backlund, M., Bremer, B. 2001. A phylogenetic analysis
of 100+ genera and 50+ families of euasterids based
on morphological and molecular data with notes on
possible higher level morphological synapomorphies.
Pl. Syst. Evol. 229: 137–169.
Brough, P. 1927. Studies in the Goodeniaceae. I. The life
history of Dampiera stricta R. Br. Proc. Linn. Soc. New
South Wales 52: 471–498.
Carlquist, S. 1969. Wood anatomy of Goodeniaceae and
the problem of insular woodiness. Ann. Missouri Bot.
Gard. 56: 358–390.
Carolin, R.C. 1959. Floral structure and anatomy in the
family Goodeniaceae Dumort. Proc. Linn. Soc. New
South Wales 84: 252–255.
Carolin, R.C. 1960. The structures involved in the presentation of pollen to visiting insects in the order Campanulales. Proc. Linn. Soc. New South Wales 85: 197–205.
Carolin, R.C. 1966. Seeds and fruit of the Goodeniaceae.
Proc. Linn. Soc. New South Wales 91: 58–83.
Carolin, R.C. 1967. The concept of the inflorescence in the
order Campanulales. Proc. Linn. Soc. New South Wales
92: 7–26.
Carolin, R.C. 1970. The trichomes of the Goodeniaceae.
Proc. Linn. Soc. New South Wales 96: 8–22.
Carolin, R.C. 1978. The systematic relationship of Brunonia.
Brunonia 1: 9–29.
Carolin, R.C. 1980. Pattern of the seed surface of Goodenia
and related genera. Austral. J. Bot. 28: 123–137.
Carolin, R.C. 1990. Nomenclatural notes, new taxa and systematic arrangement in the genus Scaevola and its synonyms. Telopea 3: 477–515.
597
Carolin, R.C. 1992. Brunoniaceae. In: Gorge, A. (ed.) Flora
of Australia, vol. 35. Canberra: Australian Government
Publishing Service, pp. 1–3.
Carolin, R.C., Rajput, M.T.M., Morrison, D. 1992. Goodeniaceae. In: George, A. (ed.) Flora of Australia, vol. 35.
Canberra: Australian Government Publishing Service,
pp. 4–351.
Cosner, M.E., Jansen, R.K., Lammers, T.G. 1994. Phylogenetic relationships in the Campanulales based on rbcL
sequences. Pl. Syst. Evol. 190: 79–95.
Duigan, S.L. 1961. Studies of the pollen grains of plants native to Victoria, Australia. I. Goodeniaceae (including
Brunonia). Proc. Roy. Soc. Victoria 74: 87–109.
Gustafsson, M.H.G. 1995. Petal venation in the Asterales
and related orders. Bot. J. Linn. Soc. 118: 1–18.
Gustafsson, M.H.G., Bremer, K. 1995. Morphology and phylogenetic interrelationships of the Asteraceae, Calyceraceae, Campanulaceae, Goodeniaceae, and related
families (Asterales). Amer. J. Bot. 82: 250–265.
Gustafsson, M.H.G., Backlund, A., Bremer, B. 1996. Phylogeny of the Asterales sensu lato based on rbcL sequences with particular reference to the Goodeniaceae.
Pl. Syst. Evol. 199: 217–242.
Gustafsson, M.H.G., Grafström, E., Nilsson, S. 1997. Pollen
morphology of the Goodeniaceae and comparisons
with related families. Grana 36: 185–207.
Hansen, H.V. 1997. Studies in the Goodeniaceae and the
Brunoniaceae with a discussion of their relationships
to Asteraceae and Calyceraceae. Nordic J. Bot. 17: 495–
510.
Howarth, D.G., Gustafsson, M.H.G., Baum, D.A., Motley, T.J.
2003. Phylogenetics of the genus Scaevola (Goodeniaceae): implication for dispersal patterns across the
Pacific Basin and colonization of the Hawaiian Islands.
Amer. J. Bot. 90: 915–923.
Krause, K. 1912. Goodeniaceae und Brunoniaceae.
Pflanzenreich IV, 277, 277a. Leipzig: Engelmann.
Lammers, T.G. 1992. Circumscription and phylogeny of the
Campanulales. Ann. Missouri Bot. Gard. 79: 388–413.
Leins, P., Erbar, C. 1989. Zur Blütenentwicklung und
sekundären Pollenpräsentation bei Selliera radicans
Cav. (Goodeniaceae). Flora 182: 43–56.
Lundberg, J., Bremer, K. 2003. A phylogenetic study of the
order Asterales using one morphological and three
molecular data sets. Intl J. Pl. Sci. 164: 553–578.
Morgan, D.R., Soltis, D.E. 1993. Phylogenetic relationships
among members of Saxifragaceae sensu lato based on
rbcL sequence data. Ann. Missouri Bot. Gard. 80: 631–
660.
Morrison, D.A. 1986. Notes on the fruits of Lechenaultia
(Goodeniaceae) with a new species from northern Australia. Telopea 3: 159–166.
Olmstead, R.J., Kim, K.-J., Jansen, R.K., Wagstaff, S.J. 2000.
The phylogeny of the Asteridae sensu lato based
chloroplast ndhF gene sequences. Mol. Phylog. Evol.
16: 96–112.
Peacock, W.J. 1959. Pollen tetrads in Leschenaultia (sic!).
Proc. Linn. Soc. New South Wales 84: 271–277.
Peacock, W.J. 1963. Chromosome numbers and cytoevolution in the Goodeniaceae. Proc. Linn. Soc. New South
Wales 88: 8–27.
Peacock, W.J., Smith-White, S. 1978. Cytogeography of
Brunonia australis Sm. ex R. Br. Brunonia 1: 31–43.
598
R.C. Carolin
Rajput, M.T.M., Carolin, R.C. 1984. Stem structure and vascularisation in Dampiera (Goodeniaceae). Proc. Linn.
Soc. New South Wales 107: 479–485.
Rajput, M.T.M., Carolin, R.C. 1988. The genus Dampiera
(Goodeniaceae): systematic arrangement, nomenclatural notes and new taxa. Telopea 3: 183–216.
Rosén, W. 1937. Beiträge zur Kenntnis der Embryologie der
Goodeniaceen. Acta Hort. Göteb. 12: 1–10.
Rosén, W. 1946. Further notes on the embryology of the
Goodeniaceae. Acta Hort. Göteb. 16: 235–249.
Soltis, D.E., Soltis, P.S., Chase, M.W., Mort, M.E., Albach, D.C., Zanis, M., Savolainen, V., Hahn, W.H.,
Hoot, S.B., Fay, M.F., Axtell, M., Swensen, S.M.,
Prince, L.M., Kress, W.J., Nixon, K.C., Farris, J.S. 2000.
Angiosperm phylogeny inferred from 18S rDNA, rbcL,
and atpB sequences. Bot. J. Linn. Soc. 133: 381–461.
Tobe, H., Morin, N.R. 1996. Embryology and circumscription of Campanulaceae and Campanulales: a review of
literature. J. Pl. Res. 109: 425–435.
Menyanthaceae
Menyanthaceae Bercht. & J. Presl, Prir. Rostlin 1, 115: 1 (1823), nom. cons.
G. Kadereit
Glabrous perennial or rarely annual herbs with
tufted rootstocks or horizontal creeping rhizomes.
Leaves monomorphic or dimorphic, alternate,
often forming a rosette, simple, linear to cordate
or reniform, rarely 3-foliolate; margin entire, undulate, regularly or irregularly dentate or crenate;
petioles mostly long, sometimes sheathing, stipules
absent. Inflorescence a simple or branched cyme,
raceme, panicle, umbel, head or cluster, or flowers
in pairs or solitary. Flowers actinomorphic, (4)5merous, either hermaphrodite and monomorphic
or heterostylous, or in some species monoecious,
dioecious, or gynodioecious. Sepals persistent, at
least basally united. Corolla sympetalous, often
deeply lobed, valvate or imbricate in bud, inner
surface of petal lobes often fimbricate or crested,
margins often fringed; petals white, pinkish or yellow, caducous. Stamens (4)5, alternating with the
corolla lobes, filaments inserted near base of the
corolla tube; anthers dorsifixed, bilocular, tetrasporangiate, mostly sagittate, opening by longitudinal
slits; sometimes flowers with fringed scales alternating with the stamens. Ovary superior to semiinferior, of two united carpels, unilocular; style of
varying length or absent in one morph, 2-lobed,
stigma papillate, lacerate or with secondary lobes.
Separate nectaries or disc present at base of ovary.
Ovules numerous, on two parietal, often intruding
placentae, anatropous, unitegmic, tenuinucellate.
Fruit a regularly or irregularly dehiscent or
indehiscent capsule, or rarely a berry. Seeds few to
many, sometimes winged, endosperm abundant.
An almost cosmopolitan family with five genera and c. 60 species.
Vegetative Morphology. All Menyanthaceae
are herbs adapted to wet or aquatic habitats. They
have either creeping rhizomes or tufted rootstocks
bearing leaves, stolons, adventitious roots and one
to several fertile culms. The adventitious roots
are thick and spur-like, or slender and more or
less straight. Leaves are monomorphic or, in many
species of Nymphoides, dimorphic with small
leaves arising from the tufted rootstocks and large
floating leaves on the fertile culms. Petioles are
erect or flexuose. In aquatic species, the leaf blades
are raised above the water surface by erect, long
petioles, or the petioles are flexuose and the leaf
blades float on the water surface. The blades are
mostly elliptic, ovate, rotund, cordate or reniform.
Menyanthes trifoliata has trifoliolate leaves and
Liparophyllum gunnii has linear leaves. Leaf blades
are thin, thick, or slightly succulent. The margin is
mostly entire, sometimes it is minutely (irregularly
or regularly) dentate or crenate. Nephrophyllidium
crista-galli has deeply crenate, reniform leaf
blades. The petioles of some species are winged,
and several species have membranous sheaths
towards the base of the petioles.
Vegetative Anatomy. Leaves of Menyanthaceae
are dorsiventral or isobilateral. On non-floating
leaves, stomata of the Ranunculus type are usually
present on both surfaces but they are confined to
the upper surface of floating leaves. Hydathodes
are often located at enlarged ends of veins at leaf
margins. The mesophyll of the leaves and the cortex of the petioles and flower-bearing culms show
characteristic systems of intercellular canals and
spaces. Some species have branched, sclerenchymatous idioblasts (astrosclereids) which project
into these air cavaties. The vascular bundles in the
petiole are normally widely spaced and are either
arranged in a circle, are scattered, or form one or
two arcs with the median vascular bundle being
considerably larger than the peripheral ones. The
vascular bundles of the fertile culms and rhizomes
are also widely spaced and isolated. Intraxylary
phloem is absent in Menyanthaceae (Metcalfe and
Chalk 1950, 1979).
Inflorescence and Flower Morphology.
The fertile culms are leafless, uniphyllous or
have several leaves. Many species have simple or
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G. Kadereit
branched cymes. The flowers of most species of
Nymphoides are aggregated in umbels, heads or
clusters. Nephrophyllidium and several species of
Villarsia have paniculate inflorescences. Menyanthes has racemes, and Liparophyllum has solitary
flowers in the leaf axils. Bracts are present but
small and inconspicuous whereas the corollas
are large and attractive. Their display is often
enhanced by a large array of outgrowths, e.g.
membranous extensions, fringes of emergences,
or ruffles on the margin or on the inner surface. In
Menyanthes trifoliata, the emergences are located
on the upper part of the adaxial side of the petals,
and fill the space between petals and anthers.
In Nymphoides peltata, the margin of the petals
is extended by undulated stretches of thinner,
translucent tissue (Erbar 1997). Armstrong (2002)
found that the fringe of trichomes in some species
of Nymphoides significantly increases the upward
force of the flower when pulled under water or
opened under water (e.g. Nymphoides geminata),
and also increases the buoyancy of the flower
when supporting the weight of a pollinator. In
several species, emergences are also present above
the nectary glands where they might protect
the secretory tissue. In some dioecious species,
clusters of yellow trichomes alternate with the
non-polliniferous stamens of the female flowers
(see below).
Embryology. Menyanthaceae have a secretory
(glandular) anther tapetum, and pollen grains have
three nuclei when shed. The ovary contains many
anatropous, tenuinucellate ovules on parietal placentae. The development of the megaspore follows
the Polygonum type. The innermost layer of the
integument develops into a tapetum. The polar nuclei fuse early to form a central nucleus which is
located in the centre of the embryosac. There are
always three antipodes which degenerate early, often before the embryosac is fully developed. The
endosperm is cellular, and there is no endosperm
haustorium at either the micropylar or the chalazal end. Endosperm in mature seeds is copious,
and embryogenesis follows the Asterid type. The
testa is one cell thick and formed by the exotesta
whereas the mesotesta and endotesta are crushed
(Stolt 1921; Erbar 1997; Tobe and Morin 1996 and
ref. therein).
Pollen Morphology. Nilsson (1973) described
two pollen types for the family, these being the
Menyanthes type and the Villarsia type. Nephro-
phyllidium and Menyanthes have the Menyanthes
type, which is subprolate to prolate, tricolpate with
each furrow with a single germ pore, and a striate
to rugulose exine. Liparophyllum, Nymphoides
and Villarsia have the Villarsia pollen type,
which is oblate to suboblate, parasyncolpate with
an isolated apocolpical area, and a spinulose,
striate, rugulose or smooth exine. In heterostylous
species, the number and size of pollen grains differ
between morphs. For example, in Menyanthes
trifoliata the pin anthers produce c. 2 × 103 pollen
grains whereas the thrum anthers produce only
half as many (0. 96 × 103 ). Pin and thrum pollen
grains measure c. 29.0 and c. 33.1 µm in diameter,
respectively. Nic Lughadha and Parnell (1989)
reported that, on average, 26.6% of thrum pollen
was inviable. By contrast, only 5.6% of pollen
grains were inviable in pin anthers.
Karyology. Postulated base numbers are x = 9
(Menyanthes, Nymphoides, Villarsia) and x = 17
(Nephrophyllidium). The chromosome number of
Liparophyllum is unknown. Chromosome counts
for Villarsia (11 species studied) and Nymphoides
(ten species studied) show different ploidy levels,
and diploid, tetraploid and hexaploid species occur
in both genera (Ornduff 1970, 1974; Ornduff and
Chuang 1988). Li et al. (2002) reported a triploid
chromosome number (2n = 27) for Nymphoides
lungtanensis.
Pollination and Reproductive Systems.
Flowers of Menyanthaceae are large and showy
with white, pinkish or yellow corollas. Their
attractiveness is increased by fimbriae or crests on
the inner surfaces of the petals, by fringed petal
margins, and by fringed scales alternating with the
stamens. Some species have sap marks. The flowers
reward the visitors with nectar which is produced
by five nectar glands or a nectar disc at the base
of the ovary. As far as is known, Menyanthaceae
flowers are visited by short-tongued insects,
mainly various Apidae. In all Nymphoides species
studied by Ornduff (1966), maturation of floral
buds is underwater. Before anthesis, the buds are
raised above the water level. The corolla withers
after a few hours of flowering, the pedicels deflect,
and fruit development takes place under water.
Dioecy occurs only in a few species of
Nymphoides. Female flowers have vestigial stamens
which alternate with clusters of staminode-like
trichomes. Male flowers have an ovary which
contains ovules but lacks a style or stigma.
Menyanthaceae
Heterostyly is found in four of the five genera. Menyanthes trifoliata is self-incompatible with
dimorphic heterostyly (Nic Lughadha and Parnell
1989), although at least one homostylous population is known from Greenland. Nephrophyllidium
crista-galli is also distylous. In Nymphoides, distyly
is common but also homostyly, dioecy and gynodioecy occur in several species. Villarsia shows
homostyly and distyly, the latter combined with
self-incompatibility in at least two species (Ornduff 1988), and with strongly dimorphic stigmas
in pin and thrum flowers in most species (Dulberger and Ornduff 2000). Liparophyllum gunnii
has monomorphic flowers and is self-compatible
(Ornduff 1982).
Vegetative reproduction is common by branching and subsequent decay of older parts of the rhizome resulting in independent ramets. Plants can
also regenerate from pieces of rhizome broken off
mature plants and washed ashore. Stoloniferous
species, e.g. many species of Villarsia, reproduce by
rooting stolons which break off the mother plant.
Fruit and Seed. Most species of Menyanthaceae
have dry fruits, only Liparophyllum has a fleshy
berry. The dry fruits are capsules which open regularly or irregularly, or they are indehiscent. The
latter is found in Nymphoides. In this genus, the
infructescence is submerged and the fruits break
off by decay of the pedicels. Seeds are released after
fruit wall decay. Villarsia has four-valved capsules
in which the valves can split secondarily. The seeds
are orbicular, elliptic, ovate or discoid. Seed size
varies in the range 0.4–5.2 mm, and seed colour
from straw yellow, light brown, brown, dark brown
to black. Seed surfaces are smooth, tuberculate or
armed with trichomes of different shape. Outlines
of epidermal cells, if not obscure, are narrowly rectangular, penta- or hexagonal, or interdigitate. The
hilum is terminal or subterminal, and in some
ant-dispersed species surrounded by a caruncle
which is rich in lipids. Size, shape and ornamentation of mature seeds and caruncle characters have
been shown to be useful for distinguishing species
(Sivarajan et al. 1989; Aston 2003) but to be too variable to characterize genera (Chuang and Ornduff
1992).
Seed Dispersal. Most species are adapted to
dispersal by water. The air-filled seed hairs of
Nymphoides and Villarsia and the air-filled testa
cells of Menyanthes as well as their hydrophobic
seed surface result in good buoyancy. For example,
601
the seeds of Menyanthes trifoliata can float for at
least 15 months, remaining viable. The hooked
hairs on the seeds of some species may serve
epizoochory mainly by birds. Endozoochory may
also play a certain role. It has been shown that the
seeds of Menyanthes trifoliata are readily eaten by
reindeer, snow bunting, mallard, various species
of fish, and probably ducks (Ridley 1930; Chuang
and Ornduff 1992). At least four Australian species
of Villarsia are reported to be dispersed by ants
(Berg 1975).
Phytochemistry. The chemical profile of
Menyanthaceae includes iridoids (loganin),
secoiridoids (seco-loganin, sweroside), iridoid
alkaloids, and inulin in roots (Lammers 1992;
Grayer et al. 1999). Bohm et al. (1986) studied the
flavonoid chemistry of 23 species including all
genera of the family. The 38 flavonoid glycosides
observed in this study are derived mainly from
kaempferol and quercetin. The majority of compounds occur as 3-O-glycosides involving glucose,
galactose, arabinose or rhamnose. Nephrophyllidium is unique in having galactosylglucosides of
kaempferol and quercetin, and the corresponding acylated derivatives. Liparophyllum has the
same, simple aglycone profiles as some species of
Nymphoides and Villarsia. Arabonosylglucosides
occur only in Liparophyllum and nine species of
Nymphoides.
Intergeneric Relationships. Lundberg and
Bremer (2003), in a molecular and morphological
analysis of Asterales, found that the genera of
Menyanthaceae (except Liparophyllum, which was
not included) form two well-supported lineages,
one comprising Villarsia and Nymphoides and
the other comprising the two monotypic genera
Nephrophyllidium and Menyanthes. Villarsia and
Nymphoides are also very similar to each other
in morphological and pollen characters. They are
separated by their different capsule morphology
(and dispersal ecology). Nephrophyllidium and
Menyanthes have the same pollen type. Flower,
fruit and leaves of Nephrophyllidium, however,
are similar to certain species of Villarsia. The
flavonoid profile of Nephrophyllidium is unique in
the family. Menyanthes is distinct from all other
genera of the family by its trifoliolate leaves. Leaf
and fruit morphology of Liparophyllum is different
from that of all other genera. However, it is similar
to Villarsia and Nymphoides in pollen morphology
and flavonoid chemistry.
602
G. Kadereit
Affinities. The placement of Menyanthaceae
has long been controversial (Bohm et al. 1986;
Erbar 1997 and ref. therein). Traditionally, the
family has been included in either Gentianales
(e.g. Takhtajan 1987), Solanales (e.g. Cronquist
1981) or Cornales (Dahlgren 1989). Pollard and
Amuti (1981) first recognized a similarity to Asterales/Campanulales on the basis of the presence
of inulin as a storage compound. Lundberg and
Bremer (2003, and ref. therein) substantiated
the monophyly of Menyanthaceae and their
placement in Asterales as sister group to a clade
consisting of Goodeniaceae, Calyceraceae and
Asteraceae. The monophyly of Menyanthaceae,
Goodeniaceae, Calyceraceae and Asteraceae is
further supported by the presence of petal lateral
veins and of a thick and multilayered integument, and the absence of micropylar endosperm
haustoria (Lundberg and Bremer 2003, and ref.
therein).
Distribution and Habitat. The family contains aquatic and marshland herbs. Menyanthes is
found in bogs, fens, lakes and rivers between sea
level and 3,200 m altitude, where it grows along
the margins of open water or in places flushed
by telluric waters, e.g. peat with surface water
flow. The main distribution area is circumpolar
between 40°N and the Arctic Circle. It grows north
of the Arctic Circle in Greenland, Norway, Alaska
and probably Siberia. Nymphoides is distributed
primarily in the tropics (13 spp. in Africa and
Madagascar, 12 in Australia, nine in India, five
in South America, three in Meso-America and
the West Indies). Two species are indigenous to
eastern North America, and Nymphoides peltata
is the only Eurasian species. Most species of
Nymphoides are true aquatics growing in ponds,
lakes and rivers. Nephrophyllidium is distributed
along the west coast of boreal North America
(from British Columbia to Alaska), in mountain
swamps of Japan and Korea, and in snowbeds
of high mountains along the coast of the Sea
of Japan. Twelve species of Villarsia are found
in southern Australia. Only two species are
distributed outside Australia, the one in the
south-western Cape Province of South Africa,
the other in south-eastern Asia. Villarsia occurs
predominantly in wet, moist or boggy habitats
such as swamps or swampy soils, lakes, stream
edges, wetlands and shallow (seasonal) waters. It
can rarely be found in aquatic habitats typical for
Nymphoides.
Economic Importance. After studying nitrogen and phosphorus accumulation and cycling by
Nymphoides peltata, Brock et al. (1983) concluded
that the species has the potential to function as
a nitrogen and phosphorus pump which regenerates sediment nutrients. Some species are cultivated as ornamentals, e.g. Nymphoides cordata.
Several species of Nymphoides have been recorded
as weeds in ricefields and irrigation channels.
Key to the Genera
1. Leaves trifoliolate, leaflets obovate or elliptic
2. Menyanthes
– Leaves simple, rounded, cordate, elliptic, ovate, reniform or linear
2
2. Leaves linear, slightly succulent; fruit a globose fleshy
berry
1. Liparophyllum
– Leaves rounded, cordate, elliptic, ovate, kidneyshaped; fruit a capsule, regularly (septicidal or loculicidal) or irregularly dehiscent, or fruit indehiscent,
dry
3
3. Leaves reniform with a deeply crenate margin
3. Nephrophyllidium
– Leaves rounded, cordate, elliptic, ovate, rarely reniform, margin entire, rarely minutely dentate-crenate
4
4. Fruit a valvate capsule
5. Villarsia
– Fruit indehiscent or opening irregularly 4. Nymphoides
Genera of Menyanthaceae
1. Liparophyllum Hook. f.
Liparophyllum Hook. f., London J. Bot. 6: 473 (1847).
Small creeping herbs with linear or spathulate,
slightly succulent leaves aggregated at nodes.
Flowers solitary in leaf axils on short and stout
pedicels. Flowers hermaphrodite, 5-merous,
monomorphic. Calyx with long, linear, acute
lobes. Corolla subrotate, deeply lobed. Corolla
lobes ovate, slightly hairy within, undulate. Ovary
bottle-shaped, stigma subsessile, two-lobed,
nectary glands absent. Fruit a globose, fleshy
berry. Seeds broadly elliptic, compressed laterally,
surface minutely wrinkled. One species, L. gunnii
Hook. f., New Zealand and Tasmania in wetland
communities at various elevations.
2. Menyanthes L.
Fig. 127
Menyanthes L., Sp. Pl. 1: 145 (1753).
Perennial herbs with submersed sympodial rhizomes with adventitious roots and a tuft of leaves
at the apex. Leaves trifoliolate, leaflets obovate or
elliptic, petiole with membranous sheaths towards
Menyanthaceae
603
Perennial herbs with stout, creeping, leafy rhizome.
Leaves erect, blades reniform, margin deeply crenate, petioles sheathing at base. One terminal culm
bearing a paniculate cyme with 10–30 flowers.
Flowers hermaphrodite, 5-merous, distylous. Calyx adnate to the ovary, with five lanceolate lobes.
Corolla campanulate, deeply lobed, lobes induplicate in bud, midrib with distinct crest. Ovary
semi-inferior, one-locular with five glands at the
base. Style short or absent, stigma with two large
semicircular lobes. Capsule cylindric, two-valved,
valves two-lobed. Seeds broadly and asymmetrically elliptic, compressed laterally, surface smooth.
2n = 34. One species, F. crista-galli (Menzies)
Makino, north-eastern Asia (Japan, Korea) and
Pacific coast of north-western North America.
4. Nymphoides Séguier
Nymphoides Séguier, Pl. Veron. 3: 121 (1754); Raynal, Adansonia II, 14: 227–270, 405–458 (1974), reg. rev.; Ornduff,
Brittonia 21: 346–352 (1969), reg. rev.; Sivarajan & Joseph,
Aquat. Bot. 45: 145–170 (1993), reg. rev.; Wood, J. Arnold
Arb. 64: 431–445 (1983), reg. rev; Li et al., Taiwania 47: 246–
258 (2002), reg. rev.
Limnanthemum S.G. Gmelin, p.p. (1769).
Fig. 127. Menyanthaceae. Menyanthes trifoliata. A Habit.
B Inflorescence and leaf. C Adaxial view of part of corolla
and androecium. D Calyx (one lobe removed) and ovary. E
Part of infructescence. F Seeds. (Ross-Craig 1964)
the base. One racemose inflorescence on leafless
scape from terminal bud of rhizome. Flowers
hermaphrodite, 5-merous, distylous. Calyx lobed.
Corolla broadly campanulate, deeply lobed, inside
of lobes bearded with fimbriae. Ovary superior,
with 5 nectary glands at the base, stigma capitate.
Fruit a two-valved loculicidal capsule. Seeds
numerous, broadly elliptic, compressed laterally,
surface smooth. 2n = 54. One species, M. trifoliata
L., circumboreal mainly between 40°N and the
Arctic Circle in lowland to montane bogs, fens,
rivers and lakes.
Perennial or rarely annual herbs with short, stout
rhizome producing clusters of spur-like adventitious roots, stolons and leaves. Leaves monomorphic or dimorphic, rhizomatous leaves small, often
spathulate with sheating petioles, floating leaves
arising from fertile culms, blades rounded to cordate, margin entire. Flowers in cymose, umbel-like
clusters. Flowers (4)5-merous, hermaphrodite and
distylous or homostylous, or flowers unisexual and
plants dioecious or gynodioecious. Calyx deeply
lobed. Corolla rotate, deeply lobed, with five glandular, staminode-like tufts or fringes of trichomes
near the base. Ovary superior or basally adnate to
the perianth tube with five nectar glands at the base.
Fruit a capsule, indehiscent or irregularly dehiscent. Seeds lenticular or globose, wingless, smooth
or variously ornamented. 2n = 18, 36, 54. About
40 species, cosmopolitan, mainly tropical and subtropical areas, aquatic plants in ponds, lakes and
slow-flowing streams.
5. Villarsia Vent.
3. Nephrophyllidium Gilg
Nephrophyllidium Gilg, in Engler & Prantl, Nat. Pflanzenfam. IV, 2: 105 (1895).
Fauria Makino (1904).
Villarsia Vent., Choix Plant: 9 (1803), nom. cons.; Aston,
Muelleria 2: 3–63 (1969), reg. rev.
Perennial or annual herbs, with rootstock or creeping rhizome bearing leaves, sometimes stolons,
604
G. Kadereit
and one to several fertile culms. Leaves erect or
floating, blades elliptic, ovate, rotund or reniform,
margin entire to crenate-dentate, petioles winged
or semi-sheathing towards the base. Inflorescence
paniculate or flowers in clusters or heads. Flowers
hermaphrodite, homo- or distylous. Calyx lobes
distinct almost to the base, persistent. Corolla
lobed, throat bearded with fimbriae, lobes with
a broad margin and sometimes with a vertical
keel on the inner surface. Ovary superior to
semi-inferior, with 5 nectary glands at the base.
Fruit a few- to many-seeded capsule, opening by
four (or secondarily more) apical valves. Seeds
smooth to tuberculate. 2n = 18, 36, 54. About 14
species, Australia, Cape region of South Africa
(1 sp.) and south-eastern Asia (1–2 spp.).
Selected Bibliography
Armstrong, J.E. 2002. Fringe science: are the corollas of
Nymphoides (Menyanthaceae) flowers adapted for surface tension interactions? Amer. J. Bot. 89: 362–365.
Aston, H.I. 2003. Seed morphology of Australian species of
Nymphoides (Menyanthaceae). Muelleria 18: 33–65.
Berg, R.Y. 1975. Myrmecochorous plants in Australia and
their dispersal by ants. Austral. J. Bot. 23: 475–508.
Bohm, B.A., Nicholls K.W., Ornduff, R. 1986. Flavonoids of
the Menyanthaceae: intra- and interfamilial relationships. Amer. J. Bot. 73: 204–213.
Brock, Th.C.M., Bongaerts, M.C.M., Heijnen, G.J.M.A., Heijthuijsen, J.H.F.G. 1983. Nitrogen and phosphorus accumulation and cycling by Nymphoides peltata (Gmel.)
O. Kuntze (Menyanthaceae). Aquat. Bot. 17: 189–214.
Chuang, T.I., Ornduff, R. 1992. Seed morphology and systematics of Menyanthaceae. Amer. J. Bot. 79: 1396–
1406.
Cronquist, A. 1981. An integrated system of classification
of flowering plants. New York: Columbia University
Press.
Dahlgren, G. 1989. The last Dahlgrenogram. System of classification of dicotyledons. In: Tan, K. (ed.) The Davis
and Hedge Festschrift. Edinburgh: University Press,
pp. 249–260.
Dulberger, R., Ornduff, R. 2000. Stigma morphology in
distylous and non-heterostylous species of Villarsia
(Menyanthaceae). Pl. Syst. Evol. 225: 171–184.
Erbar, C. 1997. Fieberklee und Seekanne – Enzian- oder
Aster-verwandt? Zur Blütenentwicklung und systematischen Stellung der Menyanthaceae. Bot. Jahrb. Syst.
119: 115–135.
Grayer, R.J., Chase, M.W., Simmonds M.S.J. 1999. A comparison between chemical and molecular characters
for the determination of phylogenetic relationships
among plant families: an appreciation of Hegnauer’s
“Chemotaxonomie der Pflanzen”. Biochem. Syst. Ecol.
27: 369–393.
Lammers, T.G. 1992. Circumscription and phylogeny of the
Campanulales. Ann. Missouri Bot. Gard. 79: 388–413.
Li, S.-P., Hsieh, T.-H., Lin, C.-C. 2002. The genus Nymphoides
Séguir (Menyanthaceae) in Taiwan. Taiwania 47: 246–
258.
Lundberg J., Bremer, K. 2003. A phylogenetic study of the
order Asterales using one morphological and three
molecular data sets. Intl J. Pl. Sci. 164: 553–578.
Metcalfe, C.R., Chalk, L. 1950. Anatomy of the dicotyledons,
vol. II. Oxford: Clarendon Press, pp. 933–939.
Metcalfe, C.R., Chalk, L. 1979. Anatomy of the dicotyledons,
ed. 2, vol. I. Oxford: Clarendon Press, pp. 190–221.
Nic Lughadha, E.M., Parnell, J.N. 1989. Heterostyly and
gene-flow in Menyanthes trifoliata L. (Menyanthaceae). Bot. J. Linn. Soc. 100: 337–354.
Nilsson, S. 1973. Menyanthaceae. In: Nilsson, S. (ed.) World
pollen and spore flora, vol. 2. Stockholm: Almquist &
Wiksell, pp. 1–19.
Ornduff, R. 1966. The origin of dioecism from heterostyly in
Nymphoides (Menyanthaceae). Evolution 20: 309–314.
Ornduff, R. 1970. Cytogeography of Nymphoides (Menyanthaceae). Taxon 19: 715–719.
Ornduff, R. 1974. Cytotaxonomic observations on Villarsia
(Menyanthaceae). Austral. J. Bot. 22: 513–516.
Ornduff, R. 1982. Heterostyly and incompatibility in Villarsia capitata (Menyanthaceae). Taxon 31: 495–497.
Ornduff, R. 1988. Distyly and monomorphism in Villarsia
(Menyanthaceae): some evolutionary considerations.
Ann. Missouri Bot. Gard. 75: 761–767.
Ornduff, R., Chuang, T.I. 1988. Chromosome numbers of Western Australian species of Villarisa
(Menyanthaceae). Pl. Syst. Evol. 161: 49–52.
Pollard, C.J., Amuti, K.S. 1981. Fructose oligosaccharides:
possible markers of phylogenetic relationships among
dicotyledonous plant families. Biochem. Syst. Evol. 9:
69–78.
Ridley, H.N. 1930. The dispersal of plants throughout the
world. Ashford: Kent.
Ross-Craig, S. 1964. Drawings of British plants, vol. 20. London: Bell & Hyman.
Sivarajan, V.V., Chaw, S.-M., Joseph, K.T. 1989. Seed coat
micromorphology of Indian species of Nymphoides
(Menyanthaceae). Bot. Bull. Acad. Sin. 30: 275–283.
Stolt, K.A.H. 1921. Zur Embryologie der Gentianaceen
und Menyanthaceen. Kungl. Svenska Vetenskaksakad.
Handl. 61, 14: 3–56.
Takhtajan, A. 1987. Systema Magnoliophytorum.
Leningrad: Nauka.
Tobe, H., Morin, N.R. 1996. Embryology and circumscription of the Campanulaceae and Campanulales: a review
of literature. J. Pl. Res. 109: 425–435.
Pentaphragmataceae
Pentaphragmataceae J. Agardh, Theoria Syst. Pl.: 95 (1858), nom. cons.
T.G. Lammers
Perennial herbs, somewhat fleshy or succulent.
Leaves alternate, simple, asymmetric at base,
estipulate, petiolate. Inflorescences axillary or
extra-axillary sympodial helicoid cymes; bracts
conspicuous, membranous. Flowers tetracyclic,
perfect, rarely imperfect and plants dioecious.
Calyx synsepalous, adnate to the ovary only by 5
longitudinal septa opposite the sepals, leaving 5
antepetalous nectariferous pits; lobes 5, imbricate.
Corolla sympetalous or rarely choripetalous,
radially symmetric; lobes (or petals) (4)5, valvate. Stamens (4)5, epipetalous or epigynous in
choripetalous species; filaments distinct; anthers
tetrasporangiate, dithecal, basifixed, distinct. Gynoecium syncarpous, 2(3)-locular; ovary inferior;
ovules numerous; placentation axile; style solitary;
stigma capitate. Fruit a berry, crowned by the
persistent perianth. Seeds minute, numerous;
endosperm copious, starchy.
The family comprises only one genus of c. 30
species, and is endemic to south-eastern Asia, the
Malay archipelago and New Guinea.
Vegetative Morphology and Anatomy.
Species of Pentaphragma are perennial herbs, generally rather coarse and often quite fleshy or succulent. Latex is not produced. Branched trichomes
occur frequently. In the stem, xylem and phloem
typically form a continuous cylinder traversed by
parenchymatous medullary rays 1–2 cells wide,
surrounding a broad pith. A distinct endodermis
is present. Vessel elements have scalariform perforation plates and bordered pits; helical thickenings
are absent. The xylem of the root is pentarch, surrounded by both an endodermis and exodermis.
Leaves are dorsiventral, large relative to plant size,
estipulate and petiolate; the base of the lamina is
quite asymmetric, resembling Begonia. Stomata
are anomocytic and chiefly abaxial. The mesophyll
has a single palisade layer or else is not clearly
differentiated. Vascular bundles in the veins are not
associated with sclerenchyma (Cronquist 1981).
Floral Structure and Anatomy. Some
species are dioecious, producing staminate
and carpellate flowers on separate plants. The
proterandry and specialized mechanisms for
secondary pollen presentation typical of Campanulaceae are lacking. The hypanthium is unique in
that the calyx tube is adnate to the ovary only by 5
narrow longitudinal septa opposite the stamens.
This creates five intervening pits or lacunae in
which nectar is produced. The corolla is typically
fleshy or cartilaginous, rarely delicate. Venation
of the petals includes both a midvein and a pair
of marginal veins, which anastamose distally.
Though the petals are connate in most species,
they are distinct in a few (e.g. P. decurrens). In the
former, the stamens are inserted on the corolla
tube just below the lobes, due to adnation of the
basal portions of the filaments. In the latter, they
are inserted at the rim of the top of the ovary. At
least one species (P. tetrapetalum) is characterized
by a tetramerous corolla and androecium. The
style is short and thick, lacking pollen-collecting
hairs. The stigma is large relative to the style.
Embryology. Anther wall development follows
the Basic pattern, forming a double middle layer.
Fibrous thickenings are found in the endothecium.
The tapetum is glandular (secretory), and tapetal
cells are binucleate. Pollen grains are binucleate
when shed.
Ovules are anatropous, unitegmic and tenuinucellar. Embryo sac formation is monosporic and
of the Polygonum type, and the sac protrudes from
the micropyle. Upon coming in contact with the
embryo sac, the inner layer of the integument develops as an endothelium (integumentary tapetum).
Embryogenesis follows the Solanad pattern. Endosperm formation is cellular ab initio; the endosperm typically is copious and starchy. A haustorium forms only at the micropylar end and is
single-celled. No hypostase forms (Kapil and Vijayaraghavan 1965).
606
T.G. Lammers
Pollen Morphology. The grains are trilobate
in outline and tricolporate or tricolpate. The surface is smooth and the exine nearly solid (Dunbar
1978).
Pollination, Reproductive Systems, Dispersal. Nothing is known of the reproductive biology of this family, aside from its clear lack of any
mechanism of secondary pollen presentation.
Phytochemistry. Nothing is known of the secondary chemistry of the family, aside from its lack
of alkaloids.
Affinities. Almost all authors have recognized a relationship between Pentaphragma
and Campanulaceae. The genus was included
among Campanuloideae by Bentham (1876) and
Hutchinson (1973). Schönland (1894) treated
the genus as a tribe within Campanulaceae
(Pentaphragmateae) whereas Takhtajan (1980)
treated it as a subfamily (“Pentaphragmatoideae”,
nom. invalid. sub Art. 41.1). However, the genus
differs from Campanulaceae as circumscribed
here in its lack of articulated lactifers, wood
anatomy (Carlquist 1997), asymmetric leaf bases,
sympodial helicoid cymes, absence of a specialized
mechanism for proterandrous secondary pollen
presentation, trilobate binucleate pollen (Dunbar
1978), basic pattern of anther wall development
and endosperm with micropylar haustorium only
(Kapil and Vijayaraghavan 1965), ovary adnate to
hypanthium only by 5 narrow longitudinal septa,
style without pollen-collecting hairs, and capitate
stigma. For these reasons, most modern authors
(Wagenitz 1964; Cronquist 1981; Dahlgren 1989;
Lammers 1992; Thorne 1992; Gustafsson and
Bremer 1995; Takhtajan 1997) have assigned the
genus to a distinct family.
Among those who have accorded Pentaphragma familial rank, the majority still considered it to be allied at some level to Campanulaceae.
Only Airy Shaw (1942, 1954, 1973) questioned this
relationship, suggesting instead (largely on the
basis of its asymmetric leaf base) that the family is
allied to Begoniaceae. However, that family (which
belongs to Malpighiales) differs from Pentaphragmataceae not only in numerous floral features but
also in its bitegmic crassinucellar ovules, Onagrad
embryogeny, and nuclear endosperm formation
(Kapil and Vijayaraghavan 1965).
A close relationship between Pentaphragmataceae and Campanulaceae is supported by recent
molecular data. Although an analysis based on rbcL
sequences (Cosner et al. 1994) suggested that Pentaphragmataceae are sister to a clade comprising
Asteraceae, Calyceraceae, Goodeniaceae, Menyanthaceae and Argophyllaceae, subsequent analyses
using two genes (Kårehed et al. 1999) and three
genes plus phenotypic data (Lundberg and Bremer
2003) have placed Pentaphragmataceae as the sister group of Campanulaceae as circumscribed here.
At the very least, Pentaphragmataceae are clearly
a member of Asterales s.l.
Distribution and Habitat. The family is
distributed in wet forest habitats from Myanmar,
southern China and Indochina, south through
Fig. 128. Pentaphragmataceae. Pentaphragma begonifolium. A Leaf. B Inflorescence. C Flower. D Corolla.
E Stamen. F Top of ovary with style. (Schönland 1894)
Pentaphragmataceae
most of the Malay archipelago (excluding Java and
Nusa Tenggara) to New Guinea.
Only one genus:
1. Pentaphragma
Fig. 128
Pentaphragma Wallich ex G. Don, Gen. Hist. 3: 731 (1834);
Airy Shaw, Fl. Malesiana I, 4: 517–528 (1954), part. rev.
See family description. Thirty species, southeastern Asia to New Guinea.
Selected Bibliography
Airy Shaw, H.K. 1942. Additions to the flora of Borneo and
other Malay islands: XIX. The Pentaphragmataceae of
the Oxford University expedition to Sarawak, 1932.
Bull. Misc. Inform. 1941: 233–236.
Airy Shaw, H.K. 1954. Pentaphragmataceae. In: van
Steenis, C.G.G.J. (ed.) Flora Malesiana I, 4. Djakarta:
Noordhoff-Kolff, pp. 517–528.
Airy Shaw, H.K. 1973. A dictionary of the flowering plants
and ferns, ed. 8. Cambridge: Cambridge University
Press.
Bentham, G. 1876. Campanulaceae. In: Bentham, G.,
Hooker, J.D., Genera Plantarum, vol. II. London:
Reeve, pp. 541–564.
Carlquist, S. 1997. Pentaphragma: a unique wood and its
significance. IAWA J. 18: 3–12.
Cosner, M.E., Jansen, R.K., Lammers, T.G. 1994. Phylogenetic relationships in the Campanulales based on rbcL
sequences. Pl. Syst. Evol. 190: 79–95.
Cronquist, A. 1981. An integrated system of classification
of flowering plants. New York: Columbia University
Press.
607
Dahlgren, G. 1989. The last Dahlrenogram. System of classification of the dicotyledons. In: Tan, K. (ed.) Plant
taxonomy, phytogeography, and related subjects. The
Davis and Hedge Festschrift. Edinburgh: Edinburgh
University Press, pp. 249–260.
Dunbar, A. 1978. Pollen morphology and taxonomic position of the genus Pentaphragm Wall. (Pentaphragmataceae). Grana 17: 141–147.
Gustafsson, M.H.G., Bremer, K. 1995. Morphology and phylogenetic interrelationships of the Asteraceae, Calyceraceae, Campanulaceae, Goodeniaceae, and related
families (Asterales). Amer. J. Bot. 82: 250–265.
Hutchinson, J. 1973. The families of flowering plants, ed. 3.
Oxford: Clarendon Press.
Kapil, R.N., Vijayaraghavan, M.R. 1965. Embryology of Pentaphragma horsfieldii (Miq.) Airy Shaw with a discussion on the systematic position of the genus. Phytomorphology 15: 93–102.
Kårehed, J., Lundberg, J., Bremer, B., Bremer, K. 1999.
Evolution of the Australasian families Alseuosmiaceae, Argophyllaceae, and Phellinaceae. Syst. Bot. 24:
660–682.
Lammers, T.G. 1992. Circumscription and phylogeny of the
Campanulales. Ann. Missouri Bot. Gard. 79: 388–413.
Lundberg, J., Bremer, K. 2003. A phylogenetic study of the
order Asterales using one morphological and three
molecular data sets. Intl J. Pl. Sci. 164: 553–578.
Schönland, S. 1894. Campanulaceae. In: Engler, A.,
Prantl, K., Die natürlichen Pflanzenfamilien, IV, 5.
Leipzig: W. Engelmann, pp. 40–70.
Takhtajan, A. 1980. Outline of the classification of flowering
plants (Magnoliophyta). Bot. Rev. 46: 225–359.
Takhtajan, A. 1997. Diversity and classification of flowering
plants. New York: Columbia University Press.
Thorne, R.F. 1992. An updated phylogenetic classification
of the flowering plants. Aliso 13: 365–389.
Wagenitz, G. 1964. Reihe Campanulales. In: Melchior, H.
(ed.) A. Engler’s Syllabus der Pflanzenfamilien, ed. 12,
Band 2. Berlin: Bornträger, pp. 478–497.
Phellinaceae
Phellinaceae (Loes.) Takht., Sist. Filogen. Cvetk. Rast.: 374 (1967).
G. Barriera, V. Savolainen and R. Spichiger
Small trees or shrubs, evergreen, dioecious. Leaves
alternate, spirally arranged or in pseudo-whorls,
clustered at the top of the branches, estipulate;
blades simple, usually papyraceous, rarely coriaceous or membranaceous, obovate, oblong,
glabrous; margin entire or crenate. Flowers small,
unisexual, regular, hypogynous, 4–5(6)-merous,
arranged in axillary or subterminal racemes or
panicles. Sepals small, connate at the base. Petals
valvate, free. Stamens isomerous, alternating with
the petals. Carpels isomerous, united into a plurilocular superior ovary. Fruit a polyspermous
drupe, lobed or not, greenish or black, containing
1–5 pyrenes. Seeds small with copious endosperm.
One genus comprising 11 species, all endemic
to New Caledonia.
Vegetative Morphology. Phellinaceae contain small trees or shrubs which are poorly
branched and 1–8 m high. Branches are hollow
in Phelline comosa. The cortex is thick, smooth
or furrowed, rugose or rugose-cracked in P. confertifolia, generally ochre but rarely greyish.
Large petiolary scars are present on the branches.
They are triangulate or trilobed, and concave in
P. comosa. The younger branches are dark brown
(P. macrophylla) or blackish (P. lucida). The leaves
are simple, evergreen, alternate or subopposite,
and clustered in loose spirals or in pseudo-whorls
at the top of the branches. They are generally
papyraceous, coriaceous in P. lucida or membranous in P. macrophylla, and obovate, oblong, or
sometimes elliptic or spathulate in shape. The
apex of the lamina is rounded to trilobed, rarely
shortly acuminate (less than 1 cm), and the margin
is entire or more or less crenate-sinuate. The base
can be cuneate, decurrent or not.
Vegetative Anatomy. Wood anatomy has been
extensively studied by Baas (1975). The wood is
fairly light and soft, and of yellowish colour. Growth
rings are indistinct, and the texture of the wood
is coarse through very conspicuous broad rays.
Growth rings are very faint to absent. Vessels are
mainly solitary, occasionally in radial pairs, angular to rounded. Perforations are scalariform in
almost vertical end-walls, resulting in strong overlap of adjacent vessel-members. Inter-vessel pits
are in a single vertical row, they are oval and variously flattened. Vessel-parenchyma pits are similar
but half-bordered. Vessel-ray pits are transitional
to scalariform, half-bordered, occasionally unilaterally compound. The ground tissue is composed of
fairly thick-walled fibre-tracheids with numerous
bordered pits on the radial and tangential walls,
the pits with slit-like apertures extending to the pit
border margins. Parenchyma is diffused and often
touching on the vessels. There are four uniseriate
rays per mm, the broad rays are extremely large,
occupying about 40% of the volume of the wood,
up to 14 cells wide and often over 2 cm high. They
are composed of large cells with numerous sheath
cells. No heartwood is differentiated. Baas (1975)
concluded that the wood anatomy of Phelline is one
of the most primitive among dicotyledons with vessels. Tufts of short reddish-brown hairs, hiding the
axillary buds, are usually present.
Inflorescence Structure. Inflorescences are
axillary racemes or simple or compound panicles.
Pedicels are generally alternate, but often opposite
in P. lucida. Bracts and prophylls are triangular or
rarely linear, deciduous or persistent, and coriaceous or papyraceous. Short, reddish-brown hairs
(long and stiff in P. comosa) are generally present
at the base of the bracts.
Flower Morphology. Sepals are small and
more or less persistent in fruit. The corolla is often
white, red in P. lucida, with valvate petals which
are 2–4.5 mm long. Petals are mostly fleshy and
mucronate or acuminate. The acumen is often
reflexed and up to 2 mm long (P. dumbeensis). The
stamens are isomerous, alternipetalous and free.
Phellinaceae
609
are between 17–23 × 17–24 µm. They are isopolar
and tricolporate. The ecto-apertures are long and
narrower in the equatorial region. The membrane
is granular and the endo-aperture is usually
horizontally 8-shaped. The exine is echinulate or
slightly rugulate and the foot layer is thick. The
endexine s. str. is very thin in the intercolpium and
very thick near the apertures. Two pollen types
can be distinguished, based on the ornamentation
and the location of the endexine (s.l.) thickening:
a microechinulate type in P. billardierei, P. brachyphylla, P. comosa, P. dumbeensis, P. erubescens,
P. indivisa and P. macrophylla, and a microrugulate
type in P. confertifolia and P. lucida.
Fruit and Seed, Dispersal. The fruits, being
polyspermous drupes, are probably adapted to dispersal by birds belonging to Turdidea. The only
species of this family present in New Caledonia is
Turdus poliocephalus.
Fig. 129. Phellinaceae A Branch of Phelline lucida. B–G
Phelline erubescens. B Flower bud. C Male flower. D Male
flower (stamens and petals removed) showing calyx and
pistillode. E Female flower. F Scale-shaped staminode. G
Fruiting branch and top view of fruit. (after Loesener 1942)
Anthers are basidorsifixed, oblong and introrse.
They usually are shorter than the filament and
show longitudinal dehiscence. In pistillate flowers,
the staminodes are similar to the stamens of the
male flowers (Loesener 1942) or are reduced to
small scales which are sometimes persistent in
fruit. The syncarpous ovary is globose and composed of isomerous carpels with as many locules
as carpels. The short style is terminal and has
distinct multilobed to multipartite stigma. There
is one axile, pendulous, hemitropous or slightly
campylotropous, unitegmic ovule per locule (Baas
1975). In staminate flowers, the pistillode is coneor bottle-shaped, and sometimes toothed at the
apex.
Pollen Morphology. Erdtman (1952) described the pollen of P. comosa, and Lobreau-Callen
(1977) examined nine species. The pollen grains
Phytochemistry. The leaves and wood of
Phelline are rich in alkaloids. Four species have
been particularly well studied, P. billardierei
(Seguineau and Langlois 1980), P. brachyphylla,
P. comosa (Debourges and Langlois 1982) and P.
aff. lucida (Langlois and Razafimbelo 1988). Alkaloids such as homoerythrin, homoerythroidin,
homoazaerythrin and holidin have been isolated.
Distribution and Habitat. Phelline is endemic to New Caledonia where it is found from c.
20 to 1,550 m elevation in mesophile to hygrophile
forests on schisteous, siliceous or serpentinous
soils (Hürlimann 1964).
Affinities. Ideas about the systematic position
of Phelline have changed several times, although
it mostly was included in rosids. In 1824, Labillardière considered Phelline as closely related to
Pouteria (Ebenales). Later, Bentham and Hooker
(1862) included Phelline in Rutaceae. Loesener
(1901, 1942) included it in Aquifoliaceae, but
as a distinct tribe characterized by valvate and
apiculate petals. Takhtajan (1966) accommodated
the genus in its own family, Phellinaceae, which
he considered to be related to Aquifoliaceae
(within Icacinineae) "from which it differs by
having valvate petals, by the character of the
inflorescence, the hemitropous or campylotropous
ovules, the wood anatomy, the quite different
sporoderm morphology, and the leaf venation".
Considered as close to Aquifoliaceae, Phellinaceae
610
G. Barriera, V. Savolainen and R. Spichiger
were placed next to this family in other systems
within Cornales (Dahlgren 1983), Theales (Thorne
1992) or Celastrales (Takhtajan 1980). Cronquist
(1981) did not accept family status for Phelline
and kept it within Aquifoliaceae (Celastrales).
Phylogenetic analyses based on molecular data
undoubtedly place Phellinaceae within Asterales
(Savolainen et al. 2000; Soltis et al. 2000; APG II
2003), with their most likely sister families being
Alseuosmiaceae and Argophyllaceae (Kårehed
et al. 1999; Lundberg and Bremer 2003).
Only one genus:
1. Phelline Labill.
Fig. 129
Phelline Labill., Sert. Austro-Caledon.: 35 (1824); Baillon,
Bull. Mens. Soc. Linn. Paris 1: 937–939 (1891), rev.; Loesener,
Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 78:
504–516 (1901), rev.
Characters as for the family. Eleven species in New
Caledonia.
Selected Bibliography
APG II. 2003. An update of the Angiosperm Phylogeny
Group classification for the orders and families of flowering plants: APG II. Bot. J. Linn. Soc. 141: 399-436.
Baas, P. 1975. Vegetative anatomy and affinities of Aquifoliaceae, Sphenostemon, Phelline and Oncotheca. Blumea
22: 311–407.
Baillon, H.E. 1891. Les Phelline de la Nouvelle-Calédonie.
Bull. Mens. Soc. Linn. Paris 1: 937–939.
Bentham, G., Hooker, J.D. 1862. Rutaceae. Genera Plantarum, vol. I. London: Reeve, Williams & Norgate, pp.
302–303.
Cronquist, A. 1981. An integrated system of classification
of flowering plants. New York: Columbia University
Press.
Dahlgren, R. 1983. General aspects of angiosperm evolution
and macrosystematics. Nordic J. Bot. 3: 119–149.
Debourges, D., Langlois, N. 1982. Nouveaux alcaloïdes
du groupe de l’homoérythrinane isolés de Phelline
brachyphylla. J. Nat. Prod. 45: 163–167.
Erdtman, G. 1952. Pollen morphology and plant taxonomy.
Angiosperms. Stockholm: Almquist & Wiksell.
Hürlimann, H. 1964. Phelline. In: Guillaumin, A. (ed.)
Résultats scientifiques de la mission franco-suisse de
botanique en Nouvelle-Calédonie (1950–1952) III.
Mém. Mus. Natl Hist. Nat., B, Bot. 15: 63–65.
Kårehed, J., Lundberg, J., Bremer, B., Bremer, K. 1999. Evolution of the Australasian families Alseuosmiaceae, Argophyllaceae, and Phellinaceae. Syst. Bot. 24: 660–682.
Labillardière, J.J.H. de 1824. Phelline. Sertum AustroCaledonicum. Paris: Huzard, pp. 35–36.
Langlois, N., Razafimbelo, J. 1988. Alcaloïdes de Phelline sp.
aff. Phelline lucida: origine de l’holidine et du phellinamide; configuration de la comosidine, de la lucidinine, et de leurs dérivés. J. Nat. Prod. 51: 499–503.
Lobreau-Callen, D. 1977. Famille des Phellinaceae. Les pollens des Célastrales. Montpellier: Ecole Pratique des
Hautes Etudes, Institut de Montpellier, pp. 42–43.
Loesener, T. 1901. Monographia Aquifoliacearum. Pars I.
Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur.
78: 504–516.
Loesener, T. 1942. Aquifoliaceae. In: Engler, A., Prantl, K.,
Die natürlichen Pflanzenfamilien, ed. 2, 20b. Leipzig:
W. Engelmann, pp. 36–86.
Lundberg, J., Bremer, K. 2003. A phylogenetic study of the
order Asterales using one morphological and three
molecular data sets. Intl J. Pl. Sci. 164: 553–578.
Savolainen, V., Fay, M.F., Albach, D.C., Backlund, A.,
van der Bank, M., Cameron, K.M., Johnson, S.A.,
Lledó, M.D., Pintaud, J.-C., Powell, M., Sheahan, M.C.,
Soltis, D.E., Soltis, P.S., Weston, P., Whitten, M.W.,
Wurdack, K.J., Chase, M.W. 2000. Phylogeny of the
eudicots: a nearly complete familial analysis based on
rbcL gene sequences. Kew Bull. 55: 257–309.
Seguineau, M.-F., Langlois, N. 1980. Phellibilidine, nouvel
alcaloïde de Phelline billiardieri. Phytochemistry 19:
1279–1281.
Soltis, D.E., Soltis, P.S., Chase, M.W., Mort, M.E., Albach, D.C., Zanis, M., Savolainen, V., Hahn, W.H.,
Hoot, S.B., Fay, M.F., Axtell, M., Swenson, S.M.,
Prince, L.M., Kress, W.J., Nixon, K.C., Farris, J.A. 2000.
Angiosperm phylogeny inferred from 18S rDN, rbcL
and atpB sequences. Ann. Missouri Bot. Gard. 84:
1–49.
Takhtajan, A.L. 1966 [1967]. Phellinaceae. Sistema i Filogenija Cvetkovyh Rastenij. Moskva: Nauka, pp. 374.
Takhtajan, A. 1980. Outline of the classification of flowering
plants (Magnoliophyta). Bot. Rev. (Lancaster) 46: 225–
359.
Thorne, R.F. 1992. Classification and geography of the flowering plants. Bot. Rev. (Lancaster) 58: 225–348.
Rousseaceae
Rousseaceae A.P. de Candolle, Prodr. 7: 521 (1839).
J.A. Koontz, J. Lundberg and D.E. Soltis
Climbing shrub, up to 4 m in height. Leaves opposite, simple, obovate-lanceolate, glandular serrate, without stipules. Flowers solitary to few, axillary, nodding, bisexual, actinomorphic, hypogynous. Sepals 4–6, reflexed, connate at base, persistent in fruit. Petals 4–5, valvate in bud, thick
and fleshy, connate into a lobed tube, lobes valvate,
revolute at apex, persistent. Stamens 4–5, inserted
on a nectary disk, anthers oblong-sagittate, extrorse. Ovary superior, 5–7-locular with numerous
unitegmic, tenuinucellate ovules. Placentation axile. Style thick and persistent, stigma globose with
5–7 lobes. Fruit a 4–7-angled berry. Seeds numerous, with small embryo and copious endosperm.
The family is monotypic. Roussea simplex is
endemic to the island of Mauritius.
Vegetative Morphology and Anatomy. The
wood contains vessels with very oblique end walls
and scalariform perforation plates with an average
of 20 (up to 49) bars, lateral scalariform pitting,
and lacks spiral thickenings. Tracheids and fibretracheids are imperforate with bordered pits and
lack septa. Wood-rays are tri- to multiseriate and
heterogeneous. The axial parenchyma are scanty
paratracheal. Nodes are trilacunar.
Leaves are opposite, but sometimes are
arranged in pseudo-whorls. They have peltate
glandular hairs on the abaxial surface as well as on
the petioles. Leaf venation is craspedodromous,
and the petiole and lamina often have radially
elongated, schizogynous resin canals. Stomata are
anomocytic.
Inflorescence, Floral Structure and
Flower Anatomy. The flowers are borne singly
(sometimes 3–4) in the leaf axils, and the pedicels
have peltate glandular hairs. The floral tube is
4–10 mm long, with lobes 10–20 mm long, and
villous outside. The calyx lobes are light green
and thick, have peltate glandular hairs but become
glabrous with age. The petals are valvate in bud,
and yellow to orange in colour. The stamens
alternate with the petals and the anthers dehisce
by longitudinal slits. The style is glabrous, thick,
unbranched and persistent. The 4–5-lobed stigma
is capitate and revolute at the margins. The
4–7-locular ovary has distinctly two-ranked ovules
on thick, axile placentas.
Embryology. Roussea is similar to most other
members of Asterales in its embryological features
(H. Tobe, J. Lundberg and P. Raven, unpubl. data).
The anthers are tetrasporangiate, with a persistent
epidermis, a glandular and binucleate tapetum,
but with an unspecialized endothecium and persistent middle layers. The ovules are anatropous,
unitegmic and tenuinucellate, with 1-celled archesporium and Polygonum-type embryo sac development. Antipodal cells are ephemeral, and no
hypostase is formed. Fertilization is porous, and
endosperm development is ab initio cellular. Micropylar haustoria and a filamentous suspensor are
developed. The seed coat is exotestal, and the exotesta is thick-walled and cuboidal. Mesotesta and
endotesta are crushed.
Pollen Morphology. The pollen has been described in some detail by Straka and Friedrich
(1988). The pollen grains are isopolar, more or less
spheroidal with a diameter up to 36.5 µm, and dispersed as monads. They are penta- to hexapanporate or -panbrachycolpate, with irregular aperture
border. The nexine is one-layered. The sexine has
a smooth or foveolate and complete tectum, and is
of about the same thickness as the nexine; the exine
is c. 2 µm thick.
Pollination and Reproductive System. Copious amounts of nectar with low sugar concentration have been reported to be produced in flowers
of Roussea (Dennis Hansen, pers. comm.). Hansen
(pers. comm.) also reports visits of the passerine
Mauritius Grey White-eye (Zosterops borbonicus
612
J.A. Koontz, J. Lundberg and D.E. Soltis
Fig. 130. Rousseaceae. Roussea simplex. A Flowering
branch. B Flower (perianth removed) showing the lobed
disc. C Ovary, longitudinal section. D Seed, longitudinal
section. (Engler 1891)
mauritianus) to Roussea, feeding on its nectar but
apparently not pollinating it.
Fruit and Seed. The fruit is a pale green angular berry with a dilated base. Seeds are numerous,
ovoid and flattened.
Phytochemistry. Very little is known about the
phytochemical constituents of Roussea. Tannins
are reported as being absent (Ramamonjiarisoa
1980).
Distribution and Habitats. Endemic to the
mountain forests of Mauritius, once locally abundant but now becoming increasingly rare.
Affinities. Historically, Roussea has often been
associated with the superficially similar genera
Brexia (East Africa, including Madagascar) and
Ixerba (New Zealand). Engler (1930), for example,
placed Roussea together with Brexia and Ixerba
in subfamily Brexioideae of Saxifragaceae, while
Dahlgren (1983), following basically the same
approach, placed the three genera in Brexiaceae of
Saxifragales, as did Takhtajan (1997) who treated
the three genera as three monogeneric families
in Brexiales. A somewhat different treatment
was given by Cronquist (1981), placing Roussea
(together with Ixerba and Brexia) in Grossulariaceae, and by Thorne (1992) who, although
recognizing Brexiaceae with Roussea, Ixerba and
Brexia, placed them not in Saxifragales but in
Hydrangeales. By contrast, various anatomical data
(e.g. trichomes and nodal vasculature) suggested
Roussea to be close to Quintinia (Paracryphiaceae), Pittosporaceae or Forgesia (Escalloniaceae)
(Al-Shammary and Gornall 1994). However, phylogenetic analyses of DNA sequence data strongly
suggest that Roussea is part of the well-supported
Asteridae s.s. (Savolainen et al. 1997; Soltis and
Soltis 1997; Koontz and Soltis 1999; Soltis et al.
2000). Roussea is thus only distantly related to
Brexia and Ixerba, which both appear as part of
a large rosid clade. Using ndhF and rbcL data,
Lundberg (2001) reanalysed the relationships of
Roussea, and found Carpodetaceae s.s. in Asterales
to be sister group to Roussea, thus confirming
the result of Savolainen et al. (2000). This was
later confirmed in an expanded analysis, using
atpB, ndhF, rbcL and morphological data, by
Lundberg and Bremer (2003). Based on the strong
support for the sister-group relationship between
Rousseaceae and Carpodetaceae, Lundberg (2001)
suggested that the two families should be treated
as subfamilies (Rousseoideae and Carpodetoideae)
in an expanded Rousseaceae s.l. However, the
monophyly of each subfamily is likewise well
supported, and they are therefore kept separate
in this treatment (see Carpodetaceae treatment in
this volume). A suggested synapomorphy for the
expanded Rousseaceae s.l. is the increased number
of carpels and ovary locules (Lundberg and
Bremer 2003). However, the basal relationships in
Asterales, including the possible sister relationship
between Rousseaceae s.l. and Campanulaceae, is
uncertain and any unequivocal synapomorphies
for Rousseaceae s.l. are still lacking.
Only one genus:
Rousseaceae
1. Roussea Smith.
Fig. 130
Roussea Smith, Pl. Icon. Ined. 1: 6 (1789).
Characters as for the family. The genus includes
one species, Roussea simplex Smith, on Mauritius.
Selected Bibliography
Al-Shammary, K.I.A., Gornall, R.J. 1994. Trichome anatomy
of the Saxifragaceae s. l. from the southern hemisphere.
Bot. J. Linn. Soc. 114: 99–131.
Cronquist, A.J. 1981. An integrated system of classification
of flowering plants. New York: Columbia University
Press.
Dahlgren, R. 1983. General aspects of angiosperm evolution
and macrosystematics. Nordic J. Bot. 3: 119–149.
Engler, A. 1891. Saxifragaceae. In: Engler, A., Prantl, K.,
Die natürlichen Pflanzenfamilien, III, 2a. Leipzig: W.
Engelmann, pp. 41–93.
Engler, A. 1930. Brexioideae. In: Engler, A., Prantl, K., Die
natürlichen Pflanzenfamilien, ed. 2, 18a. Leipzig: W.
Engelmann, pp. 185–187.
Koontz, J.A., Soltis, D.E. 1999. DNA sequence data reveal
polyphyly of Brexioideae (Brexiaceae; Saxifragaceae
sensu lato). Pl. Syst. Evol. 219: 199–208.
Lundberg, J. 2001. The asteralean affinity of the Mauritian
Roussea (Rousseaceae). Bot. J. Linn. Soc. 137: 267–276.
Lundberg, J., Bremer, K. 2003. A phylogenetic study of the
order Asterales using one morphological and three
molecular data sets. Intl J. Pl. Sci. 164: 553–578.
613
Ramamonjiarisoa, B.A. 1980. Comparative anatomy and
systematics of African and Malagasy woody Saxifragaceae sensu lato. Ph.D. Thesis, University of Massachusetts.
Savolainen, V., Spichiger, R., Manen, J.-F. 1997. Polyphyletism of Celastrales deduced from a chloroplast
noncoding DNA region. Mol. Phylog. Evol. 3: 27–37.
Savolainen, V., Fay, M.F., Albach, D.C., Backlund, A.,
van der Bank, M., Cameron, K.M., Johnson, S.A.,
Lledó, M.D., Pintaud, J.-C., Powell, M., Sheahan, M.C.,
Soltis, D.E., Soltis, P.S., Weston, P. Whitten, W.M.,
Wurdack, K.J., Chase, M.W. 2000. Phylogeny of the
eudicots: a nearly complete familial analysis based on
rbcL gene sequences. Kew Bull. 55: 257–309.
Soltis, D.E., Soltis, P.S. 1997. Phylogenetic relationships in
Saxifragaceae sensu lato: a comparison of topologies
based on 18S rDNA and rbcL sequences. Amer. J. Bot.
84: 504–522.
Soltis, D.E., Soltis, P.S., Chase, M.W., Mort, M.E., Albach, D.C., Zanis, M., Savolainen, V., Hahn, W.H.,
Hoot, S.B., Fay, M.F., Axtell, M., Swensen, S.M.,
Prince, L.M., Kress, W.J., Nixon, K.C., Farris, J.S. 2000.
Angiosperm phylogeny inferred from 18S rDNA,
rbcL, and atpB sequences. Bot. J. Linn. Soc. 133:
381–461.
Straka, H., Friedrich, B. 1988. Palynologica Madagassica
et Mascarenica Familien 65 bis 97. Trop. subtrop.
Pflanzenwelt 61: 1–117.
Takhtajan, A. 1997. Diversity and classification of flowering
plants. New York: Columbia University Press.
Thorne, R.F. 1992. Classification and geography of the flowering plants. Bot. Rev. 58: 225–348.
Stylidiaceae1
Stylidiaceae R. Br., Prodr.: 565 (1810), nom. cons.
Donatiaceae B. Chandler (1911).
R.C. Carolin
Herbs or sometimes subshrubs or cushion plants.
Leaves simple, spirally arranged, without stipules,
often in rosettes which may be basal, arranged
along a stem or on stolons. Aerial roots sometimes
present. Flowers in racemes or cymes, unisexual or
bisexual, actinomorphic or zygomorphic. Sepals 5
(rarely 3–7), connate. Petals 5(–10), free (Donatia)
or connate into a tube, abaxial lobe often differentiated into a small labellum. Stamens rarely 3
(Donatia p.p.), mostly 2, epigynous, free (Donatia) or adnate to the style to form a gynostemium.
Ovary inferior, 2–3-locular, sometimes 1-locular
by abortion of one locule or through reduction of
the septum; style simple or (Donatia) 2–3 stylodia,
stigmas 2–3; placentation axile or free central. Fruit
a capsule. Seeds small, with endosperm.
Six genera and about 245 species, mostly
Australia and New Zealand, few species in
south-eastern Asia and South America.
Vegetative Morphology. The cushion plant
genera Donatia and Phyllachne have a characteristic growth pattern. They are much-branched, each
branch is held closely against its neighbours, and
the older parts of the plant decay and new roots are
produced at the upper nodes. Carlquist (1969) postulated that the habit of Stylidiaceae is determined
by the fact that none have any significant secondary
thickening except, for example, the relatively weak,
anomalous type found in Stylidium laricifolium.
A large number of species produce rosettes (e.g.
S. graminifolium) with scapose inflorescences but
no other stems. Some of these, as they age, bear
the rosettes on substantial, thick stocks. In some
Stylidium spp., the rosettes are scattered along
the stems or the leaves appear to be whorled (e.g.
S. striatum). Levenhookia spp. and some Stylidium
spp. (e.g. S. alsinoides, S. laricifolium) bear the
leaves scattered along short aerial stems. A number
of species produce small bulbs at the base of the
1
Updated by J.W. Kadereit and J. Lundberg.
stem (e.g. S. asteroideum, S. petiolare) which may
enable them to resist very dry summer conditions.
Some, e.g. S. bulbiferum, have a branched creeping
habit where aerial ‘stilt’ roots hold the plants above
the soil surface. Yet others may be held ± erect
by aerial stilt roots which apparently survive for
only a few seasons and are replaced by others on
the branches of the original stem. This results in
vegetative fragmentation.
Carlquist (1969) suggested that the leaves are
basically linear with the broader leaves showing flabellate or striate venation, indicating an expansion
of a single mid-vein.
Vegetative Anatomy. The family shows
anomalous secondary thickening where the cambium is formed outside the vascular bundles and
where cambial activity is essentially unilateral and
tissue is produced only towards the inside. Three
different types of cambial activity are found. In
Stylidium subg. Nitrangium sect. Rhynchangium,
a cambium forms beneath the endodermis and
produces a determinate amount of fibres and
vessels with some intraxylary phloem strands
towards the inside but nothing towards the outside.
In lignotubers of the same section, a similar xylem
with intraxylary phloem strands is formed inside
a cambium inside the endodermis but the xylem
elements are distorted, and outside phellem is
formed. In some species, e.g. S. laricifolium,
a cambium is formed inside the endodermis and
an indeterminate xylary system with intraxylary
phloem is produced inside but nothing outside.
This weak, anomalous cambial activity may
indicate that the family is basically herbaceous
(Carlquist 1981). Rays are always absent (Carlquist
1992). The stomata are anomocytic, except in
Donatia where they are paracytic. Sieve tubes have
S-type plastids.
Inflorescence Structure. The inflorescence
is basically a thyrse which in many species is re-
Stylidiaceae
615
duced to a raceme or spike and, in Donatia, Phyllachne and Oreostylidium, to solitary terminal flowers (Carolin 1967).
Flower Structure. The flowers of Stylidiaceae
are unusual in the order for having imbricate petal
aestivation and extrorse anthers with divergent
thecae. They often are semi-resupinate and split
monosymmetrically. The floral construction of
Donatia differs strongly from the remainder of
the family. The petals, stamens and stylodia in
this genus are free from each other. Donatia
fascicularis has three stamens and three stylodia
but, in the other species of this genus, there are
only two stamens and stylodia. In all other genera,
the two stamens are fused with the style. Erbar
(1992) presented ontogenetic evidence that the
stamens are fused into a tube, and the tube is fused
to the style to form the gynostemium or column.
The ovary basically has a lower 2-locular region,
an upper 1-locular region and sometimes a small,
uppermost 2-locular region. In most species, the
1-locular region has the ovules attached on a free
central placenta, although in Donatia the ovules are
attached on axile placentas in the 2-locular region.
Two sterile strands of parenchyma connect the
free central placentas with the top of the loculus in
Phyllachne and some Stylidium spp. whilst in most
Stylidium spp. these two parenchymatous strands
project from the placenta but do not connect with
the top of the loculus (Mildbraed 1908; Carolin
1960). Donatia has a disc-like nectary but in the
other genera it is 2-lobed, one lobe being adaxial
and one abaxial, or sometimes just a single lobe on
one side only is present (Carolin 1960; Erbar 1992).
The nectary is extrastaminal. In Stylidium, there
are various outgrowths at the top of the corolla
tube which may function as pollinator guides or
to prevent access to non-pollinating insects. These
outgrowths are united into an erect sheath in
Levenhookia spp. (see below).
In Stylidium, the column is sensitive to touch
and executes a very fast movement. This explains
the vernacular name for these plants, i.e. ‘trigger
plants’. The movement, through an angle of 2–5
radians, occurs at the main bend in the column in
a time span of 10–30 ms, probably the fastest movement known in plants (Findlay and Findlay 1975).
The column at the bend consists of an epidermis,
with layers of parenchyma on either side with thickwalled cells inside referred to as the anterior layer
on the abaxial side and the posterior layer on the
adaxial side. Both these layers are rich in starch
Fig. 131. Stylidiaceae. A, B Donatia novae-zelandiae.
A Habit. B Flower. C, D Phyllachne colensoi. C Habit.
D Flower. E, F Stylidium graminifolium. E Habit. F Flower.
G, H Forstera bidwillii. G Habit. H Flower. I, J Oreostylidium
subulatum. I Habit. J Flower. K, L Levenhookia stipitata.
K Habit. L Flower. (Illustrations adapted from Mildbraed
1908 by Rebecca Wagstaff; C–J from Wagstaff and Wege
2002)
grains and have conspicuous vacuoles and mitochondria, and both layers have cell walls which are
rich in pectin and have almost no longitudinal cellulose fibrils but only circumferential ones. There
is also a central layer of cells (Findlay and Findlay 1989). In the firing position, the anterior layer
has high turgor and high concentrations of KCl.
Findlay (1982) considered that this turgor is balanced by the stretched cells of the posterior layer.
Stimulation causes the cell walls to buckle trans-
616
R.C. Carolin
versely (presumably, the lack of longitudinal fibrils is critical in this respect), thus unbalancing the
system and initiating the movement. Immediately
after firing, KCl moves into the posterior layer and
gradually returns to the anterior layer as the firing
position is resumed. It seems that the column has
to lie against the labellum to attain firing potential.
Recovery takes 200–700 s. Those species which do
not show movement do not show the characteristic anatomy of the bend anywhere in the column
(Findlay and Findlay 1989). In Levenhookia, the
labellum is sensitive to touch when mature.
Embryology. The ovule is anatropous, unitegmic, tenuinucellate and endotheliate. The
development of the endosperm is ab initio cellular
and, at the 8-tier stage, each tier consists of two
cells. The upper and lower tiers remain two-celled
and form intrusive haustoria at each end of the
embryo sac (Subramanyam 1952; Philipson and
Philipson 1973). Embryo sac formation follows the
Polygonum type, and the embryo develops according to the Solanad type. Anther wall formation
is dicotyledonous, the tapetum is glandular, and
pollen grains are 2- or 3-celled when shed (Tobe
and Morin 1996).
Fruit and Seed, Dispersal. Fruits are capsular, although those of Donatia do not appear to
dehisce. Seeds are very small but little is known
about their dispersal. Carlquist (1969) suggested
that they could easily be carried in mud on the feet
of birds.
Pollen Morphology. Pollen grains are fairly
uniform in the family. They are spherical and colpate. The number of colpi varies in the range of 2–8,
Phyllachne and Forstera having 3–4 (very rarely 5)
and Stylidium and Levenhookia 2–8 colpi (Bronkers
and Stainer 1972).
Karyology. Haploid chromosome numbers of
n = 5–16, 18, 24, 26, 28 and 30 have been reported
(Lammers 1992). James (1979) considered that the
base number of Stylidium is x = 15. Donatia fascicularis has a haploid number of n = 24 (Moore
1983). From this, Lammers (1992) speculated that
the base number of the family is x = 6 or 8.
Reproductive Biology. The flowers of Stylidium and Levenhookia have very precise pollination
mechanisms which often appear to make use of
specific pollinators (Erickson 1958). In both gen-
era, one of the petals is modified into a small labellum. This has a different function in the two
genera but, in both, the effect is that the flowers
appear to be tetramerous. In Stylidium the column is bent over the labellum, obscuring it in most
species. Mildbraed (1908) suggested that the labellum holds the column in position. Carlquist (1969)
believed that the glossy swelling on the labellum
of many Stylidium spp. is a false nectary. Except
in two species, S. insensitivum and S. beaugleholei,
the column is sensitive to the touch of the pollinator seeking nectar at the base of the corolla tube.
The column moves very rapidly, articulating at the
bend of the column (see above), and strikes the pollinator either on the back or on the undersurface,
thus depositing pollen there, brushing pollen onto
the stigmas, or both. Many species are protandrous,
thereby excluding self-pollination within an individual flower. By contrast, the ephemeral species do
not appear to be protandrous. The stigmas here,
however, seem to touch the insect before pollen
from the same flower is deposited (Erickson 1958).
Both Erickson (1958) and Carlquist (1969) note that
in some species, e.g. S. sacculatum, the stigmas and
anthers are closely appressed to an expansion of
part of the column before it is triggered. Carlquist
considered this to be a mechanism to ensure selfpollination when no triggering occurs. Nothing is
recorded about self-incompatibility in the family.
In Levenhoookia, the column is tightly enclosed by
the labellum. It is released only when mature, either
by the pressure of the growing column or by the pollinator touching a sensitive region on the labellum.
At release, the labellum springs downwards and
the column moves away from the labellum across
the flower and touches a sheathing outgrowth of
the top of the corolla tube. The jerk brought about
by this contact sprays pollen over the pollinator.
The column recovers, and the labellum returns to
the erect position but does not enclose the column
any more. Subsequent visits by pollinators deposit
pollen on the mature stigmas (Erickson 1958).
Phytochemistry. The storage carbohydrate of
the family is inulin. Members of the family produce
a carbocyclic iridoid similar to monotropein. No
alkaloids have been recorded from the family but
caffeic acid may be present (Lammers 1992).
Subdivision of and Relationships Within the
Family. It is yet not finally settled whether Donatia is closest relative to the remainder of the family and should be included in it. This approach,
Stylidiaceae
first taken by Mildbraed (1908), is supported by
a morphological study by Gustafsson and Bremer
(1995) and the most recent, combined molecularmorphological analysis of the order by Lundberg
and Bremer (2003), but not by some other molecular analyses where Stylidiaceae and Donatia occupy
quite separate positions in the order (Albach et al.
2001; Bremer et al. 2002; see below). Taken as one
family, Donatia is sister to the remainder (Laurent
et al. 1998), as also suggested by Mildbraed (1908).
Among the remaining genera, Forstera/Phyllachne
are sister to Levenhookia/Stylidium, and Oreostylidium appears to be nested within Stylidium and
should perhaps be included in that genus (Laurent
et al. 1998; Wagstaff and Wege 2002). Oreostylidium, with almost actinomorphic flowers, has been
interpreted as a case of extreme paedomorphic reduction (Laurent et. 1998). The exact relationship
between Phyllachne and Forstera is still uncertain
but it is likely that they are nested within each
other, and it has recently been suggested that Phyllachne should be merged with Forstera (Wagstaff
and Wege 2002).
Affinities. Although sometimes classified
outside Asterales (for discussion, see Lammers
1992; Lundberg and Bremer 2003), there is broad
consensus that the family (incl. Donatia) is
part of this order (APG II 2003). A sister-group
relationship of Stylidiaceae to the Menyanthaceae/Goodeniaceae/Calyceraceae/Compositae
clade (MGCA clade) of Asterales was resolved by
Lundberg and Bremer (2003), although with low
support. Transition from woody to herbaceous
habits, and from scalariform to simple vessel
perforation plates were identified as characters in
support of such relationships by these authors.
By contrast, Stylidiaceae (excl. Donatia) were
sister to Campanulaceae in the analyses by Albach
et al. (2001) and Bremer et al. (2002), again
with low support in the former study. In these
two analyses, Donatia was found to be closest
relative, with low support, of either Alseuosmiaceae/Argophyllaceae/Phellinaceae
(Bremer
et al. 2002), or to all families of Asterales except
Stylidiaceae/Campanulaceae (Albach et. al. 2001).
Considering the lack of sufficiently good support
for many nodes in all these analyses, the exact
relationships of Stylidiaceae should be regarded
as unclear, although the evidence presented by
Lundberg and Bremer (2003) makes a relationship
of the family to Campanulaceae a little less
likely.
617
Size, Distribution and Habitats. The family contains six genera and about 170 species, of
which most belong to Stylidium. Stylidiaceae are almost entirely a southern hemisphere family, most
species being confined to Australia. One species
of Stylidium extends into south-eastern Asia as
far as Sri Lanka. With the exception of Stylidium
and Levenhookia, all genera occur in New Zealand.
Phyllachne and Donatia are also found in southern South America. The species occur in a variety
of habitats, from seasonally wet areas in the tropics where they may be ephemerals, to the sclerophyll communities of southern Australia and the
exposed alpine habitats of the higher latitudes of
the southern continents. Donatia is the major component of the very peculiar, hard-cushion bogs in
South America and New Zealand.
Palaeobotany. The fossil record of Stylidiaceae
is scanty, but fossil pollen (Tricolpites stylidioides)
similar to pollen from Forstera has been reported
from the south-eastern Australian Oligocene (Macphail 1997).
Key to the Genera
1. Petals free; stamens free from the stylodia 1. Donatia
– Petals connate; stamens adnate to the style into a column
2
2. One corolla lobe (the labellum) smaller than the others, usually lying beneath the bent column or enclosing
the erect or slightly bent column
3
– Corolla lobes ± equal
4
3. Column usually bent, rarely almost straight and then
never enclosed at top by labellum
6. Stylidium
– Column straight or slightly bent, top enclosed by labellum before triggering
5. Levenhookia
4. Peduncles longer than leaves
3. Forstera
– Peduncles, if present, shorter than leaves
5
5. Flowers almost sessile; glabrous cushion plant
2. Phyllachne
– Flowers shortly pedicellate; stoloniferous herb with
glandular hairs on the calyx
4. Oreostylidium
Genera of Stylidiaceae
1. Donatia J.R. Forst.
Fig. 131A, B
Donatia J.R. Forst., Char. Gen.: 9, t. 5 (1776).
Much-branched short herbs, forming dense, often
large, cushion-like expanses. Leaves small, linear,
hard, crowded on the stem. Flowers solitary,
terminal, sessile, regular, bi- or unisexual. Sepals
3–7. Petals 5–10, free. Stamens 2–3, epigynous
inside a disc, extrorse. Ovary inferior, 2–3-locular
618
R.C. Carolin
with axile placentation towards the top of the
loculi. Fruit an indehiscent capsule. 2n =24. Two
species, Australia, New Zealand, South America.
2. Phyllachne J.R. Forst.
Fig. 131C, D
Phyllachne J.R. Forst., Char. Gen.: 115, t. 58 (1776).
Much-branched short herbs, forming dense,
often large, cushion-like expanses. Leaves small,
lanceolate-subulate, hard, crowded on the stems.
Flowers solitary, terminal, sessile, regular, bi- or
unisexual. Sepals 2–9, usually 5, subequal. Petals
3–9, usually 5, united into a short tube. Stamens
2, united with the style into a short erect column.
Nectaries 2, reniform. Ovary incompletely 2locular. Fruit an indehiscent capsule. Four species,
New Zealand (3 spp.), Tasmania, South America.
3. Forstera L. f.
Fig. 131G, H
Forstera L. f., Nov. Act. Upsal., 3: 184, t. 9 (1780).
Erect to decumbent herbs. Leaves scattered along
the stems. Flowers almost regular, solitary on long
peduncles, bisexual. Sepals 5–6, subequal. Petals
5–9, united into a short tube, sometimes with appendages in the throat. Stamens 2, united with the
style into a short erect column. Ovary imperfectly
2-locular. Nectaries 2, reniform. Fruit a capsule.
Five species, New Zealand, Tasmania.
4. Oreostylidium Berggren
Fig. 131I, J
Oreostylidium Berggren, Minnesskr. Fysiogr. Sällsk. Lund.
8: 1, t. 1 (1878).
Small herbs. Leaves basal, narrow. Flowers almost
sessile in the basal leaf axils, regular, bisexual, with
glandular hairs. Sepals 5, subequal. Petals 5, united
into a tube with short lobes. Stamens 2, united with
the style into a short erect column which is exserted
from the corolla tube. Nectaries 2, subglobose. Fruit
a capsule. Seeds small. 2n = 30. One species, O. subulatum Berggren, New Zealand.
The genus has been included in Stylidium
Swartz ex Willd. by Laurent et al. (1998).
5. Levenhookia R. Br.
Fig. 131K, L
Levenhookia R. Br., Prodr. 572 (1810).
Small erect herbs, often ephemeral. Leaves spirally
arranged along the stem. Flowers arranged in
racemes, irregular, bisexual. Sepals 5, free. Petals
5, connate into a tube, with outgrowths usually
forming a sheath at the throat; abaxial petal (la-
bellum) erect, sensitive and enclosing the column
until triggered. Stamens 2, united with the style
into a ± bent, slightly sensitive column. Ovary
1-locular with a free-central placentation. Nectary
on top of the ovary. Fruit a capsule. Seeds small.
Ten species, Australia.
6. Stylidium Swartz ex Willd.
Fig. 131E, F
Stylidium Swartz ex Willd., Sp. Pl. 4: 7, 146 (1805). nom.
cons., non Stylidium Lour.
Herbs, sometimes ephemerals, or undershrubs,
sometimes with bulbs. Leaves spirally arranged
along the stems or in a basal rosette or in rosettes
along the stems or stolons. Stems erect, sometimes
very short and thick as a basal stock or creeping,
often raised above the soil surface by aerial roots.
Flowers arranged in thyrses, cymes or racemes,
irregular, bisexual. Sepals 5, free or variously
connate. Petals 5, connate into a tube, with
various outgrowths in the throat; abaxial petal
smaller (labellum). Stamens 2, united with the
style into a bent, usually sensitive column. Ovary
2-locular with axile placentation or 1-locular
with free-central placentation or 1-locular (by
abortion of the other loculus) with apparently one
parietal placenta. Nectaries 1–2, hemispherical or
reniform. Fruit a capsule. Seeds numerous, usually
small. 2n = 10–32, 36, 52, 56, 60. About 220 species,
mostly Australia, a few species in south-eastern
Asia, reaching as far as Sri Lanka (1 sp.)
The following subgenera are usually recognised. Subg. Centridium: column short, bent just
below the anthers with an elongated stigma, or
not bent and not sensitive; subg. Forsteriopsis:
leaves short, almost scale-like, tightly overlapping;
subg. Andersonia: capsule linear, septum of ovary
almost complete; subg. Alsinoides: capsule linear,
septum of ovary narrow; subg. Stylidium (Tolypongium): capsule ovoid-oblong, rarely linear,
septum of ovary usually incomplete or absent;
subg. Nitrangium: capsule linear, rarely ovate,
septum of ovary very reduced, throat usually
without appendages.
Selected Bibliography
Albach, D.C., Soltis, P.S., Soltis, D.E., Olmstead, R.G. 2001.
Phylogenetic analysis of asterids based on sequences
of four genes. Ann. Missouri Bot. Gard. 88: 163–212.
APG II. 2003. An update of the Angiosperm Phylogeny
Group classification for the orders and families of flowering plants: APG II. Bot. J. Linn. Soc. 141: 399–436.
Stylidiaceae
Bremer, B., Bremer, K., Heidari, N., Erixon, P., Olmstead, R.G., Anderberg, A.A., Källersjö, M., Barkhordarian, E. 2002. Phylogenetics of asterids based
on 3 coding and 3 non-coding chloroplast DNA
markers and the utility of non-coding DNA at higher
taxonomic levels. Mol. Phylog. Evol. 24: 274–301.
Bronckers, F., Stainer, F. 1972. A l’étude morphologique du
pollen de la famille des Stylidiaceae. Grana 12: 1–22.
Carlquist, S. 1969. Studies in Stylidiaceae: new taxa, field
observations, evolutionary tendencies. Aliso 7: 13–64.
Carlquist, S. 1981. Types of cambial activity and wood
anatomy of Stylidium (Stylidiaceae). Amer. J. Bot. 68:
778–785.
Carlquist, S. 1992. Wood anatomy of sympetalous dicotyledon families: a summary, with comments on the systematic relationships and evolution of the woody habit.
Ann. Missouri Bot. Gard. 79: 303–332.
Carolin, R.C. 1960. Floral structure and anatomy in the
family Stylidiaceae Swartz. Proc. Linn. Soc. New South
Wales 85: 189–196.
Carolin, R.C. 1967. The concept of the inflorescence in the
order Campanulales. Proc. Linn. Soc. New South Wales
92: 7–26.
Erbar, C. 1992. Floral development of two species of Stylidium (Stylidiaceae) and some remarks on the systematic
position of the family Stylidiaceae. Canad. J. Bot. 70:
258–271.
Erickson, R. 1958. Triggerplants. Perth: Paterson Brokensha.
Findlay, G.P. 1982. Generation of torque by the column of
Stylidium. Austral. J. Pl. Physiol. 9: 271–286.
Findlay, G.P., Findlay, N. 1975. Anatomy and movement of
the column in Stylidium. Austral. J. Pl. Physiol. 2: 597–
621.
Findlay, G.P., Findlay, N. 1989. The structure of the column
in Stylidium. Austral. J. Bot. 37: 81–101.
619
Gustafsson, M.H.G., Bremer, K. 1995. Morphology and phylogenetic interrelationships of the Asteraceae, Calyceraceae, Campanulaceae, Goodeniaceae, and related
families (Asterales). Amer. J. Bot. 82: 250–265.
James, S.H. 1979. Chromosome numbers and genetic systems in the trigger plants of Western Australia (Stylidium; Stylidiaceae). Austral. J. Bot. 27: 17–25.
Lammers, T.G. 1992. Circumscription and phylogeny of
the Campanulales. Ann. Missouri Bot. Gard. 79: 388–
413.
Laurent, N., Bremer, B., Bremer, K. 1998. Phylogeny and
generic interrelationships of the Stylidiaceae (Asterales), with a possible extreme case of floral paedomorphosis. Syst. Bot. 23: 289–304.
Lundberg, J., Bremer, K. 2003. A phylogenetic study of the
order Asterales using one morphological and three
molecular data sets. Intl J. Pl. Sci. 164: 553–558.
Macphail, M.K. 1997. Comment on M. Pole (1994): ’The New
Zealand flora – entirely long-dispersal?’ J. Biogeogr. 22:
625–635.
Mildbraed, J. 1908. Stylidiaceae. In: Pflanzenreich IV: 278.
Leipzig: W. Engelmann.
Moore, D.M. 1983. Flora of Tierra del Fuego. Oswestry:
Anthony Nelson.
Philipson, W.R., Philipson, M.N. 1973. A comparison of the
embryology of Forstera L. and Donatia Forst. et f. N.
Z. J. Bot. 11: 449–460.
Subramanyam, K. 1952. The nutritional mechanism of
embryo sac and embryo in the families Campanulaceae, Lobeliaceae and Stylidiaceae. J. Mysore Univ.
sect. B 13: 1.
Tobe, H., Morin, N.R. 1996. Embryology and circumscription of Campanulaceae and Campanulales: a review of
literature. J. Pl. Res. 109: 425–435.
Wagstaff, S.J., Wege, J. 2002. Patterns of diversification in
New Zealand Stylidiaceae. Amer. J. Bot. 89: 865–874.
Index to Scientific Names
References to main entries in bold-faced print, to illustrations in italics.
Aaronsohnia 368
Abrophyllaceae 57
Abrophyllum 59
A. ornans 59
Abrotanella 214
A. linearis 215
Acamptopappus 321
Acanthocephalus 184
Acanthocladium 252
Acanthodesmos 152
Acantholepis 128
Acanthospermum 488
Acanthostyles 546
Acanthotheca 245
Acasma 132
Achaetogeron 340
Achillea 364
A. group 364
Achnophora 295
Achnopogon 98
Achyrachaena 499
Achyrocline 252
Achyrocome 263
Achyropappus 435
Achyroseris 198
Achyrothalamus 121
Acicarpha 23
Acilepidopsis 166
Acilepis 166
Acmella 471
Acomis 252
Acosta 146
Acourtia 103
Acrisione 223
Acritopappus 522
A. connatifolius 522
Acroclinium 279
Acroptilon 143
Actinoseris 117
Actionbole 252
A. oldfieldiana 254
Actites 191
Adelostigma 387
Adenachaena 356
Adenanthellum 350
Adenanthemum 350
Adenocaulon 108
Adenocritonia 562
Adenocyclus 159
Adenoglossa 354
Adenoon 171
Adenopappus 429
Adenophora 43
Adenophyllum 424
Adenostemma 518
A. viscosum 519
Adenostemmatinae 518
Adenostyles 240
Adenothamnus 499
Aedesia 171
Aegialophila 145
Aegopordon 137
Aequatorium 223
A. jamesonii 223
A. subg. Praegynoxys 223
Aetheolaena 234
Aetheopappus 140
Aetheorhiza 190
Ageratella 555
Ageratina 167, 514
Ageratinae 520
Ageratinastrum 167
Ageratum 522
Agiabampoa 466
Agnorhiza 464
Agoseris 191
Agrianthus 542
A. group 542
Ainsliaea 123
Ajania 357
A. group 357
Ajaniopsis 357
Akeassia 304
Akylopsis 368
Alatoseta 253
Albertinia 154
Alcantara 163
Alciope 225
Aldama 465
Alepidocline 484
Alfredia 134
Alibum 180
Aliella 253
Allagopappus 379
Allardia 360
Allendea 178
Allittia 341
Alloispermum 484
Allopterigeron 390
Almutaster 334
Alomia 556
Alomiella 551
Alomiinae 552
Alseuosmia 10
A. banksii 9
A. macrophylla 9
Alseuosmiaceae 3, 7
Alvordia 466
Amauria 509
Amauriinae 509
Amauriopsis 435
Ambassa 167
Amberboa 142
Amblyocarpum 383
Amblyolepis 404
Amblyopappus 495
Amblysperma 115
Amboroa 571
Ambrosia 444
A. canescens 444
A. polystachya 444
Ambrosiaceae 443
Ambrosieae 443
Ambrosiinae 441, 443
Ameghinoa 106
Amellus 291
A. asteroides 291
Ammanthus 365
Ammobium 253
A. craspedioides 254
Amolinia 572
Ampelaster 334
Ampherephis 160
Amphiachyris 321
Amphidoxa 266
Amphiglossa 253
Amphipappus 321
Amphoricarpos 132
Anacantha 137
Anacyclus 364
Anaglypha 275
Anaphalioides 253
A. bellidioides 252
Anaphalis 253
A. margaritacea 252
Anaxeton 254, 281
Ancathia 134
Ancistrocarphus 254
Anderbergia 255
Andromachia 178
622
A. sect. Chrysactinium 179
A. sect. Oligactis 178
Andryala 194
Anemocarpa 255
A. saxatilis 254
Angelianthus 178
Angelphytum 450
Angianthus 255
Anisocarpus 499
Anisochaeta 255
Anisocoma 193
Anisopappinae 397
Anisopappus 397
A. chinensis 397
Anisothrix 255
Annaea 42
Antennaria 255
A. pulchella 256
Anteremanthus 161
Antheidosorus 265
Anthemideae 342
Anthemis 365
A. group 365
Antheropeas 495
Anthocerastes 282
Anthotium 594
Antillanthus 227
Antillia 562
Antiphiona 384
Antithrixia 256
Antunesia 173
Anura 136
Anvillea 381
A. garcinii 381
Anvilleina 381
Apalochlamys 256
Apargidium 191
Apetahia 52
Aphanactis 484
Aphanostephus 339
Aphylloclados 111
Aplopappus 321, 338
A. sect. Acamptopappus 321
A. sect. Leucopsis 338
Apopyros 340
Aposeris 196
Apostates 435
Arbelaezaster 234
Archibaccharis 311
Arctanthemum 357
Arctium 136
A. group 135
A. lappa 136
Arctogeron 317
Arctotheca 203
Arctotideae 200
Arctotidinae 202
Arctotis 203
A. stoechadifolia 203
Arcyna 132
Argentipallium 256
Argophyllaceae 3, 13
Argophyllum 16
A. ellipticum 14
Index to Scientific Names
Argyranthemum 369
A. foeniculaceum 369
A. group 369
Argyrocalymma 59
Argyroglottis 256
Argyrophanes 260
Argyrotegium 284
Argyrovernonia 160
Argyroxiphium 500
A. sandwicense 500
Arida 329
Aristeguietia 567
Arnaldoa 90
Arnica 494
A. dealbata 494
A. mollis 494
Arnicastrum 430
Arnicinae 493
Arnoglossum 220
Arnoseris 196
Arrhenechthites 228
Arrojadocharis 542
Arrowsmithia 257
Artanacetum 358
Artemisia 358
A. absinthium 358
A. group 358
Artemisiella 360
Artemisiopsis 257
Arthrolepis 364
Asaemia 351
Asanthus 555
Ascaricida 172
Ascidiogyne 524
Aspilia 459
Asplundianthus 568
Astephania 397
Aster 317
A. subg. Eurybia 287
A. subg. Galatea 318
A. subg. Kalimeris 318
A. sect. Oritrophium 300
Asteraceae 61
Astereae 284
Asteridea 257
Asterinae 316
Asteriscus 382
Asteroideae 78
Asteropsis 313
Asterothamnus 317
Astradelphus 340
Astranthiinae 336
Astranthium 336
Astrocodon 42
Asyneuma 44
Asyneumopsis 44
Atalanthus 191
Athanasia 350
A. dentata 351
A. group 350
Athrixia 257
Athroisma 398
A. boranense 399
Athroismeae 392, 395
Athroisminae 398
Atractylis 127
Atractylodes 127
Atrichantha 257
Atrichoseris 193
Aucklandia 137
Austrobrickellia 557
Austrocritonia 568
A. angulicaulis 568
Austroeupatorium 529
A. inulifolium 529
Austroliabum 178
Austrosynotis 240
Avellara 198
Axiniphyllum 489
Ayapana 549
A. amygdalina 549
Ayapaninae 548
Ayapanopsis 550
Aylacophora 295
Aynia 154
Azorina 41
Aztecaster 295
Babcockia 191
Bacasia 89
Baccharidastrum 311
Baccharidinae 310
Baccharidiopsis 311
Baccharidoideae 310
Baccharis 311
Baccharoides 172
Badilloa 568
Baeria 496
Baeriinae 494
Baeriopsis 495
Bafutia 238
Bahia 435
B. absinthifolia 435
Bahianthus 543
B. viscosus 543
Bahieae 392, 433
Bahiopsis 466
Baileya 404
B. multiradiata 405
Bajacalia 425
Balduina 401
Balsamita 366
Balsamorhiza 464
Baltimora 449
Barkleyanthus 221
Barnadesia 89
B. odorata 89
Barnadesieae 87
Barnadesioideae 77
Barroetea 554
Barrosoa 537
Bartlettia 436
Bartlettina 573
Basedowia 257
Batopilasia 327
Bebbia 480
Bechium 167
Bedfordia 224
Index to Scientific Names
Behen 157
Bejaranoa 539
Bellida 258
B. graminea 254
Bellideae 303
Bellidiastrum 317
Bellidinae 303
Bellieae 303
Bellis 303
Bellium 303
Belloa 258
Bembycodium 350
Benitoa 329
Berardia 130
Berenice 39
Berkheya 205
B. horrida 205
Berkheyopsis 206
Berlandiera 462
B. lyrata 462
Berroa 258
Berthelotia 387
Berylsimpsonia 100
Bethencourtia 229
Bidens 411
Bidentideae 406
Bielzia 141
Bigelowia 322
Bipontia 160
Bishopalea 163
Bishopanthus 177
Bishopiella 545
Bishovia 565
Blainvillea 449
Blakeanthus 523
Blakiella 295
Blanchetia 158
B. heterotricha 150
Blaxium 245
Blennosperma 215
Blennospora 258
Blepharipappus 500
Blepharispermum 398
Blepharizonia 501
Blumea 377
Blumeopsis 389
Boeberastrum 425
Boeberoides 425
Bojeria 382
Bolandia 229
Bolanosa 152
Bolocephalus 138
Boltonia 328
Boltoniinae 327
Bombycilaena 258
Boopis 24
B. bupleuroides 24
Borkonstia 319
Borrichia 462
Bothriocline 167
Brachanthemum 357
Brachionostylum 225
Brachyactis 335
Brachychaeta 325
Brachyclados 110
Brachycodon 42
Brachycodonia 42
Brachycome 302
Brachyglottis 223
Brachylaena 119
Brachymeris 356
Brachyris 321
B. sect. Amphiachyris 321
Brachyscome 302
Brachyscominae 302
Brachythrix 167
Bracteantha 283
Brasilia 419
Breea 132
Brenandendron 173
Brickellia 553
Brickelliastrum 554
Brighamia 52
Brintonia 325
Brocchia 367
Broteroa 127
Brunonia 594
B. australis 590, 594
Brunoniaceae 589
Bryomorphe 258
B. aretioides 246
Bubonium 382
Buphthalmum 382
Burkartia 103
Burmeistera 51
B. virescens 52
Caatinganthus 164
Cabereriella 235
Cabobanthus 167
Cacalia 240
Cacaliopsis 222
C. nardosmia 208
Cacosmia 177
Cadiscus 237
Caesulia 377
Calanticaria 466
Calcaratolobelia 47
Calcitrapa 146
Calea 419
C. crocinervosa 419
Calendula 243
C. arvensis 242
C. maderensis 242
Calenduleae 241
Callicephalus 143
Callilepis 259
Callistemma 318
Callistephus 318
Calocephalus 259
Calomeria 259
Calopappus 101
Calostephane 384
Calotesta 259
Calotis 303
Calycadenia 501
Calycera 23
Calyceraceae 3, 19
623
Calycocorsus 186
Calycoseris 193
Calyptocarpus 449
Camchaya 172
Campanula 41
C. edulis 42
Campanulaceae 3, 26
Campanulastrum 42
Campanuloideae 37
Campovassouria 547
Camptacra 313
Campuloclinium 540
Canadanthus 334
Canarina 38
Canariothamnus 229
Cancrinia 360
C. group 360
Cancriniella 360
Candidea 172
Capelio 225
Cardopatiinae 127
Cardopatium 127
C. corymbosum 128
Cardueae 123
Carduinae 129
Carduncellus 145
Carduoideae 78
Carduus 132
C. group 132
Carelia 522
Carlina 126
C. acaulis 126
Carlininae 126
Carlinoides 205
Carlquistia 501
Carminatia 557
Carpesium 383
Carphephorus 531
Carphobolus 159
Carphochaete 527
Carpodetaceae 3, 57
Carpodetus 59
C. serratus 59
Carramboa 482
Carterothamnus 513
Cartesia 165
Carthamus 144
C. group 144
Cassinia 259
Castalis 245
Castanedia 569
Castrilanthemum 367
Castroviejoa 284
Catamixis 119
C. group 119
Catananche 183
Catananchinae 182
Catatia 259
Catolesia 542
Caucasalia 241
Cavalcantia 524
Cavea 146
Caxamarca 234
Celmisia 225, 296
624
Cenia 351
Cenocline 368
Centaurea 141, 146
C. benedicta 146
C. group 146
C. prolongi 124
Centaureinae 138
Centaurodendron 141
Centauropsis 173
Centaurothamnus 143
Centipeda 399
C. cunninghamii 400
C. elatinoides 400
C. minima 400
C. nidiformis 400
C. thespidioides 400
Centipedinae 399
Centrapalinae 171
Centrapalus 172
Centratherinae 160
Centratherum 160
Centromadia 502
Centropappus 224
Centropogon 51
Cephalanophlos 132
Cephalipterum 259
Cephalobembix 438
Cephalopappus 107
C. sonchifolius 107
Cephalorrhynchus 187
Cephalosorus 260
Cephalostigma 38
Ceratogyne 303
Ceruana 305
Chacoa 566
Chaenactideae 392, 431
Chaenactis 432
C. douglasii 432
Chaetadelpha 192
Chaetanthera 110
Chaetopappa 335
Chaetopappinae 335
Chaetoseris 187
Chaetymenia 436
Chamaechaenactis 436
Chamaegeron 291
Chamaeleon 126
Chamaemelum 370
C. group 370
Chamaepus 260
Chamartemisia 360
Chamomilla 368
Chaptalia 114
C. chapadensis 114
Charadranaetes 235
Chardinia 131
Chartolepis 146
Cheirolepis 146
Cheirolophus 140
C. sempervirens 124
Chersodoma 216
Chevreulia 260
Chihuahuana 322
Chiliadenus 379
Index to Scientific Names
Chiliocephalum 260
Chiliophyllum 296
Chiliotrichiopsis 296
C. peruviana 296
Chiliotrichum 297
Chimantea 99
Chionolaena 260
Chionopappus 177
Chlamydophora 372
Chlamysperma 508
Chloracantha 328
Chondrilla 184
Chondropyxis 260
Chorisis 185
Chorisiva 445
Chresta 160
Chrestinae 160
Chromolaena 533
C. morii 533
Chromolepidinae 441, 446
Chromolepis 446
Chronopappus 161
Chrysactinia 425
Chrysactinium 179
C. acaule 175
C. wurdackii 179
Chrysanthellinae 407
Chrysanthellum 408
C. filiforme 408
Chrysanthemoides 244
Chrysanthemum 357, 369
Chrysanthoglossum 372
Chrysocephalum 260
C. semicalvum 254
Chrysocoma 291
Chrysocoryne 266
Chrysogonum 463
Chrysolaena 154
Chrysoma 322
Chrysopappus 146
Chrysophthalmum 383
Chrysopsidinae 337
Chrysopsis 337
Chrysothamnus 322
Chthonocephalus 261
Chucoa 113
Chuquiraga 90
C. “anomale” 88
C. erinaceae 89
C. sect. Erinesa 90
C. sect. Gymnophoranta 88
Cicerbita 187
Ciceronia 563
Cichorieae 180
Cichoriinae 182
Cichorioideae 78
Cichorium 182
C. intybus 182
Cineraria 229
Cirsium 132
C. eriophorum 133
Cissampelopsis 240
Cladanthus 370
Cladochaeta 261
Clairvillea 177
Clappia 422
Clappiinae 421
Clermontia 53
Clibadiinae 446
Clibadium 449
Cloiselia 121
Cnicothamnus 111
Cnicus 146
Codonocephalum 382
Codonopsis 37
Coespeletia 483
Coleocoma 388
Coleostephus 372
Colobanthera 305
Cololobus 154
Columbiadoria 322
Columellea 273
Colymbada 146
Comaclinium 425
Comborhiza 261
Commidendrum 286
Complaya 457
Compositae 3, 61
Comptonanthus 268
Condylidium 551
Condylopodium 559
Conocliniopsis 539
C. prasiifolia 538
Conoclinium 541
Constancea 495
Conyza 340
Conyzella 340
Conyzinae 339
Coopernookia 595
Corellia 510
Coreocarpus 411
Coreopsidaceae 406
Coreopsideae 392, 406
Coreopsidinae 409
Coreopsis 411
C. petrophiloides 411
Corethamnium 569
Corethrogyne 330
Corokia 16
Corokiaceae 13
Corymbieae 207
Corymbium 207
C. villosum 207
Cosmos 412
Cota 366
Cotula 351
Cotuleae 342
Cotulina 368
Coulterella 422
Coulterellinae 422
Cousinia 136
Cousiniopsis 128
Crantzia 160
Craspedia 261
Crassina 476
Crassocephalum 232
Craterocapsa 39
Cratystylis 385
Index to Scientific Names
Cremanthodium 218
Cremnothamnus 261
Crepidiastrum 184
Crepidinae 183
Crepis 184
C. biennis 185
Crinitaria 318
Criscia 106
Crispiloba 10
C. disperma 9
Critonia 562
Critoniadelphus 562
Critoniella 567
Critoniinae 559
Critoniopsis 158
Crocidium 215
Crociseris 231
Crockeria 496
Crocodylium 145
C. group 145
C. syriacum 145
Cronquistia 527
Cronquistianthus 570
Croptilon 338
Crossolepis 266
Crossostephium 359
Crossothamnus 559
Crupina 139
Cryptocodon 44
Cryptogyne 352
Crystallopollen 170
Ctenosperma 351
Cuatrecasanthus 159
Cuatrecasasiella 261
Cuchumatanea 484
Culcitium 235
Cullumia 205
Cuniculotinus 342
Curio 231
Cuspidia 205
Cuttsia 59
C. viburnea 59
Cyananthus 37
Cyanea 53
Cyanopsis 141, 167
Cyanthillium 167
Cyanus 146
Cyathocline 305
Cyathomone 412
Cyathopappus 266
Cyclachaena 445
Cyclocodon 38
Cyclolepis 112
Cylindrocarpa 45
Cylindrocline 385
Cymbolaena 262
Cymbonotus 204
Cymbopappus 354
Cymophora 481
Cynara 132
Cynareae 123
Cyphia 54
C. erecta 54
Cyphioideae 53
Cyphocarpoideae 53
Cyphocarpus 53
C. psammophilus 54
Cyrtocymura 155
C. scorpioides 155
Cyrtolepis 364
Dacryotrichia 305
Dahlia 412
Damnamenia 297
Damnxanthodium 449
Dampiera 595
D. sp. 591
Darwiniothamnus 340
Dasyandantha 159
Dasyanthina 155
Dasycondylus 538
Dasyphyllum 90
Dauresia 240
Daveaua 371
Decachaeta 573
Decaneurum 170, 173, 363
Decastylocarpus 168
Decazesia 262
Deinandra 502
Delairea 240
Delamerea 388
Delilia 450
Delissea 53
Dendranthema 357
Dendrocacalia 217
Dendrophorbium 233
Dendrosenecio 236
D. keniensis 208
Dendroseris 190
Denekia 262
Desmanthodiinae 479
Desmanthodium 480
D. fruticosum 480
Detris 292
Dewildemania 168
Diacantha 89
Diacranthera 538
Dialesta 159
Dialypetalum 47
Dianthoseris 184
Diaphractanthus 168
Diaspananthus 123
Diaspasis 595
Diastatea 49
Dibothrospermum 367
Dicercoclados 217
Dichaetophora 336
Dichrocephala 305
Dichromochlamys 313
Dicoma 121
D. group 120
Dicomeae 120
Dicoria 444
Dicranocarpus 413
Didelta 206
Dielitzia 262
Dielsantha 48
Dieteria 330
625
Digitacalia 220
Dillandia 177
D. perfoliata 175
Dimeresia 432
Dimerostemma 450
Dimorphocoma 313
Dimorpholepis 283, 548
Dimorphotheca 245
D. caulescens 245
D. cuneata 245
D. dregei 245
D. ecklonis 245
D. fruticosa 245
D. jucunda 245
D. montana 245
D. nudicaulis 245
D. polyptera 245
D. scabra 245
D. walliana 245
D. zeyheri 245
Dinoseris 96
Diodontium 408
Diosphaera 42
Diotis 365
Diplopappus 337
Diplostephium 297
Dipterocome 147
Dipterocypsela 155
Disparago 262
Dissothrix 557
Distasis 335
Distephanus 173
Distreptus 164
Disynaphia 547
D. praeficta 547
Disynaphiinae 546
Dithyrostegia 262
Dittrichia 380
Doellia 390
Doellingeria 286
Dolichoglottis 225
Dolichorrhiza 241
Dolichothrix 262
Doliclasium 105
Dolomiaea 138
Dominella 50
Donatia 617
D. novae-zelandiae 615
Donatiaceae 614
Doniophyton 87
D. anomalum 88
Dorobaea 230
Doronicum 215
Downingia 50
Dracopis 471
Dresslerothamnus 234
Dubautia 502
Dubyaea 185
Dugesia 446
Dugesiinae 441, 446
Duhaldea 377
Duidaea 97
Duseniella 88
D. patagonica 88
626
Dymondia 204
Dyscritogyne 556
Dyscritothamninae 480
Dyscritothamnus 481
Dyssodia 426
D. sect. Acyphyllaea 429
D. sect. Aurantiacae 429
D. sect. Boeberastrum 425
D. sect. Boeberoides 425
D. papposa 426
Dyssodiopsis 426
Eastwoodia 323
Eatonella 497
Echinacea 475
Echinocodon 38, 42
Echinocodonia 42
Echinocoryne 155
Echinops 128
E. ritro 129
Echinopsinae 128
Eclipta 450
E. prostrata 450
Ecliptinae 441, 446
Edmondia 263
Edraianthus 41
Eenia 397
Egletes 306
Eirmocephala 155
Eitenia 534
Ekmania 159
Ekmaniopappus 226
Elachanthemum 357
Elachanthus 313
Elachopappus 273
Elaphandra 451
Elcisimia 296
Elekmania 227
Elephantopinae 163
Elephantopus 164
Elephantosis 164
Eleutheranthera 451
Ellenbergia 524
Elvira 450
Elytropappus 263
Embergeria 191
Emilia 238
Emiliella 238
Encelia 460
Enceliinae 441, 460
Enceliopsis 460
Endocellion 216
Endopappus 371
Engelmannia 463
Engelmanniinae 441, 461
Engleria 292
Enydra 418
Enydrinae 418
Epallage 397
Epaltes 386
Epilasia 198
Episcothamnus 162
Epitriche 263
Erato 179
Index to Scientific Names
Erechtites 232
Eremanthus 161
E. sect. Pycnocephalum 160
Eremiastrum 336
Eremohylema 387
Eremosis 158
Eremothamnus 202
Eriachaenium 108
E. magellanicum 104
Ericameria 287
Ericentrodea 413
Ericopsis 594
Erigeron 340
Eriocarpum 333
Eriocephalus 352
Eriochlamys 263
Eriophyllum 495
Eriotrix 239
Erlangea 168
E. sect. Platylepis 168
Erlangeinae 165
Erodiophyllum 306
Eroeda 274
Erymophyllum 263
Eryngiophyllum 408
Erythradenia 573
Erythrocephalum 121
Espejoa 436
Espeletia 483
Espeletiinae 482
Espeletiopsis 483
Ethulia 168
Ethuliopsis 386
Eucephalus 287
Euchiton 263
Eumorphia 355
Eupatoriadelphus 528
Eupatoriastrum 564
Eupatorieae 392, 441, 510
Eupatoriinae 528
Eupatorina 563
Eupatoriopsis 534
Eupatorium 528
E. sect. Austrobrickellia 557
E. sect. Austroeupatorium 529
E. sect. Campovassouria 547
E. sect. Campuloclinium 540
E. sect. Chromolaena 533
E. sect. Cylindrocephala 533
E. sect. Dimorpholepis 548
E. sect. Dysinaphia 547
E. sect. Gyptis 530, 537, 569, 571
E. sect. Hebeclinium 572
E. sect. Heterocondylus 550, 571
E. sect. Laevia 564, 566
E. sect. Macropodina 540, 541
E. sect. Microstemon 520
E. sect. Osmia 533
E. sect. Praxelis 533
E. sect. Raulinoreitzia 546
E. sect. Sphaereupatorium 565
E. sect. Steyermarkina 570
E. sect. Symphyopappus 548
E. sect. Urolepis 537
Euphrosyne 445
Eurybia 287
Eurydochus 98
Euryops 236
Eutetras 509
Euthamia 323
Evacidium 264
Evacopsis 265
Evax 265
E. sect. Hesperevax 267
Ewartia 264
E. catipes 252
Ewartiothamnus 264
Exomiocarpon 451
Eyrea 387
Fabera 196
Faberia 189
Faberiopsis 188
Facelis 264
Farfugium 216
Faujasia 239
Faujasiopsis 239
Fauria 603
Favratia 42
Faxonia 485
Feddea 390
Fedorovia 43
Feeria 41
Feldstonia 264
Felicia 292
Feliciinae 290
Femeniasia 145
Fenixia 451
Ferreyranthus 177
Ferreyrella 525
Filaginella 266
Filaginopsis 265
Filago 264
Filifolium 359
Fitchia 413
Fitchiinae 409
Fitzwillia 265
Flaveria 422
Flaveriinae 422
Fleischmannia 520
Fleischmanniinae 520
Fleischmanniopsis 563
Florestina 436
Floscaldasia 297
Flosmutisia 297
Flotovia 90
Flotowia 90
Flourensia 461
Flustula 156
Flyriella 554
Fontquera 379
Forstera 618
F. bidwillii 615
Foveolina 354
Fradinia 370
Francoeuria 379
Franseria 444
Freya 485
Index to Scientific Names
Frolovia 138
Fulcaldea 88
Gadellia 42
Gaillardia 401
Gaillardiinae 401
Galactites 133
Galatella 318
Galeana 508
G. pratensis 508
Galeaninae 508
Galeatella 47
Galeomma 265
Galinsoga 485
Galinsoginae 483
Gamocarpha 23
Gamochaeta 265
Gamochaetopsis 265
Gamolepis 236, 238
Garberia 530
Garcibarrigoa 234
Gardnerina 524
Garhadiolus 196
Garuleum 245
G. bipinnatum 242
Gastrosulum 367
Gazania 206
Geigeria 384
Geissolepis 337
Gelasia 198
Geraea 461
Gerbera 116
Gerberinae 113
Geropogon 198
Gibbaria 244
G. scabra 242
Gifola 265
Gilberta 265
G. tenuifolia 254
Gilruthia 265
Githopsis 44
Gladiopappus 122
Glaziovianthus 160
Glebionis 369
Glossanthis 361
Glossarion 97
Glossocardia 409
Glossopappus 372
Glyptopleura 194
Gnaphalieae 246
Gnaphaliothamnus 266
Gnaphalium 266
G. subg. Achyrocline 252
G. sect. Eu-Gnaphalium 266
G. sect. Euchiton 263
G. sect. Gamochaeta 265
G. supinum 246
G. uliginosum 256
Gnaphalodes 252
Gnaphalon 276
Gnephosis 266
G. eriocarpa 254
G. uniflora 271
Gnomophalium 266
Gochnatia 117
G. floribunda 117
G. subg. Nardophyllum 299
Gochnatiinae 116
Goldmanella 413
Goldmania 413
Golionema 332
Gongrostylus 550
Gongrothamnus 173
Gongylolepis 98
Goniocaulon 141
Gonospermum 366
Goodenia 595
G. macroplecta 596
Goodeniaceae 3, 589
Goodenovieae 589
Gorceixia 152
Gorteria 206
Gorteriinae 204
Gossweilera 170
Goyazianthus 558
Graciela 429
Grammatotheca 48
Grangea 306
Grangeinae 304
Grangeopsis 306
Grantia 380
Graphistylis 233
Gratwickia 266
Grauanthus 306
Grazielia 548
Greenella 323
Greenmaniella 419
Grindelia 330
Grisebachianthus 565
Grossheimia 146
Grosvenoria 569
Guaicaia 97
Guardiola 486
G. mexicana 487
G. tulocarpus 487
Guardiolinae 486
Guayania 573
Guevaria 525
Guizotia 489
Gundelia 200
Gundelieae 199
Gundlachia 323
Gunillaea 40
Gutenbergia 168
Gutierrezia 323
Guynesomia 298
Gymnantheminae 173
Gymnanthemum 173
Gymnarrhena 147
Gymnarrheneae 147
Gymnaster 319
Gymnocline 366
Gymnocondylus 551
Gymnocoronis 519
Gymnodiscus 237
Gymnogyne 351
Gymnolaena 426
Gymnolomia 451
627
Gymnopentzia 355
Gymnosperma 324
Gymnostephium 292
Gymnostyles 352
Gynoxys 223
G. sect. Praegynoxys 223
Gynura 232
Gypothamnium 112
Gyptidinae 535
Gyptidium 537
Gyptis 537
G. group 537
Gyrodoma 307
Gyrostephium 280
Haarera 168
Haastia 224
Haeckeria 267
Haegiela 267
Halacsyella 41
Hanabusaya 43
Handelia 361
H. group 361
Haplocarpha 204
Haploësthes 422
Haplopappus 330
H. sect. Inulopsis 314
H. sect. Leucopsis 338
Haplostephium 162
Haptotrichion 267
H. conicum 254
Harleya 156
Harmonia 503
Harnackia 427
Harthamnus 111
Hartwrightia 532
Hasteola 230
Hatschbachiella 530
Hazardia 331
Hebecliniinae 572
Hebeclinium 572
Hecastocleideae 118
Hecastocleis 118
H. group 118
Hedosyne 445
Hedypnois 196
Helenieae 392, 400
Heleniinae 401
Helenium 402
H. mexicanum 402
Heliantheae 392, 440
Helianthella 461
Helianthinae 441, 464
Helianthopsis 467
Helianthus 466
H. laciniatus 466
Helichrysopsis 267
Helichrysum 267
H. adenophorum 254
H. aureum 254
H. calvertianum 254
H. depressum 252
H. lanceolatum 252
H. sect. Lawrencella 269
628
H. sect. Leontopodioides 270
H. miconiifolium 254
H. milforidiae 254
H. sect. Ozothamnus 275
H. subg. Ozothamnus 275
H. sect. Rhodanthe 279
Heliocauta 365
Heliomeris 467
Heliopsis 475
Helipterum 281
H. sect. Leucochrysum 270
H. sect. Pachypterum 268
Helminthotheca 196
Helogyne 559
Hemibaccharis 311
Hemipappus 366
Hemisphaera 42
Hemistepta 137
Hemizonella 503
Hemizonia 503
H. subg. Blepharizonia 501
H. [unranked] Hemizonella 503
Henricksonia 413
Heptanthinae 418
Heptanthus 418
Herderia 168
Herodotia 226
Herreranthus 227
Herrickia 287, 342
Hertia 237
Hesperastrum 331
Hesperevax 267
Hesperodoria 324
Hesperomannia 174
Heteracia 185
Heteractis 292
Heteranthemis 370
Heterochaenia 39
Heterocodon 44
Heterocoma 163
Heterocondylus 550
Heterocypsela 156
Heteroderis 185
Heterolepis 202
Heteromera 371
Heteromma 307
Heteropappus 318
Heteroplexis 287
Heterorhachis 206
Heterosperma 414
H. pinnatum 414
Heterothalaminae 310
Heterothalamus 311
Heterotheca 338
Heterotoma 50
H. lobelioides 51
Hexinia 190
Hidalgoa 414
Hieraciinae 194
Hieracium 194
H. umbellatum 195
Hilliardia 352
Hilliardiella 168
Himalaiella 137
Index to Scientific Names
Hinterhubera 298
Hinterhuberinae 294
Hippia 353
Hippobroma 50
Hippolytia 362
Hirpicium 206
Hirschia 380
Hirtellina 130
Hispidella 194
Hochstetteria 121
Hoehnelia 168
Hoehneophytum 232
Hoffmanniella 452
Hofmeisteria 513
Hofmeisteriinae 513
Holocarpha 504
Holocheilus 106
Hololeion 195
Hololepis 163
Holophyllum 350
Holoschkuhria 437
Holozonia 504
Homochaete 272
Homochroma 293
Homochrominae 290
Homocodon 44
Homognaphalium 266, 266
Homogyne 216
Homopappus 332
Hoplophyllum 202
Howellia 50
Huarpea 89
Huberopappus 159
Hubertia 239
Hughesia 570
Hullsia 342
Hulsea 497
Hulseinae 497
Hulteniella 358
Humbertacalia 239
Humea 259
H. sect. Haeckeria 267
Humeocline 268
Hyalaea 146
Hyalis 113
Hyalochaete 137
Hyalochlamys 268
Hyalolepis 273
Hyaloseris 96
Hyalosperma 268
H. cotula 254
Hybridella 474
Hydrodyssodia 427
Hydroidea 268
Hydropectis 427
Hymenocephalus 140
Hymenoclea 444
Hymenolepis 351
Hymenonema 183
Hymenopappus 437
Hymenostemma 367
Hymenostephium 467
Hymenothrix 437
Hymenoxys 404
Hyoseris 196
Hypacanthium 136
Hypericophyllum 437
Hypochaeridinae 195
Hypochaeris 196
Hypsela 47
Hyssaria 42
Hysterionica 341
Hystrichophora 169
Ianthopappus 112
Ichthyothere 490
Idiopappus 452
Idiothamnus 566
Ifloga 268
Ighermia 382
Ignurbia 228
Iltisia 528
Imeria 569
Inezia 353
Inula 382
I. sect. Cappa 377
I. sect. Chrysopsis 337
Inulanthera 353
Inuleae 374
Inuloides 244
I. tomentosa 242
Inulopsis 314
Io 232
I. ambondrombeensis 208
Iocaste 356
Iocenes 232
Iodocephalis 169
Iogeton 452
Ionactis 288
Iostephane 467
Iotasperma 314
Iphiona 380
I. anthemidifolia 380
Iphionopsis 385
Iranecio 241
Irwinia 159
Ischnea 215
Ismelia 370
Isocarpha 552
Isocoma 331
Isoetopsis 314
Isonema 167
Isopappus 338
Isostigma 409
Isotoma 49
Iva 445
Ixeridium 185
Ixeris 185
Ixiochlamys 314
Ixiolaena 268
Ixodia 268
Jacea 146
Jacmaia 228
J. incana 208
Jacobaea 231
Jacobaeastrum 236
Jaegeria 487
Index to Scientific Names
Jaegeriinae 487
Jalcophila 269
Jaliscoa 515
Jamesianthus 430
Jaramilloa 516
Jasione 41
Jasionella 41
Jasonia 379
Jaumea 423
Jaumeinae 423
Jefea 452
Jeffreya 292
Jensia 504
Jessea 228
Joannea 90
Joannesia 90
Johannia 90
Joseanthus 159
Jungia 104
J. woodii 104
Jurinea 137
J. pjatajeviae 137
Jurinella 137
Kalimeris 318
Kanimia 517
Karelinia 387
Karvandarina 142
Kaschgaria 362
Kastnera 180
Kaunia 516
Kemulariella 319
Kentrophyllum 144
Keumkangsania 43
Keysseria 308
Kinghamia 169
Kingianthus 452
Kippistia 314
Kirkianella 191
Klasea 144
Kleinia 231
K. longiflora 208
Klenzea 257
Koanophyllon 564
K. adamantium 564
Koehneola 416
Koelpinia 198
Koyamacalia 217
Koyamasia 169
Kremeria 372
Krigia 191
Krylovia 319
Kuhniinae 552
Kyhosia 504
Kyrsteniopsis 556
Lachanodes 225
Lachnophyllum 292
Lachnorhiza 172
Lachnospermum 269
Lactuca 188
L. saligna 188
Lactuceae 180
Lactucella 188
Lactucinae 187
Lactucosonchus 191
Laennecia 315
Laestadia 298
Lagascea 467
Lagasceinae 464
Lagedium 188
Lagenifera 308
Lagenithrix 309
Lagenocypsela 309
Lagenopappus 309
Lagenophora 308
Lagenophorinae 307
Laggera 389
Lagophylla 505
Lagoseriopsis 186
Lamprachaenium 170
Lamprocephalus 236
Lampropappus 174
Lamyra 133
Lamyropappus 135
Lamyropsis 134
Lancisia 351
Landtia 204
Langebergia 269
Lantanopsis 453
Lapsana 186
Lapsanastrum 186
Lasianthaea 453
Lasiocephalus 234, 234
Lasiocoma 236
Lasiolaena 543
L. morii 544
Lasiopogon 269
Lasiospermum 353
Lasiospora 198
Lasthenia 496
Latouria 594
Launaea 190
Lawrencella 269
L. davenportii 254, 271
Layia 505
Lechenaultia 594
L. sp. 591
Lecocarpus 488
Legenere 50
Legousia 44
Leibnitzia 115
Leiboldia 152
Leiboldiinae 152
Leiocarpa 284
Lembertia 496
Lemooria 269
Leonis 226
Leontodon 197
Leontonyx 267
Leontopodium 269
L. alpinum 256
Lepia 476
Lepidanthus 368
Lepidaploa 156
L. salzmannii 150
Lepidella 171
Lepidesmia 552
629
Lepidolopha 363
Lepidolopsis 361
Lepidonia 153
Lepidophorum 372
Lepidophyllum 298
Lepidospartum 222
Lepidostephium 270
Lepidotheca 368
Leptilon 340
Leptinella 352
Leptocarpha 453
Leptoclinium 558
Leptophytus 271
Leptorhynchos 270
Leptostelma 341
Leptotriche 270
Lescaillea 427
Lessingia 331
Lessingianthus 156
Leucactinia 427
Leucanthemella 363
Leucanthemopsis 367
L. group 367
Leucanthemum 373
L. group 372
Leucheria 104
Leuciva 445
Leucoblepharis 399
Leucochrysum 270
L. albicans 246
L. stipitatum 254
Leucocyclus 365
Leucogenes 270
L. grandiceps 246
Leucoglossum 373
Leucomeris 119
Leucophyta 270
Leucopsis 338
Leucoptera 354
Leunisia 106
Leuzea 143
Levenhookia 618
L. stipitata 615
Leysera 271
Liabeae 175
Liabellum 178
Liabinae 177
Liabum 178
L. subg. Chrysastrum 180
L. sect. Paranephelius 179
Liatrinae 530
Liatris 531
Libanothamnus 483
Lidbeckia 353
Lifago 381
Lightfootia 38
Ligularia 218
Ligulariopsis 218
Limbarda 381
Limnanthemum 603
Lindheimera 463
Linosyris 318
Linzia 172
Liparophyllum 602
630
Lipochaeta 453
Lipschitziella 137
Lipskyella 136
Litogyne 386
Litothamnus 545
L. group 545
Litrisa 532
Llerasia 298
Lobelia 47
L. welwitschii 48
Lobelioideae 47
Logfia 271
L. gallica 256
Lomatozona 534
Lonas 373
Lopholaena 237
Lophopappus 101
Lorandersonia 342
Lordhowea 228
L. insularis 208
Lorentzianthus 566
Loricaria 271
L. leptothamna 271
L. thuyoides 271
Lourteigia 541
Loxothysanus 437
Lucilia 272
L. acutifolia 254
Luciliocline 272
Luciliopsis 110
Lugoa 366
Luina 222
Lulia 113
Lundellianthus 453
Lundinia 227
Luteidiscus 316
Lycapsinae 509
Lycapsus 509
Lychnocephalus 162
Lychnophora 162
Lychnophorinae 161
Lychnophoriopsis 162
Lycoseris 111
Lygodesmia 192
Lyonnetia 365
Lysichlamys 236
Lysipomia 50
Lysistemma 166
Machaeranthera 331
M. sect. Arida 329
M. subg. Dieteria 330
M. subg. Sideranthus 333
Machaerantherinae 328
Machlis 351
Macledium 121
Macowania 272
M. hamata 254
Macrachaenium 102
Macroclinidium 123
Macropodina 541
Macvaughiella 528
Madagaster 299
Madaractis 231
Index to Scientific Names
Madia 505
Madieae 392, 441, 492
Madiinae 497
Mairia 299
Malacothricinae 193
Malacothrix 194
Mallotopus 494
Malmeanthus 569
Malperia 555
Mandonia 481
Mantisalca 141
Manyonia 156
Marasmodes 354
Marcelia 370
Marshallia 402
Marshalliinae 402
Marshalljohnstonia 192
Marticorenia 106
Maruta 365
Matamoria 164
Matricaria 368
Mattfeldanthus 156
Mattfeldia 226
Matudina 574
Mauranthemum 373
Mausolea 359
Mecomischus 370
Medranoa 324
Megaliabum 178
Megalodonta 414
Melampodiinae 487
Melampodium 488
M. dicoelocarpum 488
M. divaricatum 488
M. nutans 488
M. perfoliatum 488
M. sericeum 488
Melanodendron 288
Melanoloma 146
Melanthera 454
Memecylanthus 11
Menyanthaceae 3, 599
Menyanthes 602
M. trifoliata 603
Merciera 40
Merrittia 378
Mesadenia 220
Mesanthophora 169
Mesogramma 229
Metalasia 272
Metastevia 527
Mexerion 272
Mexianthus 566
Michauxia 46
Micractis 490
Microcephala 363
Microchaete 233
Microcodon 40
Microglossa 288
Microgyne 315
Microgynella 315
Microliabum 178
M. humile 175
Microlonchus 142
Micropsis 272
Micropus 273
M. sect. Bombycilaena 258
Microseridinae 191
Microseris 191
Microspermum 528
Mikania 517
M. grazielae 517
Mikaniinae 516
Mikaniopsis 240
Millefolium 364
Milleria 490
M. quinqueflora 490
Millerieae 392, 441, 477
Milleriinae 488
Millotia 273
Minasia 162
Minuria 315
Miricacalia 217
Misbrookea 235
Miyamayomena 319
Mniodes 273
M. pulvinulata 246
Modestia 137
Molina 311
Mollera 384
Monactis 454
Monanthemum 159
Monarrhenus 388
Monenteles 385
Monochlaena 352
Monoculus 243
M. monstrosus 242
Monogereion 551
Monolopia 496
Monopsis 49
Monoptera 369
Monoptilon 336
Monosis 173
M. sect. Eremosis 158
Montanoa 470
Montanoinae 441, 469
Monticalia 233
Moonia 415
Moquinia 148
M. racemosa 149
Moquinieae 148
Morithamnus 546
Morysia 350
Moscharia 107
Msuata 169
Mtonia 307
Muehlbergella 43
Mulgedium 188
Munnozia 180
M. subg. Erato 179
M. subg. Kastnera 180
M. tenera 175
Munnoziinae 179
Munzothamnus 193
Muschleria 169
Musschia 41
Mutisia 108
M. subspinosa 109
Index to Scientific Names
Mutisieae 90
Mutisiinae 108
Mutisioideae 77
Myanmaria 174
Mycelis 188
Myconella 372
Myconia 372
Myopordon 143
Myriactis 309
Myriocephalus 273
Myriogyne 399
Myripnois 123
Myxopappus 355
Mzymtella 42
Nabalus 188
Nablonium 253
Namacodon 39
Nananthea 366
Nannoglottis 288
Nannoseris 184
Nanothamnus 390
Nardophyllum 299
Nardosmia 216
Narvalina 415
Nassauvia 101
N. dentata 101
Nassauviinae 100
Nauplius 382
Neblinaea 98
Neesia 365
Neja 341
Nelsonianthus 221
Nemacladoideae 46
Nemacladus 46
N. rigidus 46
Nemosenecio 218
N. yunnanensis 219
Neocabreria 571
Neocodon 42
Neocuatrecasia 540
Neogoodenia 595
Neohintonia 567
Neojeffreya 388
Neomirandea 574
Neomirandeinae 574
Neomolina 311
Neonesomia 324
Neopallasia 359
Neothymopsis 439
Neotysonia 273
Neowimmeria 47
Nephrophyllidium 603
Nephrotheca 245
N. ilicifolia 242
Nesampelos 226
Nesocodon 38
Nesomia 525
Nestlera 273
Nestotus 342
Neurolaena 420
Neurolaeneae 392, 417
Neurolaeninae 419
Neurolakis 172
Newtonia 173
Niclouxia 381
Nicolasia 389
Nicolletia 428
Nidorella 307
Nigromnia 595
Nikitinia 144
Nipponanthemum 363
Nivellea 374
Nolletia 293
Nordenstamia 223
Norlindhia 243
N. amplectens 242
N. aptera 242
N. breviradiata 242
Nothobaccharis 565
Nothocalais 191
Noticastrum 338
Notobasis 134
Notonia 231
Notoniopsis 231
Notoseris 188
Nouelia 119
N. group 118
Novaguinea 342
Novenia 299
Nymphoides 603
Oaxacania 513
Oaxacaniinae 513
Oblivia 454
Ochrocephala 143
Oclemena 288
Odixia 274
O. achlaena 274
Odontocline 227
O. tercentenariae 208
Odontoloma 159
Odontospermum 382
Odontotrichum 219
Oedera 274
O. genistifolia 274
Oglifa 271
Oiospermum 161
Oldenburgia 120
Oldfeltia 227
Olearia 299
Olgaea 134
Oligactis 178, 325
Oligandra 272
Oliganthes 174
Oligocarpus 243
O. calendulaceus 242
Oligochaeta 143
Oligodora 351
Oligonema 332
Oligoneuron 324
Oligosporus 358
Oligothrix 238
Olivaea 332
Omalanthus 366
Omalotes 366
Omalotheca 266
Omphalopappus 170
631
Oncosiphon 355
Ondetia 385
Onopordum 134
O. group 134
Onoseris 112
Oocephala 170
Oonopsis 332
Oparanthus 415
Ophryosporus 570
Opisthopappus 363
Orbivestus 170
Oreastrum 289
Oreochrysum 325
Oreoleysera 274
Oreostemma 289
Oreostylidium 618
O. subulatum 615
Oresbia 236
Orithrophium 300
Ormenis 370
Orochaenactis 433
Orthopappus 164
Osbertia 338
Osmadenia 506
Osmiopsis 535
Osmitopsis 353
Osteospermum 244
O. sect. Blaxium 245
O. burttianum 244
O. ciliatum 242
O. grandidentatum 242
O. herbaceum 242
Ostrowskia 38
Otanthus 365
Oteiza 485
Othonna 237
O. brandbergensis 208
Othonnopsis 237
Otocarpum 374
Otochlamys 351
Otopappus 454
Otospermum 374
Outreya 137
Oxycarpha 472
Oxylaena 275
Oxylobinae 513
Oxylobus 514
Oxypappus 430
Oxyphyllum 102
Oxytenia 445
Oyedaea 455
Ozothamnus 275
O. bidwillii 254
O. lepidophyllus 254
O. leptophyllus 252
Pachydiscus 11
Pachylaena 109
P. atriplicifolia 110
Pachystegia 300
Pachythamnus 515
Pacifigeron 300
Packera 230
Pacourina 165
632
Pacourininae 165
Palafoxia 438
Paleaepappus 300
Paleocyanus 140
Pallenis 382
Palmerella 49
Pamphalea 107
Panaetia 276
Pappobolus 467
Pappochroma 309
Pappothrix 510
Papuacalia 225
Paracalia 222
Parachionolaena 260
Parafaujasia 239
Paragynoxys 222
Paraixeris 184
Paralychnophora 161
Paramicrorhynchus 190
Paramiflos 483
Paranepheliinae 178
Paranephelius 179
Parantennaria 275
Parapiqueria 552
Paraprenanthes 188
Parasenecio 217
Parastrephia 301
Parishella 47
Parthenice 445
Parthenium 445
Pasaccardoa 120
Pascalia 455
Paurolepis 170
Pechuel-loeschea 386
Pectidinae 423
Pectis 428
P. decemcarinata 428
Pegolettia 384
Pelucha 403
Pembertonia 342
Pentacalia 233
Pentachaeta 327
Pentachaetinae 326
Pentalepis 455
Pentanema 383
Pentaphorus 117
Pentaphragma 607
P. begonifolium 606
Pentaphragmataceae 3, 605
Pentaptilon 596
Pentatrichia 275
Penthea 89
Pentzia 355
P. group 354
Peracarpa 42
Perdicium 116
Perezia 103
Pericalia 220
Pericallis 229
Pericome 510
Perideraea 370
Periomphale 11
P. balansae 9
Peripleura 315
Index to Scientific Names
Perityle 510
P. feddemae 510
Perityleae 392, 441, 507
Peritylinae 509
Perplexia 137
Perralderia 379
Perralderiopsis 380
Pertya 122
P. group 122
Pertyeae 122
Perymeniopsis 455
Perymenium 456
Petalacte 275
P. sect. Ampilasia 269
Petasites 216
Peteravenia 567
Petradoria 325
Petrobiinae 409
Petrobium 415
Petromarula 45
Peucephyllum 438
Peyrousea 356
Phaenocoma 275
P. prolifera 274
Phaeocephalus 351
Phaeopappus 140, 146
Phaeostigma 358
Phagnalon 276
Phalacrachena 140
Phalacraea 523
Phalacrocarpum 368
Phalacrodiscus 373
Phalacroseris 192
Phaneroglossa 236
Phanerostylis 554
Phania 523
Phellinaceae 3, 608
Phelline 610
P. erubescens 609
P. lucida 609
Philactis 476
Philoglossa 180
Philyrophyllum 276
Phitosia 186
Phoebanthus 468
Phonus 145
Phyllachne 618
P. colensoi 615
Phyllocalymma 255
Phyllocephalum 170
P. scabridum 150
Phyllocharis 49
Phymaspermum 356
P. group 355
Physoplexis 45
Phyteuma 45
P. comosa 45
Picnomon 132
Picris 197
P. hieracoides 197
Picrosia 192
Picrothamnus 359
Pietrosia 194
Pilosella 195
Piloselloides 116
Pilostemon 137
Pinardia 370
Pinaropappus 194
Pingraea 311
Pinillosia 416
Pinillosiinae 416
Piora 309
Pippenalia 220
Piptocarpha 90, 159
Piptocarphinae 158
Piptocoma 159
Piptolepis 162
Piptostemma 260
Piptothrix 515
Piqueria 526
Piqueriella 525
Piqueriinae 520
Piqueriopsis 525
Pithecoseris 160
Pithocarpa 276
Pithosillum 238
Pittocaulon 221
Pityopsis 339
Pladaroxylon 225
Plagiobasis 142
Plagiocheilus 307
Plagiolophus 456
Plagius 373
Planaltoa 558
Planea 276
Plateilema 403
Plateileminae 403
Platycarpha 202
Platychaete 379
Platycodon 37
Platypodanthera 540
Platyschkuhria 438
Platyspermatiaceae 7
Platyspermation 11
Plazia 111
Plectocephalus 140
Plectostachys 276
Plectreca 173
Pleiacanthus 193
Pleiogyne 351
Pleiotaxis 122
Pleocarphus 105
Pleurocarpaea 172
Pleurocoronis 555
Pleuropappus 276
Pleurophyllum 301
Pluchea 387
P. microcephala 387
Plucheeae 374
Podachaenium 473
Podanthus 456
Podocoma 315
Podocominae 312
Podolepis 276
Podopappus 315
Podosperma 277
Podospermum 198
Podotheca 277
Index to Scientific Names
Poecilolepis 293
Pogonolepis 277
Pojarkovia 240
Poljakovia 366
Pollalesta 159
Polyacantha 134
Polyachyrus 102
Polyactis 340
Polyanthina 550
Polyarrhena 293
Polycalymma 277
Polychaetia 279
Polychrysum 362
Polydora 170
Polygyne 307
Polymnia 440
Polymniastrum 440
Polymnieae 392, 439
Polymniinae 439
Polytaxis 138
Popoviocodonia 42
Porophyllum 428
Porphyrostemma 390
Porterella 49
Postia 378
Pratia 47
Praxeliinae 532
Praxeliopsis 534
Praxelis 533
Preauxia 369
Prenanthella 192
Prenanthes s.str. 189
Prestelia 162
Printzia 301
Prionolepis 180
Prionopsis 330
Prismatocarpus 40
Pristocarpha 350
Prolobus 539
Prolongoa 368
Proteopsis 162
Protoedraianthus 41
Proustia 100
Psacaliopsis 219
Psacalium 219
Psanacetum 366
Psathyrotes 403
Psathyrotinae 403
Psathyrotopsis 438
Psednotrichia 238
P. xyridopsis 208
Psephellus 140
Pseudactis 238
Pseudelephantopus 164
Pseudobahia 496
Pseudobartlettia 438
Pseudoblepharispermum 389
Pseudobrickellia 557
P. brasiliensis 558
Pseudocadiscus 237
Pseudocampanula 42
Pseudoclappia 431
Pseudoconyza 389
Pseudognaphalium 277
P. luteoalbum 252, 254
Pseudogynoxys 234
Pseudohandelia 362
Pseudokyrsteniopsis 556
Pseudoligandra 260
Pseudolinosyris 318
Pseudonemacladus 46
Pseudonoseris 179
Pseudopiptocarpha 156
Pseudostifftia 149
P. kingii 148
Pseudostifftiinae 148
Psiadia 289
Psiadiella 289
Psilactis 335
Psilocarphus 277
Psilostrophe 405
Psilostrophinae 404
Psilothonna 238
Psychrogeton 319
Psychrophyton 279
Ptarmica 364
Pterachaenia 198
Pterigeron 390
Pterocaulon 385
P. virgatum 386
Pterochaeta 277
Pterocypsela 189
Pteronia 301
Pteropogon 279
P. sect. Pteropogonopsis 263
Pterothrix 278
Pterygopappus 278
Ptilostemon 133
Ptilostephium 481
Ptosimopappus 146
Pulicaria 379
Punduana 174
Pycnocephalum 160
Pycnosorus 278
Pyrethrum 366
Pyrrhopappus 192
Pyrrocoma 332
Pytinicarpa 309
Quechualia 157
Quelchia 99
Quinetia 278
Quinqueremulus 278
Rachelia 278
Radlkoferotoma 522
Rafinesquia 193
Raillardella 506
Raillardellinae 497
Raillardia 502
Raillardiopsis 499
Railliarda 502
Railliardia 502
R. [unranked] Raillardella 506
Rainiera 221
Raoulia 278
R. eximia 246
Raouliopsis 279
633
Rastrophyllum 170
Ratibida 470
Raulinoreitzia 546
Rayjacksonia 332
Reichardia 190
Relhania 279
R. fruticosa 254
Remya 302
Rennera 355
Rensonia 456
Revealia 527
Rhagadiolus 197
Rhamphogyne 309
Rhanteriopsis 378
Rhanterium 378
Rhaponticum 143
R. group 142
Rhaponticoides 140
Rhetinodendron 230
Rhetinolepis 371
Rhigiophyllum 41
Rhinactina 319
Rhinactinidia 319
Rhizocephalum 50
Rhodanthe 279
R. anthemoides 254
R. frenchii 254
R. moschata 254
Rhodanthemum 373
Rhodogeron 385
Rhynchocarpus 279
Rhynchopsidium 279
Rhynchospermum 310
Rhynea 282
Rhysolepis 468
Rhytidanthe 270
Rhytidospermum 367
Richea 261
Richterago 117
R. discoidea 118
Richteria 361
Riddelliinae 404
Riencourtia 456
Rigiopappus 327
Robinsonecio 220
Robinsonia 230
Roccardia 279, 281
Rochonia 302
Roella 40
Rojasianthe 470
Rojasianthinae 441, 470
Rolandra 164
Rolandrinae 164
Roldana 220
Roodebergia 293
Rosenia 279
Rothmaleria 183
Roucela 42
Roussea 613
R. simplex 612
Rousseaceae 3, 611
Ruckeria 236
Rudbeckia 471
Rudbeckiinae 441, 470
634
Rugelia 220
Ruilopezia 483
Rumfordia 491
Russowia 142
Ruthiella 49
Rutidosis 280
R. leptorrhynchoides 254
Sabazia 485
Sachokiella 42
Sachsia 385
Saintmorysia 350
Salcedoa 100
Salmea 472
Santolina 371
Santosia 565
Sanvitalia 476
Sarcanthemum 289
Sartorina 520
Sartwellia 423
Saussurea 137
S. group 137
Scabrethia 464
Scaevola 595
Scalesia 468
Scariola 188
Scherya 523
Schischkinia 141
Schistocarpha 486
Schistostephium 356
Schizogyne 378
Schizoptera 457
Schizotrichia 429
Schkuhria 438, 491
S. pinnata 439
Schlechtendalia 87
S. luzulifolia 88
Schmalhausenia 136
Schoenia 280
Schumeria 144
Sciadocephala 519
Sclerocarpus 468
Sclerolepis 518
Sclerorhachis 362
Sclerostephane 379
Sclerotheca 52
Scolyminae 182
Scolymus 182
Scorzonera 198
Scorzonerinae 198
Scrobicaria 226
Scyphocoronis 280
Scyphopappus 369
Selleophytum 415
Selliera 596
S. radicans 590
Selloa 324, 486
Semiria 543
S. viscosa 544
Senecio 230
S. sect. Bethencourtii 229
S. sect. Dendrophorbium 233
S. eligulatus 208
S. sect. Nemosenecio 218
Index to Scientific Names
S. sect. Praegynoxys 223
S. vernalis 231
Seneciodes 167
Senecioneae 208
Sergia 45
Sergilus 311
Sericocarpus 325
Seridia 146
Seriphidium 358
Seriphium 281
Seris 117
Serratula 144
S. group 144
Shafera 226
Sheareria 310
Shinnersia 518
Shinnersoseris 193
Siapaea 551
Sideranthus 333
Siebera 131
Sigesbeckia 491
Siloxerus 280
Silphium 463
Silybum 133
Simsia 468
Sinacalia 217
Sinclairia 178
Sinclairiopsis 178
Sinoleotopodium 280
Sinosenecio 218
Siphocampylus 51
Siphocodon 41
Sipolisia 163
Sipolisiinae 162
Skirrhophorus 255
S. sect. Pogonolepis 277
Smallanthus 491
Soaresia 160
Solanecio 231
Solenogyne 310
Solenopsis 48
Solidagininae 320
Solidago 325
Soliva 352
Sommerfeltia 316
Sonchinae 189
Sonchus 190
S. arvensis 190
Sondottia 280
Soroseris 186
Spadonia 148
Spaniopappus 515
Sparganophoros 157
Spathipappus 366
Specularia 44
Sphacophyllum 397
Sphaeranthus 389
Sphaereupatorium 564
Sphaeroclinium 351
Sphaeromeria 360
Sphaeromorphaea 386
Sphaerophora 161
Sphagneticola 457
Sphenogyne 356
Spilanthes 472
Spilanthinae 441, 471
Spiracantha 164
Spirochaeta 164
Spiroseris 186
Squamopappus 473
Stachyanthus 160
Stachycephalum 491
Staehelina 130
Standleyanthus 516
Starkea 178
Staurochlamys 417
Stebbinsia 186
Stebbinsoseris 191
Steetzia 299
Steiractinia 457
Steirodiscus 238
Stemmacantha 143
Stenachaenium 384
Stengelia 172
Stenocarpha 485
Stenocephalum 157
Stenocline 281
Stenopadus 99
S. group 97
S. subg. Stomatochaeta 99
Stenophalium 281
Stenops 237
Stenoseris 189
Stenotheca 194
Stenotus 325
S. sect. Oonopsis 332
Stephanochilus 146
Stephanodoria 333
Stephanolepis 168
Stephanomeria 193
Stephanomeriinae 192
Stephanopappus 273
Steptorhamphus 189
Stera 385
Stereosanthus 288
Stevia 526
S. morii 526
Steviopsis 556
Steyermarkina 570
Stifftia 96
S. group 96
Stifftieae 96
Stigmatotheca 369
Stilpnogyne 229
Stilpnolepis 364
Stilpnopappus 157
Stilpnophyton 350
Stizolophus 140
Stoebe 281
Stokesia 165
Stokesiinae 165
Stomatanthes 529
Stomatochaeta 99
Stramentopappus 153
Streptoglossa 390
Strobocalyx 174
Strongylosperma 351
Strophopappus 157
Index to Scientific Names
Strotheria 429
Struchium 157
Stuartina 281
Stuckertiella 281
Stuessya 469
Stylidiaceae 3, 614
Stylidium 618
S. graminifolium 615
Stylocline 281
Stylolepis 276
Styloncerus 280
Stylotrichium 544
S. rotundifolium 545
Sventenia 191
Symblomeria 154
Symphyandra 42
Symphyllocarpus 574
Symphyochaeta 230
Symphyomera 352
Symphyopappus 548
S. decussatus 548
Symphyotrichinae 334
Symphyotrichum 335
Syncalathium 189
Syncarpha 281
Syncephalum 282
Syncretocarpus 469
Synedrella 457
Synedrellopsis 457
Syneilesis 217
Synosma 230
Synotis 239
Synotoma 45
Syntrichopappus 496
Synurus 135
Syreitschikovia 135
Taeckholmia 191
Tageteae 392, 420
Tagetes 429
Takeikadzuchia 135
Takhtajaniantha 198
Talamancalia 233
Tamananthus 483
Tamania 483
Tamaulipa 541
Tanacetopsis 364
Tanacetum 366
Taplinia 282
Taraxacum 186
Tarchonanthinae 119
Tarchonanthus 120
Tecmarsis 387
Tehuana 476
Teixeiranthus 524
Telanthophora 221
Telekia 383
Telmatophila 171
Temnolepis 397
Tenrhynea 282
Tephroseris 219
Tephrothamnus 158
Tessaria 388
Tetrachyron 473
Tetradymia 222
Tetragonotheca 481
Tetramolopium 316
Tetraneuriinae 404
Tetraneuris 405
Tetranthus 472
Tetraperone 417
Thaminophyllum 356
Thamnoseris 191
Theilera 39
Thelechitonia 457
Thelesperma 416
Theodorovia 43
Thespidium 388
Thespis 310
Thevenotia 127
Thiseltonia 282
Thodaya 236
Thorelia 172
Thoreliella 172
Thurovia 326
Thymophylla 429
Thymopsis 439
Tiarocarpus 136
Tietkensia 282
Tilesia 458
Tisserantia 389
Tithonia 469
Toiyabea 342
Tolpis 195
Tomanthea 146
Tomentaurum 339
Tonestus 289
Tostimontia 104
Tourneuxia 199
Townsendia 337
Toxanthes 282
Tracheliopsis 42
Trachelium 46
Tracyina 327
Tragoceros 476
Tragopogon 199
T. pratensis 199
Tragopogonoides 197
Trallesia 367
Traversia 224
Treichelia 40
Trematolobelia 52
Trepadonia 157
Trianthaea 174
Tricarpha 485
Trichanthemis 361
Trichanthodium 282
Trichapium 449
Trichocline 114
Trichocoroninae 517
Trichocoronis 517
Trichocoryne 476
Trichogonia 539
Trichogoniopsis 540
Trichogyne 283
T. sect. Ifloga 268
Tricholepis 142
Trichoptilium 403
635
Trichospira 174
Trichospirinae 174
Tridactylina 358
Tridax 481
T. mexicana 482
Trigonopterum 458
Trigonospermum 492
Trilisa 531
Trimeris 47
Trimorpha 340
Triniteurybia 342
Triodanis 44
Trioncinia 409
Tripleurospermum 367
Triplocephalum 389
Triplotaxis 167
Tripolium 319
T. subg. Astropolium 335
T. sect. Oxytripolium 335
Tripteris 244
T. nervosa 242
Triptilion 102
Triptilodiscus 283
T. pygmaeus 254
Trixis 105
T. vauthieri 105
Troglophyton 283
Trommsdorffia 196
Tuberculocarpus 458
Tuberostylis 571
Tugarinovia 127
Turaniphytum 364
Turpinia 158
Tursenia 311
Tussilago 216
Tuxtla 458
Tyleropappus 419
Tyrimnus 133
Tysonia 273
Uechtritzia 115
Ugamia 361
Uleophytum 571
Unamia 324
Unigenes 49
Unxia 492
Urbananthus 562
Urbinella 430
Urmenetia 109
Urolepis 537
Uropappus 191
Urospermum 197
Urostemon 224
Ursinia 356
Ursiniopsis 356
Ursinieae 342
Vanclevea 326
Vanillosmopsis 161
Varilla 430
Varillinae 430
Varthemia 384
Vasquezia 509
Velleia 596