Nordic Journal of Botany
A new species of Anisopappus (Asteraceae, Inuleae)
and its phylogenetic relationships within the genus
S. Ortiz, J. A. R. Paiva and I. Pulgar
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Ortiz, S., Paiva, J. A. R. & Pulgar, I. 2001. A new species of Anisopappus
(Asteraceae, Inuleae) and its phylogenetic relationships within the genus. - Nord. J.
Bot. 21: 55-62. Copenhagen. ISSN 0107-055X.
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A new species of the genus Anisopappus (Asteraceae, Inuleae) from Angola, A .
fruticosus, is described. The principal differences with respect to A . chinensis are
discussed, and phylogenetic relationships with the other species of the genus are
investigated by means of a cladistic analysis using morphological characters.
S. Ortiz, Laboratorio de Botanica, Facultade de Farmacia, Universidade de Santiago, 15706 Santiago de Compostela. Galicia-Spain. - 1 A . R. Paiva, Departamento de Botcinica, Faculdade de Citncias e Tecnologia. Universidade de
Coimbra, 3049 Coimbra, Portugal. - I. Pulgar. Laboratorio de Botanica. Facultade
de Farmacia. Universidade de Santiago. I5706 Suntiago de Compostela. GaliciaSpain.
Introduction
Anisopappus Hook. & Am. is a genus traditionally
included in the tribe Inuleae (Asteraceae) (Hoffmann
1890; Wild 1964; Anderberg 1991; Eldenas &
Anderberg 1996; Eldenas et al. 1998), though
recently Eldenas et al. (1999) concluded that this
genus is more closely related to the tribe Heliantheae
s. 1. The distribution area of Anisopappus covers
southern Asia (where only Anisopappus chinensis
Hook. & Am. is present), tropical and southern
Africa, and Madagascar. In his revision of the genus,
Wild (1964) accepted 29 species. More recently, 10
new species have been described (Lawalrke 1974;
Lisowski 1986a, 1986b, 1988; Ortiz & Paiva 1995),
and Eldenas & Anderberg (1996) accepted about 40
species. However, in our synopsis of the genus (Ortiz
et al. 1996) we accepted only 17 species, 10
subspecies and 4 varieties in view of our conclusion
that pappus morphology has limited diagnostic
value. Many previous workers (Humbert 1923; Wild
1964; Lisowski 1986a, 1986b, 1988) have used
pappus morphology as a diagnostic character, and
indeed several species of Anisopappus have been
described entirely or almost entirely on the basis of
pappus morphology alone.
Following recent studies on Anisopappus, we
describe a new species from Angola and analyse the
phylogenetic relationships between this species and
the other species of the genus.
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Materials and methods
Description of the new species Anisopappus fmti-
cosus is based on the morphological study of ma-
terial from the herbaria of COI, K, LISC, LUAI, and
PRE (abbreviations based on Holmgren et al. 1990).
Styles and cypselas of dried material were mounted on aluminium stubs, and coated with a c. 30 nm
layer of Au for study with a scanning electron
microscope (SEM) (LEO 435VP), operating at 15 kV.
The phylogenetic analysis of Anisopappus, for
investigation of phylogenetic relationships between
Accepted 25-01-2001
Nord. J. Bot. 21(2)2001
55
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A. fmticosus and the other species of the genus, was
based on the morphological analysis of Eldenas &
Anderberg (l996), though with some modifications.
In accordance with our synopsis of the genus (Ortiz
et al. 1996), we removed A. bumpsianus Lisowski
(considered a synonym of A. davyi S. Moore), A.
burundensis Lisowski (considered a synonym of A.
ubercornensis G. Taylor), and A. buchwaldii (0.
Hoffm.) Wild, A. oliverunus Wild, A. discolor Wild,
and A. scrophularifolius (Baker) Wild (considered
Table 1. Characters and states used in phylogenetic analysis.
1. Habit herbaceous (0). Habit woody (1).
2. Anisopuppus-type septate hair (0). Asteriscus-type septate hair (I). A. pinnatifdus-type septate hair (2)
(see Eldenas & Anderberg, 1996).
3. Wooly hair present (0). Absent (1).
4. Stems without wings (0). Stems with more or less prominent decurrent wings (1).
5. Petioles minute or completely absent (0). Petioles long, distinct (1).
6. Leaves lobed to pinnatipartite (0). Leaves simple, undivided (1).
7. Leaf venation palmate (0). Leaf venation pinnate (1). Veining costate (2).
8. Leaf main veins distintc, veinlets diffuse (0). The entire network of veins is sunken (1).
9. Leaves toothed (0). Leaves entire (1).
10. Capitula solitary on long branches (0). Few capitula together in loose corymbs (1). Capitula numerous,
in more or less dense corymbs (2).
11. Capitula campanulate to hemispheric (0). Capitula obconic to cyathiform (1).
12. Receptacle more or less flat (0). Receptacle hemisphaerical or conical (1). Receptacle narrowly conical (2).
13. Receptacle epaleate (0). Receptacle paleate (1).
14. Staminodia present in ray-florets (0). Staminodia absent in ray-florets (1).
15. Disc-floret corolla-lobes short (I0.5 mm long) (0). Disc-floret corolla-lobes long (> 0.5 mm long) (1).
16. Disc-floret corolla-lobes without sclerenchymatous cells accompanying the vascular strands (0). With
more or less narrow, submarginal sclerenchyme (1). With broad, marginal sclerenchymatous bands (2).
17. Disc-floret corolla without bristly hairs (0). Disc-floret corolla with bristly hairs (I).
18. Endothecial tissue with radial thickenings (0). Endothecial tissue with polarized thickenings (1).
19. Filament collar cells longer than wide (0). Filament collar cells wider than long (1).
20. Anther thecae long calcarate (0). Anther thecae shortly calcarate (1). Anther theca ecalcarate (2).
21. Anther tails minute (0). Anther tails long (I).
22. Anther tails completely unbranched (0). Anther tails with short branches (I). Anther tails with long
branches (2).
23. Style with crystals rectangular-prismatic or without crystal deposits (0). Style with crystals star-shaped (1).
24. Style branch apex obtuse, rounded (0). Style branch apex acute (1).
25. Style branch vein narrow (0). Style branch vein thick (1).
26. Sweeping-hairs acute (0). Sweeping-hairs obtuse (1).
27. Cypsela straight (0). Cypsela curved (1).
28. Cypsela yellowish-brown (0). Cypsela dark reddish-brown (1).
29. Cypsela trilocular (0). Cypsela more or less terete (I). Cypsela triquetrous (2).
30. Cypsela with sclerenchymatic tissue (0). Cypsela without sclerenchymatic tissue (1).
31. Cypsela surface smooth (0). Cypsela surface with distinctly raised ribs (1).
32. Cypsela epidermis without crystals (0). Cypsela with one elongated crystal in each epidermal cell (2).
Crystals or crystal fragments in parenchyma (2).
33. Epidermal cells smooth (1). Epidermal cells papillose (I).
34. Epidermal cells equal in size to subepidermal cells (0). Epidermal cells enlarged (I).
35. Carpopodium absent (0). Carpopodium present (1).
36. Pappus bristles absent (0). Pappus bristles present (1).
37. Pappus scales present (0). Pappus scales absent (1).
38. Pappus scales larger, regular (0). Pappus scales large, serrate, some with elongate midportion (I). Pappus
scales irregular, serrate (2). Pappus scales reduced into a minute rim (3).
zyxwvu
56
Nord. J. Bor. 21(2) 2001
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synonyms of A . chinensis). However, we kept A .
rhombifolius Wild, a species included by us (Ortiz et
al. 1996) in the A . chinensis complex, but probably
different in accordance with the data hrnished by
Eldenas & Anderberg (1 996) principally regarding
the type of cypsela hairs, unique in the genus. We
also included A . longipes (Comm. ex Cass.) Wild, a
prostrate endemic species from Madagascar probably
related to A . chinenis, and A . pseudopinnatifdus, a
species described by us (Ortiz & Paiva 1995) from
Namibia; neither species was included by Eldenas &
Anderberg (1996) in their phylogenetic analysis. As
regards Anisopappus athanasioides S. Ortiz & Paiva,
a species described from somewhat immature
material (Ortiz & Paiva 1995), we agree with Eldenas
& Anderberg (1996: 174), who consider that the
specimen from which it was described is “significantly different from all other species of Anisopappus”. We have subsequently found more complete material of this taxon, indicating that besides
its distinct capitula and inflorescences it has mature
ray-florets very different to those of Anisopappus.
This species was therefore not included in the
present analysis.
In relation to these modifications we coded ‘-‘
instead of 2 for character 35 (pappus scale shape) of
A . abercornensis and A . chinensis since it is variable
in the different synonyms listed above.
Like Eldenas & Anderberg (1996), we included
six other genera of the Inuleae in the analysis to
assess whether Anisopappus is monophyletic. These
genera were Asteriscus Tourn. ex Mill., Buphthalmum L., Calostephane Benth., Geigeria Griess., Inula L., and Pulicaria Gaertn. The former four genera
are representatives of the clades most closely related
to Anisopappus, which is basal within the tribe,
whereas Inula and Pulicaria are representatives of
the most evolved clade (see Anderberg 1991).
A total of 23 taxa and 38 morphological characters were studied (Tables 1 and 2). Polarization of
Table 2. Data matrix. The characters are numbered in accordance with Table 1 . Character states unknown or inapplicable are
indicated with a dash.
12345
Arctotis
Asteriscus
Bupthalmum
Calostephane
Geigeria
Inula
Pulicaria
Anisopappus abercornensis
Anisopappus chinensis
Anisopappus corymbosus
Anisopappus davyi
Anisopappus fruticosus
Anisopappus grangeoides
Anisopappus holstii
Anisopappus junodii
Anisopappus kirkii
Anisopappus latifolius
Anisopappus longipes
Anisopappus marianus
Anisopappus pinnati’dus
Anisopappus pseudopinnatifdus
Anisopappus rhombifolius
Anisopappus salvifolius
Anisopappus smutsii
Anisopappus sylvaticus
Nord. 1. Bot. 21(2) 2001
0000-
01 100
01 100
001 11
001 10
01 100
-1 100
00101
00101
00101
00 100
10101
00101
00101
o-oo 1
00101
00 100
00101
00101
02 1002 1000101
10101
o-oo 1
00101
1
6789
11111
12345
11112
67890
22222
1234.5
22223
67890
33333
12345
333
678
00000
11010
11010
10000
110111001
11011
000 10
10000
1000 1
12010
11001
10000
1000 1
10102
10102
10000
10000
10100
0101 1
0101 1
10000
1 1 101
10102
1000 1
00000
00111
00111
00001
00-01
-0011
00011
00100
00100
01100
00100
00100
00100
02100
11101
00100
00000
00100
00100
001-1
101-1
00100
00100
11101
00100
00000
10110
10110
11101
21110
00002
10011
00111
00111
00111
00111
00111
00111
00111
00111
00111
00111
00111
00111
00111
00111
00111
00111
00111
00111
00000
00001
00101
11011
1101 1
12001
12101
00000
00000
11001
00000
00000
00000
10000
11001
10000
10000
00000
00000
00000
00-00
00000
-1001
-1001
1001 1
0001 1
01001
01001
12000
12000
10101
12000
12-00
12000
10101
1000 1
12000
-001 1
12000
000
002
002
00 1
00 1
11102
0000002
002
102
0000-
00000
11000
11001
10000
0002 1
00020
00110
00- 10
000 10
000 10
11110
11110
101 10
11110
01-10
11110
101 10
01-10
11110
101 11
11110
1--111110
11110
11110
-1 110
01-10
11110
01-
003
002
003
003
01002
002
002
01003
002
01-
zy
--o--
12000
12000
12000
10101
1000 1
10101
57
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Fig. 1. Anisopappus fruticosus. - A. habit. - B. capitulum. - C. disc (left) and ray floret (right). - D. leaf.
and receptacular palea (right). - F. cypsela. - Based on holotype (Santos & Henriques 1146, LISC).
58
-
E. phyllary (left)
Nord. J. BOI. ZI(2) 2001
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characters was determined by the outgroup comparison method with Arctotis L. as outgroup, as in the
cladistic analysis of Eldenas & Anderberg (1996).
For microscopic examination of floral parts, these
were first boiled in water with a surfactant, then
mounted in Hoyer’s solution (Anderson 1954).
Wagner parsimony analysis of the data matrix
(Table 2) was performed on a PC with the aid of the
program HENNIG86 (Farris 1988). We used the
multiple hennig option (mh*, for constructing several initial cladograms by adding taxa in several different sequences, then retaining the shortest) followed
by the branch-breaker option (bb*, for generating
multiple equally parsimonious cladograms). A successive weighting procedure (Farris 1969) was used
to generate cladograms in which the relative weight
of homoplasious characters was reduced. All multistate characters were treated as nonadditive. Character state unknown or inapplicable was indicated
with a dash (-) in the matrix.
Besides the addition of the 2 species, we added 3
new characters to the 35 used by Eldenas &
Anderberg (1996): characteres 1, 11, and 15 of Table
1. As regards character I , it should be pointed out
that on the label of the type of A . fruticosus it is
described as suffrutex, but from the texture of its
branches there is no doubt that it should be considered woody.
Results and discussion
Anisopappus fruticosus S. Ortiz & Paiva sp.
nov.
Frutex ut plurimum 80 cm altus. Folia elliptica,
margine serrata, petiolis (5-)lo- 13 mm longis.
Capitula dense corymbosa, pedunculis brevibus, 1525(-35) mm longis. Flores radii ut plurimum 14 x 3
mm. Cypsela squamosa et setis 3-5(-7) praedita,
barbellatis, (0.5-)0.8-1(-1.5) mm longis. - Fig. 1.
Typus: Angola, Huila, Lubango (Sa da Bandeira),
Fenda de Tunda-Vala, high plateau. Rocky outcrops.
4.vii.1965. Santos & Henriques 1146 (holotype:
LISC, isotypes: K, LUAI, PRE).
Shrub 70-80 cm high. Branches striate, with dense
pilosity of spreading, multicellular, Anisopappustype hairs (see Eldenas & Anderberg, 1996), and
sessile to subsessile yellow glands; internodes 4-8
mm long. Leaves 40-55 x 13-20 mm, elliptic to
elliptic-lanceolate; lamina (30-)35-45 mm long, venation pinnate; petiole (5)
10-13 mm long; margins
serrate, with subobtuse teeth 1.3-13 - 3 ) mm long;
base cuneate; apex acute; both surfaces greenish,
with a moderate to dense pilosity of pluricellular
hairs like those of the stem, and sessile yellow
glands, these conspicuously more abundant on the
adaxial surface. Capitula numerous, (3-)6-12 arranged in corymbiform synflorescences, on erect-patent
peduncles 15-25(-35) mm long, radiate, heterogamous. Involucre 10-13 x 4-5 mm, campanulatehemispheric, with c. 40-60 phyllaries arranged in 3(4) rows, greeenish, with multicellular hairs like those
of the stem, the margins entire; outermost phyllaries
3-4 x 0.5-1 mm, linear; middle phyllaries 4-5 x 1.31.5 mm, lanceolate, with apex acute to acuminate,
slightly scarious; innermost phyllaries (4-)5-5.5 x
1.3-1.5 mm, lanceolate, with apex acute to acuminate; receptacle flat; receptacular paleae 3.5-4(4.5) x 0.5-0.8 mm, somewhat folded, with acute and
fimbriate apex. Florets c. 500-800 per capitulum.
Ray florets c. 40, female; corolla yellow; tube 1.31.8 mm long; limb (8-)lo-12 x ( 1 - ) I 5 3 mm, with
short glandular hairs, 3 apical lobes c. 0.3 mm long;
style 2.5-3 mm long, style branches c. 0.5 mm long;
immature cypselas 1-1.5 x 0.3 mm, slightly obconic,
slightly 4-5 angled, longitudinally striate, with
erect-patent hairs c. 0.2 mm long, carpopodium
absent; pappus of several short scales 0.3-0.4 x 0.3
mm, irregularly dentate-fimbriate, and 3(-4) short
bristles 0.8-1 mm long, about half or slightly more
than half the length of the corolla tube, barbellate.
Disc florets hermaphrodite, actinomorphic; corolla
3-4.5 mm long, apical lobes 0.3-0.5 mm long;
anthers 3-3.5 mm long, apical appendage c. 0.5 mm
long, lanceolate, endothecial tissue polarized, endothecial cells longer than wide, anther base
minutely caudate, tails 0.3 mm long, calcarate,
filament collar 0.2-0.4 mm long; style 3-4.5 mm
long, style branches c. 0.5 mm long, with sweeping
hairs acute (Fig. 2A); immature cypselas (0.8-)1-1.2 x
0.2-0.3 mm, slightly obconic, like those of the ray
florets; pappus of several short scales 0.2-0.4 x 0.20.3 mm, irregulary dentate-fimbriate, and 3-5(-7)
short bristles (0.5-)0.8-1(-1.5) mm long, about one
third or slightly more than one third of the length of
the corolla, barbellate. - Fig. 1.
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Note, The species most morphologically similar to
Anisopappus fruticosus is probably A . chinensis
Hook. & Am., which can be readly differentiated
from the new species by its herbaceous habit, its
stems rarely more than 50 cm tall, its ovatelanceolate leaves with palmate venation, its often
solitary capitula on peduncles generally 50-120 mm
long, its ray florets less than 8 mm long, its obtuse
style sweeping hairs (Fig. 2B) and its pappus without
bristles conspicuously longer than the scales.
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Nord. 1. BOI. 21(2) 2001
59
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Fig. 2. SEM micrographs of the of style branch sweeping hairs. - A. acute sweeping hairs in Anisopappus fruricosus (Santos &
Henriques 1146, LISC). - B. obtuse sweeping hairs in Anisopappus chinensis subsp. chinensis (Exell & Mendonca 1168, COI).
The specimen Gossweiler 11137 from Angola,
Huila, Ungueria, near Chaento River, 1340 m alt.,
2.vi.1937 (COI, PRE) has similar leaves and a
somewhat similar pappus to A . fruticosus, but shows
certain differences from A . fruticosus as regards the
remaining characters cited above, so we have not
included it in A . fruticosus (though this option
cannot be ruled out).
Indeed, although Anisopappus latifolius was
positioned as a sister group of the rest of the genus
Anisopappus in almost all of the 106 most
parsimonious trees (e.g. Fig 3B), as in Eldenas &
Anderberg’s analysis, in one tree (Fig. 3C) it was
positioned as a sister group of the GeigeriaBuphthalmum group, and thus separated from the
genus Anisopappus, which would make this genus
paraphyletic. However, in the more recent
phylogenetic analysis of Eldenas et al. (1999) based
on nucleotide sequences of the chloroplast gene
ndhF, the two species of Anisopappus analysed, A .
latifolius and A . smutsii Hutch., form a monophyletic
group with 100 % jackknife support .
As expected, and except in the single tree in
which A. latifolius was positioned as a sister group
of the Geigeria-Buphthalmum group, the topology
of this latter group was identical to that of Eldenas &
Anderberg’s strict consensus tree.
In our strict consensus tree (Fig. 3A), the
remaining Anisopappus species formed an 1 1-clade
polytomy. However, most of the most parsimonious
trees contained two major clades.
The first of these clades (monophyletic in 75% of
the 106 trees) comprised A . holstii (0. Hoffm.) Wild,
A . sylvaticus (Humbert) Wild, A . salvifolius (DC.)
Wild, A . corymbosus Wild, A . smutsii and A. junodii
Hutch. (as for example in Fig. 3B). Within this clade,
A. holstii was a sister group to the rest of the species
in almost all trees, while A . smutsii and A . junodii
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Phylogenetic relationships
The cladistic analysis yielded 106 equally
parsimonious cladograms, each 97 steps long. The
consistency index (CI) was 0.50, and the retention
index (RI) 0.70.
Examination of the strict consensus tree of the
106 most parsimonious trees (Fig 3A) reveals poor
resolution, as in Eldenas & Anderberg’s (1996)
analysis, though the number of unresolved
polytomies is even higher in our analysis.
Of particular relevance with respect to Eldenas &
Anderberg’s analysis is the fact that Anisopappus
latifolius ( S . Moore) B.L. Burtt (a species with
somewhat deviant characters in comparison to the
rest of the genus, notably the lack of receptacular
paleae, lack of sclerenchyma in the cypselae, and
presence of enlarged cypsela epidermis cells) forms a
polytomy both with the Geigeria-Buphthalmum
group and the remaining species of Anisopappus.
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Fig. 3. A. strict consensus tree derived from the 106 equally most parsimonious trees. - B. one of the 106 equally most
parsimonious trees. - C. the single most parsimonious tree in which A . lurifolius was positioned as sister group to the GeigeriuBuphrhulmum group. - D. one of the two cladograms resulting from the successive weighting analysis.
60
Nord. J. Bot. ZI(2) 2001
II
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C
Nard. 1. Bat. 21(2) 2001
D
61
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were the most advanced species. Note that topology
within the group comprising A . salvifolius, A .
corymbosus, A . smutsii and A . junodii was invariable
in the 106 most parsimonious trees, and is thus
reflected in the strict consensus tree (Fig. 3A).
The other major clade comprised the remaining
species of the genus, which formed a monophyletic
group in all trees except that in which A . latifolius
was positioned as a sister group of the GeigeriaBuphthalmum group. Within this clade, A . kirkii
(Oliv.) Brenan was sister group to the remaining
species (Fig. 3B), as in Eldenas & Anderberg’s
analysis. Within the clade, A . pinnutifidus (Klatt) 0 .
Hoffm. ex Hutch. and A. pseudopinnatifidus were
the most advanced species. This latter species, which
was not included in Eldenas & Anderberg’s analysis,
forms a clade with A . pinnatijidus, with which it
shows great similarity (Ortiz & Paiva 1995). Again as
in Eldenas & Anderberg’s analysis, in our strict
consensus tree A . rhombifolius and A . grangeoides
(Vatke & Hopfner ex Klatt) Merxm. formed a clade,
since these species are identical as regards the
characters considered in the analysis; notably, both
species lack a pappus.
The position of A . fruticosus was not well defined
in the strict consensus tree (Fig 3A). In all individual
most parsimonious trees, however - except in the tree
in which A . latifolius was positioned as a sister
group of the Geigeria-Buphthalmum group (Fig. 3C),
in which A . fruticosus was positioned as a sister
group to the remaining species of Anisopappus - A .
fruticosus was included in the A . kirkii / A .
pseudopinnatijidus group (Fig. 3B). Its position
within this clade was variable, although in 65% of
the 106 trees it was positioned as a sister group of
the most advanced subclade, comprising A .
pinnutifidus and A . pseudopinnatijidus, which like
A . fruticosus have capitula arranged in loose
corymbs (though this character is not as clear in A .
pseudopinnatijidus as in the other two taxa).
Analysis using a successive weighting process
gave two cladograms (one of which is illustrated in
Fig. 3D) with a somewhat similar topology to the
most equally parsimonious cladograms in which A .
latifolius was placed as the sister group of the
remaining species of Anisopappus. However, these
two cladograms contained two major clades somewhat different to those cited above.
Acknowledgements - Our thanks go to Manuel Lainz
for the Latin diagnosis, to Alfredo Lbpez “Tokio” for
the illustrations, to Guy Norman for the English
translation, and to the keepers of the herbaria
mentioned for the loan of study material.
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