Aliso: A Journal of Systematic and Evolutionary Botany
Volume 32 | Issue 1
Article 2
2014
Nelsonioideae (Lamiales: Acanthaceae): Revision
of Genera and Catalog of Species
Thomas F. Daniel
Department of Botany, California Academy of Sciences, San Francisco
Lucinda A. McDade
Rancho Santa Ana Botanic Garden, Claremont, California
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Recommended Citation
Daniel, Thomas F. and McDade, Lucinda A. (2014) "Nelsonioideae (Lamiales: Acanthaceae): Revision of Genera and Catalog of
Species," Aliso: A Journal of Systematic and Evolutionary Botany: Vol. 32: Iss. 1, Article 2.
Available at: http://scholarship.claremont.edu/aliso/vol32/iss1/2
Aliso, 32(1), pp. 1–45
ISSN 0065-6275 (print), 2327-2929 (online)
NELSONIOIDEAE (LAMIALES: ACANTHACEAE): REVISION OF GENERA AND CATALOG OF SPECIES
THOMAS F. DANIEL1,3
1
AND
LUCINDA A. MCDADE2
Department of Botany, California Academy of Sciences, 55 Music Concourse Drive, Golden Gate Park, San Francisco,
California 94118; 2Rancho Santa Ana Botanic Garden, 1500 N. College Avenue, Claremont, California 91711
3
Corresponding author (tdaniel@calacademy.org)
ABSTRACT
A taxonomic account of Acanthaceae subfamily Nelsonioideae based on morphological and
phylogenetic data treats five genera with 172 species: Anisosepalum (3), Elytraria (21), Nelsonia (2),
Saintpauliopsis (1), and Staurogyne (145). Two other currently recognized genera, Gynocraterium and
Ophiorrhiziphyllon, are included within Staurogyne, and the new combinations, Staurogyne guianensis and
S. macrobotrya, are proposed. Probable apomorphic and other diagnostic macro- and micromorphological
characters are discussed relative to the subfamily and genera. Characters of the inflorescence, androecium
(especially pollen), and seed show important phylogenetic and diagnostic signal. A key to genera, generic
descriptions and discussions, illustrations, and distribution maps are provided. Lists of currently
recognized species for each genus include synonymies and distributions by country.
Key words: Acanthaceae, Anisosepalum, Elytraria, morphology, Nelsonia, Nelsonioideae, phylogeny,
Saintpauliopsis, Staurogyne.
INTRODUCTION
Acanthaceae consist of about 190 genera and 4750 species
(Daniel unpubl.; vs. 212 genera and 3175 species fide
Mabberley 2008). The family is pantropical in distribution,
although most genera are restricted to either the New World or
the Old World. Recent phylogenetic studies (e.g., McDade
et al. 2008; Fig. 1) reveal the family to be monophyletic and
divisible into at least three subfamilies that are related as
follows: Nelsonioideae (Thunbergioideae + Acanthoideae).
Nelsonioideae, the smallest among the currently recognized
subfamilies, are here treated as consisting of 172 species in five
genera: Anisosepalum (3), Elytraria (21), Nelsonia (2), Saintpauliopsis (1), and Staurogyne (145). Geographic occurrence
and morphological diversity of Nelsonioideae reflect that of
the entire family: they are nearly worldwide in distribution and
show considerable variation in habit, foliar characteristics, and
floral features.
Nelsonioideae have had a convoluted taxonomic history
with respect to constituent genera, familial affinities, and
taxonomic rank. Nees (1832, 1847a,b) treated genera now
known as Elytraria, Nelsonia, and Staurogyne as tribe
Nelsonieae of Acanthaceae; these were characterized by having
small seeds that were not supported by conspicuous retinacula.
Bentham and Hooker (1876) recognized this tribe and also
included Ophiorrhiziphyllon in it. In the most comprehensive
and influential account of Acanthaceae to date, Lindau (1895)
treated these four genera in subfamily Nelsonioideae, which
was characterized by having numerous seeds, papilliform
retinacula, and tricolporate pollen. Van Tieghem (1908)
segregated all genera lacking retinacula from Acanthaceae as
a new family, Thunbergiaceae, with the nelsonioid assemblage
remaining a subfamily. Wettstein (1924) returned Thunbergiaceae to Acanthaceae as a subfamily with the nelsonioids as a
tribe of Thunbergioideae. In sequence, Bremekamp treated the
nelsonioid genera as a subtribe of Acanthoideae: Acantheae
(1938), as a tribe of Acanthoideae (1944), and finally, based
primarily on the presence of persistent endosperm in their
seeds, as a tribe of Scrophulariaceae: Rhinanthoideae (1953,
1955). In the most extensive treatment of the nelsonioid
assemblage to date, Hossain (1971, 1972, 1984, 2004) treated
these genera (with the addition of Anisosepalum, Saintpauliopsis and Gynocraterium) as a tribe of Acanthaceae:
Acanthoideae. Sreemadhavan (1977) and Lu (1990) have
recognized nelsonioids as a distinct family, Nelsoniaceae.
Treatment outside Acanthaceae (either as Scrophulariaceae
sensu traditionalem or as a distinct family) never received
widespread acceptance, and subfamilial rank has been
generally utilized for the taxon (e.g., Cronquist 1981; Scotland and Vollesen 2000). Recent molecular phylogenetic
studies (e.g., McDade et al. 2008, 2012) confirmed the
subfamily as monophyletic, supported recognition of the five
genera noted above, and placed the clade as sister to all other
Acanthaceae.
Some of the genera now treated as Nelsonioideae have been
placed outside Acanthaceae. Neozenkerina, Ophiorrhiziphyllon,
and Zenkerina (all 5 Staurogyne) were originally described in
Scrophulariaceae, and Saintpauliopsis was originally described
in Gesneriaceae. A further taxonomic complication is that
both Nees (1847a) and Lindau (1895) included some genera
now treated as Plantaginaceae (Adenosma R. Br.) and
Orobanchaceae (Hiernia S. Moore) in their nelsonioid taxa.
PHYLOGENY
’ 2014, The Author(s), CC-BY. This open access article is
distributed under a Creative Commons Attribution License, which
allows unrestricted use, distribution, and reproduction in any medium,
provided that the original author(s) and source are credited. Articles
can be downloaded at http://scholarship.claremont.edu/aliso/.
Because patterns of variation in macro- and micromorphological characters are best contextualized by phylogenetic relationships, we summarize recent phylogenetic results
(Wenk and Daniel 2009; McDade et al. 2012) and then discuss
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Fig. 1. Summary phylogeny of Acanthaceae showing subfamilies (and Avicennia) and relationships among genera of Nelsonioideae based on
McDade et al. (2008, 2012); OW 5 Old World, NW 5 New World.
plant characteristics. The subfamily is monophyletic and sister
to all other Acanthaceae. Monophyly is also supported by at
least one shared morphological character (possibly synapomorphic; this trait cannot be polarized with certainty until the
closest relatives of Acanthaceae are clarified), descendingcochlear aestivation of corolla lobes. Aestivation patterns have
been shown to be phylogenetically informative among
Acanthaceae and, within the family this pattern is known
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Revision of Nelsonioideae
only among Nelsonioideae (Scotland et al. 1994; Scotland and
Vollesen 2000; additional observations by us for each genus).
Therefore, there exists a basis for treating Nelsonioideae as
a distinct and circumscribable (both morphologically and
molecularly) family, as argued by Sreemadhavan (1977).
McDade et al. (2012) provided data that support the
recognition of at least five of the seven traditionally recognized
genera of Nelsonioideae: Anisosepalum, Elytraria, Nelsonia,
Saintpauliopsis, and Staurogyne. Relationships among these
genera are summarized in Fig. 1. Nelsonia is monophyletic and
sister to all other nelsonioids. We propose loss of bracteoles as
a morphological synapomorphy for Nelsonia (versus paired
bracteoles subtending each flower in other nelsonioids and,
more broadly, Acanthaceae). Elytraria is also monophyletic
and is sister to a ‘‘staurogynoid’’ clade consisting of all other
genera of Nelsonioideae except Nelsonia. Elytraria can be
delimited by at least three morphological synapomorphies:
scaly peduncles, coriaceous to sclerophyllous bracts, and
touch-sensitive stigmas. The existence of the ‘‘staurogynoid’’
clade was predicted by Hossain (2004) based on morphological similarities.Within this clade, monotypic Saintpauliopsis
is sister to Anisosepalum and these together are sister to
Staurogyne. Although no morphological synapomorphy for
this ‘‘staurogynoid’’ clade has been identified, there is a
tendency toward adnation of bracts to dichasial peduncles
(cryptic epiphylly) in all three genera. Adnation of bracteoles
to the base of flowers (i.e., receptacle and calyx tube) is likely
synapomorphic for the clade consisting of Anisosepalum and
Saintpauliopsis. Hossain (1972) indicated that Anisosepalum
and Saintpauliopsis (unlike other genera of Nelsonioideae)
lack acicular fibers in roots, stems, petioles, and veins; this
might represent an anatomical synapomorphy for these two
genera. The Asian genus Ophiorrhiziphyllon was shown to be
nested among Asian taxa of Staurogyne, and the unispecific
American genus Gynocraterium is sister to all of the sampled
American species of Staurogyne. Although a case could be
made for treating Gynocraterium as a distinct genus based on
existing molecular data, this would involve considerable
taxonomic renovations (see McDade et al. 2012), and there
do not appear to be sufficient morphological grounds for such
(see below under Staurogyne). As a result, we treat both
Ophiorrhiziphyllon and Gynocraterium in an expanded Staurogyne. Thus, all genera recognized here are monophyletic
based on molecular data and each is morphologically
distinguishable. Morphological synapomorphies are known
for all genera except for Staurogyne.
MORPHOLOGY
The subfamily can be characterized morphologically by
several presumed symplesiomorphies and at least one probable
synapomorphy (descending-cochlear aestivation of corolla
lobes as described below). Traditionally, the lack of cystoliths
and of indurate, hooklike structures (retinacula) subtending
the seeds have been important in distinguishing Nelsonioideae
(e.g., Lindau 1895). These presumably symplesiomorphic
attributes are shared with both Avicennia and genera of
Thunbergioideae. The absence of cystoliths is also shared with
the basalmost tribe (Acantheae) of subfamily Acanthoideae.
An additional distinguishing character usually associated with
Nelsonioideae is the presence of persistent endosperm in their
3
seeds (versus exalbuminous seeds in other acanths). Discussion
of this and some additional morphological characters,
highlighting phylogenetic patterns and variation among
Nelsonioideae is presented below.
Habit
Species of Nelsonioideae are either herbs or shrubs. They are
mostly terrestrial, but epiphytic or epipetric plants are known
in Anisosepalum and Saintpauliopsis and probably occur
sporadically in other genera. Herbs vary in duration from
annual to perennial. Annual herbs are apparently not
common. Both Elytraria marginata and Nelsonia gracilis are
commonly noted to be annuals (e.g., Morton 1956; Dokosi
1971; Vollesen 2013). Cultivated plants of E. marginata (Daniel
11158cv at CAS) persisted only for one year and produced
viable seeds that yielded plants lasting only a single year. While
most collections of N. gracilis consist of plants that appear to
be annual, some (see below) have larger and somewhat woody
roots suggesting that other plants may also become perennial.
Perennial herbs, with woody roots, caudices, or stems, are a
more common growth form among the subfamily. Herbs vary
from acaulescent (i.e., with a basal rosette of leaves) to
subcaulescent (i.e., with slight internodal elongation at or just
above ground level) to caulescent (up to 1 m or more high).
Most herbaceous species are consistent in growth form, but
some vary from acaulescent to subcaulescent (e.g., Elytraria
acaulis, seen in Thwaites 240 at P), some from subcaulescent to
caulescent (e.g., Elytraria marginata, seen in Adam 2058 at P),
and some from acaulescent to caulescent (e.g., Nelsonia
canescens, seen in Duthie s.n. from India at UC). Shrubs are
uncommon in Anisosepalum (e.g., A. lewallei), Elytraria (e.g.,
E. nodosa), and Staurogyne (e.g., S. minarum), and do not
occur at all in Nelsonia or Saintpauliopsis. The habit of plants
in at least one species (E. imbricata) varies from acaulescent
herbs to weak shrubs. Thus, there is no clear pattern of
evolution of plant habits at the level of genus but the trait
merits further examination once a densely sampled phylogeny
of the largest genus, Staurogyne, is available.
Leaves
Nelsonioideae exhibit greater variation in foliar disposition
and arrangement than the other main lineages of Acanthaceae.
Plants with leaves 6 evenly disposed along stems occur in all
genera; basal clusters of leaves occur in all genera except
Anisosepalum; and apical/aerial clusters of leaves occur in some
Elytraria. Arrangement of the leaves varies from alternate
(some Elytraria, Nelsonia, and Staurogyne) to subopposite
(some Staurogyne) to opposite (all genera except Elytraria) to
whorled (Anisosepalum, Elytraria).
Leaves of Nelsonioideae are either sessile or petiolate. If
petiolate, then the petioles (bladeless portion only) are nearly
always shorter than the blade, except in Saintpauliopsis and at
least one species of Staurogyne (S. chapaensis), in which the
petioles are conspicuously longer than the blades.
Leaf variation is particularly evident in Elytraria. In
addition to variation in disposition and arrangement as noted
above, length of leaf blades in the genus varies from 4 mm (E.
filicaulis) to 310 mm (E. caroliniensis) and the margin varies
from entire to sinuate to pinnatisect. Some of the variation in
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Fig. 2. Leaf variation in Elytraria acaulis from India and Sri Lanka.—A. Drummond 26727 (UC).—B. Diraviam & Xavier 20726 (A).—C.
Amrita s.n. (ASU).—D. Fagerlind 3925 (S).—E. Kostermans & Wirawan 824 (G).—F. van Beusekom 1653 (US).—G. Dittus WD70012102 (US).—
H. Jayasuriy et al. 622 (US).—I. Fosberg 50210 (US).—J. Davidse & Sumithraarachchi 8131 (US).—K. Wight 2174 (NY).—L. Wight 2174 (GH).
Drawn by N. Pugh.
shape, size, and margin among leaves of E. acaulis from India
and Sri Lanka is shown in Fig. 2.
Apical/aerial whorls or clusters of leaves may be synapomorphic for some species within Elytraria, but phylogenetic
patterns for foliar characteristics are otherwise unclear.
Inflorescences
Inflorescences of Acanthaceae are often complex or highly
modified. The basic unit of inflorescences in Nelsonioideae, as
in most other Acanthaceae, is a dichasium. The dichasial
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Revision of Nelsonioideae
nature is reflected by the paired bracteoles that subtend each
flower. Throughout the subfamily the dichasium is modified
by reduction to a single flower. Dichasia of Nelsonia have
undergone further reduction by the loss of both bracteoles
(although they are very rarely [teratologically?] present below
some flowers). Dichasia may be sessile or pedunculate; flowers
may be sessile or pedicellate. The dichasia sometimes occur in
leaf axils, but more commonly they are subtended by a bract
and grouped into sessile to pedunculate spikes (i.e., both
dichasia and flowers sessile to subsessile), racemes (i.e., dichasia
sessile to subsessile and flowers pedicellate), or thyrses (i.e.,
dichasia pedunculate and flowers sessile or pedicellate). Each of
the spikes, racemes, and thyrses are uncommonly branched to
form a panicle. Arrangement of bracts varies from alternate
(spirally arranged) in all genera to sometimes subopposite in
Anisosepalum to sometimes opposite in Staurogyne.
Unlike variation in plant habit and leaf disposition,
inflorescence traits show considerable phylogenetic pattern.
In addition to the synapomorphic absence of bracteoles in
Nelsonia: 1) scales on inflorescence peduncles are synapomorphic for Elytraria; 2) bracts with paired, subapical,
winglike teeth are autapomorphic for E. imbricata; 3) adnation
of bracts to dichasial peduncles may be synapomorphic for
Anisosepalum, Saintpauliopsis, and Staurogyne (although this
trait is difficult to assess owing to the absence of dichasial
peduncles and pedicels in the spicate inflorescences of Elytraria
and Nelsonia); and 4) adnation of bracteoles to the floral
receptacle and calyx tube may be synapomorphic for
Saintpauliopsis + Anisosepalum.
Flowers
As in other major lineages of Acanthaceae, floral diversity
among Nelsonioideae is particularly evident (Fig. 3). The calyx
varies from deeply four- to five-parted. Although calyces of
some species of Staurogyne have five homomorphic lobes, those
of most species are conspicuously heteromorphically lobed (1 +
2 + 2). The posterior lobe is often the largest. The two lateral
lobes are often the smallest, internal to the posterior and
anterior lobes, and obscured by the adjacent bracteoles (when
present). In most plants, the anterior lobes are connate to a
greater extent than the other calyx lobes; more rarely they are
completely fused into a single lobe or divided nearly to the base.
In Elytraria, the anterior lobes (irrespective of the degree of
fusion) show a conspicuous constriction proximally.
Corollas of all Nelsonioideae appear to share a unique,
putatively synapomorphic aestivation pattern, descendingcochlear, with the corolla lobes arranged in bud such that
the lower-central lobe is innermost (overlapped on each side by
the lateral lobes of the lower lip) and the two lobes of the upper
lip are outermost (with one overlapping the other, and each
overlapping the lateral lobes of the lower lip; see McDade et al.
2012: Fig. 1A–D). We observed this pattern in buds representing each genus of the subfamily. The corolla tube varies
from subcylindric to funnelform and is sometimes divisible
into a narrowly cylindric proximal portion and an expanded
(6 triangular) throat. Much of the floral diversity among
Nelsonioideae results from variation in the orientation and
form of the limb of the corolla. The limb is usually strongly
zygomorphic, but is nearly actinomorphic in some species (e.g.,
Elytraria caroliniensis; Fig. 3B). It consists of a bilobate upper
5
lip and a trilobate lower lip. The lips may project forward (in
line with the tube) or be reflexed at angles to 90u with respect
to the tube. Colored markings sometimes occur on some
portion of—or throughout—the limb. The variation in size,
shape, and color of corollas undoubtedly reflects adaptations
to diverse pollinators. Braz (2005) noted two groups of species
of Staurogyne based on floral morphology and suggested that
these corresponded to melitophily and ornithophily. Although
she did not provide evidence of visitation to these flowers by
bees or hummingbirds, we agree with her assessments based on
floral morphology.
Variation in features of the androecium provide important
diagnostic characters for distinguishing taxa of Nelsonioideae,
but few synapomorphic traits are evident. Two stamens are
present in Nelsonia, most Elytraria, and a few species of
Staurogyne. Four stamens characterize Anisosepalum, Malagasy species of Elytraria, Saintpauliopsis, and most species of
Staurogyne. Short and usually inconspicuous staminodes have
been noted in all genera except Nelsonia. Although up to 3
staminodes have been reported in species of Nelsonioideae,
usually only a single staminode (in posterior position) is
present. Anthers are bithecous with the thecae equally inserted
on the filament in all genera. The connective of the filament is
sometimes extended beyond the anthers and appears as an
apical appendage (e.g., in some Elytraria from the Old World).
The thecae are glabrous in all genera except Anisosepalum and
some Staurogyne, in which they are pubescent with eglandular
and/or glandular trichomes. The occasional presence of widely
divergent anther thecae (i.e., forming a right angle with the
filament/connective so that the pair is horizontally positioned
with the apex of each theca adjacent, i.e., with stamen Tshaped), as in some Nelsonia and Staurogyne, is not otherwise
common among Acanthaceae. In some Staurogyne (e.g., S.
capitata and S. macrobotrya) the thecae are apparently
divergent nearly 180u so that they appear upside down (i.e.,
with stamen Y-shaped). Basal or subbasal appendages are
found in at least some species of each genus of Nelsonioideae
except Elytraria (Fig. 4A–C). Appendages are often flaplike
and appear to be associated with dehiscence. Appendages in
Saintpauliopsis differ from those in other genera by being
bifurcate, a probable autapomorphy for the genus. Dehiscence
of the anthers appears to be nototribic (i.e., toward the lower
lip) in all genera except Nelsonia, in which it is pleurotribic
(i.e., toward each other).
The gynoecium consists of a cylindric to conic ovary with a
terminal, filiform style bearing an apical, 6 two-parted stigma.
Manifestations of the stigma, which can be difficult to observe
in some dried and pressed specimens, offer important
diagnostic characters among Nelsonioideae. Variation in
stigma form for the subfamily was illustrated by Hossain
(2004). Stigmas vary from (equally to) subequally to unequally
bilobed, or rarely crateriform (Fig. 4D). In all species of
Anisosepalum the longer of the unequal lobes is equally
bifurcate, a probable synapomorphy for the genus although
similar bifurcation of stigma lobe(s) occurs in some Nelsonia
and Staurogyne. Stigmas of Elytraria are remarkably distinctive. They consist of a broad, flattened lobe that folds over the
anthers, rapidly unfolds when touched, and slowly refolds over
several minutes. These touch-sensitive stigmas appear to play a
role in pollination, as they are located at the mouth of the
corolla or in the corolla tube, and must be touched before the
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Fig. 3. Flowers of Nelsonioideae.—A. Anisosepalum alboviolaceum (Congo-Brazzaville), photo courtesy of Chrisje Horions and Quentin
Groom.—B. Elytraria caroliniensis (USA), photo by Thomas Daniel.—C. Elytraria macrophylla (Mexico), photo by Thomas Daniel.—D.
Elytraria imbricata (Mexico), photo courtesy of Jon Rebman.—E. Elytraria marginata (São Tomé and Prı́ncipe), photo courtesy of Wes
Eckerman.—F. Nelsonia canescens (Australia), photo courtesy of Russell Cumming.—G. Nelsonia canescens (India), photo courtesy of Dinesh
Valke.—H. Nelsonia canescens (Zambia), photo courtesy of Helen Pickering.—I. Nelsonia canescens (Costa Rica), photo courtesy of Barry
Hammel.—J. Staurogyne stolonifera (South America), photo courtesy of Kris Weinhold.—K. Staurogyne flava (Brazil), photo courtesy of Mauro
Peixoto—www.brazilplants.com.—L. Staurogyne (Ophiorrhiziphyllon) macrobotrya (China), photo courtesy of Erin Tripp.
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Fig. 4. Reproductive structures (SEM) of Nelsonioideae.—A. Sagittate anther thecae of Nelsonia canescens (Daniel 11176) showing
eglandular trichomes and basal appendages.—B. Basal portion of theca of N. canescens (Daniel 11176) showing basal appendage (arrow).—C.
Subsagittate anther thecae of Staurogyne spatulata (Toroes 4795) showing flap-like basal appendages.—D. Distal portion of gynoecium of
Staurogyne guianensis showing unequal lobes of subcrateriform stigma (Jansen-Jacobs 2852).—E. Seed of Nelsonia gracilis (Bingham & Mpundu
122660) showing prominent hylar depression.
pollen from their flower is revealed. A minute and inconspicuous second lobe is often evident near the base of the broad
lobe on stigmas of Elytraria. These traits of the stigma provide
clear synapomorphies for Elytraria.
Pollen
Palynological characters have been widely employed in
classification of Acanthaceae since the late 19th century (e.g.,
Radlkofer 1883; Lindau 1895); indeed, the family apparently
exhibits more diversity in pollen morphology than any other
family of plants (Lindau 1895; Raj 1961). Phylogenetic signal
at varying taxonomic levels has been associated with pollen
sculptural characters of Acanthaceae (e.g., Scotland 1993;
Scotland and Vollesen 2000; McDade et al. 2000; Manktelow
et al. 2001; Kiel et al. 2006; Daniel et al. 2008). Published
studies comparing pollen from all currently treated genera of
Nelsonioideae are lacking.
We sampled pollen of 28 species of Nelsonioideae as follows
(see Appendix I): Anisosepalum (3), Elytraria (13), Nelsonia (2),
Saintpauliopsis (1), and Staurogyne (9). Size, shape, and
sculpturing of grains were observed in non-acetolyzed samples
of dry pollen using scanning electron and some light
microscopy. Additional data have been derived from Raj
(1961, 1973), Hossain (1971), Scotland (1990), and Braz
(2005). Descriptive terminology for pollen generally follows
that of Walker and Doyle (1975), with additional insights from
Hesse et al. (2009). Results of our observations are provided
in Appendix 1, summarized for the subfamily here, and
summarized for each genus in the accounts below.
Pollen of Nelsonioideae (Fig. 5–8) varies in shape from
oblate spheroidal to spherical to perprolate (P:E 5 0.82–2.43)
in equatorial views and from round to triangular (planaperturate) in polar view. Variation in size encompasses three size
classes, small to large. The polar axis varies from 13–58 mm,
and the equatorial axis varies from 11–36 mm. Based on
sampling to date, the smallest known grains among Nelsonioideae are in several species of Staurogyne (e.g., S. repens; P
5 13–15 mm, E 5 15–18 mm; Braz 2005); Elytraria caroliniensis
has the largest grains (P 5 56–58 mm, E 5 28–36 mm).
All grains are triaperturate. The interapertural exine is
(microreticulate to) reticulate, but in some grains (nonacetolyzed) it appears 6 psilate to microfoveolate to foveolate.
The muri of the reticulum vary from psilate to microverrucate.
Apertures of Nelsonioideae consist of elongate colpi, which
may or may not contain a centrally located endoaperture (os).
In a developmental study, Johri and Singh (1959: 233)
indicated the presence of ‘‘three germ pores’’ each situated in
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Fig. 5. Pollen of Anisosepalum and Nelsonia.—A. Anisosepalum humbertii (de Witte 7121), apertural view.—B. Anisosepalum humbertii (de
Witte 7121), subpolar view.—C. Anisosepalum alboviolaceum (Louis 12208), polar view.—D. Anisosepalum alboviolaceum (Louis 12208), subpolar
view, note two differentiated bands along margins of colpi here and in other views of Anisosepalum pollen.—E. Anisosepalum lewallii (Reekmans
3610), subpolar view.—F. Anisosepalum lewallii (Reekmans 3610), interapertural view.—G. Nelsonia canescens (Daniel 11186), apertural view.—
H. Nelsonia gracilis (Bingham & Mpundu 122660), interapertural view.—I. Nelsonia gracilis (Bingham & Mpundu 122660), apertural view.
a colpus in young pollen grains of Elytraria acaulis. Using light
microscopy, Raj (1961) determined that pollen of Elytraria (1
sp.), Nelsonia (1 sp.), and Staurogyne (3 spp.) was tricolpate. In
a subsequent examination of additional species (Raj 1973: 96),
S. spatulata (as S. glauca) was described as tricolpate, but with
‘‘indication of a faint os at each aperture,’’ and E. acaulis was
described as tricolpate. Based on acetolyzed grains, Hossain
(1971) noted that Elytraria (including E. acaulis), Saintpau-
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Fig. 6. Pollen of Elytraria.—A. Elytraria imbricata (Daniel 10184), apertural view.—B. Elytraria imbricata (Daniel 10184), interapertural
view.—C. Elytraria caroliniensis (Daniel s.n.cv-cultivated), apertural view.—D. Elytraria caroliniensis (Daniel s.n.cv), interapertural view.—E.
Elytraria marginata (Daniel 11158), apertural view.—F. Elytraria marginata (Daniel 11158), interapertural view.—G. Elytraria nodosa (Almeda
9232), apertural view.—H. Elytraria nodosa (Almeda 9232), interapertural view.—I. Elytraria macrophylla (Dressler 2619), polar view.—J.
Elytraria marginata (Daniel 11158).—K. Elytraria nodosa (Almeda 9232), polar view.
10
Daniel and McDade
ALISO
Fig. 7. Pollen of Saintpauliopsis and Staurogyne.—A. Saintpauliopsis lebrunii (Bamps 3270), interapertural view.—B. Saintpauliopsis lebrunii (Bamps
3270), apertural view.—C. Staurogyne macrobotrya (Hansen & Smitinand 12817), apertural view.—D. Staurogyne mandioccana (Cordeiro & Silva 108),
apertural view.—E. Staurogyne concinnula (Bartholomew & Boufford 6215), apertural view.—F. Staurogyne concinnula (Bartholomew & Boufford 6215),
interapertural view.—G. Staurogyne spatulata (Toroes 4795), apertural view, note prominent endoaperture.—H. Saintpauliopsis lebrunii (Gautier et al.
3598), polar view.—I. Staurogyne spatulata (Maxwell 94-76), polar view.—J. Staurogyne major (Kerr 10264), polar view, note prominent endoapertures.
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Revision of Nelsonioideae
11
Fig. 8. Interapertural exine surfaces of pollen of Nelsonioideae.—A. Anisosepalum alboviolaceum (Champluvier 5295).—B. Anisosepalum
humbertii (de Witte 7121).—C. Anisosepalum lewallei (Reekmans 3610).—D. Elytraria imbricata (Daniel 10184), note microverrucate muri.—E.
Elytraria caroliniensis (Daniel s.n.cv-cultivated), note microverrucate muri.—F. Elytraria marginata (Malaisse & Claes 14802).—G. Elytraria
nodosa (Almeda 9232).—H. Nelsonia canescens (Daniel 11186).—I. Saintpauliopsis lebrunii (Bamps 3270), note microverrucate muri.—J.
Staurogyne concinnula (Bartholomew & Boufford 6215).—K. Staurogyne guianensis (Jansen-Jacobs et al. 2346).—L. Staurogyne mandioccana
(Cordeiro & Silva 108).
liopsis, and Gynocraterium have tricolpate pollen whereas the
other genera of Nelsonioideae also possess either some—or
clear evidence of—endoapertures. He concluded that pollen of
Nelsonioideae is basically tricolpate with a trend to tricolporate (via ‘‘tricolporoidate’’) grains in several genera. Scotland
(1990) contended that in Nelsonioideae colpate pollen was
restricted to Elytraria. He characterized pollen of three species
of Elytraria as tricolpate, E. acaulis as tricolporate (but with
the endoapertures often indistinct with light microscopy),
Nelsonia canescens and Saintpauliopsis lebrunii as tricolporate,
12
Daniel and McDade
and 17 species of Staurogyne (including Gynocraterium
macrobotryum) from both the Old World and the New World
as tricolporate (but with endoapertures often indistinct).
Indeed, the indistinctness of endoapertures had led Raj
(1961) to describe some of the same species as tricolpate.
Twenty-seven Neotropical species of Staurogyne were characterized as tricolporate (Braz 2005; Braz and Monteiro 2011),
and endoapertures are evident in most of the images provided
for these taxa by these authors.
Contradictory reports for the type of apertures (colpi vs.
colpori) among taxa of Nelsonioideae undoubtedly reflect 1)
the difficulty of seeing endoapertures in grains where the
flanks of the colpi are ‘‘closed’’ and do not reveal the interior
contents (e.g., Fig. 5G, 6A,E,G, 7B,C,E), and/or 2) their rather
weak manifestation, even in ‘‘open’’ colpi compared to most
other Acanthaceae with compound apertures (at least in
mature grains; e.g., Fig. 5A–F, 7H,I). It is evident from the
studies noted above, which have used various techniques (e.g.,
acetolyzed and non-acetolyzed pollen, whole grains and
sectioned grains) and both light and scanning electron
microscopy, that acetolysis does not always help to reveal
the presence or absence of endoapertures in Nelsonioideae. It
is possible that all Nelsonioideae have endoapertures, but that
these are cryptic (or become so in later stages of development)
in some taxa. Our observations, using non-acetolyzed pollen
and both light and scanning electron microscopy, largely agree
with those of Hossain (1971). Thus, endoapertures among
Nelsonioideae vary from absent (or imperceptible) to inconspicuous to conspicuous.
The apparent taxonomic pattern of aperture type among
genera of Nelsonioideae (including colpate and colporate pollen,
which also both occur among other Acanthaceae and related
families) does not appear to be phylogenetically informative.
However, the lack of subsidiary apertures (e.g., pseudocolpi) and
other modifications (e.g., bireticulate exine, bifurcate colpi,
sexine lips) implicate Nelsonioideae (as well as Thunbergioideae,
Avicennia, and Acantheae) as likely more basal in the family.
This implication has been confirmed by several molecular
phylogenetic analyses (e.g., McDade et al. 2012).
Johri and Singh (1959) indicated that pollen of Elytraria
acaulis used in their developmental studies was syncolpate.
Pollen with colpi fused at one or both poles was not
encountered in our studies of Nelsonioideae; however, the
syncolpate condition is sometimes variable within species of
Acanthaceae (e.g., Aphelandra; McDade 1984; Daniel 1991).
Some variation in pollen sculpturing among Nelsonioideae
appears to have phylogenetic and/or taxonomic significance.
Pollen of all species of Anisosepalum differs from that of other
genera of Nelsonioideae in two features: conspicuous ridges
near margins of colpi (Fig. 5A–F), and microreticulate (i.e.,
lumina ,1 mm in diameter) exine (Fig. 8A–C). In Elytraria
caroliniensis the reticulum of the exine has much broader muri
(Fig. 6C,D, 8E) than encountered elsewhere in the subfamily.
Based on this feature and its large size, pollen of E. caroliniensis
is the most distinctive in the genus. Microverrucate muri
(Fig. 8D,E,I) occur only in Saintpauliopsis and in Elytraria from
the New World. Although microverrucate muri are present in
all pollen samples of Elytraria from the New World examined to
date, pollen has yet to be examined for any species from the
West Indies (flowers for most species there remain unknown).
All species of Elytraria from the Old World have psilate muri.
ALISO
Because phylogenetic work to date indicates that New World
species of Elytraria (inclusive of the one West Indian species
sampled) are monophyletic, we predict that West Indian species
will also have pollen with microverrucate muri.
Fruits
Fruits of Nelsonioideae, like most other Acanthaceae,
consist of indurate and loculicidal capsules that dehisce
explosively into two valves. They show only minor diversity of
size and shape among or within genera, unlike other major
lineages of Acanthaceae that encompass considerably more
variation. Nelsonioid capsules are estipitate, usually ovoid to
subconic to linear-ellipsoid in shape, fertile from the base, and
sterile apically (Fig. 9). They lack the indurate and hooklike
retinacula subtending the seeds that are synapomorphic for
subfamily Acanthoideae. The presence of papilliform retinacula
has often been noted for genera of Nelsonioideae. This
presumably refers to the discolored regions of septum where a
papilliform connection (to 0.1 mm long) to the seed sometimes
remains (Fig. 9A–D). Hossain (1972) also noted the presence of
non-indurate and caducous retinacula in Anisosepalum, but he did
not describe or illustrate these. In Anisosepalum alboviolaceum, we
observed coarse, threadlike connections (0.1–0.4 mm long) from
discolored regions of septum to ovules and seeds (Fig. 9E).
Whether these structures are homologous to the retinacula
encountered in Acanthoideae remains to be determined.
Hossain (1971, 2004) noted the rare presence of tardily
dehiscent or indehiscent capsules in some Staurogyne, and he
noted that this characteristic had been the basis for the
African genus Staurogynopsis. Champluvier (1991) described
the African species of Staurogyne section Zenkerina as having
capsules with a membranous (vs. indurate in other species)
wall and both delayed and non-explosive dehiscence. Tardily
dehiscent or indehiscent capsules are also known elsewhere in
the family in genera with otherwise regularly dehiscent
capsules (e.g., Justicia; Hedrén 1989), and likely represent a
specialization that has occurred numerous times in the family.
Seeds, Hygroscopic Trichomes, and Endosperm
Ovules and seeds among Nelsonioideae tend to be more
numerous than elsewhere in the family. The number of seeds
that develop per capsule varies from six (e.g., Elytraria spp.,
Staurogyne carvalhoi) to 68 (e.g., Staurogyne spp.). Mature
seeds are generally smaller than in other Acanthaceae. The
longest axis varies from 0.3 to 2 mm, but in most species is less
than 1 mm. Anisosepalum has the largest seeds, with the
longest axis varying from 1.1 to 2 mm. Dimensions of seeds
that have been expelled from the mature capsule can be
difficult to attribute to length, width, or height because the
hylar scar is not always evident and the seeds are usually
irregular in shape (frequently somewhat blocky to oblong to
ellipsoid to subhemispheric to cochlear). However, the hylar
scar forms a conspicuous depression in Anisosepalum and to a
lesser extent in Nelsonia (Fig. 4E). The surface of seeds varies
from smooth to reticulate to foveolate, and sometimes includes
minute protrusions (e.g., verrucae, gemmae, baculae, etc.;
Fig. 10–12). The presence or absence of seminal trichomes was
found to be useful in characterizing the five genera of
Nelsonioideae. They are apparently universal in all genera
except Elytraria and Anisosepalum, in which they do not occur.
VOLUME 32, NUMBER 1
Revision of Nelsonioideae
13
Fig. 9. Capsules and seeds of Nelsonioideae.—A. Capsule valve of Elytraria caroliniensis (Cooley et al. 9135) showing darkened funicular
regions where nine seeds were attached.—B. Capsule valve of Staurogyne merguensis (Maxwell 94-234) showing darkened funicular regions where
24 seeds were attached (with one aborted seed remaining).—C. Capsule valve of Staurogyne obtusa (Maxwell 90-362) showing both unejected
seeds and funicular regions of ejected seeds.—D. Capsule valve of Elytraria marginata (Daniel 11158 cv-cultivated) showing slight funicular
elongations from some funicular regions.—E. Anisosepalum alboviolaceum (Champluvier 5295) showing five immature seeds with non-indurate
funicular elongations below each.—F. Dry seed of Nelsonia canescens (Daniel 11186) with hygroscopic trichomes appressed.—G. Portion of
hydrated seed of N. canescens (Daniel 11186) showing strong hygroscopic reaction of trichomes.—H. Portion of hydrated seed of S. concinnula
(Peng 6777) showing moderate hygroscopic reaction of trichomes.—I. Dry seed of Staurogyne concinnula (Peng 6777) with hygroscopic trichomes
appressed. See text for details.
Seminal trichomes of Nelsonioideae are simple or usually
branched; the number of branches per trichome can vary on a
seed. The trichomes are also hygroscopic.
Hygroscopic trichomes in Acanthaceae are usually appressed to the seed surface and expand and/or become erect
on being wetted with water; they often discharge mucilage.
Such trichomes have long been known among various
Acanthaceae (Kippist 1845). They occur on seeds of some
taxa in two of the three currently recognized subfamilies
(Nelsonioideae and Acanthoideae) and in six of the seven
tribes of Acanthoideae: Ruellieae (Tripp et al. 2013), Justicieae
(e.g., Daniel 1990; Daniel and Figueiredo 2009), Barlerieae
(Manktelow et al. 2001; McDade et al. 2008), Acantheae
(Vollesen 2000; McDade et al. 2005), Whitfieldieae (Manktelow et al. 2001; McDade et al. 2008), and Neuracantheae
(Bidgood and Brummitt 1998; McDade et al. 2008). Whether
hygroscopic trichomes in all of these taxa are homologous
remains to be determined. To our knowledge, hygroscopic
trichomes are not known to occur on seeds of Avicennia,
Thunbergioideae, or Acanthoideae: Andrographideae.
Hygroscopic trichomes on seeds of Nelsonioideae were
illustrated by Oersted (1855) and Lindau (1895). In this
subfamily apically hooked, double-hooked (i.e., anchor
shaped), or multi-branched trichomes that lack copious
mucilage are appressed to the dry seed surface (Fig. 9F,I,
11G–I, 12); these usually become erect (and clearly visible)
upon hydration (Fig. 9G,H). Kippist (1845: 73) noted that
whereas trichomes of Nelsonia ‘‘expand very slowly when
wetted,’’ in two species now treated in Staurogyne, a similar
reaction to water was inconclusive ‘‘owing to [the trichomes’]
extreme minuteness, and the very slight action which water has
upon them.’’ Our results (Appendix 2) were similar. Trichomes
14
Daniel and McDade
ALISO
Fig. 10. Seeds of Anisosepalum and Saintpauliopsis.—A–C. Anisosepalum alboviolaceum (Champluvier 5295, CAS).—A. Seed.—B–C. Closeups of surface.—D–F. Anisosepalulm humbertii (Raynal 20630, K).—D. Seed.—E–F. Close-ups of surface.—G–I. Saintpauliopsis lebrunii
(Ndangalasi 0449, C).—G. Seed.—H–I. Close-ups of surface.
are not present on seeds of Anisosepalum or Elytraria based
on SEM observations (Fig. 10A–F, 11A–F) and hydration
studies. Trichomes were observed on seeds in all species of
Nelsonia, Saintpauliopsis, and Staurogyne examined. Most or
all trichomes on seeds of some samples of Nelsonia and
Staurogyne became erect en masse immediately and conspicuously (i.e., trichomes to 0.1 mm long) on hydration (Fig. 9G).
Less dramatic reactions observed in other samples of Nelsonia,
Saintpauliopsis, and Staurogyne included: some or most of the
trichomes expanded, but the trichomes are short (,0.05 mm
long, e.g., Staurogyne obtusa; Fig. 9H) and inconspicuous,
and the reaction was not immediate (e.g., both samples of
Saintpauliopsis lebrunii); only some of the trichomes became
erect after a minute or more (e.g., some Nelsonia canescens);
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Revision of Nelsonioideae
15
Fig. 11. Seeds of Elytraria and Nelsonia.—A–C. Elytraria acaulis (Iwarsson 664, UPS).—A. Seed.—B–C. Close-ups of surface.—D–F.
Elytraria bromoides (Daniel 3683, CAS).—D. Seed.—E–F. Close-ups of surface.—G–I. Nelsonia canescens (Lewalle 3921, UC).—G. Seed.—H–I.
Close-ups of surface (note apically uncinate, anchor-shaped and appressed hygroscopic trichomes).
and no apparent reaction of the trichomes within three
minutes (e.g., some seeds of Staurogyne spp.). Thus not all
of the seminal trichomes were always reactive. Although
hygroscopy was observed in all species of Staurogyne tested, all
trichomes on some, but not all, seeds showed no reactivity.
Possible causes of non-reactivity include: age of seed, method
of preservation of seed, immaturity of seed, and innate lack of
reactivity in some trichomes.
Bremekamp (1953) noted well developed endosperm in
Nelsonioideae, and it was largely based on this trait that he
considered these plants more closely related to Scrophulariaceae than Acanthaceae (Bremekamp 1955). Hossain (1971)
characterized endosperm of Nelsonioideae as usually persistent, oily, and alveolate. Borg et al. (2008) indicated that the
presence of persistent (vs. non-persistent) endosperm was one
of the characteristics by which the subfamily differs from other
16
Daniel and McDade
ALISO
Fig. 12. Seeds of Staurogyne.—A–C. Staurogyne miqueliana (Lott et al. 3226, CAS).—A. Seed.—B–C. Close-ups of surface (note appressed
hygroscopic trichomes).—D–F. Staurogyne spatulata (Clemens & Clemens 4006, UC).—D. Seed.—E–H. Close-ups of surface (note appressed
hygroscopic trichomes).—G–I. Staurogyne macrobotrya (Poilane 20703, P).—G. Seed.—H–I. Close-ups of surface (note short
hygroscopic trichomes).
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Revision of Nelsonioideae
Acanthaceae. Several species of Andrographis (Acanthoideae:
Andrographideae) have both persistent and ruminate endosperm, like that described for species of Elytraria and Nelsonia
(Mohan Ram and Wadhi 1965), and the presence of one or
more layers of persistent endosperm in mature seeds was noted
for diverse taxa throughout the family by Mohan Ram and
Wadhi (1964). However, as noted by Scotland and Vollesen
(2000), this trait remains poorly documented in the family and
merits further study.
CHROMOSOME NUMBERS
Knowledge of chromosome numbers has been helpful in
understanding the systematics, evolution, and biogeography of
some taxa of Acanthaceae (e.g., Daniel 2000, 2006; McDade
et al. 2000, 2005). To date, chromosome numbers have been
reported for seven species of Elytraria (Appendix 3). Among
them, the three Mexican taxa sampled have relatively low
(presumably diploid) numbers of n 5 (11) 12 whereas taxa
from the southeastern United States and the Old World have
higher (presumably polyploid) numbers of n 5 (17) 19, 22, 23,
and 25. The only other genus of Nelsonioideae for which
counts have been reported is Nelsonia (Appendix 3), with all
counts (n 5 14, 16–18) presumably attributable to N.
canescens. It remains to be seen how useful chromosome data
will be for understanding patterns of relationships among these
taxa, but the diversity of numbers reported for the small
sample of Elytraria is encouraging. Additional counts for
species of Elytraria and Nelsonia, and initial counts for all
other genera of Nelsonioideae are highly desirable.
DISTRIBUTIONS, BIOTIC COMMUNITIES, AND BIOGEOGRAPHY
The distribution of Nelsonioideae is nearly pantropical with
incursions into temperate regions in North America (northern
Mexico, southern United States to ca. lat. 33uN), South
America (southeastern Brazil), Africa/Madagascar (northern
South Africa to ca. lat. 27.5uS, southern Madagascar), and
Asia (China, India, Japan-southern Ryukyu Islands, Nepal).
The subfamily does not appear to be present on tropical
Pacific islands located east of the Solomon Islands. All genera
occur in Africa, with Anisosepalum restricted to that continent
and Saintpauliopsis occurring only in Africa and Madagascar.
Three genera (Elytraria, Nelsonia, Staurogyne) also occur in
Asia, with Nelsonia and Staurogyne also distributed eastward
to Australia. Two genera (Elytraria and Staurogyne) are native
to—and have radiated in—the New World. The native range
of Nelsonia is not known with certainty; the genus may not be
native to the New World, but the non-native range of N.
canescens there appears to be expanding (cf. Franck and
Daniel 2012).
A comparative study of biotic communities in which
Nelsonioideae occur has not been attempted based on the
information available in floristic accounts and on herbarium
specimens. However, it is clear from those sources and from
our own field observations that some species occur in very wet
communities (e.g., wet evergreen forest) and others occur in
dry communities (e.g., tropical deciduous forest). Species of
Anisosepalum and Saintpauliopsis appear to occur only in wet
evergreen forests. Plants treated herein as Nelsonia canescens
occur in a wide range of biotic communities from wet
evergreen forest (in western Africa) to grasslands (in Southern
17
Africa) to arid scrub (in southern Madagascar). Most species
of Staurogyne appear to occur in moist to wet communities
(e.g., all African species occur in wet evergreen forest), but
Mexican plants of S. miqueliana (Daniel and Lott 1993; as S.
agrestis) were collected along a dry streambed in a region of
dry forest (tropical subdeciduous forest). Elytraria contains
species that are restricted to wet evergreen forest (e.g., E.
marginata) and others that occur only in seasonally arid
tropical deciduous forest (e.g., E. mexicana). The widespread
E. imbricata occurs in diverse biotic communities from desert
to mesophytic montane forest and at elevations from sea level
to 2200 m.
Biogeographic analyses of Elytraria (Wenk and Daniel 2009;
McDade et al. 2012) optimized Africa as the area of origin for
the genus with dispersals to Madagascar, the Indian subcontinent, and the New World. At least three subsequent dispersal
events within the New World could account for the current
distribution of species of Elytraria there. Wenk and Daniel
(2009) proposed rafting as the mode of dispersal for seeds or
plants of the genus between landmasses. Dispersal by rafting
may be most plausible for genera that lack the apically hooked
and hygroscopic trichomes (e.g., Elytraria), which possibly
play a role in zoochoric dispersal. Saintpauliopsis + Anisosepalum retain the symplesiomorphic African distribution, with
dispersal to Madagascar within Saintpauliopsis lebrunii.
The richest genus of Nelsonioideae, Staurogyne, apparently
dispersed to the New World early in its evolutionary history as
the New World clade is sister to a clade comprising all other
species. A subsequent dispersal from Africa to Asia resulted in a
major radiation that is today represented on the southeastern
mainland of that continent and islands to the east. Interestingly,
only five species are present in Africa today, far fewer than in
the New World or in Asia. Whether this represents a decrease in
richness over time due to climatic changes (e.g., increasing
aridity) in Africa or relatively little speciation during its history
on that continent remains an interesting question.
There have been at least three colonizations of Madagascar
by Nelsonioideae. Presumably because of Madagascar’s
lengthy isolation from Africa (separation of Indo-Madagascar
began ca. 165 mybp; Krause 2003), there are relatively few
species of Acanthaceae common to both landmasses (Benoist
1967; Daniel unpubl. data). It is therefore of interest that
whereas Elytraria is represented on the island by two very
distinctive endemic species, the other genera of Nelsonioideae
are each represented by a species that also occurs in Africa:
Nelsonia canescens and Saintpauliopsis lebrunii. We hypothesize that future phylogenetic work to date these dispersal
events will show that Elytraria arrived considerably earlier that
the two other Nelsonioideae.
FUTURE STUDIES
With Braz’s (2005) account of the American species and
Champluvier’s (1991) treatment of the African species of
Staurogyne, a modern revision of the Asian species of that
genus remains the most urgent taxonomic priority among
Nelsonioideae. Bremekamp’s (1955) detailed study of the
Malesian species serves as a useful starting point in this regard.
Additional molecular studies of Nelsonia on a worldwide basis
will likely be useful in resolving both the number and rank of
taxa that would be appropriate to recognize in that genus.
18
Daniel and McDade
Such studies linked to a time-calibrated phylogeny should
determine whether putatively native populations of the genus
in the New World are indeed pre-Columbian. Collections of
Elytraria with flowers and fruits from Cuba and Haiti are
much desired to better resolve the taxonomy of the genus in
the West Indies.
TAXONOMY
The following account is intended to delimit monophyletic
genera of nelsonioids, to characterize each one, and to tabulate
currently recognized species. It provides the known geographic
distribution (by country), synonyms, and relevant taxonomic
or nomenclatural notes for each genus and species.
NELSONIOIDEAE Pfeiff., Nomencl. Bot. 1(1): 10 (1871) (as
‘‘Nelsonieae’’). Nelsonieae Nees in N. Wallich, Pl. Asiat.
Rar. 3: 74 (1832). Nelsoniinae Bremek., Recueil Trav. Bot.
Néerl. 35: 134 (1938). Nelsoniaceae Sreem., Phytologia 37:
412 (1977).—TYPE. Nelsonia R. Br.
Elytrarieae Dumort., Anal. Fam. Pl. 23 (1829).—TYPE.
Elytraria Michx.
Annual or perennial, acaulescent to caulescent, and terrestrial (to epiphytic or epipetric) herbs or shrubs, cystoliths
absent. Leaves opposite or whorled or alternate, sometimes
clustered at ground level or at apex of branches or along stems,
sessile to petiolate, margin of blade entire to sinuate-crenate to
shallowly dentate to pinnately lobed. Inflorescence of dichasia
in leaf axils or more commonly consisting of sessile or
pedunculate, axillary or terminal dichasiate spikes or racemes
or thyrses (or if branched then panicles of these), peduncles
naked or with alternate (spirally disposed) clasping scales;
dichasia opposite or alternate (sometimes spirally disposed),
sessile or pedunculate, 1-flowered, subtended by a (leaf or)
bract. Bracts subfoliose or usually greatly reduced in size
relative to leaves, not overlapping to densely imbricate, borne
along rachis or sometimes adnate at base to dichasial peduncle
and becoming free along peduncle. Flowers sessile to
pedicellate, homostylous, flower or flower + pedicel subtended
by paired bracteoles (except in most Nelsonia). Bracteoles of a
pair homomorphic, sometimes with basal portion partially
adnate to calyx tube. Calyx deeply 4–5-parted, lobes heteromorphic (1 + 2 + 2) or rarely subequal, posterior lobe often
largest, lateral lobes usually shortest, interior to posterior and
anterior lobes, and often obscured by bracteoles, anterior lobes
sometimes longest and sometimes connate to a greater extent
than other lobes. Corolla entirely white or variously colored,
tube subcylindric to funnelform, narrow proximal portion 6
cylindric, throat (if present) expanded toward mouth of
corolla, limb (subactinomorphic to) strongly zygomorphic (2labiate), upper lip entire or 2-lobed, lower lip 3-lobed, central
lobe usually largest, corolla lobes with descending cochlear
aestivation. Stamens 2 or 4, if 4 then didynamous (or if 6
equal in length then inserted at different heights), included in
or exserted from mouth of corolla, anthers not surpassing lips
or extended well beyond lips, 2-thecous, thecae dehiscing
toward lower lip (i.e., flower nototribic) or toward each other
(i.e., flower pleurotribic), those of a pair equally inserted, with
or without a basal appendage; pollen oblate spheroidal to
spherical to perprolate (P:E 5 0.82–2.43), round to triangular
ALISO
in polar view, 3-colpate (endoapertures, if present, not evident)
to 3-colporoidate (endoapertures evident but indistinct) to 3colporate (endoapertures distinct), exine reticulate (sometimes
appearing foveolate in non-acetolyzed grains), muri smooth or
microverrucate; staminodes 0–2 (–3). Style included in or
exserted from mouth of corolla. Capsule estipitate, ovoid to
subconic to linear-ellipsoid, lacking retinacula. Seeds (6–) 12–
68 per capsule, variable in shape (often 6 blocky or irregularly
shaped), hygroscopic trichomes present or absent, endosperm
present.
We recognize five genera with 172 species in the subfamily.
They are nearly pantropical in distribution, and occur to a
lesser extent in temperate regions. All genera occur in Africa;
Elytraria, Nelsonia, and Staurogyne are also native to Asia;
and at least Elytraria and Staurogyne are native in the New
World.
Key to Genera of Nelsonioideae
1. Peduncles of inflorescence (if present) bearing alternate
(spirally arranged) clasping scales; bracts coriaceous to
sclerophyllous; abaxial surface of calyx lobes with veins
obscured or not evident; seeds lacking hygroscopic
trichomes; stigma inconspicuously and scarcely divided (if
at all), folded over and enclosing anthers in undisturbed
flowers, unfolding and becoming erect when touched . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . Elytraria
1. Peduncles of inflorescences (if present) lacking clasping
scales; bracts membranaceous; abaxial surface of calyx
lobes with veins evident to prominent; seeds (except for
Anisosepalum) with hygroscopic trichomes; stigma usually
conspicuously 2 (or more)-lobed, neither folded over nor
enclosing anthers, not touch-sensitive.
2. Bracteoles absent; inflorescence of dense, cylindric spikes
5–14 mm in diameter near midpoint (excluding corollas),
with bracts imbricate and corollas 2.5–7 mm long;
androecium of 2 stamens, staminodes absent; bracts
borne on inflorescence rachis only . . . . . . . . . Nelsonia
2. Bracteoles present; inflorescence not with the combination of characters noted above; androecium of 4
stamens (rarely with 2 stamens in some Staurogyne),
staminodes absent or 1 (–3); bracts borne on inflorescence rachis or fused to and emerging from dichasial
peduncle along its length or at its apex.
3. Anther thecae with a bifurcate (forked) appendage
at base; petiole usually longer than blade of leaves;
corolla 6 pendant and appearing 6 campanulate;
bracts adnate to (and indistinguishable from)
dichasial peduncles for 91–98% the length of the
peduncle (emerging near apex of peduncle); pollen
with muri of reticulate exine beset with microverrucae . . . . . . . . . . . . . . . . . . Saintpauliopsis
3. Anther thecae lacking an appendage or (rarely)
with a short, non-forked subbasal or basal appendage; petiole usually shorter than blade of leaves;
corolla neither pendant nor campanulate; bracts
borne on inflorescence rachis or adnate to (and
distinguishable or indistinguishable from) dichasial
peduncle for 11–100% the length of the peduncle
(usually emerging well below apex of peduncle);
pollen with muri of reticulate exine smooth.
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Revision of Nelsonioideae
4. Base of bracteoles adnate (for up to 2 mm) to
floral receptacle and sometimes base of calyx
(or sometimes free in A. lewallei); corolla with a
conspicuous bullate palate on lower lip; seeds
with longest axis 1.1–2 mm, shortest axis 0.9–
1.2 mm, lacking trichomes . . . Anisosepalum
4. Base of bracteoles not adnate to floral receptacle or calyx; corolla lacking a palate on lower
lip; seeds with longest axis 0.3–1 mm, shortest
axis 0.2–0.8 mm, pubescent with hygroscopic
trichomes . . . . . . . . . . . . . . . . . Staurogyne
ANISOSEPALUM E. Hossain, Notes Roy. Bot. Gard. Edinburgh
31: 377 (1972).—TYPE. Anisosepalum humbertii (Mildbr.)
E. Hossain (Staurogyne humbertii Mildbr.)
Caulescent, erect to spreading perennial herbs or shrubs
(sometimes epiphytic) to 1.5 (–3) m long or tall. Young stems
subterete to subquadrate to quadrate-sulcate or becoming 6
flattened on drying, internodes glabrous or pubescent with
eglandular trichomes or glabrate. Leaves opposite or ternate,
6 evenly disposed along younger stems, subsessile to petiolate,
petiole shorter than blade, margin of blade entire to sinuate to
sinuate-dentate. Inflorescence usually of subsessile to pedunculate loose to dense terminal raceme-like thyrses, peduncles
lacking scales, fertile portion of spike neither dense nor
cylindric or dense and 6 cylindric, 9–30 mm in diameter
(excluding corollas) near midpoint, not branching, rachis
clearly visible to not visible; dichasia (subopposite to)
alternate, pedunculate. Bracts (subopposite to) alternate, not
imbricate to 6 densely imbricate, green, membranaceous,
basal portion adnate to dichasial peduncle for 50–96% the
length of the peduncle, free portion linear to elliptic to
oblanceolate to obovate or subfoliose, 4.3–17 mm long, 1–
4 mm wide, 3-veined. Bracteoles borne at base of flower and
usually basally adnate to receptacle and calyx tube for up to
2 mm (sometimes free in A. lewallei) and appearing much like
the longer calyx lobes, each positioned exterior to and usually
obscuring the short lateral calyx lobes, green to at least
partially hyaline, subulate to lanceolate, 3.1–16.5 mm long,
0.3–2 mm wide. Flowers sessile (i.e., pedicels absent). Calyx
deeply 5-lobed, 3.5–19 mm long, green to hyaline, accrescent
and becoming somewhat brownish and bony at or following
fruiting, lobes heteromorphic (1 + 2 + 2), posterior lobe 2.5–
15 mm long, 3-veined (at least basally), lateral lobes shortest,
1.1–9 mm long, usually #K the length of anterior lobes, 1veined, anterior lobes longest, 3.5–18 mm long, 3-veined (at
least basally). Corolla dark reddish or greenish white with
brownish pink markings or white with purplish markings or
blue or purplish or maroon internally and pale purple with
violet striations externally, 6.5–39 mm long, externally
pubescent with stipitate glandular (sometimes sparse) and/or
eglandular trichomes or glabrous, tube cylindric for K or more
of its length, expanded into a throat for distal J–K its length,
limb 2-labiate, upper lip entire to shallowly bifid, lower lip
with a conspicuous bullate palate proximally. Stamens 4,
didynamous, inserted at or distal to base of throat, 1 or both
pairs exserted from mouth of corolla, anthers not extending
beyond lips, thecae dehiscing toward lower lip (i.e., flower
nototribic), elongate, ellipsoid, 0.7–2 mm long, those of a pair
19
subsagittate to sagittate, equal in length, dorsally pubescent
with flexuose eglandular and glandular trichomes, each with a
subtriangular and flaplike basal appendage; staminode 0–1;
pollen oblate spheroidal to spherical to euprolate (P:E 5 0.90–
1.36; polar axis 22–31 mm, equatorial axis 22–32 mm), 3-colpate
to 3-colporoidate, exine microreticulate (or appearing 6
psilate to microreticulate to microfoveolate in A. alboviolaceum), muri smooth. Style exserted from mouth of corolla, not
extending beyond lips of corolla, stigma not touch-sensitive,
subequally to unequally 2-lobed, lobes linear (to inrolled), 0.3–
2 mm long, 0.2–0.4 mm wide, longer lobe equally 2-furcate at
apex into lobes 0.1–0.7 mm long. Capsule ovoid, 4–13.5 mm
long, glabrous or sometimes glandular punctate. Seeds mostly
12–16 per capsule, oblong to ellipsoid to subhemispheric,
longest axis 1.1–2 mm, shortest axis 0.9–1.2 mm, deeply
excavated on one side, surface coarsely foveolate- to rugosereticulate, microverrucate to microgemmate, and usually
microreticulate as well, lacking trichomes; (Fig. 13).
Three species from tropical Africa (Fig. 14) are recognized
in this genus. All of them are well collected and morphologically documented. Detailed taxonomic accounts of the genus
were provided by Champluvier (1991) and Vollesen (2008).
The two species initially treated by Hossain (1972: 378) were
segregated from Staurogyne based on differences in the calyx
(‘‘form of calyx … approaches Saintpauliopsis’’), corolla
(palate on lower lip), number of ovules (6 to 8 vs. 12 or
more), and seeds (with a large hilar excavation and rugose
testa, and lacking ‘‘hooks on their testa cells’’). We recognize
Anisosepalum as morphologically distinct among Nelsonioideae by a unique combination of characters: bracteoles fused
to the base of the calyx (like Saintpauliopsis), corollas with a
conspicuous palate (likely synapomorphic), thecae with an
undivided and flaplike basal appendage (like Nelsonia), seeds
lacking hygroscopic trichomes (like Elytraria) and with a
conspicuous and deep hylar excavation (likely synapomorphic), and pollen with conspicuous ridges near the colpal
margins (synapomorphic).
Currently Recognized Species of Anisosepalum
1. Anisosepalum alboviolaceum (Benoist) E. Hossain (Central
African Republic, Congo-Brazzaville, Congo-Kinshasa,
Gabon, Rwanda, Uganda, Zambia)
Staurogyne alboviolacea Benoist; synonyms fide Vollesen
(2008, 2013): Anisosepalum alboviolaceum subsp. grandiflorum
(Napper) E. Hossain, Staurogyne alboviolacea subsp. grandiflora Napper
Champluvier (1991) recognized two subspecies, alboviolaceum and gracilius (Heine) Champluvier, based on habit and
pubescence of the external surface of the corolla. Within subsp.
alboviolaceum she treated five informal groups based on
geographic distribution and differences in shape, size, and
pubescence of the corolla. Vollesen (2008) noted variation in
size of the corollas in different parts of the geographic range of
the species, but he did not recognize formal infraspecific taxa;
he also did not specifically include subsp. gracilius as a
synonym. However, he did include the distribution of subsp.
gracilius within the geographic range of the single taxon
recognized.
20
Daniel and McDade
ALISO
Fig. 13. Anisosepalum lewallei.—A. Habit (Bidgood et al. 4570; Tanzania).—B. Inflorescence node (Bidgood et al. 4570).—C. Calyx subtended
by paired bracteoles (on sides) and bract (background) (Bidgood et al. 4728; Tanzania).—D. Flower (Bidgood et al. 4570).—E. Apex of stamen
with anthers (Bidgood et al. 4570).—F. Apex of style with stigma (Bidgood et al. 4570).—G. Capsule (exterior view on left, interior view on right)
(Bidgood et al. 4728).—H. Seed (Bidgood et al. 4728). Drawn by S. Adler.
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Revision of Nelsonioideae
21
Fig. 14. Map of Africa showing the distribution by country of Anisosepalum.
2. Anisosepalum humbertii (Mildbr.) E. Hossain (Burundi,
Congo-Kinshasa, Rwanda, Tanzania, Uganda, Zambia)
minor and not mutually exclusive), but their geographic ranges
apparently do not overlap.
Staurogyne humbertii Mildbr.
Champluvier (1991) recognized two subspecies: humbertii
and zambiense Champluvier. These were based on position of
the ring of trichomes inside the corolla tube and the width of
the corolla tube below the lips. Vollesen (2008, 2013)
maintained these subspecies and noted that subsp. zambiense
has a longer corolla tube with the ring of hairs on its internal
surface located more distally. Based on our observations, the
subspecies appear very similar (the differences being relatively
3. Anisosepalum lewallei P. Bamps (Burundi, Tanzania)
This is a highly distinctive species with generally ternate
leaves. Although opposite leaves occur, at least some
nodes (usually most) per shoot bear three leaves. Corollas
of this species (3–3.9 cm long; Fig. 13D) are the largest in the
genus.
ELYTRARIA Michx., Fl. Bor.-Amer. 1: 8 (1803), nomen
conserv.—TYPE. Elytraria virgata Michx., nomen illegit.
22
Daniel and McDade
(E. caroliniensis (Walter ex J.F. Gmel.) Pers.; Tubiflora
caroliniensis Walter ex J.F. Gmel.)
Anonymos Walter, Fl. Carol. 91 (1788), excluded name
(McNeill et al. 2012: Article 20.4(a)).—TYPE. Anonymos
caroliniensis Walter
Tubiflora J.F. Gmel., Syst. Nat., ed. 13[bis]. 2(1): 27 (1791),
nomen rej.—TYPE. Tubiflora caroliniensis J.F. Gmel.
Acaulescent to caulescent, erect to spreading annual or
perennial herbs to shrubs to 0.5 (–1) m tall. Young stems
subterete to 3-angulate to subquadrate, sometimes 6 flat on
drying, glabrous, pubescent with eglandular trichomes, or
glabrate. Leaves alternate or whorled, 6 diffuse along stems or
borne in basal rosettes (usually from a subterranean caudex or
rhizome) or aerial whorls or aerial clusters (up to 11 leaves per
cluster), sessile to petiolate, petiole (if present) usually shorter
than blade, margin of blade entire to crenate to pinnatifid.
Inflorescence of (sessile to) pedunculate axillary and terminal
densely bracteate spikes, peduncles (if present) covered with
imbricate or remote coriaceous clasping scales, fertile portion
of spike cylindric, 1.8–17 mm in diameter (excluding corollas)
near midpoint, simple or sometimes branched, rachis not or
only partially visible; dichasia alternate (spirally disposed),
sessile. Bracts alternate (spirally disposed), imbricate, green or
partially hyaline, not fused to other structures, coriaceous to
sclerophyllous, lanceolate to ovate to broadly ovate to elliptic
to hour-glass shaped to obovate, 2.5–11 mm long, 1–5.8 mm
wide, 1- or 3-veined or veins not evident, sometimes apically
toothed and/or winged. Bracteoles borne at base of (but not
adnate to) flower, positioned near edges of posterior calyx lobe
and 6 conduplicate there, often mostly hyaline, narrowly
elliptic to linear to lanceolate to lance-lunate to subulate, 1.6–
7.5 mm long, 0.2–1.7 mm wide. Flowers sessile. Calyx usually
appearing deeply 4-lobed, mostly hyaline, 1.8–8.5 mm long,
lobes heteromorphic, 6 equal in length, posterior lobe 1.8–
7.5 mm long, lateral lobes 1.7–7 mm long, anterior lobe 1.9–
8 mm long, 2-dentate to 6 deeply 2-cleft from apex, all lobes
obscurely veined abaxially. Corolla pinkish or blue or yellow
or entirely white or dark purple and white, sometimes with
colored markings near mouth, 3–17 mm long, externally
glabrous or pubescent, tube cylindric or slightly expanded near
mouth, throat indistinct or evident only near mouth, limb
(subactinomorphic to) 2-labiate, upper lip 2-lobed, lower lip
lacking a bullate palate, corolla lobes often apically divided or
2-cleft. Stamens 2 (or 4 in species native to Madagascar),
inserted at/near apex of corolla tube or near midpoint of tube,
anthers included in corolla tube or partially exserted from
mouth of corolla, not extending beyond lips of corolla, thecae
(covered by folded stigma during anthesis) dehiscing toward
lower lip (i.e., flower nototribic), oblong to broadly ellipsoid,
0.3–1.7 mm long, those of a pair parallel, equal to subequal in
size, glabrous, lacking basal appendages, connective sometimes
with an apical appendage (in some Old World spp.);
staminodes 0–2, minute; pollen suboblate to spherical to
perprolate (P:E 5 0.82–2.43; polar axis 25–58 mm, equatorial
axis 14–36 mm), 3-colpate to 3-colporoidate, exine reticulate
(sometimes appearing 6 foveolate in non-acetolyzed grains),
muri psilate or microverrucate. Style included in corolla tube
or if exserted from mouth of corolla then not extending
beyond lips of corolla, stigma folded over anthers during
ALISO
anthesis and straightening when touched (touch-sensitive) and
gradually refolding, appearing non-lobed but usually with a
minute and inconspicuous lobe (to 0.3 mm long) borne on an
expanded, flat, subelliptic to subspatulate lobe (rarely subcrateriform), 0.4–2.5 mm long, 0.2–1.2 mm wide. Capsule
conic to ovoid to linear-ellipsoid, sometimes irregularly
constricted proximally, 2.1–9 mm long, glabrous (rarely
inconspicuously glandular punctate, but lacking elongate
trichomes). Seeds to 26 per capsule, irregularly shaped (often
blocky or reniform or ellipsoid), longest axis 0.3–1.7 mm,
shortest axis 0.3–1.5 mm, surface (smooth to) lumpy- to
rugose- to foveolate-reticulate, microverrucate to microbaculate and with smaller rounded to pointed protrusions that
sometimes form chains or a microreticulum, lacking trichomes;
x 5 11 or 12?; (Fig. 15).
The genus consists of 21 species occurring in the tropics,
subtropics, and warm deserts of both the Old World and the
New World (Fig. 16). The majority of species are American
(14), but species also occur in Africa (5), Madagascar (2), and
the Indian subcontinent (1). Except for the naturalized
presence of weedy Elytraria imbricata in Madagascar, southeastern Asia, and Malesia, there are no species common to the
Old World and the New World. North American and African
species are well collected and fully documented morphologically (e.g., Morton 1956; Dokosi 1970, 1971, 1979; Daniel and
Acosta C. 2003; Vollesen 2008; Daniel in press). Species
endemic to South America and Madagascar are poorly
collected but reasonably well known (e.g., Benoist 1967; Wenk
2008). The seven species endemic to the West Indies remain
very poorly collected, morphologically documented, and
taxonomically resolved; indeed, corollas remain unknown for
most of those species (e.g., Borhidi and Muñiz 1978). A
monograph of the entire genus is currently in preparation.
Hossain (1972) recognized two subgenera: Tetrandra E.
Hossain consisting of the two species endemic to Madagascar
(characterized by woody habit, whorled leaves, and four
stamens), and Elytraria consisting of all other species
(characterized by mostly herbaceous habit; alternate, whorled,
or clustered leaves; and two stamens). Molecular phylogenetic
results (Wenk and Daniel 2009; McDade et al. 2012) reveal
that the subgenera are not monophyletic.
Elytraria is distinctive among Nelsonioideae by having
peduncles (absent only in E. madagascariensis) covered with
alternate (spirally arranged) scales, coriaceous to sclerophyllous bracts, and touch-sensitive stigmas. All three traits appear
to be synapomorphic for the genus. Another significant
character of Elytraria is the absence of minute, hooklike,
and hygroscopic trichomes on its seeds (shared with Anisosepalum). Variation in several morphological attributes is
notable for the genus. Habit varies from acaulescent herbs
from a woody caudex or rhizome (most species) to monocaulous or branched herbs (e.g., E. marginata, E. imbricata) to
shrubs (e.g., E. nodosa). Most species have either a taproot or
fibrous roots, but one (E. tuberosa) has fibrous roots that bear
elongate and conspicuously swollen tubers. Tuberous roots are
otherwise unknown among Nelsonioideae, but they are known
elsewhere among Acanthaceae (e.g., Ruellia tuberosa). Leaf
margin varies (sometimes intraspecifically) from entire to
crenate to lobed to subpinnatisect. Floral diversity in Elytraria
encompasses size, color, and form of the limb of corollas. The
West Indian endemic, E. shaferi, has the smallest known
VOLUME 32, NUMBER 1
Revision of Nelsonioideae
23
Fig. 15. Elytraria caroliniensis (Daniel s.n.cv, cultivated from plants from the USA, except as noted).—A. Habit (based on Daniel s.n.cv and
Curtis 5947; USA).—B. Leaf (Wiggins 19812; USA).—C. Inflorescence with flower (bract, bracteoles, and calyx subtending flower removed).—D.
Bract.—E. Bracteole.—F. Calyx subtended by bracteoles.—G. Limb of corolla from above.—H. Corolla opened to show androecium.—I.
Stamen.—J. Gynoecium.—K. Variation in stigmas (one lobe highly reduced at top, crateriform at bottom).—L. Capsule valves (internal view on
left, external view on right).—M. Seeds (top view to left of leaf, side view to right of leaf). Drawn by N. Bollinger.
24
Daniel and McDade
ALISO
Fig. 16. Map showing the worldwide distribution by country of Elytraria.
corollas (3 to 3.7 mm long), and the continental American
species E. caroliniensis and E. mexicana have the largest (up to
17 mm long). Although the color(s) of corollas for several
species in South America and the West Indies remains
unknown, so far as is known, all species endemic to the Old
Word (with possible exception of E. madagascariensis, which is
reported to have white-pink corollas fide Benoist 1967), South
America, and West Indies have white to cream corollas.
Among North American (United States, Mexico, and Central
America) species, corolla color varies from white (E. caroliniensis) to pink (E. bromoides) to blue (E. imbricata) to yellow
(E. macrophylla) to white and purple (E. mexicana). Limb form
varies from bilabiate with lips little opened (e.g., E. marginata)
to bilabiate with lips widely spreading (e.g., E. macrophylla) to
subactinomorphic (e.g., E. caroliniensis). Androecial diversity
in Elytraria is greater than in any other genus of Nelsonioideae
(except perhaps Staurogyne in which various numbers of
staminodes have been reported). The following combinations
of number of stamens and staminodes are represented: two
stamens + no staminodes, two stamens + two staminodes, and
four stamens + no staminodes.
Currently Recognized Species of Elytraria
This list is based on Wenk and Daniel (2009) and on a
revision of the genus that is currently in preparation.
1. Elytraria acaulis (L.f.) Lindau (Angola, Botswana, CongoKinshasa, Ghana, India, Kenya, Malawi, Mozambique,
Nigeria, Somalia, South Africa, Sri Lanka, Tanzania,
Uganda, Zambia, Zimbabwe)
This is the most widely distributed species of Elytraria in
Africa, the southernmost-occurring species of Nelsonioideae
(to ca. lat. 27.5uS), and the only Old World species of the genus
with an apparently natural intercontinental (Africa and Asia),
disjunct distribution.
2. Elytraria bissei H. Dietr. (Cuba)
3. Elytraria bromoides Oerst. (Guatemala, Mexico, USA)
Synonyms fide Daniel (1995): Elytraria acuminata (Small)
Cory, Tubiflora acuminata Small
4. Elytraria caroliniensis (Walter ex J.F. Gmel.) Pers. (USA)
Tubiflora caroliniensis Walter ex J.F. Gmel., Anonymos
caroliniensis Walter; synonyms fide Daniel (in press): Elytraria
angustifolia (Fernald) Leonard, E. caroliniensis var. angustifolia
(Fernald) S.F. Blake, E. caroliniensis var. vahliana (Nees) D.B.
Ward, E. virgata Michx., E. virgata var. angustifolia Fernald, E.
virgata var. latifolia Nees, E. virgata var. vahliana Nees,
Tubiflora caroliniensis var. angustifolia (Fernald) Small
This species, which occurs in subtropical (southern Florida)
and temperate regions of the southeastern USA, is the
northernmost-occurring species of Nelsonioideae (to ca.
33uN latitude). Its flowers are unusual in their subsalverform
shape with a subactinomorphic limb.
5. Elytraria cubana Alain (Cuba)
Synonym fide Alain (1956): Elytraria crenata Leonard
6. Elytraria filicaulis Borhidi & O. Muñiz (Cuba)
Justicia acaulis L.f., Tubiflora acaulis (L.f.) Kuntze; synonyms fide Vollesen (2008): Elytraria acaulis var. lyrata (Vahl)
Bremek., E. crenata Vahl, E. crenata var. lyrata (Vahl) Nees,
E. indica Pers., E. lyrata (Vahl) Vahl, Justicia acaulis var.
lyrata Vahl
7. Elytraria imbricata (Vahl) Pers. (Argentina, Belize, Bolivia,
Brazil, Colombia, Costa Rica, Ecuador, El Salvador, Guatemala, Honduras, Mexico, Netherlands Antilles-Curaçao, Nicaragua, Panama, Peru, Trinidad and Tobago, USA, Venezuela)
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Revision of Nelsonioideae
Justicia imbricata Vahl; synonyms fide Daniel (1995):
Elytraria apargiifolia Nees, E. fasciculata Kunth, E. frondosa
Kunth, E. microstachya Oerst., E. pachystachya Oerst., E.
ramosa Kunth, E. scorpioides Roem. & Schult., E. squamosa
(Jacq.) Lindau, E. tridentata Vahl, Stachytarpheta squamosa
(Jacq.) Vahl, Tubiflora pachystachya (Oerst.) Kuntze, Tubiflora squamosa (Jacq.) Kuntze, Verbena squamosa Jacq.
Although native to, and widespread in, the New World, this
weedy species has been introduced into several Paleotropical
regions (e.g., Madagascar, Philippines, Vietnam). It is the only
species in the New World to occur in both North America and
South America.
8. Elytraria ivorensis Dokosi (Ghana, Ivory Coast)
9. Elytraria klugii Leonard (Peru)
10. Elytraria macrophylla Leonard (Mexico)
11. Elytraria madagascariensis (Benoist) E. Hossain (Madagascar)
Tubiflora madagascariensis Benoist
12. Elytraria marginata Vahl (Angola, Congo-Kinshasa,
Cameroon, Equatorial Guinea-Bioko and Rı́o Muni,
Gabon, Ghana, Ivory Coast, Liberia, Nigeria, São Tomé
and Prı́ncipe, Sierra Leone, South Sudan, Togo, Uganda)
Synonym fide Vollesen (2008): Tubiflora paucisquamosa De
Wild. & T. Durand
This is the most widespread, and a rather common, species
in western Africa.
13. Elytraria maritima J.K. Morton (Ghana, Ivory Coast)
14. Elytraria mexicana Fryxell & S.D. Koch (Mexico)
15. Elytraria minor Dokosi (Kenya, Tanzania)
16. Elytraria nodosa E. Hossain (Madagascar)
17. Elytraria planifolia Leonard (Cuba)
Borhidi and Muñiz (1978) recognized Elytraria planifolia
subsp. acunae Borhidi, the status of which remains unresolved.
18. Elytraria prolifera Leonard (Haiti)
19. Elytraria shaferi (P. Wilson) Leonard (Cuba)
Tubiflora shaferi P. Wilson; synonym fide León and Alain
(1974): Elytraria tridentata var. wrightii Gomez
20. Elytraria spathulifolia Borhidi & O. Muñiz (Cuba)
21. Elytraria tuberosa Leonard (Ecuador)
So far as is known, this is the only species of Elytraria that
has conspicuously tuberous roots.
NELSONIA R. Br., Prodr. Fl. Nov. Holland. 480 (1810).—
TYPE. Nelsonia campestris R. Br. (lectotype designated in
1955 by Bremekamp, ING card 00749)
Acaulescent to caulescent (usually low and often matforming and rooting at nodes), annual to perennial herbs to 2.5
25
dm tall and up to 7 dm long. Young stems subterete or 6
flattened on drying, pubescent with glandular and/or eglandular (often silky and up to 4.5 mm long) trichomes. Leaves
opposite (to alternate), 6 clustered in a basal rosette to evenly
disposed, sessile to petiolate, petiole (if present) shorter than
blade, margin of blade entire. Inflorescence of sessile to
pedunculate axillary and terminal densely bracteate spikes,
peduncles (if present) lacking scales, fertile portion of spike
dense and cylindric, 5–14 mm in diameter (excluding corollas)
near midpoint, not branching, rachis mostly not visible, often
bearing elongate, silky trichomes, especially at base of
dichasia; dichasia alternate (6 spirally disposed), sessile.
Bracts alternate (6 spirally disposed), imbricate, green,
membranaceous, not fused to other structures, broadly ovate
to ovate to elliptic to subcircular to oblate to obovate, 4–8
(–10) mm long, 2.5–5.2 (–6) mm wide, (1– or) 3– or 5–veined
(lateral veins sometimes indistinct). Bracteoles absent (see
discussion). Flowers sessile to subsessile (i.e., with pedicels to
1 mm long). Calyx usually deeply 4-lobed, green to hyaline,
3.3–7 mm long, lobes heteromorphic (sometimes equal in
length, but differing in shape and width), posterior lobe 3–
6 mm long, sometimes largest, 3– or 5– or 7–veined, lateral
lobes 2.5–5 mm long, 1– or 3– or 5–veined, anterior lobe 2.5–
5 mm long, 2– (or 5–) veined, (entire) 2– (or more-) cleft at
apex (or rarely divided almost to tube). Corolla whitish
(usually with lobes suffused with shades of pink or purple) or
pink or blue or purple, sometimes with differently colored
striations or spots, (2.5–) 4–7 mm long, externally glabrous,
tube 6 funnelform, narrow proximal portion 6 constricted (or
narrowest) at apex thence expanded into a short throat, limb 2labiate, upper lip 2-lobed, lower lip lacking a bullate palate.
Stamens 2, inserted near base of corolla throat, anthers
included in throat or partially exserted from mouth of corolla,
not extending beyond lips of corolla, thecae dehiscing toward
each other (i.e., flowers pleurotribic), subellipsoid, 0.2–0.5 (–1)
mm long, those of a pair parallel to widely divergent (i.e.,
thecae 6 end to end and horizontal), 6 equal in size, glabrous,
with a subbasal flaplike appendage associated with dehiscence;
staminode 0; pollen subprolate to perprolate (P:E 5 1.28–2.18;
polar axis 20–39 mm, equatorial axis 15–21 mm), 3-colpate to 3colporoidate, exine reticulate, muri smooth. Style included in
corolla tube or if exserted from mouth of corolla then not
extending beyond lips of corolla, stigma not touch-sensitive,
unequally 2-lobed (sometimes with at least one lobe again
divided), lobes 0.1–0.2 mm long, 0.1–0.2 mm wide. Capsule
ovoid-conic, 2.8–5 mm long, lacking trichomes or with
inconspicuous sessile to subsessile glands (especially near
apex). Seeds to 30 per capsule, subglobose to subhemispheric
to subellipsoid, one face somewhat flattened and with a
shallow pore, longest axis 0.3–0.6 mm, shortest axis 0.3–
0.5 mm, surface coarsely lumpy to nearly smooth, microreticulate and microverrucate, pubescent with inconspicuous,
hygroscopic, anchor-shaped trichomes (expanding to 0.2 mm
long on moistening); (Fig. 17).
Opinion varies as to whether a single variable species (e.g.,
Hossain 1984; McDade et al. 2012) or several species (e.g.,
Bremekamp 1964; Morton 1979; Vollesen 1994) should be
recognized. Although the genus appears to be native in
Africa, Asia, and Australia, it remains unknown whether it is
native or introduced in tropical America (Fig. 18). Bremekamp (1955) indicated that Nelsonia was a Paleotropical
26
Daniel and McDade
ALISO
Fig. 17. Nelsonia canescens.—A. Habit (Palmer 583; Mexico).—B. Spike with flower (Daniel et al. 5452gh; cultivated from plants from
Panama).—C. Calyx flattened and with anterior lobe lowermost.—D. Calyx with anterior lobe bent to show gynoecium.—E. Corolla (view from
above).—F. Corolla cut open to show androecium and gynoecium.—G. Apex of stamen with anthers.—H. Apex of style with stigma.—I.
Dehisced capsule (Palmer 583).—J. Seed (Palmer 583). Drawn by M. Tenorio.
VOLUME 32, NUMBER 1
Revision of Nelsonioideae
27
Fig. 18. Map showing the worldwide distribution by country of Nelsonia (some or all of the New World distribution represents
naturalized occurrences).
genus that has spread to the New World in post-Columbian
times. Vollesen (1994) indicated that there are likely five
species of Nelsonia, three in Africa (N. canescens, N. gracilis,
and N. smithii), one in southern Asia (N. canescens), one in
Australia (N. campestris), and two in the New World (N.
canescens-introduced and N. pohlii-native). In addition to
species that have been described, Champluvier has annotated
a large number of specimens at BR as a new species, ‘‘N.
feracissima Champl.,’’ from Burundi, Cameroon, CongoKinshasa, Equatorial Guinea-Bioko, Malawi, Mozambique,
Rwanda, and Tanzania. Molecular phylogenetic studies that
included plants sampled from multiple continents led
McDade et al. (2012) to suggest that only a single species
was likely represented. However, these authors did not
include N. gracilis, and that species appears rather distinctive
morphologically. Major questions that remain to be resolved
in this genus include the number of species deserving
recognition and whether any of the occurrences in the New
World represent native (i.e., pre-Columbian) populations.
Nelsonia exhibits a unique synapomorphy within Nelsonioideae, the absence of bracteoles below the flowers. The rare
(teratological?) presence of bracteoles in some Nelsonia
(Daniel and Figueiredo 2009) permits recognition of the
homology of its flowers to those of the one-flowered dichasial
units in other nelsonioids. The inflorescences of the genus (i.e.,
with peduncles, if present, lacking scales and with dense,
cylindric spikes bearing imbricate bracts and small corollas)
are also unlike those of other Nelsonioideae. With Staurogyne
and Saintpauliopsis, Nelsonia shares the characteristic of seeds
bearing hooked or branched hygroscopic trichomes.
Currently Recognized Species of Nelsonia
1. Nelsonia canescens (Lam.) Spreng. (Angola, Australia,
Benin, Botswana, Burkina Faso, Burundi, Cambodia,
Cameroon, Central African Republic, Chad, Congo-Brazzaville, Congo-Kinshasa, Equatorial Guinea-Bioko, Eritrea,
Ethiopia, Gabon, Gambia, Ghana, Guinea, Guinea-Bissau,
India, Indonesia-Papua, Ivory Coast, Kenya, Laos, Liberia,
Madagascar, Malawi, Malaysia, Mali, Mozambique, Myanmar, Nepal, Niger, Nigeria, Papua New Guinea, São Tomé
& Prı́ncipe, Senegal, Sierra Leone, South Sudan, Sudan,
Tanzania, Thailand, Togo, Uganda, Vietnam, Zambia,
Zimbabwe).
Justicia canescens Lam.; synonyms fide Vollesen (1994):
Justicia bengalensis Spreng., J. hirsuta Vahl, J. lamiifolia
Roxb., J. nummulariifolia Vahl, J. origanoides Vahl, J.
tomentosa Roxb. ex C.B. Clarke, J. vestita Roem. & Schult.,
N. albicans Kunth, N. campestris R. Br. var. vestita (Roem. &
Schult.) C.B. Clarke, N. canescens var. vestita (Roem. &
Schult.) E. Hossain, N. hirsuta (Vahl) Roem. & Schult., N.
lamiifolia (Roxb.) Spreng., N. nummulariifolia (Vahl) Roem. &
Schult., N. organoides (Vahl) Roem. & Schult., N. pohlii Nees,
N. senegalensis Oerst., N. tomentosa (Roxb.) Dietr., N. villosa
Oerst.; synonyms fide Hossain (1984): N. campestris R. Br., N.
rotundifolia R. Br.; synonyms fide Daniel (in press): N.
canescens var. smithii (Oerst.) E. Hossain, N. smithii Oerst.
The species is native to several regions in the Old World
(Africa, Asia, Australia), but due to its propensity to spread
into disturbed habitats, its overall natural distribution is not
known with certainty and may be less extensive than the list
of nations, noted above, in which it is currently found. Its
numerous occurrences in the New World (e.g., Bolivia,
Brazil, Colombia, Costa Rica, El Salvador, French Guiana,
Honduras, Mexico, Nicaragua, Panama, Puerto Rico,
United States) possibly contain some native populations
(e.g., Colombia, where the species was collected near
Cartagena in 1801 [without collector at P], and Brazil, where
the species was collected in Goiás as early as 1821 [Saint-
28
Daniel and McDade
Hilaire cat. num. 766 at K and P]). The species was collected
in southern Central America (Costa Rica and Panama) and
the West Indies (Puerto Rico) during the 1850s. Other
collections from the New World are clearly of recent origin
(e.g., Florida in the USA; Franck and Daniel 2012). Both
Morton (1979) and Hossain (1984) noted the likely adventive
occurrence of N. canescens on many islands in the Pacific.
We have not been able to locate records of Nelsonia from
Pacific islands (other than New Guinea), nor have such been
reported in numerous floras covering the region (e.g.,
Yuncker 1959; Heine 1976; Smith 1991; Fosberg et al.
1993; Meyer and Lavergne 2004).
Hossain (1984) noted considerable variation in habit, leaves,
bracts, and pubescence for this species, but he reported
insignificant correlations among the major morphological
characters. We observed additional variation in the color of
corollas as noted on herbarium specimen labels and observed by
us in the field or in photographs. Plants may have corollas with
the limb entirely white (Australia), white with pink to maroon
coloration on the distal portion of the lobes (Africa, America),
blue to purple (Africa, America, Asia), purple with dark purple
striations on the upper lip (Africa, Asia), and purple with dark
purple spots on the lower lip (America; Fig. 3F–I). A similar case
among Acanthaceae of a widespread species with multiple color
patterns on corollas was noted for Henrya insularis Nees ex
Benth. (Daniel 1990). We also noted variation in the division of
the anterior calyx lobe. Most plants have calyces with the anterior
lobe shallowly cleft or divided up to M the length of the lobe. In
some plants from Australia the anterior lobe is entire (Barker
1986) whereas in others it is divided more than L its length (e.g.,
Lazarides 9080 at L from Australia: Northern Territory).
Discussions about whether to recognize N. smithii as distinct
from N. canescens include those of Morton (1979), Hossain
(1984), Vollesen (1994), Daniel and Figueiredo (2009), Franck
and Daniel (2012), and McDade et al. (2012). Observations
made on plants occurring in three different habitats in the
vicinity of Lagos, Nigeria showed slight differences in
characters, but it was not determined whether those differences
were genetically fixed or correlated with the different habitats
(Hossain and Emumwen 1981). Morton (1979) noted that
presumably native plants from Brazil that were described as N.
pohlii are intermediate between N. smithii and N. canescens in
most of the morphological characters he used to distinguish
the two species. Until additional studies provide more
convincing insights, Hossain’s (1984) broad interpretation of
this species is utilized here.
Fongod et al. (2010) reported efficacy of N. canescens as a
beneficial cover crop (especially in weed suppression) in
banana plantations.
2. Nelsonia gracilis Vollesen (Angola, Malawi, Zambia)
This species of annual duration with sessile to subsessile
leaves and bluish flowers appears rather distinctive, but some
plants (e.g., Bingham 10092 at K from Zambia) appear to be
perennial, and flowers of some collections of N. canescens are
also described as ‘‘blue.’’ Some apparently annual (or first-year
perennial) plants of the latter species (e.g., Henty & Foreman
NGF 49370 from Papua New Guinea at L) are similar to N.
gracilis with their wiry stems and small, elliptic, and subsessile
to short-petiolate leaves.
ALISO
Excluded Name
Nelsonia brunelloides (Lam.) Kuntze
Justicia brunelloides Lam.
Although it is often cited as a synonym of N. canescens, this
name pertains to Hemigraphis (Bremekamp 1944).
SAINTPAULIOPSIS Staner, Bull. Jard. Bot. Brux. 13: 8 (1934).—
TYPE. Saintpauliopsis lebrunii Staner (Staurogyne lebrunii
(Staner) B.L. Burtt)
Terrestrial, epiphytic, or epipetric acaulescent or caulescent
stoloniferous perennial herbs to 4 dm tall or long, often
trailing, rooting at nodes and forming mats on forest floor or
rosellate clumps on rocky walls. Young stems subquadrate to
quadrate-sulcate, evenly pubescent with conspicuously (or
inconspicuously) multi-septate flexuose eglandular trichomes, septae maroon. Leaves opposite, evenly disposed
along young stems or sometimes 6 congested into subrosettes, petiolate, petiole usually longer than blade, margin of
blade entire to sinuate-crenate. Inflorescence of pedunculate
loose terminal raceme-like modified thyrses (or panicles of
modified thyrses; see discussion), peduncles lacking scales,
fertile portion of inflorescence neither dense nor cylindric, 8–
15 mm in diameter (excluding corollas) near midpoint, not
branching, rachis clearly visible; dichasia alternate (sometimes resulting in a zig-zag inflorescence), pedunculate.
Bracts alternate, not imbricate, greenish, membranaceous,
adnate to (and indistinguishable from) dichasial peduncle for
91–98% the length of the peduncle, free portion of bract
(arising near apex of peduncle) linear to lance-linear, 2–
3.7 mm long, 0.3–0.5 mm wide, 1-veined. Bracteoles borne at
base of flower where partially adnate basally to receptacle
and calyx tube, greenish, lance-linear to lanceolate to elliptic
(to obovate), 1.8–5 mm long, 0.3–0.7 mm wide. Flowers
sessile. Calyx deeply 5-lobed, greenish (or lateral lobes
hyaline), 3.3–5 (–6.5 in fruit) mm long, lobes heteromorphic
(1 + 2 + 2), 3 larger lobes (posterior and 2 anterior) lanceolate
to linear to elliptic (to obovate), 3.2–4.2 (–6) mm long, 3veined, 2 smaller (lateral) lobes often hidden by bracteoles,
lanceolate to subulate, 1.1–2 mm long, 1-veined. Corolla
white to pink with darker pink striations to pale blue, 6
pendant, 4.5–8.5 mm long, externally glabrous (margins of
lobes ciliolate), tube 6 campanulate, narrow proximal
portion 6 cylindric, distally widely spreading into throat,
limb 2-labiate, upper lip 2-lobed, shorter than lower lip,
lower lip lacking a bullate palate. Stamens 4, didynamous,
inserted at 6 different heights or equally inserted, presented
at different heights, longer (anterior) pair inserted near base
of throat, exserted from mouth of corolla but anthers not
extending beyond limb, shorter (posterior) pair inserted near
base of throat (often distal to anterior pair) and with anthers
included in throat, thecae apparently dehiscing toward lower
lip (i.e., flower nototribic), ellipsoid, 0.7–1 mm long, those of
a pair parallel to subsagittate, equal in length, glabrous, each
with a slender, straight or curved, bifurcate (forked) basal
appendage 0.2–0.4 mm long; staminode not evident or 1, to
0.5 mm long; pollen oblate spheroidal to euprolate (P:E 5
0.88–1.87; polar axis 21–43 mm, equatorial axis 21–26 mm),
3-colpate to 3-colporoidate, exine reticulate (sometimes
appearing 6 foveolate in non-acetolyzed grains), muri
VOLUME 32, NUMBER 1
Revision of Nelsonioideae
29
Fig. 19. Saintpauliopsis lebrunii.—A. Habit (Reekmans 10955; Burundi).—B. Inflorescence nodes with two flowers in fruit (Lewalle 753;
Burundi).—C. Dichasium with bract, bracteoles, and calyx (Lewalle 753).—D. Corolla opened to show androecium (redrawn from sketch on
d’Alleizette 1107 at P; Madagascar).—E. Capsule (Lewalle 753). Drawn by S. Adler and N. Pugh.
verrucate. Style exserted from mouth of corolla, equaling or
extending beyond upper lip, usually not exceeding lower lip,
stigma not touch-sensitive, subequally to unequally 2-lobed,
lobes somewhat flattened and spreading, 0.1–0.6 mm long,
0.1–0.3 mm wide. Capsule ovoid, 3.5–5.3 mm long, glabrous
proximally, sparsely pubescent distally with flexuose to
antrorse eglandular trichomes. Seeds up to 26 per capsule
(ovules (12–) 28–36 per capsule), 6 blocky to oblong, longest
axis 0.5–1 mm, shortest axis 0.4–0.7 mm, surfaces coarsely
rugose- to lumpy- to foveolate-reticulate, pubescent with
hygroscopic, uncinate to anchor- or Y-shaped trichomes
,0.05 mm long; (Fig. 19).
A single species from tropical Africa and Madagascar
(Fig. 20) is recognized in this genus. Plants are often mistaken
for Gesneriaceae (indeed, the genus was originally treated in
that family; Staner 1934), and often grow in habitats (e.g.,
mossy banks) typical of many gesneriads. The species shows
similarities to, and has been treated in, Staurogyne (Burtt 1958).
Indeed, leaves and inflorescences of Saintpauliopsis lebrunii
greatly resemble those of Staurogyne chapaensis from southeastern Asia (although the bracteoles are not fused to the flower
in the latter species). If Saintpauliopsis were to be treated in an
expanded concept of Staurogyne, Anisosepalum also would have
to be included in that genus in order to maintain monophyly of
genera. Saintpauliopsis differs from Staurogyne and Anisosepalum by the characters noted in the key above.
In addition to the distinctive habit and leaves of Saintpauliopsis, the inflorescence is unusual among Nelsonioideae. It is
30
Daniel and McDade
ALISO
Fig. 20. Map of Africa showing the distribution by country of Saintpauliopsis.
sometimes noted to be a raceme. It is not treated here as such
because the lateral branches are presumed to be modified
dichasia on peduncles. The combination of alternate dichasia
(sometimes resulting in a zig-zag rachis), a thin (,0.5 mm in
diameter on flattened specimens) and wiry rachis, partial
adnation of the bract with the dichasial peduncle and its
emergence near the apex of the peduncle on the anterior side
(i.e., farthest from rachis) of the dichasium, and partial adnation
of the bracteoles to the floral receptacle and calyx tube is unique
among Nelsonioideae. According to Dickinson (1978; where
Saintpauliopsis is cited as an example), flower-bract associations
of this type are examples of cryptic epiphylly. Adnation of
bracteoles with the base of the flower also occurs in
Anisosepalum, and appears to represent a morphological
synapomorphy for the clade consisting of these two genera.
A bifurcate basal appendage on each anther theca was
reported by Hossain (2004) and was observed on specimens
with visible stamens that we examined. The bifurcate nature
of the appendage was not noted by Staner (1934),
Champluvier (1991), or Vollesen (2008). Whether it is
indeed ubiquitous in plants of the genus or not remains
unknown.
VOLUME 32, NUMBER 1
Revision of Nelsonioideae
Currently Recognized Species of Saintpauliopsis
1. Saintpauliopsis lebrunii Staner (Burundi, Congo-Kinshasa,
Gabon, Madagascar, Rwanda, Tanzania)
Synonyms fide Vollesen (2008): Saintpauliopsis lebrunii var.
obtusa Staner, Staurogyne lebrunii (Staner) B.L. Burtt
We found no morphological differences between plants
from Madagascar and those from the African continent.
Detailed label notes on Perrier de la Bâthie 9257 (P) from
Madagascar indicated flowers as having two stamens and
two staminodes. It is unknown whether this represents an
error of observation (no corollas are extant on the specimen
at P) or variation in the androecium of this species. Other
collections from Madagascar have four stamens and a single
staminode.
STAUROGYNE Wall., Pl. Asiat. Rar. 2: 80 (1831).—LECTOTYPE (designated by Leonard, Contr. U.S. Natl. Herb. 31:
5 (1951). Staurogyne argentea Wall.
Ancistrostylis T. Yamaz., J. Jap. Bot. 55: 1 (1980).—TYPE.
Ancistrostylis harmandii (G. Bonati) T. Yamaz. (Herpestis
harmandii G. Bonati)
Ebermaiera Nees in Wall., Pl. Asiat. Rar. 3: 75 (1832).—
LECTOTYPE (designated by Leonard, Contr. U.S. Natl.
Herb. 31: 5 (1951). Ebermaiera humilis Nees
Erythracanthus Nees in Wall., Pl. Asiat. Rar. 3: 75 (1832).—
TYPE. Erythracanthus racemosus (Roxb.) Nees (Ruellia
racemosa Roxb.)
Gynocraterium Bremek. in Sandwith, Kew Bull. 1939: 557
(1939).—TYPE. Gynocraterium guianense Bremek.
Neozenkerina Mildbr., Notizbl. Bot. Gart. Berlin-Dahlem 7:
491 (1921).—TYPE. Neozenkerina bicolor Mildbr.
Ophiorrhiziphyllon Kurz, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist.
40: 76 (1871).—TYPE. Ophiorrhiziphyllon macrobotryum
Kurz
Phyllophiorhiza Kuntze in Post and O. Kuntze, Lex. Gen.
Phan. 435 (1903).—TYPE. Ophiorrhiziphyllon macrobotryum Kurz
Staurogynopsis Mangenot & Aké Assi, Bull. Jard. Bot. État 29:
27 (1959).—TYPE. Staurogynopsis paludosa Mangenot &
Aké Assi
Stiftia Pohl ex Nees in Alph. de Candolle, Prodr. 11: 70 (1847),
nomen non rite publicatum
Zenkerina Engl., Bot. Jahrb. Syst. 23: 497 (1897).—TYPE.
Zenkerina kamerunensis Engler
Acaulescent to caulescent, prostrate to decumbent or
ascendant to erect annual or perennial herbs or rarely shrubs.
Younger stems subterete to subquadrate or becoming 6
flattened on drying, subglabrous or pubescent with eglandular
and/or glandular trichomes. Leaves opposite (or distal ones
sometimes subopposite or alternate), evenly disposed along
stems or in a basal cluster, sessile to petiolate, petiole usually
shorter than blade, margin of blade entire to sinuate-crenate or
shallowly dentate. Inflorescence of dichasia in leaf axils
or usually of pedunculate axillary and/or terminal spikes,
racemes, or thyrses, these sometimes branched and forming
panicles, fertile portion of inflorescence loose or dense, not
cylindric to cylindric, peduncles lacking clasping scales (rarely
31
with a few scattered non-clasping sterile bracts); dichasia
opposite or alternate, sessile to pedunculate. Bracts opposite or
alternate, imbricate or not, green or otherwise colored,
membranaceous, borne along inflorescence rachis or adnate
to dichasial peduncle for 11–80 (–100%) the length of the
peduncle, subfoliose or triangular to subulate to elliptic to
subcircular to obovate-elliptic to oblanceolate, 2–35 mm long,
0.5–11 mm wide, 1-, 3-, 5-, or 7-veined. Bracteoles borne at
apex of dichasial peduncle or at base of flower (but not adnate
to receptacle or calyx), green or otherwise colored, subulate to
narrowly elliptic to elliptic to oblanceolate, 1.5–19 mm long,
0.4–5.7 mm wide. Flowers sessile to pedicellate. Calyx deeply
5-lobed, green or otherwise colored or at least partially
hyaline, 3–36 mm long, lobes 6 heteromorphic (1 + 2 + 2;
rarely subequal), posterior lobe 3–35 mm long, 1-, 3-, 5-, or 7veined, usually larger than others, lateral lobes 1–22 mm long,
1- or 3-veined, anterior lobes sometimes fused to a greater
extent than other lobes, 2–31 mm long, 1-, 3-, or 5-veined.
Corolla white, yellow, pinkish, red, bluish, or purple,
sometimes with colored markings within limb, 3.8–47 mm
long, externally glabrous or pubescent, tube subcylindric to
funnelform, throat inconspicuous or conspicuous and 6 equal
to or longer than cylindric narrow proximal portion of tube,
limb subactinomorphic to 2-labiate, upper lip 2-lobed,
sometimes shorter than lower lip, lower lip lacking a bullate
palate. Stamens 4 (rarely 2), usually didynamous, inserted in
corolla tube, anthers included in tube or if slightly exserted
from mouth of corolla then usually not (or but barely)
extending beyond lips of corolla (rarely long-exserted, see
discussion of S. macrobotrya below), thecae dehiscing toward
lower lip (i.e., flower nototribic), broadly ellipsoid, 0.3–2.9 mm
long, those of a pair parallel to widely divergent (i.e., thecae 6
end to end and horizontal [stamen T-shaped] to ‘‘upsidedown’’ [stamen Y-shaped]), equal to subequal in length,
glabrous or pubescent, sometimes minutely mucronate near
base (with mucro on side of theca opposite dehiscing side),
connective sometimes expanded; staminode 0 or 1 (posterior)
or rarely 2–3, inconspicuous; pollen oblate spheroidal to
perprolate (P:E 5 0.95–2.31; polar axis 13–47 mm, equatorial
axis 11–36 mm), 3-colpate to 3-colporoidate to 3-colporate,
exine reticulate (sometimes appearing 6 foveolate in nonacetolyzed grains), muri smooth. Style included in corolla tube
or if exserted from mouth of corolla then usually not extended
beyond lips of corolla (except in S. macrobotrya), stigma
not touch-sensitive, equally to unequally 2-lobed (sometimes
appearing subcrateriform), 1 lobe again 2-parted (in some
species), lobes linear (to broadly ovate-triangular), 0.2–2 mm
long, 0.05–0.8 mm wide. Capsule subellipsoid to subcylindric
to ovoid, 3–12 (–20) mm long, glabrous or pubescent (usually
with glandular trichomes). Seeds up to 68 per capsule, spheric
to ellipsoid to oblong to blocky or irregularly shaped, longest
axis 0.3–1 mm, shortest axis 0.2–0.8 mm, surface coarsely
lumpy- to foveolate- to rugulate-reticulate, microverrucate
to microbaculate, pubescent with 6 hygroscopic trichomes,
trichomes anchor-shaped to apically multibranched or sometimes trichomes branched throughout; (Fig. 21, 22).
This, the richest genus of Nelsonioideae, contains 145
currently recognized species in tropical regions (Fig. 23) of
the Americas (30 spp.), Africa (5 spp.), and mainland Asia to
the Solomon Islands (110 spp.). The genus is especially well
represented in southeastern Asia, Malesia, and Brazil. Due to a
32
Daniel and McDade
ALISO
Fig. 21. Staurogyne macrobotrya (Ophiorrhiziphyllon macrobotryum).—A. Habit (Maxwell 97-170; Thailand).—B. Inflorescence nodes with
flower (Maxwell 97-170).—C. Apex of stamen with anther (Maxwell 97-170).—D. Apex of style with stigma (Garrett 654; Thailand).—E. Capsule
with aborted seeds (Maxwell 95-211; Thailand). Drawn by S. Adler.
lack of recent studies of Acanthaceae from throughout Asia,
the distributions of species occurring there, as noted in the
catalog below, is not as complete as for species from other
regions. The Malesian species were treated by Bremekamp
(1955); African species, which are restricted to the Gulf of
Guinea region, were treated by Champluvier (1991); and the
New World species were revised by Braz (2005). No species are
common to both the New World and the Old World; and no
species occurs in both Africa and the Asia-Malesian region.
Given the size of this genus, which is about four times as
species rich as all other nelsonioid genera combined, and its
broad distribution, the morphological diversity noted in the
description above is not surprising. Diversity in habit, leaf
shape, pubescence, inflorescence type, and floral characters
has resulted in the recognition of numerous genera in a
staurogynoid alliance. Only two of these are recognized here.
Both Anisosepalum and Saintpauliopsis have been, and could
with some justification yet be, included in Staurogyne;
together, the former genera are sister to the geographically
wide-ranging sample of 25 species of Staurogyne studied by
McDade et al. (2012). Each of those genera also shares some
morphological characteristics with Staurogyne, but each also
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Revision of Nelsonioideae
33
Fig. 22. Staurogyne miqueliana.—A. Habit (Lott et al. 3396; Mexico).—B. Inflorescence node with bract, bracteoles, and calyx (Lott et al.
3396).—C. Corolla opened to show androecium (Lott et al. 3226; Mexico).—D. Stamen (Lott et al. 3226).—E. Gynoecium (Lott et al. 3226).—F.
Capsule (external view at top, internal view at bottom) (Lott et al. 3396).—G. Seed (Lott et al. 3396). Drawn by K. Douthit (copyright reserved to
University of Michigan Herbarium, used with permission).
34
Daniel and McDade
ALISO
Fig. 23. Map showing the worldwide distribution by country of Staurogyne.
differs in several diagnostic features noted in the key above.
Because of these diagnostic and unusual features, we prefer to
recognize both Anisosepalum and Saintpauliopsis. Older names
that are now treated as synonymous with Staurogyne were
discussed by Bremekamp (1955) and Hossain (2004). Ancistrostylis T. Yamaz., a unispecific genus originally described in
Scrophulariaceae based on a collection from Laos (Yamazaki
1980), was treated as congeneric with Staurogyne by Scotland
and Vollesen (2000). Morphological characters enumerated in
the protologue agree well with those of Staurogyne. The sole
species has not been transferred to Staurogyne; whether it
corresponds to a previously described species of that genus
from southeastern Asia remains to be determined.
Bremekamp (1955) noted that Gynocraterium was very
similar to Staurogyne, from which it differed by its crateriform
stigma and fewer number of ovules. Braz and Monteiro (2011)
also noted the distinctive, elongate, and subulate bracts,
bracteoles, and calyx lobes of the sole species, G. guianense.
Subulate bracts and bracteoles are known elsewhere among
Staurogyne (e.g., S. setigera). Braz and Monteiro (2011)
noted between six and ten ovules per locule in their
description of G. guianense, a range that does not distinguish
the genus from other genera of Nelsonioideae in South
America. Form of the stigma would appear to be the primary
morphological character for distinguishing Gynocraterium
from Staurogyne. Stigmas of Jansen-Jacobs 2852 (at U) that
were hydrated and studied have two unequal (0.4 and 0.7 mm
long) and broadly (1–1.2 mm wide) ovate-triangular lobes
that result in a subcrateriform or funnel-shaped aspect.
Although form of the stigma is not well documented among
species of Staurogyne, and is often not well preserved on
herbarium specimens, bilobed stigmas occur in the genus.
Indeed, Hossain’s (2004: Fig. 1L17) illustration of the stigma
of S. subcordata showing two broad and flattened lobes are
similar to those of G. guianense. Subcrateriform stigmas
sometimes also occur in at least one species of Elytraria (i.e.,
E. caroliniensis; Fig. 15K). Intermediates or variations
between funnel-shaped stigmas and bilobed stigmas in
another genus of Acanthaceae (Thunbergia Retz.) are
illustrated by Retief and Reyneke (1984: Fig. 3) and
Schönenberger (1999: Fig. 118). Given other similarities
between G. guianense and species of Staurogyne and the
molecular phylogenetic results of McDade et al. (2012), this
putative morphological distinction would not seem sufficient
to be the basis of a separate genus. Further, because current
phylogenetic results (McDade et al. 2012) indicate that
Gynocraterium is sister to all species of Staurogyne in the
New World, if the former genus were to be recognized as
distinct from the latter, then either all Neotropical species of
Staurogyne would have to be transferred to Gynocraterium or
a new genus would have to be recognized for them. Because
morphological synapomorphies remain unknown for species
of Staurogyne in the New World, neither of these alternatives
seems prudent at present.
Bremekamp (1955) also noted a close alliance between
Ophiorrhiziphyllon and Staurogyne. He contended that the sole
distinction for Ophiorrhiziphyllon was the exsertion of the
stamens, and he noted ‘‘this can hardly be regarded as sufficient
to justify the maintenance of the genus’’ (Bremekamp 1955: 160).
Hossain (2004) further noted that several species of Staurogyne
have shortly exserted stamens. Another unusual character of O.
macrobotryum, the type of Ophiorrhiziphyllon, is the relationship
between the anther connective and the two thecae. Among
Nelsonioideae, the thecae are attached to the connective distally
and are sometimes free from it (to a lesser or greater extent)
proximally. Among species of Staurogyne both narrow and
broad connectives are encountered, and the pair of thecae varies
from parallel to sagittate to horizontal (i.e., each perpendicular to
the connective and oriented apex to apex, thus stamen T-shaped).
In O. macrobotryum the thecae are attached only at one end
(presumably the distal end), and are divergent and erect (stamen
Y-shaped) so that they appear ‘‘upside-down.’’ Somewhat similar
VOLUME 32, NUMBER 1
Revision of Nelsonioideae
thecae among species of Staurogyne were observed by us in S.
setosa (Toroes 2320 at MICH), illustrated by Hossain (2004: 22,
Fig. 1A) in S. subcordata, and illustrated by Champluvier (1991:
107, Fig. 3G) in S. capitata. Although the shapes and sizes of
connectives and thecae differ among these four species, in all of
them the thecae appear to be attached only at the apex and
oriented ‘‘upside-down.’’ Because anthers are rarely described for
species of Staurogyne, this unusual arrangement of thecae might
be more common than is known. Considering phylogenetic
relationships (McDade et al. 2012) and the apparent lack of
significant morphological distinctions, there appears to be
insufficient basis for recognition of Ophiorrhiziphyllon as distinct
from Staurogyne.
Bremekamp (1955) created an infrageneric classification for
the Asian species of the Malesian region. He indicated that the
classification of African and American species would have to
await further studies, but suggested that they were not
affiliated with the subgenera he recognized for Asian species.
He recognized two subgenera, Staurogyne and Tetrastichum
Bremek., based on the number of ovules, number of rows of
ovules in each ovary cell, and inflorescence characters.
Subgenus Staurogyne was divided into sections, subsections,
and series based on differences in habit, inflorescence, bract,
and calyces. Hossain (1971, 1972) dealt with the species on a
worldwide basis, and proposed (but only partially published) a
more extensive infrageneric treatment. Hossain (1972) published sect. Zenkerina (Engl.) E. Hossain in subgen. Tetrastichum, and he proposed three unnamed series in subgen.
Staurogyne, sect. Staurogyne, subsect. Staurogyne (as ‘‘Macrosepalae Bremek.’’) to account for species not treated by
Bremekamp (1955) and/or that could not be accommodated in
his classification for Malesian taxa.
Champluvier (1991) provided descriptions of, and a key to,
the five African species of Staurogyne. These all occur in
tropical western Africa, primarily around the Gulf of Guinea
in wet evergreen forests. Champluvier (1991) also discussed
the disposition of the African species relative to Hossain’s
classification. She treated S. letestuana in subgenus Staurogyne
and noted problems and inconsistencies in characters of other
African species vis-à-vis their treatment by Hossain in sect.
Zenkerina. She questioned the recognition of sect. Zenkerina
based on its circumscription, but noted that the African species
assigned to this section differ from S. letestuana by having: 1)
capsules with a membranous wall, and that are tardily and
apparently not explosively dehiscent, and 2) calyx lobes that
converge and rest upon the capsule (even after dehiscence). At
least the membranous wall of capsules in these species is
apparently unique in the genus. Thus, additional studies are
necessary to determine an appropriate taxonomic placement
for most of the African species.
Braz (2005) provided descriptions of, and a key to, 28
species of Staurogyne in the Neotropics. Profice (2000)
described an additional Neotropical species, Staurogyne
carvalhoi Profice, which according to Braz (2005) likely
represents an undescribed genus of Acanthaceae with hypocrateriform corollas and three seeds per locule. This species
was excluded from Braz’s study, and we have not seen
specimens of it. Braz (2005) noted two centers of richness for
New World Staurogyne: southeastern Brazil (especially in the
mata Atlântica biome where 19 of the Neotropical species are
restricted) and northern South America (especially in the
35
Amazon forest biome of northern Brazil, the Guianas, and
Venezuela). Braz (2005) concluded that based on Hossain’s
(1972) infrageneric taxa, all New World species of Staurogyne
pertain to subsect. Staurogyne (as ‘‘Macrosepalae’’).
Hossain’s (1971) mostly unpublished infrageneric account
of Staurogyne is particularly informative because he assigned
most species to infrageneric taxa, which permits testing of his
taxonomic hypotheses using molecular sequence data. Also,
Braz (2005) used morphological data of mostly Neotropical
species to construct a phylogeny for the species she studied. She
concluded that Staurogyne was monophyletic, but that the
Neotropical species were not (i.e., two Asian species were nested
among those from the New World). Infrageneric relationships
based on molecular phylogenetic data of McDade et al. (2012)
confirm that Staurogyne is monophyletic (inclusive of Ophiorrhiziphyllon and Gynocraterium), and reveal the Neotropical
species to be monophyletic and sister to those from the Old
World. Although additional phylogenetic work is required to
fully understand the relationships of species in the Old World,
there is some support for those sampled from Asia as also being
monophyletic (McDade et al. 2012).
Currently Recognized Species of Staurogyne
For this list the following sources provided most of the
information about species currently recognized and their
synonymies: Benoist (1935; southeastern mainland Asia),
Bremekamp (1955; Malesia), Hossain (1971, 1972; worldwide),
Champluvier (1991; Africa), Hô (1993; Vietnam), Braz (2005;
New World), and Hu et al. (2011; China). Numerous
additional local floras in Asia, Malesia, Papuasia, and
Australia were also consulted. Where there were conflicts,
the most recent taxonomic interpretations were generally given
preference. The list undoubtedly lacks some synonyms.
Although all African (Champluvier 1991) and New World
(Braz 2005) species have been revised recently, the majority of
species occur in southern Asia and Malesia, and there is no
recent comprehensive account of the genus in either of those
regions.
1. Staurogyne alba Braz & R. Monteiro (Brazil)
2. Staurogyne amboinica Bremek. (Indonesia)
3. Staurogyne amoena Benoist (Vietnam)
4. Staurogyne anigozanthus (Nees) Kuntze (Brazil)
Ebermaiera anigozanthus Nees
5. Staurogyne anomala Bremek. (Indonesia)
6. Staurogyne arcuata C.B. Clarke (Malaysia)
7. Staurogyne argentea Wall. (India, Myanmar)
Synonyms fide Hossain (1971): Ebermaiera argentea (Wall.)
Nees, E. staurogyne Nees
Hossain (1972) recognized Staurogyne argentea var. angustifolia (Wall.) E. Hossain based on S. angustifolia Wall.
(5 Ebermaiera staurogyne Nees var. angustifolia (Wall.) Nees,
E. angustifolia (Wall.) T. Anderson).
8. Staurogyne aristata E. Hossain (Thailand)
36
Daniel and McDade
9. Staurogyne athroantha Bremek. (Philippines)
ALISO
Hossain (1971) indicated that this taxon should be treated as
a variety of S. debilis, but did not publish a combination for it.
10. Staurogyne atropurpurea E. Hossain (Thailand)
28. Staurogyne citrina Ridl. (Indonesia)
11. Staurogyne axillaris S. Moore (Malaysia)
29. Staurogyne comosa Kuntze (Malaysia)
12. Staurogyne balansae Benoist (Malaysia, Vietnam)
13. Staurogyne batuensis Bremek. (Indonesia)
14. Staurogyne beddomei (C.B. Clarke) Kuntze (Myanmar)
Ebermaiera beddomei C.B. Clarke
Bremekamp’s (1961) combination for this name would
appear to be superfluous.
15. Staurogyne bella Bremek. (Thailand, Vietnam)
16. Staurogyne bicolor (Mildbr.) Champluvier (Cameroon)
Neozenkerina bicolor Mildbr.
17. Staurogyne brachiata (Hiern) Leonard (Brazil)
Ebermaiera brachiata Hiern; synonyms fide Braz (2005):
Ebermaiera gracilis Wawra, S. wawrana Leonard
18. Staurogyne brachystachya Benoist (China, Vietnam)
19. Staurogyne brevicaulis Benoist (Laos, Vietnam)
20. Staurogyne bullata Bremek. (Indonesia)
Didissandra parviflora Ridl. (non S. parviflora (T. Anderson
ex C.B. Clarke) Kuntze)
Ebermaiera axillaris Nees (non S. axillaris S. Moore);
synonym fide Hossain (1971): Staurogyne cremostachya Bremek.
Hossain (1971) indicated that Staurogyne comosa (Wall.)
Kuntze, based on Ruellia comosa Wall., is illegitimate, and he
treated it as a synonym of S. cremostachya. Although R. comosa
Wall. (non Vell. 1829) is illegitimate, Kuntze’s name, S. comosa
Kuntze, which he equated with both R. comosa Wall. and
Ebermaiera axillaris, dates from 1891 and is legitimate.
30. Staurogyne concinnula (Hance) Kuntze (China, Japansouthern Ryukyu Islands, Taiwan)
Ebermaiera concinnula Hance
Matsumura’s (1912) combination for this name would
appear to be superfluous.
31. Staurogyne condensata (Ridl.) Bremek. (Malaysia)
Staurogyne longifolia var. condensata Ridl.
32. Staurogyne coriacea (T. Anderson ex C.B. Clarke) Kuntze
(Myanmar)
Ebermaiera coriacea T. Anderson ex C.B. Clarke
Bremekamp’s (1961) combination for this name would
appear to be superfluous.
33. Staurogyne cuneata Imlay (Thailand)
21. Staurogyne burbidgei C.B. Clarke ex Bremek. (Malaysia)
34. Staurogyne dasyphylla Bremek. (Malaysia)
22. Staurogyne cambodiana (Benoist) E. Hossain (Cambodia,
Laos)
35. Staurogyne debilis (T. Anderson) C.B. Clarke ex Merr.
(Indonesia, Malaysia, Philippines, Solomon Islands, Taiwan, Vietnam)
Staurogyne polybotrya Nees var. cambodiana Benoist
23. Staurogyne capillipes Bremek. (Philippines)
24. Staurogyne capitata E.A. Bruce (Ghana, Ivory Coast,
Liberia, Sierra Leone)
Synonyms fide Champluvier (1991): Staurogyne paludosa
(Mangenot & Aké Assi) Heine, Staurogynopsis capitata (E.A.
Bruce) Mangenot & Aké Assi, Staurogynopsis maiana Mangenot & Aké Assi, Staurogynopsis paludosa Mangenot & Aké
Assi
25. Staurogyne carvalhoi Profice (Brazil)
Braz (2005) noted that this species likely represented an
undescribed genus of Acanthaceae, near or pertaining to
Nelsonioideae. She noted several putative distinctions with
respect to Staurogyne (e.g., corollas hypocrateriform and seeds
three per locule). We have not seen specimens of it.
26. Staurogyne chapaensis Benoist (China, Vietnam)
27. Staurogyne ciliata Elm. (Philippines)
Ebermaiera debilis T. Anderson
Bremekamp (1955) recognized Staurogyne debilis vars.
angustifolia Bremek., longifolia Bremek., pubifolia Bremek.,
and tomentosa Bremek.
36. Staurogyne densifolia Bremek. (Thailand)
37. Staurogyne diandra E. Hossain (Vietnam)
Hossain (1972) noted that this species was unique among
Staurogyne by having only two stamens and two or three
staminodes, and that its taxonomic relationship was uncertain.
It was treated by Xin et al. (2010) as Ophiorrhiziphyllon
diandrum (E. Hossain) H.J. Xin, D.V. Hai & Y.F. Deng. This
taxon has not been included in phylogenetic work and further
studies of it are warranted.
38. Staurogyne diantheroides Lindau (Bolivia, Brazil)
39. Staurogyne dispar Imlay (Thailand)
40. Staurogyne elegans (Nees) Kuntze (Brazil)
Ebermaiera elegans Nees
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Revision of Nelsonioideae
41. Staurogyne elongata (Blume) Kuntze (Indonesia)
Adenosma elongata Blume, Ebermaiera elongata (Blume)
Hassk., Erythracanthus elongatus (Blume) Nees; synonym fide
Bremekamp (1955): Ebermaiera subpaniculata Hassk.
Bremekamp (1955) recognized Staurogyne elongata var.
sumatrana Bremek.
37
Ebermaiera hirsuta Nees; synonym fide Braz (2005):
Staurogyne glutinosa Lindau (non Kuntze)
58. Staurogyne humifusa Bremek. (Indonesia)
59. Staurogyne hypoleuca Benoist (China, Vietnam)
Ophiorrhiziphyllon hypoleucum (Benoist) Benoist
42. Staurogyne ericoides Lindau (Brazil)
60. Staurogyne inaequalis E. Hossain (Thailand)
Synonym fide Braz (2005): Ebermaiera minarum var.
microphylla Nees
43. Staurogyne euryphylla E. Hossain (Brazil)
Synonym fide Hossain (1972): Ebermaiera riedeliana Nees
var. latifolia Nees (non S. latifolia Bremek., 1969)
44. Staurogyne eustachya Lindau (Brazil)
61. Staurogyne incana (Blume) Kuntze (Indonesia, Myanmar)
Ruellia incana Blume, Ebermaiera incana (Blume) Hassk.,
Lepidagathis incana (Blume) Nees
62. Staurogyne itatiaiae (Wawra) Leonard (Brazil)
Ebermaiera itatiaiae Wawra
45. Staurogyne expansa Bremek. (Thailand)
63. Staurogyne jaherii Bremek. (Indonesia, Malaysia)
46. Staurogyne fastigiata (Nees) Kuntze (Brazil, Venezuela)
47. Staurogyne filipes E. Hossain (Laos, Thailand)
Synonym fide Hossain (1971): Staurogyne maschalostachys
Bremek.
Bremekamp (1955) recognized Staurogyne jaherii var.
angustifolia Bremek.
Hossain (1972) noted two geographic variants but refrained
from naming them.
64. Staurogyne kamerunensis (Engl.) Benoist (Cameroon,
Nigeria)
48. Staurogyne flava Braz & R. Monteiro (Brazil)
Zenkerina kamerunensis Engl.
Champluvier (1991) recognized Staurogyne kamerunensis
subsp. calabarensis Champl.
Ebermaiera fastigiata Nees
49. Staurogyne glutinosa (Wall. ex C.B. Clarke) Kuntze
(Bangladesh, India)
Ebermaiera glutinosa Wall. ex C.B. Clarke
50. Staurogyne gracilis (T. Anderson) Kuntze (Myanmar)
Ebermaiera gracilis T. Anderson
51. Staurogyne grandiflora E. Hossain (Myanmar)
52. Staurogyne griffithiana (Nees) Kuntze (Indonesia, Malaysia, Thailand)
Erythracanthus griffithianus Nees, Ebermaiera griffithiana
(Nees) T. Anderson; synonym fide Hossain (1971): Staurogyne
setisepala C.B. Clarke
53. Staurogyne guianensis (Bremek.) T.F. Daniel & McDade,
comb. nov. (Brazil, French Guiana, Guyana, Suriname)
Gynocraterium guianense Bremek., Bull. Misc. Inform. Kew
1939: 557 (1940).
65. Staurogyne kerrii E. Hossain (Thailand)
66. Staurogyne kinabaluensis Bremek. (Malaysia)
Hossain (1971) indicated that this taxon should be treated as
a variety of Staurogyne jaherii, but did not publish such a
renovation. Bremekamp (1955) recognized S. kinabaluensis
var. angustifolia Bremek.
67. Staurogyne kingiana C.B. Clarke (Indonesia, Malaysia)
Bremekamp (1955) recognized Staurogyne kingiana vars.
angustifolia Bremek., magnifolia Bremek., and pubicalyx Bremek.
68. Staurogyne lanceolata (Blume) Kuntze (Cambodia, Indonesia, Laos, Malaysia, Myanmar, Vietnam)
Adenosma lanceolata Blume; Ebermaiera lanceolata (Blume)
Hassk.
Bremekamp (1955) recognized Staurogyne lanceolata var.
scabridula Bremek.
54. Staurogyne hainanensis C.Y. Wu & H.S. Lo (China)
55. Staurogyne havilandii C.B. Clarke ex Bremek. (Indonesia,
Malaysia)
69. Staurogyne lasiobotrys (Nees) Kuntze (Malaysia, Myanmar, Thailand)
Ebermaiera lasiobotrys Nees
56. Staurogyne helferi (T. Anderson) Kuntze (Myanmar)
Ebermaiera helferi T. Anderson
57. Staurogyne hirsuta (Nees) Kuntze (Brazil)
70. Staurogyne latifolia Bremek. (Thailand)
71. Staurogyne lepidagathoides Leonard (Brazil, Colombia,
Guyana, Suriname, Venezuela)
38
Daniel and McDade
72. Staurogyne letestuana Benoist (Angola, Congo-Brazzaville,
Congo-Kinshasa, Gabon)
Ebermaiera letestuana (Benoist) Benoist; synonym fide
Champluvier (1991): Staurogyne congoensis S. Moore
73. Staurogyne linearifolia Bremek. (Brazil, French Guiana,
Guyana, Suriname)
ALISO
Ebermaiera humilis Miq. (non S. humilis (Nees) Kuntze);
synonyms fide Braz (2005): Staurogyne agrestis Leonard, S.
fockeana Bremek., S. stahelii Bremek., S. wullschlaegeliana Bremek.
This is the most widespread species of the genus in the New
World.
89. Staurogyne monticola Benoist (Cambodia, Thailand)
Synonym fide Hossain (1971): S. kradengensis Bremek.
74. Staurogyne longeciliata Bremek. (Thailand)
90. Staurogyne neesii (S. Vidal) Merr. (Philippines)
75. Staurogyne longibracteata E. Hossain (Myanmar)
76. Staurogyne longicuneata H.S. Lo (China)
77. Staurogyne longifolia (Nees) Kuntze (Malaysia)
Ebermaiera longifolia Nees
78. Staurogyne longispica (Ridl.) Ridl. (Indonesia, Malaysia)
Ebermaiera longispica Ridl.
79. Staurogyne macclellandii (T. Anderson) Kuntze (Myanmar)
Ebermaiera macclellandii T. Anderson
80. Staurogyne macrobotrya (Kurz) T.F. Daniel & McDade,
comb. nov. (China, Laos, Myanmar, Thailand, Vietnam)
Ophiorrhiziphyllon macrobotryum Kurz, J. Asiat. Soc.
Bengal, pt. 2, Nat. Hist. 40: 76. 1871; synonym fide Hu et al.
(2011): Ophiorrhiziphyllon poilanei Benoist
81. Staurogyne macrophylla (T. Anderson ex C.B. Clarke)
Kuntze (Myanmar)
Ebermaiera macrophylla T. Anderson ex C.B. Clarke
82. Staurogyne major Benoist (Vietnam)
Ebermaiera neesii S. Vidal
Benoist (1913) treated this species as Staurogyne glauca var.
neesii (Vidal) Benoist.
91. Staurogyne novoguineensis (Kaneh. & Hatys.) B.L. Burtt
(Indonesia)
Didissandra novoguineensis Kaneh. & Hatys.; synonym fide
Hossain (1971): Staurogyne dasystachya Bremek.
92. Staurogyne obtusa (Nees) Kuntze (Myanmar, Thailand)
Erythracanthus obtusus Nees, Ebermaiera obtusa (Nees) T.
Anderson
93. Staurogyne ophiorrhizoides Elm. (Philippines)
94. Staurogyne panayensis Bremek. (Philippines)
95. Staurogyne paniculata (Wall. ex T. Anderson) Kuntze
(India, Myanmar)
Ebermaiera paniculata Wall. ex T. Anderson
Karthikeyan et al. (2009) recognized Staurogyne paniculata
var. glabrior (C.B. Clarke) Karthik. & Moorthy based on
Ebermaiera paniculata var. glabrior C.B. Clarke.
96. Staurogyne paotingensis C.Y. Wu & H.S. Lo (China)
83. Staurogyne malaccensis C.B. Clarke (Cambodia, Indonesia,
Laos, Malaysia, Myanmar, Thailand, Vietnam)
97. Staurogyne papuana Laut. (Indonesia)
Bremekamp (1955) recognized Staurogyne malaccensis var.
stenophylla Bremek.
Bremekamp (1955) recognized Staurogyne papuana var.
glabrifolia Bremek.
84. Staurogyne mandioccana (Nees) Kuntze (Brazil)
98. Staurogyne parva D.M. Braz & R. Monteiro (Brazil)
Ebermaiera mandioccana Nees; synonym fide Braz (2005):
Ebermaiera mandioccana var. triandra Hiern
85. Staurogyne merguensis (T. Anderson) Kuntze (Malaysia,
Myanmar, Thailand, Vietnam)
Ebermaiera merguensis T. Anderson
86. Staurogyne merrillii Bremek. (Philippines)
87. Staurogyne minarum (Nees) Kuntze (Brazil)
Ebermaiera minarum Nees
88. Staurogyne miqueliana Kuntze (Brazil, Costa Rica, French
Guiana, Guyana, Mexico, Nicaragua, Panama, Suriname,
Venezuela)
99. Staurogyne parviflora (T. Anderson ex C.B. Clarke) Kuntze
(Myanmar)
Ebermaiera parviflora T. Anderson ex C.B. Clarke
Bremekamp’s (1961) combination for this name would
appear to be superfluous.
100. Staurogyne perpusilla A.N. Henry & N.P. Balakr. (India)
101. Staurogyne petelotii Benoist (China, Vietnam)
102. Staurogyne polybotrya (Nees) Kuntze (India, Myanmar,
Thailand)
Ebermaiera polybotrya Nees
Hossain (1972) recognized Staurogyne polybotrya var.
humilis (Nees) E. Hossain (based on Ebermaiera humilis Nees
VOLUME 32, NUMBER 1
Revision of Nelsonioideae
5 S. humilis (Nees) Kuntze) and S. polybotrya var. thorelii
(Benoist) E. Hossain (based on S. neesii var. thorelii Benoist).
39
Ebermaiera simonsii T. Anderson
120. Staurogyne singularis Bremek. (Thailand)
103. Staurogyne pseudocapitata Champluvier (Cameroon,
Gabon)
Champluvier (1991) recorded this species only from
Cameroon. In 2004, she annotated a specimen at P from
Gabon (N. Hallé 424) as this species.
104. Staurogyne punctata J.B. Imlay (Thailand)
105. Staurogyne racemosa (Roxb.) Kuntze (Cambodia, Indonesia, Malaysia)
Ruellia racemosa Roxb., Ebermaiera racemosa (Roxb.) Miq.,
Erythracanthus racemosus (Roxb.) Nees
Benoist (1935) recognized Staurogyne racemosa ‘‘var.
pierrei’’ Benoist but the ‘‘name’’ was not validly published
because it lacked a Latin description. A subsequent valid
publication of this variety has not been seen.
106. Staurogyne ranaiensis Bremek. (Indonesia)
107. Staurogyne repens (Nees) Kuntze (Brazil)
Ebermaiera repens Nees
Hossain (1972) recognized Staurogyne singularis var. longipedicellata (Bremek.) E. Hossain, based on S. bella var.
longipedicellata Bremek.
121. Staurogyne sinica C.Y. Wu & H.S. Lo (China)
Synonym fide Hu et al. (2011): Titanotrichum parviflorum
X.B. Ye & W.T. Lin
122. Staurogyne spatulata (Blume) Koord. (Australia, Cambodia, China, India, Indonesia, Laos, Myanmar, Papua New
Guinea, Philippines, Thailand, Vietnam)
Adenosma spatulata Blume, Ebermaiera glauca var. spatulata
(Blume) C.B. Clarke; synonyms fide Benoist (1935): Ebermaiera glauca Nees, E. ligulata Bedd., Herpestis cochinchinensis Bonati; synonyms fide Hossain (1971): Staurogyne candelabrum Bremek., S. flexicaulis Bremek., S. glauca (Nees)
Kuntze, S. glauca var. siamensis (C.B. Clarke) Benoist, S.
latibracteata Bremek., S. leptocaulis Bremek., S. polycaulis
Bremek., S. rivularis Merr., S. siamensis C.B. Clarke
Barker (1986) recognized Staurogyne leptocaulis subsp.
decumbens R.M. Barker.
108. Staurogyne riedeliana (Nees) Kuntze (Brazil)
123. Staurogyne spiciflora (Miq.) Bremek. (Indonesia)
Ebermaiera riedeliana Nees
109. Staurogyne rosulata Bremek. (Thailand)
110. Staurogyne rubescens D.M. Braz & R. Monteiro (Brazil)
Ebermaiera spiciflora Miq.
Bremekamp (1955) recognized Staurogyne spiciflora vars.
glabricaula Bremek. and stenosepala Bremek.
124. Staurogyne spiciformis E. Hossain (Cambodia)
111. Staurogyne samarensis Bremek. (Philippines)
112. Staurogyne sandakanica Bremek. (Malaysia)
113. Staurogyne scandens Benoist (Vietnam)
114. Staurogyne scopulicola Kiew (Malaysia)
115. Staurogyne sesamoides (Hand.-Mazz.) B.L. Burtt (China,
Laos, Vietnam)
Loxostigma sesamoides Hand.-Mazz.; synonym fide Hu et
al. (2011): Staurogyne dolichocalyx E. Hossain
116. Staurogyne setigera (Nees) Kuntze (Indonesia, Malaysia,
Philippines, Singapore, Thailand)
Ebermaiera setigera Nees; synonyms fide Benoist (1935):
Ebermaiera subcapitata C.B. Clarke, E. trichocephala Miq.;
synonym fide Hossain (1971): Adenosma uliginosa R. Br. var.
erecta Nees
Bremekamp (1955) recognized Staurogyne setigera var.
grandis Bremek., based on S. balabacensis Quisumb.
125. Staurogyne spraguei Wassh. (Bolivia, Brazil, Colombia,
Guyana, Venezuela)
Synonym fide Wasshausen (1992): Staurogyne leptocaulis
Leonard (non Bremek.)
126. Staurogyne stenophylla Merr. & Chun. (China)
127. Staurogyne stolonifera (Nees) Kuntze (Brazil, Suriname)
Ebermaiera stolonifera Nees
128. Staurogyne strigosa C.Y. Wu & H.S. Lo (China)
129. Staurogyne subcapitata Bremek. (Philippines)
130. Staurogyne subcordata Benoist (Vietnam)
Staurogyne subcordata (Elm.) Bremek., based on Gymnostachyum subcordatum Elm., is a later homonym, and likely
pertains to a different species.
131. Staurogyne subglabra C.B. Clarke (Indonesia, Malaysia)
117. Staurogyne shanica W.W. Sm. (Myanmar)
118. Staurogyne sichuanica H.S. Lo (China)
119. Staurogyne simonsii (T. Anderson) Kuntze (India,
Vietnam)
Bremekamp (1955) recognized Staurogyne subglabra var.
prianganensis (Bremek.) Bremek. based on S. prianganensis
Bremek.
132. Staurogyne subrosulata E. Hossain (Thailand)
40
Daniel and McDade
133. Staurogyne sundana Bremek. (Indonesia)
134. Staurogyne sylvatica Lindau ex Braz & R. Monteiro
(Brazil)
135. Staurogyne tenera Benoist (Vietnam)
136. Staurogyne tenuispica Bremek. (Thailand)
137. Staurogyne thyrsoidea (Nees) Kuntze (Cambodia, India,
Myanmar, Vietnam)
Ebermaiera thyrsoidea Nees
138. Staurogyne trinitensis Leonard (Brazil, Guyana, Suriname, Trinidad & Tobago, Venezuela)
Synonym fide Braz (2005): Staurogyne versteegii Bremek.
139. Staurogyne vauthieriana (Nees) Kuntze (Brazil)
Ebermaiera vauthieriana Nees; synonym fide Hossain (1971)
and Braz (2005): Staurogyne macrantha Lindau (non C.B.
Clarke)
140. Staurogyne veronicifolia (Nees) Kuntze (Brazil)
Ebermaiera veronicifolia Nees
ALISO
Ebermaiera sanctae-catharinae Nees
Wasshausen and Smith (1969) treated this name, based on a
Tweedie collection from Santa Catarina, Brazil in Hooker’s
herbarium at K, as a synonym of S. mandioccana. Braz (2005)
did not place this name, and restricted S. mandioccana to the
Brazilian states of Rio de Janeiro and Minas Gerais. Its
placement remains unresolved. Hossain (1971) indicated that it
did not pertain to Staurogyne.
Ebermaiera velutina Nees
This name was validly published based on a Griffith collection
from Mergui, India in Hooker’s herbarium at K. A valid
combination for it has not been made in Staurogyne (see below),
and its taxonomic status remains unknown.
Ophiorrhiziphyllon laxum Lindau
Hossain (1971) referred this name to an unidentified species of
Justicia.
‘‘Staurogyne cf. densa Benoist’’
Hô’s (1993) use of this name for a Vietnamese plant was
undoubtedly an error for Strobilanthes densa Benoist (5
Strobilanthes atropurpurea Nees var. atropurpurea), which occurs
in Vietnam.
141. Staurogyne vicina Benoist (China, Vietnam)
‘‘Staurogyne floribunda Rizzini’’
142. Staurogyne viscida (Ridl.) Bremek. (Malaysia)
Limnophila viscida Ridl.
143. Staurogyne warmingiana (Hiern) Leonard (Brazil)
Ebermaiera warmingiana Hiern
144. Staurogyne yunanensis H.S. Lo (China)
145. Staurogyne zeylanica (Nees) Kuntze (India, Sri Lanka)
Ebermaiera zeylanica Nees; synonym fide Kramer (1998):
Ebermaiera glauca T. Anderson (non Nees)
Excluded or Unresolved Names
This list includes some nomina nuda that have appeared in
the literature cited.
This nomen nudum was used by Rizzini (1951, 1954), but was
never validly published.
‘‘Staurogyne glabrata Braz’’
This unpublished name was used by Braz (2005) for the species
subsequently published as S. alba Braz & R. Monteiro.
Staurogyne javanica Lindau
This name was treated by Hossain (1971) as a synonym of
Pararuellia napifera (Zoll.) Bremek.
Staurogyne macrantha C.B. Clarke, nomen illegit.
Clarke’s name is a later homonym. Hossain (1971) indicated that
this name pertains to a species of Gesneriaceae.
‘‘Staurogyne minor (Kraenzl.) B.L. Burtt’’
Ancistrostylis harmandii (G. Bonati) T. Yamaz.
Scotland and Vollesen (2000) included the genus in Staurogyne,
but no combination has been proposed for the sole species from
Laos. It remains to be determined if the name corresponds to a
previously described species of Staurogyne from southeastern
Asia.
Ebermaiera nelsonioides Miq.
Hossain (1971) noted that this was the basionym of Adenosma
nelsonioides (Miq.) Hallier f. ex Bremek. in Scrophulariaceae.
Ebermaiera nitida Rchb.f.
Hossain (1971) indicated that this name did not pertain to
Staurogyne.
This ‘‘name’’ was included by Hossain (1971), who indicated that
he had not seen the species, but no publication of it has been
located. It was likely to be based upon Hexatheca minor Kraenzl.
(Gesneriaceae from Borneo). Its taxonomic status remains
unknown, and the combination is not accepted here.
Staurogyne nudispica (C.B. Clarke) Bremek.
This name was treated by Hossain (1971) as a synonym of
Pararuellia nudispica (C.B. Clarke) Bremek.
Staurogyne palawanensis (Elmer) Bremek.
This name was treated by Hossain (1971) as a synonym of
Gymnostachyum palawanense Elmer.
VOLUME 32, NUMBER 1
Revision of Nelsonioideae
Staurogyne pauper C.B. Clarke
This name was treated by Hossain (1971) as a synonym of
Phialacanthus pauper (C.B. Clarke) Bremek.
‘‘Staurogyne pedicellata E. Hossain’’
A description in English of a collection from Indonesian Borneo,
Kostermans 12998, was treated as a new species under this name
in Hossain (1971), but the name was not subsequently validly
published.
Staurogyne serrulata C.B. Clarke
This name was treated by Hossain (1971) as a synonym of an
unidentified species of Didymocarpus (Gesneriaceae).
‘‘Staurogyne tenuicaulis E. Hossain’’
Hossain (1971) used, but did not publish, this ‘‘name’’ as a new
name for the illegitimate Staurogyne leptocaulis Leonard (non
Bremek.), to which Wasshausen (1992) gave the replacement
name S. spraguei.
‘‘Staurogyne velutina (Nees) E. Hossain’’
Hossain (1971) cited this ‘‘name’’ as a new combination based on
Ebermaiera velutina Nees, but did not subsequently validate it.
The combination is not accepted here.
‘‘Zahlbrucknera fruticosa Pohl ex Nees’’
This ‘‘name’’ was listed by Nees (1847a,b), Wasshausen and
Smith (1969), and Hossain (1971) as pertaining to Staurogyne
mandioccana. It was based on a drawing and specimen (Pohl
4740 at W). Because it was published as a synonym, the name
was not validly published.
ACKNOWLEDGMENTS
We are grateful to D. Champluvier and A. Franck for
supplying important specimens; D. Braz, D. Champluvier, E.
Hossain, and R. Wenk for their useful studies on Nelsonioideae; S. Adler, N. Bollinger, K. Douthit, N. Pugh, and M.
Tenorio for their excellent illustrations; R. Cumming, W.
Eckerman, Q. Groom, B. Hammel, C. Horions, M. Peixoto,
H. Pickering, J. Rebman, E. Tripp, D. Valke, and K.
Weinhold for permission to reproduce their photographs;
S. Serata for assistance with scanning electron microscopy;
J. Young for assistance with automontage photography;
D. Burge for rendering the phylogenetic tree; E. Bécquer for
assistance with herbarium studies in Cuba; R. Letsara, H.
Ranarivelo, and T. Randriambololona for assistance with field
studies in Madagascar; G. Carnevali, F. Chiang, J. Germán
A., L. Hernández S., E. Lott, M. Martı́nez D., J. Pascual, J.
Rzedowski, S. Salas, A. Sánchez, V. Steinmann, and J. Tapia
M. for assistance with field and herbarium studies in Mexico;
and N. Chumchim, F. Freund, C. Kiel, and G. Ocampo
for assistance in the molecular lab. The project was largely
funded by the U.S. National Science Foundation to TFD
(DEB0743273) and LAM (DEB0743178). Additional funding
for field activities was provided by the California Academy of
Sciences (In-House Research Fund, Lindsay Field Expedition
Fund, Lewis and Elise Rose Memorial Fund), Gulf of Guinea
Project Fund (studies in São Tomé e Prı́ncipe), National
41
Geographic Society (studies in Guatemala), and American
Philosophical Society (studies in Arizona and Mexico). For
loans of herbarium specimens and/or arranging visits we thank
the curators of the following herbaria: BM, BP, BR, BRLU, C,
CAS, CICY, CO, DS, DUKE, EAP, ENCB, F, FCME,
FLAS, FSU, G, GA, GZU, HAL, IBUG, IEB, JE, K, L,
LISC, LL, LY, MICH, MEXU, MO, NCSC, NCU, NY, P,
PH, POM, QMEX, RSA, S, SERO, STPH, TAN, TEX, U,
UC, UPS, US, USCH, USF, VSC, and WAG.
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VOLUME 32, NUMBER 1
Revision of Nelsonioideae
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APPENDIX 1
Pollen of Nelsonioideae studied with SEM. The name of each taxon
studied is followed by collection data (country, collector’s number,
herbarium of deposit), measurements (not all measurements could be
made on all grains), and observations on endoapertures (not evident,
evident but not prominent, prominent). n 5 number of grains
measured, P 5 polar diameter (mm), E 5 equatorial diameter (mm).
Pollen of additional samples and species (e.g., Elytraria klugii) was also
44
Daniel and McDade
studied, but was not in a suitable orientation or condition for accurate
measurements. Collection numbers with ‘‘cv’’ or ‘‘gh’’ refer to
cultivated collections.
Anisosepalum alboviolaceum (Congo-Brazzaville, Champluvier 5295,
CAS): n 5 2; P 5 30–31; E 5 22–23; P:E 5 1.35–1.36;
endoapertures not evident.
Anisosepalum alboviolaceum (Congo-Kinshasa, Louis 12208, CAS): n
5 5; P 5 22; E 5 23–25; P:E 5 0.92–0.96; endoapertures not
evident.
Anisosepalum humbertii (Congo-Kinshasa, de Witte 7121, CAS): n 5 2;
P 5 24; E 5 26–32; P:E 5 0.90–0.92; endoapertures evident but not
prominent.
Anisosepalum humbertii (Tanzania, Congdon 591, K): n 5 3;
P 5 28–30; E 5 22–25; P:E 5 1.27–1.36; endoapertures evident
but not prominent.
Anisosepalum lewallei (Burundi, Reekmans 3610, WAG): n 5 2; P 5
27; E 5 23–24; P:E 5 1.17; endoapertures not evident.
Elytraria acaulis (India, Haines 3435, K): n 5 2; P 5 40–41;
E 5 23–26; P:E 5 1.58–1.74; endoapertures not evident.
Elytraria acaulis (Tanzania, Iversen et al. 86369, UPS): n 5 3; P 5 44–
46; E 5 24; P:E 5 1.83–1.92; endoapertures not evident.
Elytraria bromoides (Mexico, Daniel 303, CAS): n 5 1; P 5 50; E 5 27:
P:E 5 1.85; endoapertures not evident.
Elytraria bromoides (Mexico, Daniel & Baker 3683, CAS): n 5 1; P 5
51; E 5 25; P:E 5 2.04; endoapertures not evident.
Elytraria caroliniensis (USA, Daniel s.n. cv, CAS): n 5 3; P 5 56–58;
E 5 28–36; P:E 5 1.56–2.07; endoapertures not evident.
Elytraria imbricata (Bolivia, Daniel 10184, CAS): n 5 3; P 5 42–44;
E 5 25–30; P:E 5 1.47–1.76; endoapertures not evident.
Elytraria imbricata (Mexico, Daniel & Butterwick 3259gh, CAS): n 5 2;
P 5 45; E 5 27–35; P:E 5 1.67; endoapertures not evident.
Elytraria imbricata (Mexico, Porter 297, CAS): n 5 1; P 5 44; E 5 25;
P:E 5 1.76; endoapertures not evident.
Elytraria ivorensis (Ghana, Enti GC42410, WAG): n 5 2; P 5 35;
E 5 27–31; P:E 5 1.30; endoapertures not evident.
Elytraria macrophylla (Mexico, Carranza 2397, CAS): n 5 2;
P 5 42–43; E 5 27; P:E 5 1.56–1.59; endoapertures not evident.
Elytraria macrophylla (Mexico, Dressler 2619, DUKE): n 5 1; P 5 not
determined; E 5 28; P:E 5 not determined; endoapertures not
evident.
Elytraria madagascariensis (Madagascar, Humbert & Cours 32602, P):
n 5 3; P 5 28–29; E 5 34; P:E 5 0.82; endoapertures evident but
not prominent.
Elytraria marginata (Guinea-Bissau, Malaisse & Claes 14802, CAS):
n 5 6; P 5 31–35; E 5 30–33; P:E 5 0.97–1.13; endoapertures
not evident.
Elytraria maritima (Ivory Coast, de Wilde 877, WAG): n 5 3; P 5 30–
32; E 5 29–33; P:E 5 0.91–1.10; endoapertures not evident.
Elytraria mexicana (Mexico, Daniel & Butterwick 3263gh, CAS): n 5 5;
P 5 49–53; E 5 29–31; P:E 5 1.65–1.83; endoapertures not evident.
Elytraria minor (Kenya, Greenway & Rawlins 9366, K): n 5 4; P 5 31–
33; E 5 26–29; P:E 5 1.14–1.19; endoapertures not evident.
Elytraria nodosa (Madagascar, Almeda 9232, CAS): n 5 5; P 5 44–52; E 5
23–26; P:E 5 1.69–2.26; endoapertures not evident.
Elytraria nodosa (Madagascar, Daniel 11838, CAS): n 5 4; P 5 28–49;
E 5 20–31; P:E 5 0.90–2.25; endoapertures not evident to evident
but not prominent.
Elytraria nodosa (Madagascar, Goudot s.n., P): n 5 3; P 5 33–34;
E 5 14–16; P:E 5 2.06–2.43; endoapertures not evident.
Elytraria tuberosa (Ecuador, Eggers 15405, S): n 5 4; P 5 25–31;
E 5 17–28; P:E 5 0.96–1.82; endoapertures not evident.
Nelsonia canescens (Panama, Daniel 5452, CAS): n 5 3; P 5 34–35;
E 5 17–18; P:E 5 1.94–2.00; endoapertures not evident.
Nelsonia canescens (Panama, Daniel 5452cv, CAS): n 5 3; P 5 34–39;
E 5 19–21; P:E 5 1.62–2.05; endoapertures not evident.
ALISO
Nelsonia canescens (São Tomé & Prı́ncipe, Daniel 11186, CAS): n 5 2;
P 5 37–38; E 5 17–18; P:E 5 2.11–2.18; endoapertures not evident.
Nelsonia gracilis (Zambia, Bingham & Mpundu 122660, K):
n 5 5; P 5 23–28; E 5 15–18; P:E 5 1.28–1.87; endoapertures
not evident.
Saintpauliopsis lebrunii (Congo-Kinshasa, Humbert 7625, CAS): n 5 2;
P 5 21; E 5 24–26; P:E 5 0.88; endoapertures not evident.
Saintpauliopsis lebrunii (Madagascar, Gautier et al. 3598, P): n 5 1;
P 5 not determined; E 5 26; P:E 5 not determined;
Saintpauliopsis lebrunii (Rwanda, Bamps 3270, BR): n 5 2;
P 5 38–43; E 5 21–23; P:E: 5 1.81–1.87; endoapertures not
evident.
Staurogyne concinnula (Taiwan, Bartholomew & Boufford 6215, CAS):
n 5 2; P 5 36–40; E 5 19–20; P:E 5 1.89–2.00; endoapertures not
evident.
Staurogyne guianensis (Guyana, Jansen–Jacobs et al. 2346, US): n 5 1;
P 5 42; E 5 19; P:E 5 2.21; endoapertures not evident.
Staurogyne macrobotrya (China, Henry 11627, US): n 5 1; P 5 not
determined; E 5 18; P:E 5 not determined; endoapertures not
evident.
Staurogyne macrobotrya (Thailand, Hansen & Smitinand 12817, C);
n 5 2; P 5 27–29; E 5 13–14; P:E 5 1.93–2.23; endoapertures
not evident.
Staurogyne major (Thailand, Kerr 10264): n 5 1; P 5 not determined;
E 5 26; P:E 5 not determined; endoapertures prominent.
Staurogyne mandioccana (Brazil, Cordeiro & Silva 108, CAS):
n 5 4; P 5 22–29; E 5 16–21; P:E 5 1.10–1.81; endoapertures
not evident.
Staurogyne miqueliana (Mexico, Lott et al. 3226, CAS): n 5 3;
P 5 27–30; E 5 13; P:E 5 2.08–2.31; endoapertures not evident.
Staurogyne miqueliana (Mexico, Lott et al. 3396, MEXU): n 5 1; P 5
24; E 5 25; P:E 5 0.96; endoapertures prominent.
Staurogyne spatulata (Indonesia, Toroes 4795, MICH): n 5 4; P 5 19–
25; E 5 14–18; P:E 5 1.11–1.79; endoapertures not evident to
prominent.
Staurogyne spatulata (Thailand, Maxwell 94-76, CAS): n 5 3; P 5 21;
E 5 22; P:E 5 0.95; endoapertures evident but not prominent.
Staurogyne stolonifera (unknown, Daniel s.n. cv, CAS): n 5 2; P 5 32–
36; E 5 15–17; P:E 5 2.12–2.13; endoapertures not evident.
APPENDIX 2
Results of tests for hygroscopic trichomes among Nelsonioideae by
hydration of seeds from herbarium specimens. At least two seeds per
collection were submerged in tap water and observed under a
dissecting microscope at 643 for at least 3 minutes; some samples
were photographed with an automontage camera system in both the
dry and wet states (Fig. 9); these and/or additional samples were
observed with SEM (Fig. 10–12) to observe seed surface features,
including trichomes. The name of each taxon is followed by the
collection/herbarium sampled, date of the collection, and the reaction
observed. No reaction 5 no change observed in hydrated seeds vs. dry
seeds; moderate reaction 5 only some trichomes become erect on
hydrated seeds and/or this reaction is not immediate; strong reaction 5
most or all trichomes become erect immediately upon hydration.
Collection numbers with ‘‘cv’’ or ‘‘gh’’ refer to cultivated collections.
Anisosepalum alboviolaceum (Champluvier 5295/CAS, 1995): no reaction.
Anisosepalum lewallei (Bidgood et al. 4728/CAS, 2000): no reaction.
Elytraria caroliniensis (Daniel s.n. cv/CAS, 2007): no reaction.
Elytraria imbricata (Daniel & Butterwick 3201/CAS, 1983): no reaction.
Elytraria macrophylla (Daniel 11773 cv/CAS, 2013): no reaction.
Elytraria marginata (Daniel 11158 cv/CAS, 2013): no reaction.
Nelsonia canescens (Daniel 11186/CAS, 2008): strong reaction.
Nelsonia canescens (Balakrishnan 998/L, 1974): moderate reaction.
Nelsonia canescens (Rosales 111/CAS, 2000): moderate reaction in
some seeds and strong reaction in others.
VOLUME 32, NUMBER 1
Revision of Nelsonioideae
Nelsonia gracilis (Henriquez 663/BM, 1965): strong reaction.
Saintpauliopsis lebrunii (Humbert & Capuron 28443/P, 1955): moderate
reaction.
Saintpauliopsis lebrunii (Perrier de la Bathie 17106/P, 1925): moderate
reaction.
Staurogyne concinnula (Peng 6777/CAS, 1984): moderate reaction.
Staurogyne debilis (Merrill 1215/MO, 1913): no reaction in some seeds
and moderate reaction in others.
Staurogyne diantheroides (Wasshausen 2181/CAS, 1998): moderate
reaction.
Staurogyne kamerunensis (McPherson 13913/MO, 1989): no reaction in
some seeds and moderate reaction in others.
Staurogyne letestuana (Arends et al. 358/MO, 1984): no reaction in
some seeds and moderate reaction in others.
Staurogyne longeciliata (Maxwell 76-7/MO, 1976): moderate reaction
in some seeds and strong reaction in others.
Staurogyne macrobotrya (Poilane 20703/CAS, 1932): moderate reaction.
Staurogyne merguensis (Maxwell 94-254/CAS, 1994): moderate reaction.
Staurogyne miqueliana (Lott et al. 3226/CAS, 1991): strong reaction.
Staurogyne obtusa (Maxwell 90-362/CAS, 1990): moderate reaction in
some seeds and strong reaction in others.
Staurogyne setigera (Toroes 4798/MICH, 1933): moderate reaction.
45
Staurogyne spatulata (Tsang 126/MO, 1928): no reaction in some seeds
and moderate reaction in others.
APPENDIX 3
Chromosome numbers reported for Nelsonioideae. All counts for a
taxon are shown as meiotic (i.e., haploid or n) numbers, followed by
the author(s) reporting the count.
Elytraria acaulis (17, Kaur 1969; 19, Morton 1956 [as E. lyrata]; 22,
Govindarajan and Subramanian 1983; 23, Subramanian and
Govindarajan 1980).
Elytraria bromoides (12, Daniel et al. 1990).
Elytraria caroliniensis (25, Grant 1955 [as E. virgata]).
Elytraria imbricata (11, Ward 1984; 12, Daniel et al. 1990).
Elytraria marginata (19, Morton 1956; 22, Mangenot and Mangenot
1962).
Elytraria maritima (19, Morton 1956).
Elytraria mexicana (12, Daniel et al. 1990).
Nelsonia canescens (16, Joseph 1964 [as N. campestris]; 14, Sarkar et al.
1980 [as N. campestris]; 17 + B, Saggoo and Bir 1982 [as N.
campestris]; 17 + B, Saggoo 1983; 17 + B, Saggoo and Bir 1983; 18,
Daniel and Chuang 1993).