ISSN: 0075-5974 (print)
ISSN: 1874-933X (electronic)
KEW BULLETIN
(2019) 74:39
DOI 10.1007/S12225-019-9828-Z
Two new genera of Acanthaceae from tropical Africa
Iain Darbyshire1 , Carrie A. Kiel2, Thomas F. Daniel3, Lucinda A. McDade2 & W. R. Quentin Luke4
Summary. DNA sequence data, macro-morphological evidence and pollen analysis are used to clarify the phylogenetic
placement of two African species of Acanthaceae: Schaueria populifolia C.B.Clarke and Rhinacanthus ndorensis Schweinf.
The combined data demonstrate that these species are only distantly related to the genera in which they are currently
placed and provide strong support for recognition of new genera to accommodate them. Two new genera are therefore
proposed and a taxonomic account is provided for each of these. The first, Champluviera I.Darbysh., T.F.Daniel &
C.A.Kiel, is based primarily on S. populifolia but Chlamydocardia nuda C.B.Clarke (= Justicia tigrina Heine) is also included
within this new genus based on shared morphological traits, at least one of which is synapomorphic and restricted to
these species. Based on molecular evidence, Champluviera is placed in a well-supported clade of several genera that are
together sister to the core Isoglossinae lineage of tribe Justicieae. The genus may be recognised morphologically by the
combination of a dense spiciform terminal thyrse with the bracts, bracteoles and calyx lobes all slender and closely
resembling one another; a bilabiate corolla lacking a rugula and with intricate speckling or striping on the lips; and, in
particular, the putatively synapomorphic trait of anther thecae with broad, flattened, basal appendages that have an
irregularly toothed fringe. The second new genus, Kenyacanthus I.Darbysh. & C.A.Kiel, is based on R. ndorensis and is so
far thought to be monospecific. It is placed between core Diclipterinae and Monechma Group II within the expanded
subtribe Diclipterinae in tribe Justicieae. Kenyacanthus can be recognised morphologically by having the combination of
a trailing or procumbent habit; fasciculate inflorescences; bracts, bracteoles and calyces with hyaline margins; a
bilabiate corolla with a slender cylindrical tube longer than the limb and with a shallow rugula; stenotribic flowers, with
the stamens held against the lower lip; and anthers with only slightly offset thecae that lack appendages.
Key Words. Floristics, justicioid, phylogeny, taxonomy.
Introduction
With at least 2000 species and a nearly worldwide
distribution, the Justicieae lineage of the Acanthaceae
has long challenged taxonomists. Considerable progress
has been made in the last 20 years in understanding this,
the largest of the major lineages of Acanthaceae.
However, recent phylogenetic work (e.g., Kiel et al. 2006,
2017; Daniel et al. 2008; McDade et al. 2000, 2018), as well
as floristic and taxonomic research (e.g., Côrtes et al. 2016;
Daniel 1995a, 2015, 2017; Darbyshire & Govaerts 2017;
Darbyshire & Goyder 2019), has pointed to a number of
outstanding problems that require resolution. Here, we
propose solutions for two such taxonomic problems
involving the African plants Schaueria populifolia
C.B.Clarke and Rhinacanthus ndorensis Schweinf.
Schaueria populifolia
The genus Schaueria Nees (1839: 3) was originally
described from the forests of eastern Brazil, and is best
known for the species S. calytricha (Hook.)
A.L.A.Côrtes (“golden plume”), which is popular in
botanical gardens (Côrtes et al. 2016). Schaueria is a
member of the “Tetramerium lineage” of the tribe
Justicieae (Daniel et al. 2008; Côrtes et al. 2016;
McDade et al. 2018). The genus is exclusively New
World (NW) in distribution with the notable exception
of S. populifolia C.B.Clarke (1900a: 242), which is
found in the lowland rainforests of eastern Nigeria,
Bioko and western and southern Cameroon. Daniel
et al. (2008) noted that there are otherwise no genera
of the Tetramerium lineage common to both the Old
World (OW) and NW. As such, it would be remarkable
if this African species were indeed a Schaueria.
A recent taxonomic revision of Schaueria by Côrtes
et al. (2016), using both molecular and morphological
evidence, demonstrated that Schaueria s.l. is polyphyletic. As a result, six species traditionally included in
Schaueria were excluded following a recircumscription
of the genus. The 14 species of Schaueria treated in the
revision are all restricted to Brazil (Côrtes et al. 2016).
Accepted for publication 25 June 2019.
1
Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, UK. e-mail: i.darbyshire@kew.org
2
Rancho Santa Ana Botanic Garden, Claremont Graduate University, 1500 North College Avenue, Claremont, CA, 91711, USA.
3
California Academy of Sciences, 55 Music Concourse Drive, San Francisco, CA, 94118, USA.
4
East African Herbarium, National Museums of Kenya, P.O. Box 45166, Nairobi, 00100, Kenya.
© The Author(s), 2019
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Schaueria populifolia was not included in these authors’
molecular studies, but they rejected this species from
Schaueria on the basis that then unpublished molecular data by L. McDade show that S. populifolia is more
closely related to the Isoglossinae lineage than to the
Tetramerium lineage. These data have subsequently
been published in an expanded sampling of the
Tetramerium lineage based on six gene regions
(McDade et al. 2018). These authors showed that
S. populifolia, represented by Gereau et al. 5666 (MO)
from Cameroon, was resolved with taxa of Isoglossinae
that were sampled as outgroups. However, the authors
noted that taxon sampling among outgroups was
insufficient for the results to be considered conclusive
regarding the placement of S. populifolia.
Côrtes et al. (2016) did not comment on any
morphological differences between Schaueria s.s. and
S. populifolia. The two are superficially similar, sharing
many-flowered inflorescences comprising a series of
dichasial inflorescence units; long narrow bracts, bracteoles and calyx lobes; a bilabiate corolla lacking a rugula
(stylar furrow); two bithecous stamens and stipitateclavate 4-seeded capsules with tuberculate seeds. However, these similarities are symplesiomorphic or of
uncertain phylogenetic status and thus do not
constitute evidence of a close relationship.
KEW BULLETIN
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species sampled. However, they found that two species
of the genus fall outside this clade: R. virens (Nees)
Milne-Redh. and R. ndorensis. The former, a GuineoCongolian forest species of West Africa, was placed
basally among members of the Diclipterinae lineage; it
is tentatively maintained in Rhinacanthus whilst we
await further molecular investigation of this species
and the other forest species of Rhinacanthus
(Darbyshire et al. 2018). The latter was placed sister
to core Diclipterinae, a lineage that comprises
Dicliptera Juss., Peristrophe Nees (sometimes treated as
congeneric with Dicliptera; Darbyshire & Vollesen
2007), Xerothamnella C.T.White and Hypoestes R.Br.
For this study, we combine macro-morphological
data, pollen analysis and DNA sequence data for
Schaueria populifolia and Rhinacanthus ndorensis with
the goal of clarifying the phylogenetic placement of
these two taxa. With regard to the former, we test the
hypothesis that this African species is not related to its
American congeners in the context of a more
expansive sample of OW taxa that was included in
earlier studies. Regarding Rhinacanthus, we seek to test
the hypothesis that R. ndorensis is not closely related to
Rhinacanthus s.s.
Materials & Methods
Rhinacanthus ndorensis
Rhinacanthus ndorensis was described by Schweinfurth
(1892) from Ndoro near Mt Kenya. It is restricted to
central Kenya where it occurs in grassland and open
woodland, typically on seasonally damp black clay soils
(Darbyshire et al. 2010). The species was maintained
within the genus Rhinacanthus Nees for the Flora of
Tropical East Africa account of Acanthaceae
(Darbyshire et al. 2010) and for related studies on
the genus in Africa (Darbyshire & Harris 2006),
primarily on the basis of the slender corolla tube that
is longer than the lips and the anther thecae of each
stamen being offset and basally muticous. However,
keys to the species of Rhinacanthus presented in those
publications identify R. ndorensis first due to a number
of unique characters including its procumbent or
trailing habit, fasciculate inflorescences, hyaline margins to the bracts, bracteoles and calyx lobes and
essentially glabrous capsules.
In a phylogenetic study of the ‘justicioid’ lineage of
Justicieae based on six gene regions, Kiel et al. (2017)
sampled one voucher of Rhinacanthus ndorensis (Young
3169, EA) together with 10 other accessions of
Rhinacanthus representing eight additional species.
They found that all sampled Rhinacanthus are resolved
in an expanded Diclipterinae lineage, but that
Rhinacanthus s.l. is not monophyletic (Kiel et al. 2017,
Fig. 8). These authors recognised a well-supported
Rhinacanthus clade comprising seven of the nine
© The Author(s), 2019
Morphological Analysis
Herbarium specimens of the focal taxa and related
groups held at the BM, K, MO and P herbaria were
examined, supported by study of digital images of
specimens from the BR, L, LBV and WAG herbaria
accessed via their respective online catalogues or by
individual request. Abbreviations for herbaria follow
Index Herbariorum (Thiers 2019). All duplicates of
specimens seen are marked in the “Specimens Examined” sections with an exclamation mark (!). Those for
which only online images have been seen are marked
with an asterisk (*). The herbarium research was
supplemented by field studies and photographs of wild
plants where possible.
All measurements were made on dried material
except for those of the flowers. Dried flowers were
soaked in Aerosol OT 5% solution prior to dissection
and measurement.
Pollen analysis
For the study of pollen of Schaueria populifolia,
individual grains were removed from multiple anthers
on the specimen Cheek 7771 using an insect pin. The
grains were mounted onto a SEM stub coated with
double-sided sticky tape, sputter coated with
gold/palladium and examined under SEM at CAS.
Study of pollen of Rhinacanthus ndorensis followed a
similar methodology except that a whole anther was
extracted from a flower bud on Luke 17084 and
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dissected to extract the pollen. Also in this case, the
SEM stub was sputter coated with platinum before
examination under SEM at K.
Molecular analysis
For this study, new sequence data were generated for nine
taxa (Appendix 1). These were supplemented by
downloading data for an additional 94 accessions from
GenBank (Appendix 1). Most of these latter data were
generated for previous studies in Justicieae (McDade et al.
2000, 2008, 2018; Kiel et al. 2006, 2017; Daniel et al. 2008).
To make the datasets congruent across the five loci
sampled, we sequenced an additional 94 samples
(Appendix 1) for the final dataset totalling 103 taxa. For
the newly generated sequences, leaf tissue was extracted
using Doyle & Doyle’s (1987) CTAB method. Procedures
to amplify the cp trnT-L, trnL-F and trnS-G spacers, and
rps16 intron were as described by Daniel et al. (2008).
Procedures for amplification of nrITS are found in
McDade et al. (2000) and Daniel et al. (2008). PCR
products were cleaned using PEG precipitation (Morgan
& Soltis 1993) and the purified PCR products were cycle
sequenced with both forward and reverse PCR primers
using Applied Biosystems (ABI, Foster City, California)
PRISM BigDye version 3.1, according to the manufacturer’s specifications. All sequenced reactions were
cleaned with Sephadex G50 and sequenced on an ABI
3100 genetic analyzer. Chromatograms were edited and
aligned manually in Geneious® version 9.15 (https://
www.geneious.com) and alignments were deposited in
TreeBASE, 23681.
Data matrices for the five DNA regions were
prepared as nexus files for Maximum Likelihood and
Bayesian analyses in Geneious 9.1.5. Possible incongruence between the nrITS and cp data sets was
examined by comparing the results of majority rule
trees from Bayesian inference (analyses conducted as
described below). We used a posterior probability
value of 0.90 as a conservative threshold for wellsupported conflict. Finding no evidence of incongruence between nrITS and cp data, we combined the
datasets for further analyses.
Bayesian inference (BI) of phylogenetic relationships with posterior probabilities (BPP) as the support
measure was conducted in MrBayes 3.2.6
(Huelsenbeck & Ronquist 2001; Huelsenbeck et al.
2001; Ronquist & Huelsenbeck 2003). Models of
molecular evolution for the genic regions were
selected using the Akaike information criterion (AIC)
(Akaike 1974; Posada & Crandall 2001) as implemented in JModelTest 2.1.3 (Posada 2008; Darriba et al.
2012). The selected models of nucleotide substitution
were GTR + G for the combined plastid dataset and
GTR + G + I for nrITS. Each dataset was run with data
partitions corresponding to the combined cp and
nrITS regions. Two simultaneous runs of four Monte
Carlo Markov Chains were run for 20,000,000
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generations with a random tree as the starting point
and saving trees every 1,000 generations. Results were
inspected in Tracer version 1.5 (Rambaut et al. 2014)
to determine the burn-in period. Analysis of the
nuclear + cp dataset reached stationarity (i.e., average
standard deviation of split frequencies ≤ 0.01) after
approximately 5,000,000 generations and the first 20%
of the sampled trees were discarded as burn-in before
constructing a majority rule consensus tree.
The three data sets (i.e. cp, nrITS, cp + nrITS) were
also analysed using maximum likelihood (ML) using
RAxML ver. 8.0.0 (Stamatakis 2014). For analysis, the
combined data were partitioned under the same models
of evolution as used for Bayesian inference, and 1000
bootstrap replicates were implemented. Each analysis
was repeated three times to test the consistency of tree
topology and bootstrap values (MLBS).
Species conservation (extinction risk) assessments
The species conservation assessments are based on the
Categories and Criteria of the IUCN Red List (2012).
The Extent of Occurrence (EOO) was calculated
using the Kew Geospatial Conservation Assessment
Tool GeoCAT (geocat.kew.org; Bachman et al. 2011).
Results
The topologies returned by BI and ML are congruent
and are also congruent with the results of all earlier
studies of Justicieae (e.g., McDade et al. 2000, 2018;
Kiel et al. 2006, 2017; Daniel et al. 2008; Côrtes et al.
2016). We emphasise here the placement of the two
focal taxa and do not reiterate the relationships
established in earlier studies, a more complete review
of relationships within Justicieae will be presented in a
forthcoming paper (Kiel et al. in prep).
The three accessions of Schaueria populifolia (Fig. 1)
are together monophyletic with strong support (1.00
BPP; 100 MLBS); this clade is in turn part of a wellsupported clade (1.00 BPP; 100 MLBS) of several
genera that are together sister with weak support (0.75
BBP; 51 MLBS) to the core Isoglossinae lineage (1.00
BPP; 100 MLBS) (Fig. 1). Accessions of Schaueria s.s.
are placed phylogenetically distant from S. populifolia
in the Tetramerium lineage (Fig. 1).
Two accessions of Rhinacanthus ndorensis (Fig. 2) are
sister with strong support (1.00 BPP; 99 MLBS) and
are together sister to Dicliptera s.l. + Hypoestes with
strong support for this relationship (1.00 BPP; 100
MLBS). Rhinacanthus virens is a member of a weakly
supported basal lineage (0.96 BPP; 56 MLBS) in
Diclipterinae and is sister to Justicia aff. tenuipes with
strong support (1.00 BPP; 98 MLBS). All other
sampled Rhinacanthus are members of the core
Rhinacanthus clade (1.00 BPP; 88 MLBS) and are not
closely related to R. ndorensis (Fig. 2).
© The Author(s), 2019
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KEW BULLETIN
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Justicieae
1.00
BPP
1.00
BS
90
justicioid lineage
100
Tetramerium
lineage
1.00
91
Monothecium aristatum
1.00
92
Schaueria s.s.
M. glandulosum
Jadunia biroi
1.00
1.00
100
100
1.00
-
65 0.80
62
1.00
1.00
100
Calycacanthus magnusianus
Ptyssiglottis pubisepala
Ptyssiglottis psychotriifolia
90
Marcania grandiflora
1.00
Schaueria populifolia 1336
98
0.75
1.00
51
100
0.80
71
1.00
S. populifolia 5666
S. populifolia 10153
core Isoglossinae
100
outgroups
1.00
92
Pseuderanthemum
lineage
Fig. 1. Summary of phylogenetic relationships among Justicieae highlighting Schaueria populifolia (= Champluviera populifolia)
from Maximum likelihood. BPP = values above branch and MLBS = values below branch.
Analysis of DNA sequence data thus supports recognition of two new genera to accommodate Schaueria
populifolia and Rhinacanthus ndorensis. Supporting evidence from the morphological analyses, including pollen,
will be described in detail in the Discussion below.
Discussion and taxonomic treatments
1. Champluviera — a new genus from the
forests of central Africa
Our study of specimens of Schaueria populifolia found
notable morphological differences between plants of
this African species and Brazilian congeners. Brazilian
Schaueria (henceforth Schaueria s.s.) have white or
yellow flowers lacking speckling, whilst those of
S. populifolia are dull red, red-brown or purple with
intricate speckling on at least the lower lip (Fig. 3).
More significantly, in Schaueria s.s., the anther thecae
of each stamen lack basal appendages and are
glabrous, whereas those of S. populifolia have a broad,
flattened and irregularly fringed basal appendage and
the upper theca has a fringe of trichomes on the
margin that is not adpressed to the second theca.
© The Author(s), 2019
Pollen of Schaueria populifolia is remarkable in that the
ora are markedly enlarged in comparison to the colpi
and the two pseudocolpi in each mesocolpium curve
toward one another equatorially and sometimes fuse at
the equator, forming an hour-glass shape or figure eight
(Fig. 4). In contrast, pollen of Schaueria s.s. have ora that
are less markedly enlarged and the pseudocolpi are ±
parallel to the apertures such that the two pseudocolpi of
each mesocolpium do not form an hour-glass shape (see
Côrtes et al. 2016, Fig. 6).
These morphological differences indicate that the
resemblance between Schaueria s.s. and S. populifolia is
superficial and that they are not closely related, thus
supporting the findings of the molecular analysis (Fig. 1).
On studying specimens of Schaueria populifolia
during her research on the Acanthaceae of West and
Central Africa, Dominique Champluvier (formerly of
BR) proposed the new combination Justicia populifolia
(C.B.Clarke) Champl. and annotated specimens at the
Kew herbarium as such in April 1999. This name was
adopted by Onana (2011, 2013) as an unpublished
combination in the checklist of vascular plants of
Cameroon, but has never been validly published. This
new combination was proposed on the basis that the
flowers, and in particular the unique appendages of
the anthers described above, bear a close resemblance
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Fig. 2. Summary of phylogenetic relationships among Diclipterinae in the “justicioid” lineage highlighting Rhinacanthus ndorensis
(= Kenyacanthus ndorensis) from Maximum likelihood. BPP = values above branch and MLBS= values below branch.
© The Author(s), 2019
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Fig. 3. Champluviera populifolia in the field, Cameroon.
© The Author(s), 2019
KEW BULLETIN
PHOTOS:
A J.
OSBORNE
(2019) 74:39
(collected as Osborne 97); B & C M.
CHEEK.
KEW BULLETIN
(2019) 74:39
Fig. 4. Pollen of Champluviera populifolia. A equatorial view
showing the compound aperture and pseudocolpi; B equatorial view showing the mesocolpium with equatorially fused
pseudocolpi (from Cheek 7771).
to those of Justicia tigrina Heine from Gabon (D.
Champluvier, pers. comm.). Justicia tigrina is based on
Chlamydocardia nuda C.B.Clarke, a species that Clarke
(1900b) had only tentatively placed in Chlamydocardia,
noting that it lacked the characteristic bracts of that
genus (see Darbyshire & Govaerts 2017). Heine (1966)
recorded the anther appendage of J. tigrina as
resembling a chicken’s crest (“en forme de crête de
poule”, p. 220) which is an apt description for the
flattened broad appendage with an irregularly toothed
fringe. He further noted that this appendage corresponds well to that of the other species of Justicia
treated in the Flore du Gabon account, and it was on
this basis that he transferred C. nuda to Justicia. In doing
so, Heine, was obliged to give the species a new name,
because the epithet “nuda” was already occupied in
Justicia — J. nuda (Nees) O.Schwartz (1939: 259).
We concur with Dr Champluvier that, based on
morphological evidence, Schaueria populifolia and
Justicia tigrina are closely allied. They are vegetatively
very similar, have a similar inflorescence type and the
anther morphology is closely similar except that the
thecae are more markedly offset in J. tigrina. However,
neither the morphological nor the molecular evidence
support the placement of these species within Justicia.
Contrary to Heine’s (1966) statement, the anther
appendages differ significantly from African species
of Justicia where often only the lower theca has an
appendage (but note that appendages on both thecae
is more frequent in New World Justicia — see, for
example, Daniel 1995b; Kiel et al. 2017, Fig. 4E – F)
and the appendage is usually longer than it is broad,
often markedly so, and is untoothed except sometimes
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for a forked apex. This is in marked contrast to the
broad, short and flattened appendage with an
irregularly toothed fringe in S. populifolia and
J. tigrina (Fig. 5). Furthermore, both of these latter
species lack a rugula (stylar furrow), whilst species of
Justicia have rugulate corollas (Kiel et al. 2017).
Unfortunately, suitable material for DNA analysis of
Justicia tigrina was not available for the current study
and so we have been unable to test the phylogenetic
relationship between Schaueria populifolia and J. tigrina.
However, as just described, the detailed morphological examination conducted in this study strongly
supports the hypothesis that they are closely related.
Our results place Schaueria populifolia in a wellsupported clade of several genera that are together
sister to the core Isoglossinae lineage, albeit without
strong support (Fig. 1). This result is congruent with
morphological evidence. Core Isoglossinae, a clade
that includes the large genera Isoglossa Oerst. (OW)
and Stenostephanus Nees (NW), is well-marked by
structural synapomorphies: plants have gürtelpollen,
i.e. biporate pollen that is circular in apertural view
and with a broad marginal girdle (Kiel et al. 2006),
although this can be modified to 3 (– 4)-porate
pollen in Brachystephanus Nees (Figueiredo & KeithLucas 1996; Champluvier & Darbyshire 2009). In
contrast, the clade of genera sister to Core
Isoglossinae, including S. populifolia and Ptyssiglottis
T.Anderson, have different pollen types (Kiel et al.
2006). This clade includes only one other African
representative so far as is known at present, that
being Monothecium Hochst. Monothecium comprises
two species in continental Africa one of which
extends to India; a third, Malagasy species
(M. leucopterum Benoist) is unlikely to belong to
Monothecium in view of several morphological differences (I. Darbyshire, pers. obs.). Monothecium is easily
separated from S. populifolia and J. tigrina in, amongst
other differences, having monothecous anthers lacking appendages (see Vollesen in Darbyshire et al.
2010).
The other genera allied to Schaueria populifolia are
confined to Asia: Calycacanthus K.Schum., Jadunia
Lindau, Marcania J.B.Imlay and Ptyssiglottis. Of these,
only the last could potentially be confused with the
African plants. Ptyssiglottis was most recently revised
by Hansen (1992) who recognised 33 species, occurring from Sri Lanka eastwards through Southeast
Asia to New Guinea. It differs from S. populifolia and
Justicia tigrina in anther form and inflorescence
characteristics. All species of Ptyssiglottis have nonappendaged anthers with the exception of
P. mucronata B.Hansen from Sumatra, which has a
sharply mucronate base to each theca (Hansen
1992). However, this is quite different to the broad,
irregularly fringed appendages seen in S. populifolia
and J. tigrina. In Ptyssiglottis, the inflorescence is
© The Author(s), 2019
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Champluviera; this treatment should be tested in future
via acquisition of material suitable for DNA analysis.
Taxonomic Treatment
Champluviera I. Darbysh., T.F.Daniel & C.A.Kiel genus
no v. Type species: Champluv iera populifolia
(C.B.Clarke) I.Darbysh. & T.F.Daniel
http://www.ipni.org/urn:lsid:ipni.org:names:77197847-1
Fig. 5. Anthers of Champluviera nuda (from Bates 494, BM).
PHOTO: HANNAH ARMER (NATURAL HISTORY MUSEUM, LONDON).
axillary and is a variation of a compound dichasium,
usually branched dichotomously. In some species
(e.g. P. pubisepala (Lindau) B.Hansen, sampled
here), a lax compound dichasium with elongate
internodes is fully developed, but in many species
(e.g. P. psychotriifolia (Stapf) B.Hansen, sampled
here) the dichasium is reduced and contracted,
confined to two (to four) contracted branches, these
often bearing several pairs of imbricate, scale-like
bracts (Hansen 1992). In S. populifolia and J. tigrina,
in contrast, the inflorescence is a terminal spiciform
thyrse, with the dichasial cyme units arranged along
a central, indeterminate axis. Pollen morphology is
variable in Ptyssiglottis, but some pollen types are
somewhat similar to that of S. populifolia, having the
3-colporate 6-pseudocolpate grains with enlarged
ora, although the pseudocolpi are not or only slightly
convergent in the mesocolpium (see Hansen 1992,
Figs. 10 and particularly 11). Ptyssiglottis is not
monophyletic in our results but, with only two
species sampled, this result must be viewed as
preliminary. Further sampling is warranted to investigate this further.
Based on the molecular and morphological evidence presented, our data indicate that a new genus is
warranted to accommodate Schaueria populifolia within
the lineage sister to Core Isoglossinae and we describe
the new genus Champluviera I.Darbysh., T.F.Daniel &
C.A.Kiel below. Further, based on the close morphological similarity between S. populifolia and Justicia
tigrina, we include the latter species within
© The Author(s), 2019
Perennial herbs or subshrubs, erect or decumbent; young
stems somewhat 4- or 6-angular. Leaves opposite-decussate, petiolate, pairs isophyllous, blade ovate or oblongovate to elliptic or oblong-elliptic; cystoliths present,
linear, numerous, conspicuous in dry state at least on
adaxial leaf surface. Inflorescences terminal, spiciform,
comprising a series of opposite subsessile or shortly
pedunculate cymes arranged along an indeterminate
axis, i.e. a spiciform thyrse; bracts (excluding those pairs
at base of inflorescence spike), bracteoles and calyx lobes
subequal in size and shape, linear, linear-spathulate or
linear-lanceolate; flowers subsessile or shortly pedicellate.
Calyx divided almost to base into five lobes, subequal to
somewhat unequal in length. Corolla intricately speckled
and/or striped, rugula absent; aestivation ascendingcochlear, limb bilabiate; upper lip with apex shortly
bilobed, lower lip 3-lobed. Stamens 2, inserted on interior
of corolla tube, anthers exserted, dehiscing toward lower
lip (i.e. flowers nototribic), bithecous, thecae parallel or
slightly oblique, very slightly to more markedly offset, both
thecae with a flattened, broad, irregularly toothed
appendage at the base; staminodes absent. Floral disk
annular, fleshy, 0.5 – 0.7 mm tall. Ovary oblong-ellipsoid,
4-ovulate; style filiform; stigma shortly bilobed. Capsule
(where known) 4-seeded (or fewer by abortion),
retinaculate, stipitate; seeds (only seen in immature state)
tuberculate.
RECOGNITION. Champluviera is recognised by having
the combination of a dense spiciform terminal
thyrse with the bracts, bracteoles and calyx lobes
all slender and closely resembling one another; a
bilabiate corolla lacking a rugula and with intricate
speckling or striping on the lips; and, in particular,
anthers comprising two slightly to more markedly
offset thecae each of which has a broad flattened
basal appendage with an irregular fringe, and at
least the upper theca having a line of trichomes on
the exposed side. It differs from Schaueria primarily
in having appendaged and pubescent anthers and
in having speckled or striped corolla lips (vs
anthers lacking appendages and glabrous, corollas
not speckled or striped). It differs from Justicia
primarily in the corolla lacking a rugula and in the
anther appendages being broad and irregularly
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toothed (vs corolla with a rugula, anther appendages elongate to rounded and smooth except
sometimes for a forked apex). Champluviera
populifolia also has an apparently unique pollen
type in which the two pseudocolpi of each
mesocolpium converge towards the equator and
sometimes meet, forming an hour-glass or figure
eight shape. It is not clear, however, whether the
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39
second species, C. nuda, has the same pollen type
(see Note to that species).
ETYMOLOGY. This genus is named in honour of Dr
Dominique Champluvier (previously of the BR herbarium), an expert on the Acanthaceae of Africa with a
particular interest in the Central African region. Dominique was the first to recognise the close relationship
between the two species in this new genus.
Key to species of Champluviera
Corolla tube 10 – 14 mm long, clearly longer than limb, corolla glabrous externally except for few trichomes at apex of
lower lip; anther thecae only very slightly offset; bracteoles and calyx lobes typically linear-spathulate with a blunt
tip, usually with conspicuous glandular trichomes distally [Nigeria, Bioko, Cameroon] . . . . . . . . C. populifolia
Corolla tube 6 – 8 mm long, only slightly longer than limb, corolla pubescent externally; anther thecae markedly
offset; bracteoles and calyx lobes linear-lanceolate with a sharply acute tip, glandular trichomes minute and
inconspicuous [Gabon]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. nuda
Champluviera populifolia (C.B.Clarke) I.Darbysh. &
T.F.Daniel comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:77197848-1
Schaueria populifolia C.B.Clarke, Fl. Trop. Afr. 5: 242
(1900a); Hutchinson & Dalziel (1931: 263); Heine
(1963: 423); Lebrun & Stork (1997: 503); Vollesen
in Cable & Cheek (1998: 5); Vollesen & Darbyshire
in Cheek et al. (2004: 230). Type: Equatorial
Guinea, Fernando Po [Bioko], fl. 1862, Mann
1426 (K! [K000419169] lectotype, selected here).
Additional syntypes: Equatorial Guinea, Bioko, in
bud Nov. 1841, Vogel s.n. (K! [K000419170]);
Cameroon, Bipinde, fl. 1896, Zenker 1205 (BM!
[BM000949851], K!, M* [M0186812], S* [S095904], WAG* [WAG0248378]).
Justicia populifolia (C.B.Clarke) Champl. ined. sensu
Onana (2011: 31; 2013: 64), nom. nud.
Perennial herb or subshrub, 30 – 200 cm tall, erect or
decumbent, sometimes rooting at lower nodes; young
stems somewhat 4- or 6-angular, furrowed in dry state,
markedly swollen above each node (often shrunken in
dry state), with two opposite-decussate lines of short
retrorse trichomes most numerous below each node,
trichomes multicellular with conspicuous dark cell
walls, stems also with numerous minute red-brown
sessile glands throughout; mature stems terete. Leaves
with a strong liquorice odour (fide Bos 5548); petioles
18 – 66 mm long, furrowed above, antrorsepuberulous; blade dark green above, sometimes paler
or purplish along main veins, pale grey-green to
purplish beneath, ovate, oblong-ovate or ovate-elliptic,
6.2 – 19.7 × 3.4 – 10.5 cm (length : width ratio 1.5 – 2.5
: 1), base cordate, sometimes asymmetrically so,
margin entire or obscurely undulate, apex acuminate,
adaxial surface glabrous or with scattered short
multicellular trichomes that appear deflated in dry
state, abaxial surface antrorse-puberulous along main
veins, margin with few short patent glandular trichomes in proximal half; cystoliths linear, numerous
and conspicuous on adaxial surface, most dense along
veins, both surfaces also with numerous minute redbrown sessile glands; lateral veins 6 – 9 pairs,
brochidodromous, prominent beneath, with ± scalariform tertiary veins. Inflorescence a terminal spiciform
thyrse with few flowers open at any one time, (1.8 –) 3.5 –
11 × 2 – 2.7 cm, comprising a series of opposite (sub)sessile
dichasia, these at first 3-flowered but can become manyflowered with age; rachis greenish-white or purplish,
indumentum as that of stem but trichomes can be more
dense; bracts along rachis greenish-white or purplish
towards base, apical portion darker, linear, linearspathulate or more rarely linear-lanceolate, 6.5 – 14 × 0.3
– 1.3 mm, pairs divergent, distal portion often ascending,
apex usually obtuse or rounded, surface with short
antrorse eglandular trichomes and with few to numerous
spreading glandular trichomes distally, cystoliths numerous and conspicuous in proximal half or two thirds; bract
pair at base of thyrse often somewhat larger, lanceolate,
up to 4 mm wide; bracteoles as bracts but usually linearspathulate, 7.5 – 11.5 × 0.3 – 0.8 mm, apex sometimes
slightly recurved; flowers subsessile or on stout glabrous
pedicels to 2 mm long. Calyx lobes linear or usually linearspathulate, closely resembling the bracteoles, somewhat
unequal in length, 8 – 12.5 × 0.2 – 0.5 mm, apex blunt,
often slightly recurved, indumentum like that of bracts.
Corolla 16 – 21 mm long, dull red, red-brown or purple,
lower lip white to pale yellow with red to purple speckling;
tube 10 – 14 mm long, basal cylindrical portion 7.5 –
10 mm long, c. 1.2 – 1.7 mm in diam. at midpoint,
expanded throat 3 – 4.5 mm long, glabrous externally,
minutely puberulous within with mixed eglandular and
© The Author(s), 2019
39
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glandular trichomes particularly on roof, with a diffuse
broad band of longer eglandular trichomes starting 2.5 –
4.7 mm from base, only sparsely hairy below this; upper lip
triangular, 4.5 – 7.3 × 4 – 5 mm when flattened but
margins narrowly involute, apex shortly bilobed, lobes 0.5 –
1.3 mm long, tips reflexed; lower lip 4.5 – 7 mm long,
deeply 3-lobed, lateral lobes lanceolate, 2.3 – 4.2 ×
1 – 2.2 mm when flattened, median lobe broadly rounded,
1.8 – 3 × 3 – 3.6 mm, with two raised, convergent ridges and
a central furrow and with prominent reticulate venation,
lips glabrous externally except for few short trichomes at
apices of lower lobes. Stamens inserted 6.3 – 8.7 mm from
base of corolla; filaments free for 5 – 7.5 mm, flattened
either side of raised central vein, shortly pubescent at base
and along fused portion, glabrous distally; anthers
bithecous, thecae parallel or slightly oblique, slightly offset
by c. 0.15 mm, narrowly oblong, 1.9 – 2.4 mm long, both
with broad subsessile glands on adhering surfaces, upper
(inner) theca with dense band of short multicellular
eglandular trichomes and shorter capitate glandular
trichomes dorsally along exposed side, lower (outer) theca
glabrous on exposed side, both thecae with broad
flattened, slightly protruding appendage below suture with
irregular fringe of conical teeth to 0.07 mm long. Pollen
perprolate, 3-colporate, 6-pseudocolpate, polar diameter
(P) 57 – 60 μm, equatorial diameter (E) 24 μm, P:E = 2.37 –
2.5, ora considerably wider than colpi and with membrane
echinate (to gemmate), the two pseudocolpi in each
mesocolpium curved toward one another equatorially
(and sometimes toward poles as well) and sometimes
fusing at equator forming an hour-glass or figure eight
shape, interaperatural exine reticulate. Floral disk annular,
fleshy, 0.5 – 0.7 mm tall. Ovary black, oblong-ellipsoid, 1.1 –
1.6 mm long, glabrous; style pale orange-brown in dry state,
10 – 13.5 mm long, with few pale eglandular trichomes at
base; stigma shortly bilobed. Capsule c. 17.5 mm long
including stipe 8 mm long, apex shortly attenuate, surface
glabrous; immature seeds only seen, flattened, tuberculate.
Figs 3, 4 and 6.
DISTRIBUTION. Nigeria (Cross River State), Equatorial
Guinea (Bioko), Cameroon (Southwest, Littoral and
South Regions); Map 1.
SPECIMENS EXAMINED. NIGERIA. Cross River State: Oban
Distr., fl. 1911 – 1912, Talbot s.n. (K!); Oban, fl. 1911,
Talbot 993 (BM, K!); Ogoja Prov., Ikom Distr., Cross
River North Forest Reserve, fl. 7 Dec. 1950, Keay in FHI
28157 (FHI, K!); Ogoja Prov., Ikom Distr., Afi River
Forest Reserve, fl. 13 Dec. 1950, Keay in FHI 28238 (FHI,
K!). EQUATORIAL GUINEA. Bioko (unplaced): Fernando
Po [Bioko], fl. 1862, Mann 1426 (K! lectotype). Bioko
Norte: Fernando Po [Bioko], Clarence Peak [Pico
Basilé], in bud Nov. 1841, Vogel s.n. (K!). CAMEROON.
Southwest Region: Meme Div., Southern Bakundu
Forest Reserve, near Pete, 28 Nov. 1985, Thomas &
Nemba 5022 (MO!); Ndian Div., along path from “Fabe
Road” towards Fabe, fl. 22 Nov. 1986, Manning 916 (K!,
© The Author(s), 2019
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MO!); Korup N.P., WCI study area, 35 km W of Nguti, fl.
18 Jan. 1991, Harris 2704 (K!); Mabeta-Moliwe, Diculu, fl.
16 Dec. 1993, Cable 585 (K!, SCA, YA); Ndian Div., N of
Mundemba at second crossing of Moliba R. on
Ekundu Kundu road, fl. 12 Dec. 1994, Gereau et al.
5666 (K!, MO!); Mt Kupe SW slope, main trail
leading to the summit, fl. 11 July 1995, Ensermu
3533 (K!, YA); Mt Kupe, path to Kupe rock along
Esense R., fl. 2 Nov. 1995, Cheek 7654 (BR*, K!,
MO!, P, SCA, WAG, YA); Mt Kupe, SW slope,
Muanenzum trail, fl. 6 Nov. 1995, Schoenenberger 26
(K!, YA); Mt Kupe, main trail to the top from Kupe
village, fl. 13 Nov. 1995, Schoenenberger 53 (K!, YA);
Mt Kupe, path to top of Kupe rock, fl. 14
Nov. 1995, Cheek 7771 (BR*, K!, MO!, NY, P*,
SCA, US, WAG, YA); Meme Div., Mungo River
Forest Reserve, S of Kumba-Tombel road, c. 1 km
from Mungo bridge, fl. & fr. 26 Nov. 1999, Cheek
10153 (K!, SCA, YA); Ngomboku, on path to
Mbombe hill, fl. 9 Dec. 1999, Mackinder 292 (BR*,
K!, SCA, YA); Banyang-mbo Sanctuary, Research
Station path to sanctuary via Nlowoa and Mbu river
crossings, fl. 25 Nov. 2000, Cheek 10605 (K!, MO,
WAG, YA); Mone Forest Reserve, Mokoyong to Tafu
path, 2.5 hrs E, 30 Nov. 2000, Cheek 13984 (K!).
Littoral Region: Forêt de Bakaka, 3 km E of Eboné
(km 11 Nkongsamba-Loum road), fl. 17 Dec. 1971,
Leeuwenberg 8913 (K!, WAG); près Yingui II, à 5 km
à l’ENE de Yingui, 35 km E de Yabassi, fl. & fr. 8
Jan. 1972, Letouzey 10919 (K!, P*); Ebo Proposed N.
P., Bekob, fl. 15 Feb. 2006, Tchiengue 2486, (K! YA);
Ebo Proposed N. P., Ebo Forest Research Station,
W transect, fl. 9 Oct. 2006, Osborne 97 (K!, WAG,
YA); Ebo Proposed N. P., Ebo Bekob forest, fl. 26
Nov. 2013, Tchiengue 3613 (K!, MO, P*, WAG, YA);
Ebo Proposed N. P., Bekob camp, fl. 2 Dec. 2013,
Ngansop 187 (K!, WAG, YA). South Region: Bipinde,
fl. 1896, Zenker 1205 (BM!, K!, M*, P*, S*, WAG*);
au S de Zingui, 40 km ESE de Kribi, fl. 20
Jan. 1962, Letouzey 4072 (K!, P*); Mviglimangalé,
16 km E Nayabesan, 1 km NW village, fl. 15
Feb. 1965, Raynal 13411 (P*); près de R. Kienke,
au NNW de Nkolbewa, km 35 route Kribi-Ebolowa,
fl. 12 March 1968, Letouzey 9001 (K!, YA); about
11 km from Kribi, between airfield and Kienke, fl.
27 Oct. 1969, Bos 5548 (BR*, WAG); c. 5 km SSE of
Bipindi, fl. & fr. 14 Jan. 1987, Manning 1336 (K!,
MO!, YA*); Efoulan, au sud d’Akom II (mi-chemin
entre Ebolowa et Kribi), fl. 11 May 2009, Lachenaud
et al. 737 (BR*).
HABITAT & ECOLOGY. This species is recorded from
1lowland rainforest in deep shade, sometimes being
recorded from swamp and riverine forests. It can also
persist in secondary and disturbed forest patches; 70 –
900 m elevation.
PHENOLOGY. Champluviera populifolia has been recorded in flower most frequently in the months of
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39
Fig. 6. Champluviera populifolia. A habit: rootstock and distal portion of flowering stem; B dichasial inflorescence unit showing
calyces and bracteoles (some bracteoles cut away at base); C apex of calyx lobe showing glandular trichomes; D dissected corolla
with androecium; E anther; F detail of anther base showing thecal appendages; G dehisced capsule. A from Ngansop 187 with
rootstock from Osborne 97, B from Cheek 10153; C – E from Ngansop 187; F from Gereau et al. 5666; G from Manning 1336 and
Cheek 10153. DRAWN BY HAZEL WILKS.
© The Author(s), 2019
39
Page 12 of 25
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http://www.ipni.org/urn:lsid:ipni.org:names:60478962-2
Chlamydocardia nuda C.B.Clarke, Fl. Trop. Afr. 5: 235
(1900b); Darbyshire & Govaerts (2017: 4). Type:
Gabon, Mabam, Como R., 60 miles from Gaboon, fl. 13
July 1896, Bates 494 (K! [K000419168] holotype; BM!
[BM000931152], L* [L0003054], P! [P00540805]
isotypes).
Justicia tigrina Heine, Fl. Gabon 13: 219, pl. XXXVIII 10
– 13 (1966); Lebrun & Stork (1997: 489); Sosef
(2006: 38). Type as for Chlamydocardia nuda.
Map 1. Distribution of Champluviera: C. populifolia (black
circles), C. nuda (red stars).
(October –) November – December (– January) but
with occasional records from February, March, May
and July. The peak flowering time coincides with
the onset of the dry season within this species’
range.
CONSERVATION STATUS. This species was recorded as
Near Threatened (NT) by Vollesen & Darbyshire (in
Cheek et al. 2004). It appears to be fairly frequent in
the lowland to mid-altitude forests of the Cameroon
Highlands and southwestern Cameroon but its
favoured habitat is threatened through large parts of
its range due to timber extraction and forest clearance
for subsistence and commercial agriculture. It has an
EOO of 53,797 km2 which is considerably greater than
the 20,000 km2 threshold for Vulnerable under IUCN
criterion B1 and has up to 20 locations historically but
some of these are likely to have been lost. The
assessment of Near Threatened (NT) under criterion
B is therefore upheld here.
NOTES. Mann 1462 (K) is chosen as the lectotype, as it
is the most informative of the three syntypes and, from
his annotations and illustrations on this specimen, it is
clear that Clarke (1900a) used it as the basis for his
description in the protologue.
Champluviera nuda (C.B.Clarke) I.Darbysh. &
T.F.Daniel comb. nov.
© The Author(s), 2019
Perennial herb or subshrub, 70 – 180 cm tall, basal stems
trailing and rooting at nodes, decumbent; young stems
somewhat 6-angular, ± evenly pubescent with sparse
antrorse to appressed eglandular trichomes and with
opposite lines of denser trichomes, older internodes
glabrescent. Leaves petiolate, petioles to 70 mm long,
furrowed above, antrorse-puberulous; blade ±
concolorous, green or dark glaucous-green, elliptic,
oblong-elliptic or ovate-elliptic, 8.5 – 15.5 × 3.6 – 8 cm
(length : width ratio (1.75 –) 2.1 – 2.8 : 1), base
rounded, shallowly cordate or obtuse, can be somewhat asymmetric, margin entire or irregularly undulate, apex acuminate, surfaces sparsely pubescent on
main veins, with antrorse eglandular trichomes; cystoliths linear, numerous and conspicuous on both
surfaces when dry, surfaces also with numerous
minute red-brown sessile glands; lateral veins 6 – 8
pairs, brochidodromous, pale and prominent beneath.
Inflorescence a slender erect terminal spike to 15 –
24.5 cm long including peduncle to 0.5 – 1.2 (– 3) cm
long, comprising a series of opposite 1 – 3 (potentially
more)-flowered cymes, those at base of spikes borne
on peduncles to 2.5 mm long, those in distal portion
of spike sessile or peduncle to 0.5 mm; rachis ± evenly
pubescent with antrorse eglandular trichomes like
those of young stems; bracts opposite, green, linearlanceolate, 6 – 8 mm × c. 1 mm, pubescent abaxially,
like rachis; bracteoles similar to bracts in shape and
size but can have a more puberulent indumentum and
can be purplish (drying brown); pedicels to 1.5 (– 2.5)
mm long, puberulent. Calyx purple or grey-purple in
live state at least towards base and apex, drying brown,
divided almost to base into five subequal to ± unequal
linear-lanceolate lobes resembling the bracteoles, 7.5 –
11 mm long, puberulent externally with mixed erect
glandular and eglandular trichomes to 0.05 mm long.
Corolla ± 11 – 14 mm long, yellow with dark purple
stripes on upper lip and dark purple speckling on
lower lip, pubescent externally with erect eglandular
trichomes; tube ± 6 – 8 mm long, cylindrical; limb
bilabiate with lips ± 5.5 – 6 mm; upper lip hooded,
apex shortly bilobed; lower lip pendant, shortly 3lobed. Stamens 2; filaments ± 4 mm long (fide Heine
1966), glabrous; anthers with thecae unequally
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inserted, overlapping by 0.8 – 0.9 mm, thecae narrowly
elliptic, 1.3 – 1.6 mm long, with multicellular
eglandular trichomes along the exposed side more
numerous on upper theca, and both with a flattened,
broad appendage at base below suture with irregularly
toothed fringe. Pollen not seen (see note). Floral disk
annular, ± 0.5 mm tall. Ovary ± 2 mm long, glabrous;
style ± 11 mm long (fide Heine 1966), with eglandular
trichomes towards base; stigma shortly bilobed. Capsule
not seen. Fig. 5.
DISTRIBUTION. Gabon (Estuaire, Moyen-Ogooué and
Woleu-Ntem Provinces); Map 1.
SPECIMENS EXAMINED. GABON. Estuaire Province:
Mabam, Como R., 60 miles from Gaboon, fl. 13
July 1896, Bates 494 (K! holotype; BM!, L*, P! isotypes);
Abanga Chantier C.E.T.A., fl. 3 June 1963, N. Hallé
2176 (P! [P03616770, P03616771]); idem, fl. 8
June 1963, N. Hallé 2362 (P! [P03616769]). MoyenOgooué Province: 15 km NE of Ndjolé, Exploitation
road, 31 Jan. 1991, de Wilde & Sosef 10329 (WAG
[WAG0130200], n.v.). Woleu-Ntem Province: Crystal
Mts, Tchimbélé, fl. 23 Jan. 2001, Mayombo-Nzengue 271
(LBV* [LBV0019437]).
HABITAT & ECOLOGY. This species is recorded from
primary wet forest, shaded river margins and in
alluvial silts by streams; the elevation range is estimated at 10 – 630 m.
PHENOLOGY. Phenology information on this species is
very limited; three of the four flowering specimens
seen by the current authors were collected in June or
July, during the short dry season; the fourth was
collected in January during the wet season.
CONSERVATION STATUS. This species is clearly scarce,
being known with certainty from only four herbarium
collections, despite Gabon being one of the best explored
countries botanically in Africa according to Sosef et al.
(2017). It appears to have a highly restricted range with an
EOO of 2890 km2 based on current knowledge. Of the
four known localities, Ndjolé is an area of gold and
manganese mining (https://mining-atlas.com/project/
Ndjole-Mangenese-Gold-Mine-Project.php) and the
footprint of this mining activity along the Ogooué River
is clearly visible on Google Earth™ imagery (accessed 15th
August 2018). This is likely to have impacted upon habitat
suitable for this species. There is still extensive intact forest
habitat elsewhere within its range, and this species is likely
to occur at other localities within this small range.
However, based on current knowledge, with an EOO of
less than 5000 km2, fewer than five locations and an
inferred decline in extent and quality of habitat due to
mining activities, this species is provisionally assessed as
Endangered - EN B1ab(iii).
NOTES. This species is well illustrated in Heine (1966:
pl. XXXVIII 10 – 13). The anthers are illustrated in
Fig. 5 here.
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39
Sosef (2006) also lists Mayombo-Nzengue 390 (LBV)
under this species but this collection number is listed
as a Zingiberaceae in the LBV database and only
Mayombo-Nzengue 271 is housed under Justicia tigrina at
LBV (P. Bissiengou, pers. comm.). Further, Sosef et al.
(l.c.) list Arends 570 (WAG) as Justicia cf. tigrina but this
specimen was redetermined by D. Champluvier as
J. tristis T.Anderson, and we concur with this
identification.
We have not seen the pollen of this species. Clarke
(1900b) recorded the pollen as “globose with 12
equidistant ribs not reaching the pores” (p. 235) and
illustrated it as such on a sheet of paper attached to
the holotype specimen. He thought that there were 6
equidistant pores on the grains which supported the
placement of this species in Chlamydocardia (see
Darbyshire & Govaerts 2017), but this does not seem
to fit with the presence of 12 ‘ribs.’ If the grains are
colporate and pseudocolpate, then 12 ribs would
suggest four germinal apertures — if Clarke (1900b)
was correct then these grains are quite different to
those of C. populifolia described above.
Heine (1966) notes the presence of “sterile floral
buds” in this species — these are presumably cleistogamous flowers, which are not uncommon in some
genera of Acanthaceae, for example in Isoglossa
(Darbyshire et al. 2012).
Asystasia lindauiana Hutchinson & Dalziel (1931:
257) could potentially be confused with Champluviera
on morphological grounds and has a sympatric
distribution, hence is worthy of mention here. The
two are superficially very similar both in vegetative
characters and inflorescence form, sharing a terminal
many-flowered spiciform thyrse. Indeed, Clarke
(1900b) remarked on this close superficial similarity
when describing Chlamydocardia nuda. However,
A. lindauiana is easily separated from Champluviera in
having four stamens (a character shared by all species
of Asystasia), and in the anthers having thecae
markedly offset and lacking basal appendages. Molecular evidence confirms that these genera are not
closely related, with Asystasia falling within the
“Pseuderanthemum lineage” of Justicieae (McDade
et al. 2000).
2. Kenyacanthus — a new monotypic genus
endemic to Kenya
Notable morphological differences exist between
Rhinacanthus ndorensis and species of Rhinacanthus s.s.
In particular, R. ndorensis is unique in Rhinacanthus in
the positioning of stamens, these being held against
the lower lip of the corolla with thecae dehiscing
towards the upper lip (i.e. flowers are stenotribic; Kiel
et al. 2017). In Rhinacanthus s.s., the stamens are held
between the two lips and dehisce toward the lower lip
© The Author(s), 2019
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Page 14 of 25
(i.e. flowers are nototribic). The two lips of the corolla
in R. ndorensis are subequal in size, whilst in other
Rhinacanthus the upper lip is markedly smaller and
usually much narrower than the lower lip (Darbyshire
& Harris 2006; Darbyshire et al. 2018). In addition, as
noted in the Introduction, R. ndorensis can be separated by its procumbent or trailing habit, fasciculate
inflorescences, hyaline margins to the bracts, bracteoles and calyx lobes and essentially glabrous capsules.
In other species of Rhinacanthus, plants are erect,
decumbent or straggling, the inflorescences are usually lax panicles or spikes (although fascicled in
R. rotundifolius C.B.Clarke), the bracts, bracteoles and
calyces lack hyaline margins and the capsules are
variously puberulous to pubescent with eglandular
and/or glandular trichomes (Darbyshire & Harris
2006). These morphological differences support the
findings of the molecular analysis (Fig. 2) that
R. ndorensis is not closely related to Rhinacanthus s.s.
Rhinacanthus ndorensis has 3-colporate, 6pseudocolpate pollen (Fig. 7), a pollen type that is
hypothesised to be synapomorphic for Justicieae as a
whole (Kiel et al. 2017) and that thus does not
contribute to assessing the phylogenetic position of
R. ndorensis within Justicieae.
The placement of Rhinacanthus ndorensis sister to
core Diclipterinae, as confirmed in our analysis here
(Fig. 2), is not surprising on morphological grounds,
as this species could easily be confused with a member
of the genus Dicliptera. Indeed, the general corolla
morphology, with the lips being widely divergent and
subequal in size, is very similar to that of African
Dicliptera. Inflorescences of Dicliptera are typically of
monochasial units (cymules) with conspicuous paired
clasping bracts that are often umbellately arranged
(Balkwill et al. 1996; Darbyshire 2009). However, some
species have solitary or fascicled axillary inflorescences
Fig. 7. Pollen of Kenyacanthus ndorensis. A equatorial view
showing the compound aperture and pseudocolpi; B equatorial view showing the mesocolpium (from Luke 17084).
© The Author(s), 2019
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with reduced cymule bracts (e.g. D. minutifolia
Ensermu from Ethiopia and Kenya; Ensermu 2003;
Darbyshire et al. 2010) which are similar to the
inflorescences of R. ndorensis. Furthermore, hyaline
margins on the bracts, bracteoles and calyces are fairly
widespread in Dicliptera (see, for example, Balkwill
et al. 1996; Darbyshire et al. 2010). However,
R. ndorensis differs significantly from Dicliptera in floral
morphology. In R. ndorensis the corolla tube is straight,
not twisted, and, as noted above, the flowers are
stenotribic with the stamens held against the lower, 3lobed lip of the corolla (Fig. 8). All OW species and
many NW species of Dicliptera (and all species in the
related genus Hypoestes) have a resupinate corolla, in
which the tube is twisted through ± 180°, thus the 3lobed “lower” lip is held in the upper position,
although Daniel & Figueiredo (2009) note that in
“D. verticillata (Forssk.) C.Chr.” (possibly = D. hyalina
Nees) from São Tomé the tube can be twisted by only
90° such that the limb is held on its side. In these
resupinate-flowered species, the flowers are effectively
stenotribic. However, the positionally lower 2-lobed lip
that the stamens are held against is actually the upper
lip morphologically. In his synopsis of the species of
Dicliptera in the Nueva Galicia region of western
Mexico, Daniel (2009) noted three distinct floral
forms: tube twisted through 180°, tube twisted through
360° and tube untwisted. The latter two cases both
result in non-resupinate corollas in which the 3-lobed
lip is in the “normal” lower position. In these nonresupinate forms, the flowers are nototribic with the
stamens held close to the upper corolla lip, quite
different to those of R. ndorensis. The species of
Dicliptera with non-resupinate flowers in the NW have
large, brightly coloured and unspeckled red, orangeyellow or purple corollas that appear to be adapted for
bird pollination (Daniel 2009; Kiel et al. 2017). These
are quite unlike the small, pink flowers of R. ndorensis.
The stamens of Dicliptera are usually exserted well
beyond the corolla mouth and not appressed to the
corolla lip as in R. ndorensis. Further, R. ndorensis differs
from all African members of Dicliptera in having only
slightly offset as opposed to fully superposed anther
thecae. This character is, however, more variable in
some species of NW Dicliptera (Kiel et al. 2017).
Rhinacanthus ndorensis is further separated from
most taxa in core Diclipterinae by the presence of a
rugula (stylar furrow), which in R. ndorensis comprises
two raised ciliate bands of tissue running dorsally
along the corolla tube and base of the upper lip (Fig.
9F), similar to the rugula seen in Rhinacanthus s.s.
Whilst rugulas are widespread in the justicioid lineage,
they are mainly lacking or indistinct in the core
Diclipterinae lineage including Dicliptera (Balkwill &
Welman 2000; Darbyshire & Vollesen 2007). The
rugula is hypothesised to hold the style in position
during anthesis, and it has been suggested that the loss
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of the rugula in core Diclipterinae occurred as a
correlate of corolla resupination (Kiel et al. 2017),
where the twisting of the corolla tube may hold the
style in position. That said, recent observations on
putatively bird-pollinated species of Hypoestes from
Madagascar with elongate, expanded corolla tubes
such as H. isalaensis Benoist have revealed the
presence of a shallow rugula of two raised, ciliate lines
not dissimilar to the rugula in R. ndorensis (I.
Darbyshire, pers. obs.). It may be that this character
is more widespread in taxa of Diclipterinae with
enlarged corolla tubes.
Monechma Group II is sister to (Rhinanthus ndorensis
(Hypoestes + Dicliptera s.l.)) (Fig. 2); the former is one of
two widely separated clades comprising the nonmonophyletic genus Monechma Hochst. (= Justicia L.
sect. Monechma (Hochst.) T.Anderson). However,
R. ndorensis is unlikely to be confused morphologically
with species of Monechma Group II as these plants have
broader cylindrical corolla tubes, usually subequal in
length to the limb, a markedly hooded upper corolla
lip, stamens with a conspicuous appendage on the
lower anther theca, capsules with two seeds and the
seeds having a smooth testa. Rhinacanthus ndorensis, on
the other hand, has a narrowly cylindrical tube clearly
longer than the limb, the upper corolla lip is not
hooded, the stamens lack appendages, the capsule is
4-seeded and the seeds are sculptured (Fig. 9J).
In summary, phylogenetically, Rhinacanthus
ndorensis forms an isolated clade composed of, so far
as known, a single species that is placed between
(Hypoestes + Dicliptera s.l.) and the Monechma Group II
(Fig. 2). Regarding morphological traits, R. ndorensis
appears to be intermediate between plants belonging
to the core Diclipterinae and those of the other clades
that are part of the expanded Diclipterinae as
recognised by Kiel et al. (2017), as described above
(Figs 8 and 9). Thus, morphology and phylogenetic
relationships based on DNA data are congruent.
Kiel et al. (2017) reviewed the potential taxonomic
implications of their findings on the complex relationships within the “justicioid” lineage, and five
39
potential solutions were offered. They noted that in
order to maintain a broadly circumscribed Justicia as is
currently widely accepted (see, for example, Graham
1988; Vollesen in Darbyshire et al. 2010), the entire
“justicioid” lineage would potentially have to be
treated as a single genus. This is highly undesirable
as it would mean subsuming several genera that are
easily separated by morphological differences, including those within Diclipterinae. The most credible
solution offered therefore appears to be to divide
Justicia s.l. into a number of segregate genera and to
uphold the morphologically distinct genera such as
Dicliptera, Hypoestes and Rhinacanthus s.s. Whilst the
taxon sampling in Kiel et al. (2017) was insufficiently
dense to support full revision of generic delimitations
among justicioids, the strong morphological and
molecular evidence presented here supports the
recognition of a new genus to accommodate
R. ndorensis, allied to but clearly separated from
Dicliptera, Monechma and Rhinacanthus. In consequence, the new genus Kenyacanthus I.Darbysh &
C.A.Kiel is described below.
Taxonomic Treatment
Kenyacanthus I.Darbysh. & C.A.Kiel genus nov. Type
species: Kenyacanthus ndorensis (Schweinf.) I.Darbysh.
& C.A.Kiel.
http://www.ipni.org/urn:lsid:ipni.org:names:77197855-1
Perennial herbs, with trailing or procumbent branches
radiating from a small woody base; stems markedly
angular with 2 opposite pairs of ridges. Leaves opposite-decussate, sessile or very shortly petiolate, pairs
isophyllous, blade obovate or obovate-elliptic; cystoliths present, shortly linear, numerous but often
inconspicuous. Inflorescences axillary, sessile or shortly
pedunculate fasciculate cymes of (1 –) 2 – 3 flowers,
bracts and bracteoles paired, linear-lanceolate, green
with hyaline margins. Calyx shortly tubular, deeply
Fig. 8. Kenyacanthus ndorensis. A flower, lateral view; B flower, face view; C bracts, bracteoles and calyx with corolla base
(photographed from plant cultivated by W. R. Q. Luke).
© The Author(s), 2019
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divided into five lobes, these subequal, linearlanceolate with hyaline margins, shorter than the
bracteoles. Corolla with rugula present, comprising
two slender ciliate ridges extending from near base
of corolla tube to proximal portion of upper lip; tube
slender, longer than limb, gradually widened towards
the mouth; aestivation ascending-cochlear, limb bilabiate; upper lip ovate or elliptic, apex shortly bilobed
or emarginate, lower lip conspicuously 3-lobed, palate
with faint raised venation. Stamens 2, inserted on
ventral side of corolla tube just below the corolla
mouth; filaments short; anthers shortly exserted, held
± adpressed to the lower lip, dehiscing toward upper
lip (i.e. flowers stenotribic), bithecous, thecae ±
parallel, slightly offset, lacking basal appendages;
staminodes absent. Floral disk cupular, ± 0.5 mm tall,
with unevenly undulate margin. Ovary oblong-ellipsoid, 4-ovulate; style filiform; stigma minutely bilobed,
one lobe slightly larger than the other. Capsule 4seeded (or fewer by abortion), retinaculate, stipitate
and with attenuate apex; seeds tuberculate.
RECOGNITION. Kenyacanthus is recognised by having the
combination of a trailing or procumbent habit; fasciculate inflorescences; bracts, bracteoles and calyces with
hyaline margins; a bilabiate corolla with a slender
cylindrical tube longer than the limb and with a shallow
rugula; and two stamens held against the lower corolla
lip and anthers with slightly offset thecae, the flowers
being stenotribic. It differs from Rhinacanthus primarily
in the trailing or procumbent habit; hyaline margins to
the bracts, bracteoles and calyx lobes; two corolla lips of
subequal size; stenotribic flowers and essentially glabrous
capsules (vs plants erect, decumbent or straggling; bracts
etc. lacking hyaline margins; upper corolla lip markedly
smaller and narrower than the lower lip; flowers
nototribic and capsules puberulous to pubescent in
Rhinacanthus). It differs from Dicliptera primarily in the
combination of a straight, untwisted corolla tube and
stenotribic flowers (vs corolla tube twisted through ±
180° such that the corolla is resupinate, or if corolla tube
untwisted then flowers nototribic in Dicliptera); it also
differs from all African Dicliptera in the anther thecae
being only partially offset (vs fully superposed).
ETYMOLOGY. Kenyacanthus is named in honour of the
Republic of Kenya, to which this genus is apparently
endemic based on current evidence.
Kenyacanthus ndorensis (Schweinf.) I.Darbysh. &
C.A.Kiel comb. nov.
http://www.ipni.org/urn:lsid:ipni.org:names:60478965-2
Rhinacanthus ndorensis Schweinfurth in Höhnel, Zum
Rudolph-See und Stephanie-See: 858 (1892); Engler
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(1892: 394), nom. nud.; Schweinfurth (1894: 357);
Clarke (1900a, 1900b: 225); Mildbraed (1926: 504);
Agnew (2013: 327, pl. 129C); Lebrun & Stork (1997:
500) as “sont mal connus”; Darbyshire et al. (2010:
602). Type: Kenya, Ndoro, foot of Mt Kenya, fr.
1887 – 1888, Höhnel s.n. [Höhnel 76 per Engler
1892] (B† holotype; BM! [BM001011739] lectotype,
selected here — see note); Nyeri County, NanyukiNaromoru, near pt. 866 (Ndoro?), fl. & fr. 7
Feb. 2016, Luke 17084 (EA! epitype, selected here;
K! isoepitype).
Procumbent or trailing perennial herb, branching widely
from a small woody base and rootstock, forming small
mats in the ground layer; young stems with two
prominent pairs of opposite ridges and two broader
opposite ridges, glabrous except for tufts of multicellular trichomes along the nodal lines. Leaves sessile or
on a poorly defined petiole to 3 mm long; blade
fleshy, obovate (-elliptic), 0.8 – 1 (– 1.4) × 0.45 – 0.6 (–
0.75) cm, base cuneate, margin entire, apex obtuse or
rounded, surfaces glabrous except for minute trichomes along the margin; lateral veins 3 (– 4) pairs.
Inflorescences axillary, fasciculate, (1 –) 2 – 3-flowered;
peduncle 0 – 3 mm long, glabrous; bracts linearlanceolate, 5.5 – 9 mm long, green with narrow
hyaline margin at least in the proximal half, ciliate
with trichomes of variable length, midrib prominent
abaxially, apex minutely apiculate; bracteoles resembling bracts but 5 – 8.5 mm long, more gradually
narrowed towards apex, margin more conspicuously
hyaline; flowers sessile. Calyx with short basal tube 0.7
– 0.8 mm long; lobes linear-lanceolate, 3.7 – 5.5 mm
long, margins narrowly hyaline, ciliate, surfaces with
sparse short ascending trichomes. Corolla (13 –) 15 –
18.5 mm long, bright pink to mauve, rather densely
retrorse-pubescent externally; tube (8 –) 10 – 11 mm
long, 1 – 1.5 mm in diam. centrally, widening somewhat
towards mouth, with scattered minute glandular trichomes on internal surface, rugula of two raised lines of
trichomes on dorsal side of tube and barely extending
onto upper lip; lips widely divergent, upper lip suberect,
ovate or elliptic, 5 – 7 × 3 – 3.7 mm, glabrous internally,
apex shortly bilobed or emarginate; lower lip pendant, 6 –
7.5 mm long, glabrous internally, lobes oblong, 2.5 – 3 ×
1.4 – 1.8 mm, apices obtuse or rounded with irregular
margins, median lobe slightly longer and narrower than
lateral pair. Stamens with filaments 1.2 – 2.3 mm long,
glabrous; anther thecae offset by 0.25 – 0.3 mm, each
theca 1 – 1.5 mm long, basally muticous. Pollen prolate, 3colporate, 6-pseudocolpate, polar diameter (P) 35 – 37
μm, equatorial diameter (E) 20 – 22.5 μm, P:E = 1.57 – 1.8,
ora ± the same width as the colpi, the pseudocolpi parallel
to the colpi, interaperatural exine reticulate. Ovary ±
1.5 mm long, glabrous; style ± 8.5 mm long, ± sparsely
appressed-pubescent or glabrous. Capsule 8 – 10 mm long,
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39
Fig. 9. Kenyacanthus ndorensis. A habit; B leaf, adaxial surface with cystoliths; C inflorescence, with one bracteole cut away at base
to show the calyx; D flower, lateral view; E flower, face view showing position of stamens; F upper lip and dorsal side of corolla
tube, showing the rugula; G stamen; H capsule; J seed, with detail of one tubercle. Drawn from Battiscombe 721 except J from
Luke 17084 and D & E from photos. by W. R. Q. Luke. DRAWN BY HAZEL WILKS.
© The Author(s), 2019
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glabrous or with very few inconspicuous trichomes
towards apex; seeds blackish at maturity, 1.3 – 1.8 mm in
diameter, tuberculate, the tubercles short, rounded with
minute glochidia. Figs. 8, 9 and 10.
DISTRIBUTION. Kenya (Laikipia and Nyeri Counties;
Flora of Tropical East Africa floristic regions K3 and K4);
Map 2.
SPECIMENS EXAMINED. KENYA. Unplaced: West Kenya
plains, fl. & fr. 29 July 1913, Battiscombe 721 (K! 2 sheets,
EA photo!). Laikipia County: Rumuruti, NE slope
Aberdare Mts, fl., without date [received at K: 22
Sept. 1915], Dowson 561 (EA!, K!); Nanyuki, fl.
May 1932, Napier 2093 (EA!); Solio Ranch, 20 miles SW
of Nanyuki, fl. 4 April 1973, Elliott 33 (EA!); Laikipia
Distr., Rumuruti, fl. & fr. 5 Nov. 1978, Hepper & Jaeger
6626 (EA!, K!); Mpala, 1800 m, fl. & fr. 5 Nov. 1992,
Young 3169 (EA!); Laikipia Distr., Nanyuki, Batuk,
Laikipia Airbase, 5 km NW Nanyuki town, fl. 18
March 2010, Malombe & Kimeu BATUK 1594 (EA!).
Nyeri County: Ndoro, foot of Mt Kenya, fr. 1887 – 1888,
Höhnel s.n. (BM! lectotype); Cole’s Mill, fl. 17 Jan. 1922,
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Fries 1001 (BM!); Nyeri, Amboni R., fl. & fr. 18 Jan. 1933,
Napier 2579 (EA!, K!); Nanyuki, fl. Dec. 1933, Minns s.n.
[in Coryndon Museum 5881] (EA!); 3 miles SE Nanyuki
Station, fl. 19 June 1943, Moreau & Moreau 34 (EA!);
North Nyeri Distr., Nyeri to Nanyuki, c. 40 km S Nanyuki,
fl. 3 April 1975, Hepper & Field 4838 (K!); NanyukiNaromoru, near pt. 866 (Ndoro?), fl. & fr. 7 Feb. 2016,
Luke 17084 (EA! epitype, K! isoepitype).
HABITAT & ECOLOGY. This species occurs in upland
grassland and open woodland, including Acacia
drepanolobium Harms ex Y.Sjöstedt woodland, where it
can form small mats amongst the grasses in the
ground layer of the vegetation (Fig. 10). These mats
are conspicuous during the dry season, appearing
green amongst the dried brown grasses (Odadi et al.
2013). It can also sometimes be found along road
verges. It often grows on seasonally damp black clay
soils; 1700 – 2150 m elevation.
PHENOLOGY. From the limited information available,
mainly derived from specimen data, this species
appears to have a prolonged flowering season, recorded from early November through to late July. This
Fig. 10. Habit of Kenyacanthus ndorensis in the field, collected as Luke 17084.
© The Author(s), 2019
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PHOTO: W. R. Q. LUKE.
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Map 2. Distribution of Kenyacanthus ndorensis (black circles).
spans the period of the two rains — the first rains
occur primarily in October and November whilst the
peak rains fall in April to May. The intervening period
is dry, with January being the driest month.
CONSERVATION STATUS. Darbyshire et al. (2010) considered this species to be Near Threatened (NT) based on
its very restricted range (the EOO is approx. 3050 km2),
with few collections known, and with potential loss of
habitat due to expansion of agriculture. It can be locally
frequent to abundant in suitable habitat (W. R. Q. Luke,
pers. obs.) and is regularly consumed by cattle as an
important fodder during dry periods (Odadi et al. 2013).
However, in some parts of its range it is threatened by
widespread habitat loss through conversion to commercial pastoral agriculture and the global population is
considered to be declining. This species has, therefore,
recently been provisionally assessed as Endangered (EN
B1ab(ii,iii,iv)+2ab(ii,iii,iv)) by the East African Plant Red
List Authority (W. R. Q. Luke et al., unpubl. data). This
assessment is awaiting publication on the IUCN Red List.
NOTES. From Fig. 8 and the discussion above, it is
apparent that during at least a significant portion of
anthesis (if not throughout) the stamens and anthers are
held adjacent to the lower lip whereas the style and stigma
are adjacent to the upper lip. This poses the question as to
how pollen is transferred by pollinators from the anthers
to the stigma, when these two organs are held on opposite
sides of the corolla mouth. Field observations are
required to document the position of the anthers and
stigma throughout anthesis and also to document
pollinator interactions in order to resolve this issue.
The author for Rhinacanthus ndorensis has sometimes been given as Schweinf. ex Engl. (e.g. see
Klopper et al. 2006: 18). This is based on Engler
Page 19 of 25
39
(1892: 394) who recorded “Rhinacanthus ndorensis
Schweinf. nov. spec. msc.” then cited the proposed
type specimen, which he listed as v. Höhnel 76.
However, the only description provided states
“Gebüschpfl.” (i.e. bushy plant), a phrase applied to
multiple taxa within Engler’s work and not considered
to be sufficient to constitute a description as per
Article 38.1 of the Botanical Code, hence Engler
(1892) did not validly publish the name R. ndorensis.
A short but sufficient description was provided by
Schweinfurth (1892: 858) together with the type
citation, which validated the name.
The extant material of the Höhnel type specimen at
BM is only a very small scrap with a few leaves, an
inflorescence and an immature fruit; the holotype
specimen at B is believed to have been destroyed
during World War II. An epitype is therefore selected
to supplement the lectotype, and Luke 17084 is chosen
as it was collected from a site near Naromoru which a
local informant said had previously been known as
'Mukiri wa Ndoro' (translated as “valley of mud” in
Kikuyu); this is quite likely to be the same as the
Ndoro of the original type locality.
The general habit, of trailing stems with axillary
(sub)sessile clusters of flowers, is superficially most
reminiscent of trailing members of the genus
Dyschoriste Nees which is distantly related, in tribe
Ruellieae of Acanthaceae (Tripp et al. 2013). Indeed,
both Mildbraed (1926) and Agnew (2013) remark
upon this potential confusion. However, Dyschoriste is
easily separated from Kenyacanthus in, amongst other
differences, having a markedly tubular calyx, an
androecium comprising 4 fertile stamens or 2 fertile
stamens + 2 staminodes, left-contorted corolla aestivation and seeds with hygroscopic trichomes.
Acknowledgements
We are highly indebted to Dominique Champluvier
(formerly of BR) for providing useful information on
the species of Champluviera treated here. We thank
Hazel Wilks for the line drawings of Champluviera
populifolia and Kenyacanthus ndorensis, and Martin
Cheek and Jo Osborne (both K) for allowing us to
use their photographs of C. populifolia. Heather
Lindon is thanked for her assistance with the nomenclatural history of Rhinacanthus ndorensis. Aurélie Grall
(R.B.G. Kew) assisted with the SEM analysis of pollen
grains of R. ndorensis and Hannah Armer (Manager of
the Light Microscopy Imaging Suite, Natural History
Museum, London) very kindly photographed the
stamens of Champluviera nuda presented in Fig. 5. We
thank the curators of the herbaria visited for providing
access to the collections of the study taxa, notably
Itambo Malombe (EA) and Ranee Prakesh (BM).
Pulchérie Bissiengou kindly checked for specimens
© The Author(s), 2019
39
Page 20 of 25
of J. tigrina at the LBV herbarium on our behalf and
sent scanned images of Mayombo-Nzengue 271 which
were very helpful. This work was supported by grants
from the U.S. National Science Foundation to CAK
(DEB1754845); TFD (DEB0743273, DEB 0108589,
BSR 8304790) and LAM (DEB 9707693, DEB
0108589, DEB 0743178, DEB 1754845).
Appendix 1
Taxa, sources of plant materials from which DNA was
extracted for sequencing and GenBank accession
numbers (trnS-G, trnT-L, rps16, trnL-F, nrITS; – =
sequence not obtained); when a given taxon is not
depicted by name as a terminal in Figs. 1 or 2, the last
entry indicates the clade where that taxon is placed in
our results (clade names are as given in Figs 1 and 2).
Taxa are listed in alphabetical order by genus and
species. When plants in cultivation were used, we
provide information on native range in parentheses.
GenBank numbers beginning with MF and MK are
new to this study.
Ancistranthus harpochiloides Lindau; Cuba, Shafer 13404
(US) MF670371, MF768291, MF678397, MF768343,
MF768396; Tetramerium lineage. Angkalanthus
oligophylla Balf.f.; Yemen (Socotra), Miller M10292
(UPS) EU081105, EU081172, EU087533, EU087567,
EU087478; Tetramerium lineage. Anisacanthus gracewoodiae Hammel & McDade; Costa Rica, Hammel et al.
26865 (RSA) MF670359, MF768281, MF678386,
MF768327, MF768385; Tetramerium lineage. Anisotes
formosissimus Milne-Redh.; cultivated, Lowveld National Botanical Garden, South Africa: Mpumalanga
(native to South Africa), Daniel 9382 (CAS)
KY584162, KY632232, MK282280, MK282352,
KY632576; Old World justicioid lineages. Anisotes
macrophyllus (Lindau) Heine; Uganda, Synnott 488
(EA) KY584141, KY632212, –, MK282353, MK282341;
Old World justicioid lineages. Anisotes madagascariensis
Benoist; Madagascar, Daniel & Butterwick 6736 (CAS)
KY584157, KY632227, MK282280, MK282354,
AF289772; Old World justicioid lineages. Anisotes
rogersii S.Moore; South Africa, McDade 1265 (J)
KY584140, KY632211, MK282282, MK282355,
KY632560; Old World justicioid lineages. Aphelandra
leonardii McDade; Costa Rica, McDade 310 (DUKE)
DQ059287, EU529071, DQ059205, AF063112,
AF169761; outgroup.
Barleria lupulina Lindl.; cultivated, San Francisco
Conservatory of Flowers, San Francisco, U.S.A. (native
to Madagascar), Daniel s.n. (CAS) EU528946,
EU529075, EU529010, AF289758, AF169751;
outgroup. Brachystephanus africanus S.Moore; Tanzania, Luke et al. 6704 (US) DQ372491, DQ372446,
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EU087537, MF768374, DQ372469; core Isoglossinae.
Calycacanthus magnusianus K.Schum.; cultivated, Royal
Botanic Gardens, Sydney (native to Papuasia), Daniel
10072 (CAS) EU081108, EU081175, EU087536,
MK282356, EU087481. Thyrsacanthus sulcatus (Nees)
C.Ezcurra & A.L.A.Côrtes; Paraguay, Schinini & Bordas
18098 (MO) MF670373, MF768293, MF678398,
MF7 68 34 6, MF 7 68 39 9; Tetram erium l in e age .
Cephalacanthus maculatus Lindau; Peru, Veja & Dillon
8517 (F) KY584123, KY632195, KY632029, KY632353,
KY632505, KY632545; New World justicioid lineage.
Chlamydocardia buettneri Lindau; cultivated, National
Botanic Garden of Belgium (native to Cameroun,
Gabon, Ivory Coast and Nigeria), Accession No. 950034-44 (BR) EU081107, EU081174, EU087535,
E U 0 87 56 9 , E U 0 8 74 80 ; Tetramerium l i ne ag e.
Chlamydostachya spectabilis Mildbr.; Poest Knox 88192/4
(K) KY584160, KY632230, MK282283, MK282357,
KY632574; Old World justicioid lineages. Chorisochora
transvaalensis (A.Meeuse) Vollesen; South Africa, Daniel 9379 (CAS) EU081100, EU081167, EU087528,
E U 0 8 75 6 5, EU 0 8 74 7 4 ; T e t r a me r iu m l i ne a ge.
Clinacanthus siamensis Bremek.; cultivated, National
Botanic Garden of Belgium (native to Thailand),
Accession No. 1979-0344 (BR) EU081106, EU081173,
EU087534, EU087568, EU087479; Tetramerium lineage.
Clistax brasiliensis Mart., Brazil, Lombardi 2257 (US)
KY584112, KY632186, MK282284, MK282358,
KY632537; New World justicioid lineage. Conocalyx
laxus Benoist; Madagascar, Leandri 3033 (P);
DQ372485, DQ372440, –, MK282359, MK282342; core
Isoglossinae.
Dicliptera extenta S.Moore; South Africa, McDade 1306
(J) EU528955, EU529084, EU529018, AF289724,
KC420542; Dicliptera s.l. Dicliptera insularis Benoist;
Madagascar, Daniel 9194 (CAS) KY584194, –,
MK282285, AF289723, AF289764; Dicliptera s.l.
Dicliptera magaliesbergensis K.Balkwill; South Africa,
Daniel 9357 (CAS) KY584192, MK282273, –,
AF289725, KY632607; Dicliptera s.l. Dicliptera resupinata
Juss.; United States, Boyd s.n. (ARIZ); KY584189,
EU529084, EU529019, MK282360, AF169841; Dicliptera
s.l. Dicliptera suberecta (André) Bremek., cultivated,
University of Arizona (native to Argentina, Bolivia,
Paraguay), McDade 1176 (ARIZ) KY584188, KY632257,
MK282286, AF289722, AF289763; Dicliptera s.l.
Henrya insularis Nees ex Benth; Mexico, Jenkins 89-432
(ARIZ) EU081071, EU081142, EU087507, AF063125,
AF169843; Tetramerium lineage. Hoverdenia speciosa
Nees; Mexico, Daniel & Baker 3739 (CAS) EU081089,
EU081157, EU087519, AF289738, AF289777;
Tetramerium lineage. Hypoestes sp.; Madagascar, Daniel
& McDade 10433 (CAS) KY584203, KY632274,
MK282287, MK282361, –; Hypoestes clade. Hypoestes
KEW BULLETIN
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aristata (Vahl) Sol. ex Roem. & Schult.; cultivated, San
Francisco Conservatory of Flowers (East Africa, South
Africa, Madagascar), Daniel s.n. (CAS) KY584208,
KY632279, MK282288, AF289726, KY632625; Hypoestes
clade. Hypoestes phyllostachya Baker, cultivated University of Arizona (widespread in OW tropics), McDade
1232 (ARIZ) KY584207, KY632278, EU529025,
AF195520, AF169842; Hypoestes clade.
Isoglossa gracillima Baker; Madagascar, Daniel 9106
(CAS) DQ372488, DQ372443, MK282289, AF289746,
AF289789; core Isoglossinae. Isoglossa grandiflora
C.B.Clarke; Cultivated, San Francisco Conservatory of
Flowers, San Francisco, CA, U.S.A. (Malawi, Tanzania,
Zambia), Daniel s.n. (CAS) DQ372490, DQ372445,
MK282290, AF289745, AF289788; core Isoglossinae.
Isoglossa ovata (Nees) Lindau; South Africa, Daniel 9336
(CAS) DQ372487, DQ372442, EU529027, MK282362,
DQ372468; core Isoglossinae.
Jadunia biroi (Lindau & K.Schum.) Lindau, Papua New
Guinea, Daniel et al. 6603 (CAS) MK282333,
MK282274, MK282291, MK282363, MK282343. Justicia
sp.; Cameroon, Tchiengue 2518 (K) MK282268,
MK282269, MK282270, MK282271, MK282344. Justicia
sp.; Madagascar, Daniel 9024 (CAS) KY584182,
KY632251, MK282292, AF289729, AF289768; Old
World justicioid lineages. Justicia sp.; Madagascar,
Daniel et al. 10446 (CAS) KY584138, KY632210,
MK282293, MK282365, KY632559; Old World
justicioid lineages. Justicia aconitiflora (A.Meeuse)
Cubey; cultivated, San Francisco Conservatory of
Flowers (native to South Africa), Daniel et al. 9361
(CAS) KY584161, KY632231, MK282294, AF289735,
AF289774; Old World justicioid lineages. Justicia
adhatoda L.; cultivated, University of Arizona (native
to South Africa), Barr 60-393 (ARIZ) KY584158,
EU081109, DQ059214, AF289734, AF289773; Old
World justicioid lineages. Justicia americana (L.) Vahl;
United States, Daniel & Lott 10530 (CAS) KY584096,
KY632173, MK282295, MK282366, KY632522; New
World justicioid lineage. Justicia anagalloides (Nees)
T.Anderson; cultivated NBI Garden, South Africa
(native to Africa), McDade & Balkwill 1239 (J)
KY584151, KY632221, MK282296, MK282367,
MK282345; Old World justicioid lineages. Justicia
andromeda (Lindau) J.C.Manning & Goldblatt;
South Africa, Balkwill et al. 11660 (J) KY584146,
KY632216, MK282297, MK282368, KY632563; Old
World justicioid lineages. Justicia anselliana (Nees)
T.Anderson; Tanzania, Kiel et al. 155 (RSA)
KY584181, KY632250, MK282298, MK282369,
KY632594; Old World justicioid lineages. Justicia
betonica L.; cultivated, Johannesburg, Gauteng,
South Africa (widespread tropical Africa), Daniel
9369 (CAS) KY584165, KY632235, MK282299,
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39
AF289731, AF289770; Old World justicioid lineages.
Justicia bolusii C.B.Clarke; cultivated, South Africa
(native to South Africa), Balkwill s.n. (J) KY584246,
KY632313, –, –, KY632656. Justicia brandegeeana
Wassh. & L.B.Sm.; cultivated, University Arizona
(native to Mexico), Starr c. 32 (ARIZ) KY584130,
KY632203, MK282300, MK282370, KY632553; New
World justicioid lineage. Justicia brasiliana Roth;
Paraguay, Zardini & Aguayo 9523 (CAS) KY584118,
KY632190, MK282301, MK282371, KY632542; New
World justicioid lineage. Justicia cavernarum
F.Muell.; Australia, Forster 28601 (CAS) KY584143,
KY632214, MK282302, MK282372, KY632562; Old
World justicioid lineage. Justicia comata (L.) Lam.;
Costa Rica, Kiel & McDade 138 (CAS) KY584088,
KY632165, MK282303, MK282373, KY632515; New
World justicioid lineage. Justicia flava (Forssk.)
Vahl; South Africa, McDade et al. 1256 (J)
KY584235, KY632302, MK282304, MK282374,
KY632648; Harnieria & Tyloglossa clades. Justicia
gendarussa Burm.f.; cultivated, San Francisco Conservatory of Flowers (native to tropical Asia), Daniel
s.n. (CAS) KY584166, KY632236, MK282305,
MK282375, KY632580; Old World justicioid lineages. Justicia grossa C.B.Clarke; Thailand, Von
B o u s e k o m & S a n t i s u k 2 6 9 9 ( L ) K Y 5 8 41 7 1 ,
KY632241, MF678390, MF768371, KY632585;
Tetramerium lineage. Justicia hassleri (Lindau)
V.A.W.Graham; Bolivia, Wood 19750 (CAS)
KY584131, KY632204, MK282306, MK282376,
KY632554; New World justicioid lineage. Justicia
hyssopifolia L.; cultivated Kew 1991-1486 (native to
C a n a r y I s la n d s ) ( K ) K Y 58 4 2 50 , K Y6 3 2 3 17 ,
MK282307, MK282377, KY632659; Old World
justicioid lineages. Justicia linifolia (Lindau)
V.A.W.Graham; South Africa, Balkwill 10860 (J)
KY584238, KY632305, MK282308, MK282378,
KY632651; Old World justicioid lineages. Justicia
longii Hilsenb.; United States, Arizona, Van Devender
87-307 (ARIZ) KY584107, KY632182, MK282309,
MK282379, KY632532; New World justicioid lineage.
Justicia odora (Forrsk.) Lam.; South Africa, Daniel &
Butterwick 9364 (CAS) KY584241, KY632308,
MK282310, MK282380, KY632653; Harnieria &
Tyloglossa clades. Justicia pacifica Hemsl.; Mexico,
Kiel et al. 74 (RSA) KY584103, KY632179, MF678421,
MF768367, KY632529; New World justicioid lineage.
Justicia prominens Benoist; Laos, Klackenberg 1383 (S)
KY584177, KY632246, MF678395, MF768370,
KY632590; Tetramerium lineage. Justicia rhodoptera
Baker; Madagascar, Daniel & Butterwick 6741 (CAS)
KY584136, KY632208, MK282272, MK282381,
KY632557; Old World justicioid lineages. Justicia
ruwenzoriensis C.B.Clarke; Zaire, de Witte 2862
(CAS) KY584164, KY632234, MK282311, MK282382,
KY632578; Old World justicioid lineages. Justicia
© The Author(s), 2019
39
Page 22 of 25
vagabunda Benoist; China, Tripp 1547 (COLO)
MK282334, MK282275, MK282312, MK282383, –.
Lepidagathus formosensis C.B.Clarke ex Hayata; Taiwan,
Bartholomew et al. 7654 (CAS) EU528973, EU529101,
EU529034, EU528930, EU529101; outgroup.
Marcania grandiflora J.B.Imlay; Thailand, Van Beusekom
et al. 3985 (L) MK282335, MK282276, MK282313,
MK282384, MK282346. Megalochlamys revoluta
(Lindau) Vollesen; South Africa, McDade & Balkwill
1264 (J) EU081099, EU081166, EU087527, EU087564,
EU087473; Tetramerium lineage. Megaskepasma
erythrochlamys Lindau; Costa Rica, Hammel et al. 26088
(RSA) KY584120, KY632192, MK282314, MK282385,
KY632544; New World justicioid lineage. Metarungia
galpinii (Baden) Baden; cultivated, Johannesburg, Gauteng, South Africa (native to South Africa), Daniel 9322
(CAS) KY584168, KY632238, EU529046, AF289737,
KY632582; Old World justicioid lineages. Metarungia
longistrobus Baden; cultivated, Witwatersrand National
Botanic Garden, Roodepoort, South Africa (native to
South Africa), Daniel et al. 9355 (CAS) KY584169,
KY632239, MK282315, MK282386, KY632583; Old World
justicioid lineage. Mirandea huastecensis T.F.Daniel; Mexico, Manktelow 706 (UPS) EU081099, EU081166,
EU087527, EU087564, EU087473: Tetramerium lineage.
Monechma divaricatum C.B.Clarke; South Africa, McDade
1275 (J) KY584209, KY632280, MK282316, MK282387,
KY632626; Monechma II clade. Monechma divaricatum
C.B.Clarke; South Africa, Daniel et al. 9373 (CAS)
KY584210, KY632281, MK282317, MK282388,
KY632627; Monechma II clade. Monechma mollissimum
P.G.Mey.; South Africa, Balkwill et al. 11787 (J)
KY584211, KY632282, MK282318, MK282389,
KY632628; Monechma II clade. Monothecium aristatum
(Nees) T.Anderson; Tanzania, Abdallah et al. 512 (US)
MK282336, MK282277, MK282319, MK282390,
MK282348. Monothecium glandulosum Hochst.; Ethiopia,
Friis et al. 9498 (C) MK282337, MK282278, MK282320,
MK282391, MK282347.
Odontonema tubaeforme (Bertol.) Kuntze; cultivated
Duke University greenhouse (native to Mexico and
Central America), McDade 1182 (ARIZ) KY584172,
DQ372462, DQ059215, AF063127, AF169748;
Pseuderanthemum lineage.
Pachystachys lutea Nees; cultivated, Duke University
greenhouse (native to Peru), McDade 1181 (DUKE)
EU081082, EU081151, EU087516, AF063128,
AF169844; Tetramerium lineage. Peristrophe brassii
R.M.Barker; Australia, Forster 28600 (CAS) KY584201,
KY632272, MK282321, MK282392, KY632618; Dicliptera
s.l. Peristrophe hyssopifolia (Burm.f.) Bremek., cultivated,
Missouri Botanical Garden, St. Louis, Missouri, U.S.A.,
MBG861410 (native to Indonesia), MacDougal 5047
© The Author(s), 2019
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(MO) KY584202, KY632273, MK282322, MK282393,
KY632620; Dicliptera s.l. Peristrophe speciosa (Roxb.)
Nees, cultivated San Francisco Conservatory of Flowers
(native to Asia), Daniel s.n. (CAS) KY584200,
KY632271, MK282323, MK282394, KY632617; Dicliptera
s.l. Poikilacanthus macranthus Lindau; Costa Rica, Kiel
et al. 85 (RSA) KY584127, KY632200, MK282324,
MK282395, KY632550; New World justicioid lineage.
Populina richardii Baill.; Madagascar, Kerardren 1671 (P)
EU081104, EU081171, EU087532, EU087566, EU087477;
Tetramerium lineage. Pseuderanthemum floribundum
T.F.Daniel; cultivated, San Francisco Conservatory (native
to Mexico), Daniel 5381cv (CAS) KY584173, –, MK282325,
MK282396, DQ37247; Pseuderanthemum lineage. Ptyssiglottis
psychotriifolia (Stapf) B.Hansen; Borneo, Poulsen 40 (C)
DQ372484, DQ372439, MK282326, MK282397,
DQ372466. Ptyssiglottis pubisepala (Lindau) B.Hansen;
Papua New Guinea, Daniel 6630 (CAS) DQ372483,
DQ372438, EU529055, AF289744, AF289787.
Razisea spicata Oerst; Costa Rica, Hammel 7974 (DUKE)
DQ372502, DQ372457, EU529056, AF063131, AF169848;
core Isoglossinae. Rhinacanthus angulicaulis I.Darbysh.;
Kenya, Kiel et al. 170 (RSA) KY584226, KY632293, –, –,
KY632639; core Rhinacanthus. Rhinacanthus dichotomus
(Lindau) I.Darbysh. var. dichotomus; Tanzania, Mfinanga
17 (US) KY584223, KY632290, –, –, KY632636; core
Rhinacanthus. Rhinacanthus latilabiatus (K.Balkwill)
I.Darbysh.; cultivated San Francisco Conservatory of
Flowers (native to South Africa), Daniel s.n. (CAS)
KY584222, KY632289, MK282327, MK282398, KY632635;
core Rhinacanthus. Rhinacanthus nasutus (L.) Kuntze;
Thailand, Nanakorn 880081 (US) KY584229, KY632296,
–, –, KY632642; core Rhinacanthus. Rhinacanthus ndorensis
Schweinf.; Kenya, Yovng 3169 (EA) KY584221, KY632288,
–, –, –. Rhinacanthus ndorensis Schweinf.; Kenya, Luke
17084 (EA) MK282338, MK282279, MK282328,
MK282399, MK282349. Rhinacanthus pulcher Milne.Redh.; Kenya, Gilbert & Thulin 1081 (EA) KY584225,
KY632292, –, –, KY632638; core Rhinacanthus.
Rhinacanthus rotundifolius C.B.Clarke; Kenya, Luke et al.
TPR60 (EA) KY584227, KY632294, –, –, KY632640; core
Rhinacanthus. Rhinacanthus virens (Nees) Milne-Redh.;
Uganda, Lye 3875 (EA) KY584220, –, –, –, KY632634.
Rhinacanthus xerophilus A.Meeuse; South Africa, McDade
1277 (J) KY584224, KY632291, MK282329, MK282400,
KY632637; core Rhinacanthus. Ruellia californica (Rose)
I.M.Johnst.; Cultivated, Univ. Arizona campus, Tucson,
Arizona, U.S.A. (native to northwest Mexico), McDade
1157 (ARIZ) EU528996, EU529123, EU529058,
AF063115, AF167704; outgroup. Rungia klossii S.Moore;
Papua New Guinea, Daniel et al. 6561 (CAS) KY584170,
KY632240, MK282330, MK282401, KY632584; Old World
justicioid lineages.
Schaueria gonatistachya Nees; Brazil, Paixao et al. 67
(CAS) MF670378, MF768297, MF678389, MF768352,
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MF768404; Tetramerium lineage. Schaueria populifolia
C.B.Clarke; Cameroon, Gereau et al. 5666 (MO)
MF670388, MF768309, MF678381, MF768372,
MF768415. Schaueria populifolia C.B.Clarke; Cameroon,
Cheek 10153 (K) MK282339, –, –, MK282402, MK282351.
Schaueria populifolia C.B.Clarke; Cameroon, Manning
1336 (K) MK282340, –, –, MK282403, MK282350.
“Tetramerium lineage” indet.; Malaysia, Hansen &
Smitinand 12005 (L) MF670389, MF768307,
MF678404, MF768368,–; Tetramerium lineage.
Tetramerium nervosum Nees; United States, McDade &
Jenkins 1154 (ARIZ) EU081058, EU081126, EU087493,
AF063133, AF169847; Tetramerium lineage. Trichocalyx
obovatus Balf.f.; Socotra, Yemen, Thulin & Gifri 8503
(UPS) KY584176, KY632245, MK282331, MK282404,
KY632589; Harnieria & Tyloglossa clades. Xerothamnella
herbacea C.T.White; Australia, Daniel 10066 (CAS)
KY584204, KY632275, MK282332, MK282405,
KY632621; Dicliptera s.l.
Open Access This article is distributed under the
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International License (http://creativecommons.org/
licenses/by/4.0/), which permits unrestricted use,
distribution, and reproduction in any medium, provided you give appropriate credit to the original
author(s) and the source, provide a link to the
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