The phylogeny of Linderniaceae – The new genus Linderniella, and new combinations within Bonnaya, Craterostigma, Lindernia, Micranthemum, Torenia and Vandellia

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DOI: 10.3372/wi.43.43201
Cite this publication
Abstract
The recently described Linderniaceae are a monophyletic group that emerged in the course of the disintegration of the Scrophulariaceae in the last years. First molecular studies sampled only a small fraction of the genera assigned to the Linderniaceae, but later Lindernia was shown to be non-monophyletic. Here we used a plastid trnK/matK sequence dataset to get further insights into the relationships within Linderniaceae. The genus Lindernia as accepted to date is shown to be polyphyletic. Stemodiopsis is found to be sister to the remaining Linderniaceae, and the genera Psammetes and Bryodes are found to be nested in a Lindernia s.str.clade. We present a taxonomic framework, taking account of the phylogenetic relationships in the family, and a first key to the genera. The genera Bonnaya and Vandellia are resurrected and a new genus Linderniella is described. The following eight replacement names are proposed: Craterostigma engleri, C. tanzanicum, Lindernia benthamii, L. lemuriana, Linderniella pusilla, Torenia bonatii, T. davidii and T. philcoxii. Seventy-seven new combinations are made in Bonnaya, Craterostigma, Lindernia, Linderniella, Micranthemum, Torenia and Vandellia.
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209Willdenowia 43 – 2013
EBERHARD FISCHER1*, BASTIAN SCHÄFERHOFF2 & KAI MÜLLER2
The phylogeny of LinderniaceaeThe new genus Linderniella, and new combinations
within Bonnaya, Craterostigma, Lindernia, Micranthemum, Torenia and Vandellia
Abstract
Fischer E., Schäferho B. & Müller K.: The phylogeny of Linderniaceae – The new genus Linderniella, and new
combinations within Bonnaya, Craterostigma, Lindernia, Micranthemum, Torenia and Vandellia. – Willdenowia 43:
209 238. December 2013. – ISSN 0511-9618; © 2013 BGBM Berlin-Dahlem.
Stable URL: http://dx.doi.org/10.3372/wi.43.43201
The recently described Linderniaceae are a monophyletic group that emerged in the course of the disintegration of
the Scrophulariaceae in the last years. First molecular studies sampled only a small fraction of the genera assigned
to the Linderniaceae, but later Lindernia was shown to be non-monophyletic. Here we used a plastid trnK/matK
sequence dataset to get further insights into the relationships within Linderniaceae. The genus Lindernia as accepted
to date is shown to be polyphyletic. Stemodiopsis is found to be sister to the remaining Linderniaceae, and the genera
Psammetes and Bryodes are found to be nested in a Lindernia s.str. clade. We present a taxonomic framework, taking
account of the phylogenetic relationships in the family, and a first key to the genera. The genera Bonnaya and Vandel-
lia are resurrected and a new genus Linderniella is described. The following eight replacement names are proposed:
Craterostigma engleri, C. tanzanicum, Lindernia benthamii, L. lemuriana, Linderniella pusilla, Torenia bonatii, T.
davidii and T. philcoxii. Seventy-seven new combinations are made in Bonnaya, Craterostigma, Lindernia, Linderni-
ella, Micranthemum, Torenia and Vandellia.
Additional key words: trnK, matK, Lamiales, poikilohydric plants, desiccation tolerance
Introduction
Research interest in Lindernia and related genera has in-
creased in the last decade. One focus has been on nature
conservation as the European Lindernia procumbens is a
priority species of the Convention on the Conservation of
European Wildlife and Natural Habitats (Appendix IV).
The species is threatened by the destruction and also the
management change of its habitats (i.e. ephemeral Na-
nocyperion communities in ponds) (Šumberová & al.
2012). Another focus has been the research on poikilo-
hydric taxa. Craterostigma (Fig. 1A) and Chamaegigas
(Fig. 1G) are well-known resurrection plants (e.g. Bartels
& al. 1990; Fischer 1992), and also species of Lindernia
have been shown to be poikilohydric (Phillips & al. 2008,
see Fig. 1F).
An assemblage of taxa that would currently be cir-
cumscribed as Linderniaceae was first considered by
Bentham (1846) to represent a subtribe within the
Gratio leae. Based on Lindernia, the circumscription of
genera was, however, controversially discussed result-
ing in a complex taxonomic history with proposals to
unite all species into a broadly defined genus Lindernia
or splitting them into 13 dierent genera (Fischer 1992).
Lindernia was based on L. procumbens from Europe.
Bentham (1835) recognized Vandellia (Fig. 1H) with
4 fertile stamens and Bonnaya with 2 fertile stamens
and 2 staminodia. Bentham (1846) accepted Arta nema,
1 Institut für Integrierte Naturwissenschaften Biologie, Universität Koblenz-Landau, Universitätsstr. 1, 56070 Koblenz, Ger-
many; *e-mail: efischer@uni-koblenz.de (author for correspondence).
2 Institut für Evolution and Biodiversität, Universität Münster, Hüerstr. 1, 48149 Münster, Germany; e-mail: schaeferho@uni-
muenster.de; kaimueller@uni-muenster.de
210 Fischer & al.: The phylogeny of Linderniaceae
Fig. 1. Linderniaceae – A: Craterostigma hirsutum, Rwanda; B: Linderniella pygmaea, Madagascar; C, D: Linderniella horomb-
ensis, Madagascar; E: Linderniella gracilis, Rwanda; F: Linderniella brevidens, Kenya; G: Chamaegigas intrepidus, Namibia; H:
Vandellia diusa, Gabon; I, K: Torenia stolonifera, Madagascar; L: Torenia thouarsii, Gabon; M: Torenia crustacea, Gabon. – All
photographs by E. Fischer except G by N. Jürgens.
211Willdenowia 43 – 2013
Fig. 2. LinderniaceaeA: Lindernia viguieri, Madagascar; B: Lindernia rotundifolia, Madagascar; C: Lindernia benthamii (= Bryo-
des micrantha), Madagascar; D: Lindernia madagascariensis (= Psammetes madagascariensis), Madagascar; E: Crepidorhopalon
microcarpaeoides, Madagascar; F: Crepidorhopalon tenuis, Congo-Kinshasa; G, H: Crepidorhopalon whytei, Rwanda; I: Hart-
liella capitata, Congo-Kinshasa; K: Stemodiopsis ruandensis, Rwanda. – All photographs by E. Fischer except I by M. Schaijes.
212 Fischer & al.: The phylogeny of Linderniaceae
Bonnaya, Curanga Juss. (= Picria Lour.), Hemianthus,
Ilysanthes, Lindernia, Micranthemum, Peplidium Delile,
Torenia (Fig. 1I, K) and Vandellia as genera within sub-
tribe Lindernieae. The African genus Craterostigma was
reduced to a section of Torenia (Bentham 1846). Hooker
(1885) used the size of the fruit and the leaf venation for
the distinction of Bonnaya (pinnate venation) and Ilysan-
thes (palmate venation). Urban (1884) lumped Bonnaya
and Ilysanthes together with Bazina Raf. into Ilysanthes
with 2 fertile stamens and Ilyogeton, Tittmannia and Van-
dellia with 4 fertile stamens into Lindernia. This concept
of separating Lindernia and Ilysanthes, was also ac-
cepted by Wettstein (1891) and used until the 1960s (e.g.
Hepper 1963). Other proposals, e.g. by Hance (1861) to
unite Torenia and Vandellia, or by Müller (1882) to place
Bonnaya, Ilyogeton, Ilysanthes, Tittmannia, Vandellia
and a part of Torenia with Lindernia were not accepted.
Wettstein (1891) also recognized Craterostigma because
of its rosulate habit and Torenia because of the winged
calyx as separate genera. This concept was modified by
Engler (1897) based on taxa described from tropical Afri-
ca. He published a new circumscription of Craterostigma,
where he placed species with a dense inflorescence, e.g.
C. crassifolium, C. goetzei, C. schweinfurthii (originally
described as Torenia schweinfurthii) and C. welwitschii.
Torenia was mainly characterized by the winged calyx.
Already Schlechter (1924), however, noted that Torenia
was quite heterogeneous and many African taxa appar-
ently misplaced in this genus.
A major achievement was made by Pennell (1935),
who argued that a reduction of stamens cannot be used
as a generic character. Subsequently he united Ilysan-
thes and Lindernia into a broadly circumscribed genus
Lindernia. This was accepted by the majority of au-
thors (e.g. Philcox 1968; Yamazaki 1985, 1990; Fischer
1992, 1995, 2004; Lewis 2000; Philcox 2008). However,
doubts arose due to the morphological heterogeneity
(e.g. Fischer 1992) whether Lindernia was really mono-
phyletic. Yamazaki (1954a b, 1955) already proposed a
division into Vandellia with pinnate leaf venation, serrate
leaf margin, a 1-celled chalazar haustorium, and alveo-
late endosperm (bothrospermous seeds, i.e. with rounded
pits, see Fig. 3A, B), and Lindernia with palmate leaf
venation, entire or only slightly dentate leaf margin, a
2-celled chalazar haustorium and smooth, non-alveolate
endosperm (seeds as in Fig. 3C). Later the same author
adopted the concept of Pennell (Yamazaki 1985, 1990).
The genus Craterostigma was redefined by Fischer (1986)
Fig. 3. SEM photographs of Linderniaceae seeds – A: Torenia crustacea; B: Linderniella cerastioides; C: Lindernia rotundifolia;
D: Crepidorhopalon microcarpaeoides. – Scale bar 100 µm. – Photographs by E. Fischer.
213Willdenowia 43 – 2013
comprising rosulate plants with truncate inflorescences
and bothrospermous seeds. Hepper (1987a) came to the
same conclusion but transferred species like C. gracile,
C. schweinfurthii and others to Torenia (Hepper 1987b).
Philcox (1987, 1990) described several new species of
Torenia which subsequently were transferred to Crepi-
dorhopalon (Fischer 1992). This highly unnatural con-
cept of Crate rostigma and Torenia was even maintained
later (Hepper 2008). Fischer (1989) united all former
African species of Craterostigma with aulacospermous
seeds (i.e. with longitudinal furrows, see Fig. 3D), non-
rosulate habit and anatomically highly complex clavate
hairs on the lower corolla-lip comprising a distinct mul-
ticellular base into the new genus Crepidorhopalon.
Also plants formerly placed in Lindernia, Torenia or
even Stemodiopsis (Fig. 2K) showed the above men-
tioned characters and were subsequently transferred to
Crepidorhopalon (Fischer 1992, 1995, 1997a, 1999a b).
A further new genus of suruticose metallophytes from
Central Africa (Katanga), Hartliella Eb. Fisch. (Fig. 2I),
was described and based on former species of Linder-
nia (Fischer 1992, 1999b). Morphological data already
pointed out that the family Scrophulariaceae is not
monophyletic (Fischer 1992). Molecular phylogenetic
studies (Olmstead & Reeves 1995; Olmstead & al. 2001)
confirmed the polyphyly of traditional Scrophulariaceae
and lead to its disintegration. However, Lindernia and its
relatives could not be placed. First molecular studies in-
cluding members of the former tribe Lindernieae (Rah-
manzadeh & al. 2005) revealed that they represented a
new family, Linderniaceae as a further lineage distinct
from Scrophula riaceae and Plantaginaceae. It also be-
came evident that Lindernia sensu Pennell is not mono-
phyletic, and that Crepidorhopalon and Torenia repre-
sent dierent clades. Linderniaceae as a separate lineage
was confirmed by Albach & al. (2005), who included
also Micranthemum, by Oxelman & al. (2005), who ad-
ditionally included Stemodiopsis, and by Schäferho &
al. (2010). Thus the objectives of the present paper are to
provide a first detailed phylogeny of Linderniaceae and
to elaborate the correct formal taxonomy for the revealed
lineages.
Material and methods
Taxon sampling and plant material Taxon sampling
included as many dierent genera of Linderniaceae as
we could obtain. Also several species per genus were in-
cluded where available. As outgroups for Linderniaceae,
the genus Byblis Salisb. (Byblidacaeae) was used as it
appeared as a close relative in a study of Lamiales phy-
logeny (Schäferho & al. 2010). Within Linderniaceae,
the genera Stemodiopsis and Micranthemum were in-
cluded to show their exact position. Voucher information
and GenBank accession numbers of taxa included in this
study are listed in Table 1.
DNA extraction, purification and sequencing — Total
genomic DNA was isolated using the AVE Gene Plant
Genomics DNA Mini Kit (AVE Gene, Korea) according
to the manufacturer’s protocol. As phylogenetic mark-
ers, the trnK intron including the coding matK gene was
amplified in two overlapping halves, using standard PCR
protocols. The 5´-part was amplified using trnK3914Fdi
(GGGGTTGCTAACTCAACGG, Johnson & Soltis
1995), and LindmatK1714R (CTCCAAAGAAAGYC
AGTTCCTCTT, Schäferho & al. 2010); for the 3´-
part the primers ACmatK500F (TTCTTCTTTGCATT
TATTACG, Müller & Borsch 2005), LindmatK1580F
(TCAATTCATTCAACWTTTCCC, Schäferho & al.
2010), and trnK2R (AACTAGTCGGATGGAGTAG,
Johnson & Soltis 1995) were used. Reactions were per-
formed in 50 µl volumes containing 2 µl template DNA
(10 ng/µl), 10 µl dNTP mix (1.25 mM each), 2 µl of each
forward and reverse primer (20pm/µl), and 0.25 µl Taq
polymerase (5 U/µl, Peqlab). Thermal cycling was car-
ried out on a Biometra T3 thermocycler using the fol-
lowing PCR profiles: 1: 30 min at 96°C, 1 min at 50°C,
1: 30 min at 72°C, 35 cycles of 30 sec at 96°C, 1 min at
50°C, 1: 30 min at 72°C, and a final extension time of
10 min at 72°C. Fragments were gel-purified on a 1.2 %
agarose gel (Neeo-agarose, Roth), extracted with the Gel/
PCR DNA Fragments Extraction Kit (AVE Gene, Korea)
and sequenced on an ABI3730XL automated sequencer
using the Macrogen sequencing service (Macrogen Inc.,
Seoul, Korea). Pherogram editing and contig assembly
was done manually using PhyDE (Müller & al. 2006).
Alignment, indel coding, and phylogenetic analyses —
DNA sequences were aligned manually in PhyDE (www.
phyde.de), taking microstructural changes into account
as outlined elsewhere (Kelchner 2000; Löhne & Borsch
2005). Regions of uncertain homology were excluded
from phylogenetic analyses. For maximum parsimony
(MP) analyses, indels were coded according to simple
indel coding (SIC) (Simmons & Ochoterena 2000) with
help of the program SeqState (Müller 2005a).
Searches for the shortest tree were performed using
the parsimony ratchet approach implemented in PRAP2
(Wall & al. 2008) using the following settings: 10 ran-
dom addition cycles with 200 ratchet replicates, setting
the weight for 25 % of the characters to 2. The files
generated were executed in PAUP* v4.0b10 (Swoord
1998). Tree evaluation was performed with 10 000 boot-
strap replicates (BS), each using TBR branch swapping
and holding only one tree (Müller 2005b).
The model of best fit for the dataset was found to be
the GTR+G+I model using jModelTest v.0.1.1 (Posada
2008). Bayesian inference (BI) of phylogeny was done
using MrBayes v3.1.2 (Ronquist & Huelsenbeck 2003).
Default priors were used, i.e. flat dirichlets (1.0, 1.0) for
state frequencies and instantaneous substitution rates, a
uniform prior (0.0, 50.0) for the shape parameter of the
gamma distribution, a uniform prior (0.0, 1.0) for the pro-
214 Fischer & al.: The phylogeny of Linderniaceae
Table 1. Species, voucher information, and GenBank accession numbers of the sequences used in the present study.
FamilySpecies Voucher GenBank
accession
number
Citation
Linderniaceae Artanema angustifolium Singapore, Perak, Kampong Permatang Serai, Dindings, J. Sinclair 9876 (M) FR728403 this study
Linderniaceae Artanema fimbriatum Germany, Botanical Gardens Bonn 0-BONN-15545, T. Borsch 3790 (BONN) AY667460 Rahmanzadeh & al. 2005
Linderniaceae Bonnaya ciliata (Lindernia ciliata) Thailand, Koh Samui, E. Fischer EF Th 677 (KOBL) FR728401 this study
Linderniaceae Bonnaya multiflora (Lindernia bonatii) Madagascar, Ambositra, E. Fischer 10236 (KOBL) FR728408 this study
Linderniaceae Bonnaya ruellioides (Lindernia ruellioides) Thailand, Koh-Samui, E. Fischer EF Th 690 (KOBL) FR728402 this study
Linderniaceae Chamaegigas intrepidus Südwestafrika [Namibia], Distr. Omaruru, Ohere-Oos, H. Merxmüller 1580 (M) FR728398 this study
Linderniaceae Craterostigma crassifolium (Lindernia crassifolia) Angola, Huilla, T. Leyens 347 (BONN) FR728390 this study
Linderniaceae Craterostigma engleri (Lindernia welwitschii) Angola, Huilla, Lobin & al. s.n. (BONN) FR728391 this study
Linderniaceae Craterostigma hirsutum Rwanda, Nyarubuye, 9003 (KOBL) AF531776 Rahmanzadeh & al. 2005
Linderniaceae Craterostigma newtonii (Lindernia oliveriana) Rwanda, Lac Ihema, Akagera National Park, E. Fischer s.n. (KOBL) FR728393 this study
Linderniaceae Craterostigma nummulariifolium (Lindernia
nummulariifolia)
Nyungwe National Park, Cyamundongo, E. Fischer 10256 (KOBL) FR728392 this study
Linderniaceae Craterostigma plantagineum Rwanda, Nyamata, E. Fischer s.n. (BG Bonn 19988) FR728387 this study
Linderniaceae Craterostigma pumilum Kenya, Nairobi, E. Fischer 8759 (KOBL) AY667462 Rahmanzadeh & al 2005
Linderniaceae Craterostigma pusillum (Lindernia acicularis) Rwanda, Nyarubuye, E. Fischer 4230 (KOBL) AY667465 Rahmanzadeh & al 2005
Linderniaceae Craterostigma sp. nov. Angola, Huilla, T. Leyens 401 (BONN) FR728389 this study
Linderniaceae Craterostigma yaundense (Lindernia yaundensis) Cameroon, Yaunde, S. Porembski 3858 (KOBL) FR728388 this study
Linderniaceae Crepidorhopalon bifolius Congo-Kinshasa, Katanga, Lubumbashi, M. Faucon s.n. (BRLU, KOBL) FR728417 this study
Linderniaceae Crepidorhopalon goetzei Congo-Kinshasa, Katanga, Lubumbashi, M. Faucon s.n. (BRLU, KOBL) FR728419 this study
Linderniaceae Crepidorhopalon microcarpaeoides Madagascar, Ambalavao, E. Fischer 10296 (KOBL) FR728418 this study
Linderniaceae Crepidorhopalon perennis Congo-Kinshasa, Katanga, Lubumbashi, M. Faucon s.n., E. Fischer 16 (BRLU,
KOBL)
FR728415 this study
Linderniaceae Crepidorhopalon tenuis Congo-Kinshasa, Katanga, M. Faucon s.n., E. Fischer 3 (BRLU, KOBL) FR728413 this study
Linderniaceae Crepidorhopalon whytei Rwanda, Butare, E. Fischer 7657 (KOBL) AY667463 Rahmanzadeh & al. 2005
Linderniaceae Lindernia benthamii (Bryodes micrantha) Madagascar, Antsirabé, E. Fischer 10258 (KOBL) FN773532 Schäferhoff & al. 2010
Linderniaceae Lindernia bryoides Madagascar, Ambositra, E. Fischer 10240 (KOBL) FR728404 this study
Linderniaceae Lindernia dubia Germany, F. Müller s.n. (BONN) FR728411 this study
Linderniaceae Lindernia madagascariensis (Psammetes
madagascariensis)
Madagascar, Ambositra, E. Fischer 10234 (KOBL) FR728410 this study
Linderniaceae Lindernia procumbens Portugal, Baixo Alentejo, H. Kalheber 99-1990 (BONN) FR728409 this study
Linderniaceae Lindernia rotundata Rwanda, Musanze, E. Fischer 7320 (KOBL) AY667469 Rahmanzadeh & al. 2005
continued on next page
215Willdenowia 43 – 2013
FamilySpecies Voucher GenBank
accession
number
Citation
Linderniaceae Lindernia rotundifolia Madagascar, Ambositra, E. Fischer 10241 (KOBL) FR728406 this study
Linderniaceae Lindernia microcalyx Germany, Botanical Garden Mainz, E. Fischer s.n. (BONN) AY667468 Rahmanzadeh & al. 2005
Linderniaceae Lindernia viguieri Madagascar, Antsirabe, E. Fischer 10267 (KOBL) FR728407 this study
Linderniaceae Linderniella brevidens (Lindernia brevidens) Kenya, Teita Hills, E. Fischer 8022 (KOBL) FN773545 Schäferhoff & al. 2010
Linderniaceae Linderniella gracilis (Lindernia exilis) Rwanda, Nyarubuye, E. Fischer 12225 (KOBL) FR728396 this study
Linderniaceae Linderniella horombensis (Lindernia horombensis) Madagascar, Ambalavao, E. Fischer 10298 (KOBL) FR728394 this study
Linderniaceae Linderniella pusilla (Lindernia philcoxii) Rwanda, Nyarubuye, E. Fischer 553/87 (KOBL) AY667466 Rahmanzadeh & al. 2005
Linderniaceae Linderniella pygmaea (Lindernia pygmaea) Madagascar, Ambalavao, E. Fischer s.n. (KOBL) FR728395 this study
Linderniaceae Micranthemum umbrosum Germany, Botanical Gardens Bonn, Schäferho 43 (BONN) FR773548 Schäferhoff & al. 2010
Linderniaceae Stemodiopsis ruandensis Rwanda, Rugarama, E. Fischer 10352 (KOBL) FN773559 Schäferhoff & al. 2010
Linderniaceae Torenia crustacea (Lindernia crustacea) Thailand, Koh-Samui, E. Fischer EF Th 692 (KOBL) FR728405 this study
Linderniaceae Torenia diusa (Torenia vagans) U.K., Royal Botanical Garden Edinburgh, B. L. Burtt s.n. (BONN) AF531812 Rahmanzadeh & al 2005
Linderniaceae Torenia stolonifera Madagascar, Ranomafana, E. Fischer 10249 (KOBL) FN773561 Schäferhoff & al. 2010
Linderniaceae Vandellia diusa (Lindernia diusa) Dominican Republic, T. Borsch 3837 (B) FR728399 this study
Linderniaceae Vandellia pusilla (Lindernia pusilla) Thailand, Koh-Samui, E. Fischer, EF Th 691 (KOBL) FR728397 this study
Linderniaceae Vandellia senegalensis (Lindernia senegalensis) Gabon, Makokou, E. Fischer s.n. (KOBL) FR822526 this study
Linderniaceae Vandellia subracemosa (Lindernia subracemosa) Rwanda, Uwinka, E. Fischer 1350 (BG Bonn 19990-2) (KOBL) FR728400 this study
Byblidaceae Byblis gigantea Lindl.Germany,
Botanical Gardens Bonn 0-BONN-2551
4, K. Müller 733 (MSUN) AF531774 Müller & al. 2004
Byblidaceae
Byblis lamellata Conran & Lowrie
Germany,
Botanical Gardens Bonn 0-BONN-25513, K. Müller s.n. (MSUN) FR773534
Schäferhoff & al. 2010
Byblidaceae
Byblis liniflora Salisb.
Germany,
Botanical Gardens Bonn 0-BONN-16008, K. Müller s.n. (MSUN) FR773535
Schäferhoff & al. 2010
216 Fischer & al.: The phylogeny of Linderniaceae
portion of invariable sites, and a uniform topological pri-
or. Using the formula provided by Brown & al. (2010), we
calculated an exponential rate parameter for the branch-
length prior based on an average branch length estimate
from an initial neighbor joining analysis in PAUP. Four
categories were used to approximate the gamma distribu-
tion. Two runs with 5 million generations each were run,
and four chains were run in parallel for each run, with
the temperature set to 0.2. The chains were sampled every
100th generation, and a relative burn in fraction was con-
servatively set to 25 %. Convergence was assessed via
mean and maximum potential scale reduction factors and
mean/maximum standard deviation of split frequencies,
which all clearly indicated that runs converged.
For maximum likelihood (ML) analyses, RAxML
v7.0.0 (Stamatakis 2006) was used. The GTRGAMMA
model was used during the search for the best tree, while
the slightly simpler GTRCAT model was employed in
RAxML during the 500 bootstrap replicates. Support
values from all types of analysis were mapped on the
tree topology from the Bayesian analysis and conflicting
nodes were identified with help of TreeGraph2 (Stöver &
Müller 2010).
Results
Molecular sequence data For most of the accessions,
complete trnK/matK sequences were obtained. The to-
tal alignment comprised 2742 characters; 92 characters
were excluded from phylogenetic analyses due to un-
certain homology. Sequence lengths ranged from 1173
to 2488nt (mean: 2350, SD 204; 33 % GC-content). Of
the 2650 characters used for phylogenetic analyses, 33 %
were variable, and 23 % were informative. Average se-
quences divergence was 5.63 % (S.E. 0.11).
The shortest trees from a parsimony ratchet analysis
were 1634 steps long (CI 0.75, RI 0.90, RC 0.67). The
tree topology of the Bayesian inference (BI) of phyl-
ogeny is given in Fig. 4, together with posterior prob-
abilities (pp), Maximum Likelihood bootstrap (ML BS)
and Maximum Parsimony bootstrap (MP BS) values. A
phylogram from BI indicating relative branch lengths is
shown in Fig. 5.
Relationships — In addition to the taxa included in the
study of Rahmanzadeh & al. (2005), Bryodes, Micran-
themum and Stemodiopsis (Schäferho & al. 2010), as
well as Psammetes and Chamaegigas are inferred to be
part of Linderniaceae. Stemodiopsis is found sister to all
remaining Linderniaceae with maximum support. There
is strong support (pp 1.00, 100 % ML BS, 100 % MP BS)
for two major clades within the remaining Linderniace-
ae. The first clade comprises the genus Crepidorhopalon
(pp 1.00, 100 % ML BS, 100 % MP BS) and Lindernia
s.str. including Micranthemum. The genera Bryodes and
Psammetes (Fig. 2D) nest within the Lindernia clade and
their position is well supported (pp 1.00, 98 % ML BS,
93 % MP BS and pp 1.00, 100 % ML BS, 100 % MP BS,
respectively). The second major clade in Linderniaceae
is also well supported (pp 1.00, 100 % ML BS, 99 & MP
BS). Herein, L. pusilla and a clade with Torenia including
L. crustacea (pp 1.00, 100 % ML BS, 100 % MP BS, Fig.
1M) are in unresolved positions. The clade comprised of
the remainder of the Linderniaceae taxa is only well-sup-
ported in model-based analyses (pp1.00, 100 % ML BS,
79 % MP BS); herein a group with Artanema, L. ciliata
and L. ruellioides (pp 1.00, 91 % ML BS, 83 % MP BS)
branches first. The next-branching clade with L. diusa
and related taxa is well supported (pp 1.00, 97 % ML BS,
92 % MP BS). The remaining two major clades have only
moderate support (pp 0.99, 97 % ML BS, 73 % MP BS
and pp 1.00, 82 % ML BS, 65 % MP BS, respectively).
The first branch covers Chamaegigas intrepidus and a
well-supported clade with L. horombensis (Fig. 1C, D)
and related taxa. The next moderately supported branch
(pp 0.99, 97 % ML BS, 73 % MP BS) com prises L. oliv-
eriana in a clade with L. nummulariifolia (pp 0.63, 59 %
ML BS), and a clade with L. crassifolia, L. welwitschii
and an undescribed species from Angola (pp1.00, 99 %
ML BS, 97 % MP BS). The last clade is well support-
ed (pp 1.00, 100 % ML BS, 100 % MP BS) and covers
the members of Craterostigma and L. acicularis with L.
yaundensis.
Discussion
Our results confirm the close relationship between Ste-
modiopsis and other taxa from Linderniaceae. The
abaxial filaments are curved or twisted and thus match
the general staminal morphology of Linderniaceae. Ste-
modiopsis is an African genus confined to rock outcrops
with a centre of diversity in SE Africa (Fischer 1997b).
There seems to be a progression from curved and twisted
to geniculate abaxial stamens with partial reduction to
staminodes occurring independently in several taxa (e.g.
Lindernia, Vandellia).
Our results show that Crepidorhopalon with aula-
cospermous seeds (Hartl 1959) and clavate hairs on the
lower corolla lip bearing a multicellular base (Fischer
1992) is a well-supported clade (pp 1.00, 100 % ML
BS, 100 % MP BS), and that inclusion of its members
in Lindernia (Philcox 2008) or Torenia (Hepper 2008)
would result in polyphyletic genera. Lindernia s.str. is
characterized by seeds with a smooth (non-alveolate)
endosperm (Yamazaki 1954a b, 1955; Hartl 1959;
Fischer 1992). Lindernia here forms a clade that is well
supported (pp 1.00, 100 % ML BS, 100 % MP BS) with
the exception of Lindernia dubia. Our results show that
the monotypic genera Bryodes and Psammetes (Fischer
& Hepper 1997) are nested within the Lindernia clade.
Both genera have mainly cleistogamous flowers. Howev-
er, this is a feature that regularly occurs in Lindernia and
217Willdenowia 43 – 2013
also Vandellia. The type species of Lindernia, L. procum-
bens usually bears small cleistogamous flowers, and the
form with chasmogamous flowers occurring mainly in
the tropics has even been described as a dierent species
(Vandellia erecta Benth.). Psammetes was considered to
be exclusively cleistogamous, but the first author could
discover chasmogamous plants in Madagascar. Both
Bryodes and Psammetes are included here in Lindernia.
Micranthemum (incl. Hemianthus) also bears abaxial
stamens with clavate geniculations covered with glan-
dular hairs. Their position close to Lindernia, already
proposed by Albach & al. (2005), is thus confirmed.
They share with Lindernia the non-alveolate endosperm.
They form a well-supported clade (pp 1.00, 100 % ML
BS, 100 % MP BS) with the species of Lindernia s.str.
However, since so far only one species has been stud-
ied, Micranthemum is not included in Lindernia. Also
several morphological dierences exist between the two
genera: the usually complete lack of adaxial stamina
and the strong reduction of the corolla and especially
the upper lip. The majority of species are resolved in a
well-supported clade that is characterized by bothrosper-
mous seeds with alveolate endosperm of the Torenia type
(Hartl 1959; Fischer 1992). Torenia is maintained here,
and L. crustacea, suggested to have anities to Torenia
by some previous authors, is shown to be indeed a mem-
ber of Torenia. This group of species has been called
Lindernia sect. Torenioides (e.g. Philcox 1968), and its
members are transferred here to Torenia. Lindernia cili-
ata, L. ruellioides and Artanema form a moderately sup-
ported clade (pp 1.00, 91 % ML BS, 83 % MP BS) that
can be divided into two well-supported clades. Artanema
has a unique stamen morphology with two boss-like fila-
ment knees per abaxial stamen (Fischer 1999b, 2004).
The genus comprises large herbs with usually very large
flowers. For the clade with L. ciliata and L. ruellioides,
the already existing name Bonnaya can be revived. In
most recent treatments (e.g. Philcox 1968; Yamazaki
Fig. 4. Phylogeny of Linderniaceae inferred from parsimony, likelihood and Bayesian analysis of trnK/matK sequences. Topology
from the Bayesian tree depicted, collapsing nodes not supported by > = 50% in at least one of the three analyses. Bold numbers
above branches are posterior probabilities from Bayesian inferences, italic numbers above branches are ML bootstrap values, num-
bers below branches indicate MP bootstrap proportions. Numbers in brackets indicate that the respective node was not supported by
all three methodological approaches. The bracketed number then indicates the strongest support found for any node that contradicts
the shown node. Names are according to Fischer (1992). To the right of the tree are taxonomic changes as suggested in this study.
218 Fischer & al.: The phylogeny of Linderniaceae
1990) it was used as a subgenus of Lindernia. The spe-
cies of Bonnaya share abaxial staminodes, pinnate leaf
venation and bothrospermous seeds.
The well-supported clade around Lindernia diusa
can be called Vandellia as L. diusa is the type species of
that genus. Yamazaki (1954a b, 1955) already proposed
a division into Vandellia with pinnate leaf venation, ser-
rate leaf margin, a 1-celled chalazar haustorium, and al-
veolate endosperm (bothrospermous seeds), and Linder-
nia with palmate leaf venation, entire or only slightly
dentate leaf margin, a 2-celled chalazar haustorium and
smooth (non-alveolate) endosperm. Although L. pusilla
is found unresolved in the bothrospermous clade, its mor-
phological resemblance to other members of Vandellia
is so striking that the taxon is provisionally assigned to
this genus. The poikilohydric Chamaegigas intrepidus, a
local endemic of granitic outcrops in Namibia, is a mor-
phologically very distinct plant with seeds with non-alve-
olate endosperm and basally rosulate leaves that are fused
(Fischer 1992). Its position within the bothrospermous
clade points toward a secondary loss of bothrospermy.
Due to its isolated morphology the genus Chamaegigas is
maintained here. Lindernia horombensis and L. pygmaea
(Fig. 1B) from Madagascar and L. brevidens, L. exilis and
L. philcoxii from East Africa form a well-supported clade
for which no name is available. It is described here as the
new genus, Linderniella.
The last major clade (pp 1.00, 82 % ML BS, 65 %
MP BS) includes the genus Craterostigma and several
taxa of Lindernia. The members of Craterostigma s.str.
can easily be recognized by the usually rosulate, Planta-
go-like habit with truncate synflorescence. This type of
habit is also represented in some members of the former
genus Lindernia that also belong to the Craterostigma
clade (e.g. L. gossweileri, L. welwitschii) and that also
share the poikilohydry with Craterostigma. The only aut-
apomorphy of core Craterostigma is the red to yellow-
orange color in the intercellular spaces of the root cortex
(Fischer 1992). Thus Craterostigma is included here as a
more broadly defined and well-supported genus with the
addition of L. acicularis, L. nummulariifolia, L. oliveri-
ana, L. welwitschii and L. yaundensis.
Several genera of Linderniaceae already assigned
to that family (Fischer 2004; Rahmanzadeh & al. 2005)
have not been available for study up to now mainly due
to the condition and age of herbarium specimens and the
lack of fresh material. Hartliella with aulaco spermous
seeds comprises four species endemic to the copper belt
of Katanga province in Congo-Kinshasa and N Zam-
bia (Fischer 1999b, 2004). Schizotorenia with two spe-
cies from Indo-Malesia also has aulacospermous seeds
(Fischer 2004). The monotypic Hemiarrhena plantag-
inea from NW Australia, the monotypic Legazpia poly-
gonoides from SE Asia, Micronesia and New Guinea,
the monotypic Pierranthus capitatus from SE Asia and
Scolophyllum, with three species from Thailand to Cam-
bodia and Vietnam, have bothrospermous seeds (Fischer
2004). Fresh or younger herbarium material also has
not been available for study for the eight Australian taxa
of Lindernia subg. Didymadenia (Barker 1990, 1998).
Barker (1990) characterized this subgenus by the pres-
ence of 2-celled sessile glands and 4- or 5-angled seeds,
which are probably bothrospermous as the author stated
“angles alternating with grooves” (Barker 1990: 80). As
Barker also assigned to his new subgenus L. scapigera
and L. subulata, which are placed here in Vandellia, the
eight Australian taxa are included here in Vandellia.
Taxonomy
The following list covers all accepted taxa of Linder-
niaceae genera in alphabetical order. New combinations
and replacement names are made according to the new
circumscription of taxa in the new genus Linderniella
and in Bonnaya, Crate rostigma, Lindernia, Micranthe-
mum, Torenia and Vandellia. The species not represented
in the molecular analysis are assigned to genera accord-
ing to their morphology.
Fig. 5. Phylogram from Bayesian Inference of phylogeny with
branch lengths giving the relative substitution rates using the
GTR+G+I model. Taxa are named according to the taxonomic
treatment suggested in this study.
219Willdenowia 43 – 2013
Linderniaceae Borsch, Kai Müll. & Eb. Fisch. in Pl.
Biol. (Stuttgart) 7: 76. 2005 Lindernieae Rchb., Fl.
Germ. Excurs.: 385. 1831 1832. – Type: Lindernia All.
Artificial key to genera
1. Bracteoles present; perennial prostrate to ascending
subshrubs; corolla white, palate on lower lip present;
filaments twisted or curved; fruit reflexed; tropical
Africa ...................... 15. Stemodiopsis
Bracteoles absent; corolla not as above . . . . . . . . 2
2. Plants minute, prostrate to ascending; upper lip of co-
rolla absent or present, lobes of lower lip rounded,
entire; calyx 4-lobed to middle, except for lower side,
where cleft extends to base; or calyx 5-lobed; leaves
orbicular; Central America and Ca ribbean . . . . . . .
........................ 11. Micranthemum
Plants not as above, usually more robust . . . . . . . . 3
3. Seeds without alveolate endosperm, seed surface ±
smooth or only weakly furrowed, endosperm weakly
polygonal or undulate in transverse section . . . . . 4
Seeds with alveolate endosperm, seed surface with
rounded pits (bothrospermous) or longitudinal fur-
rows (aulacospermous), endosperm star-shaped in
transverse section ........................ 5
4. Aquatic plants with basal rosette, with 2-lobed com-
plex leaves, originating by connation of two adjacent
leaves of one whorl; cleistogamous flowers absent;
Namibia .................... 3. Chamaegigas
Plants of humid places or aquatic, if with basal rosette
then leaves simple, not 2-lobed; cleistogamous flow-
ers often present, prevailing in some species (e.g. L.
procumbens, L. benthamii, L. madagascariensis) . .
.............................. 9. Lindernia
5. Seeds with longitudinal furrows (aulacospermous);
all 4 stamens fertile . . . . . . . . . . . . . . . . . . . . . . . 6
Seeds with rounded pits (bothrospermous); some-
times 2 stamens reduced to staminodes . . . . . . . . 8
6. Plants procumbent, creeping; leaf venation pinnate;
flo wers in very lax racemes; calyx deeply divided,
with 5 linear-lanceolate lobes; Asia . . . . . . . . . . . .
.......................... 13. Schizotorenia
Plants erect or ascending (except Crepidorhopalon
scaettae, a creeper); leaf venation palmate; flowers
generally in dense racemes or in capitula, only occa-
sionally lax racemes; calyx with long tube and 5 short
teeth, if deeply divided then corolla less than 1 cm
long; Africa and Madagascar . . . . . . . . . . . . . . . . . 7
7. Lower lip of corolla with yellow club-shaped hairs on
a pluricellular base; leaves linear-lanceolate to ovate,
smooth; filaments of the abaxial stamens with a ba-
sal spur-like appendage, tip of spur rounded and with
glandular hairs; plants mostly annuals, only 2 species
perennial ................ 5. Crepidorhopalon
Lower lip of corolla lacking yellow club-shaped hairs
with pluricellular base; leaves largely ovoid, leathery,
somewhat shining; filament of abaxial stamens only
weakly curved with a small basal knee-like append-
age present or absent; perennial plants with large sub-
terraneous woody rhizomes on heavy metal soil . . .
............................... 6. Hartliella
8. Rosulate herbs; fertile stamens 2, one pair reduced to
filiform staminodes, anthers with 2 unequal mucro-
nate thecae or monothecous; Australia . . . . . . . . . .
........................... 7. Hemiarrhena
Rosulate or non-rosulate herbs, fertile stamens 2 or 4,
anthers equal (except Torenia sylvicola with unequal
but rounded anthers) . . . . . . . . . . . . . . . . . . . . . . . 9
9. Plants with a basal, decussate and distinct leaf rosette,
lacking the terminal main inflorescence; flowering
shoots all axillary ......................... 10
Plants without basal leaf rosette, or leaf rosette small,
inconspicuous, the stem with decussate leaves and of-
ten long internodes; terminal main inflorescence and
paracladia generally well developed . . . . . . . . . . 11
10. Leaves large, broadly ovoid to lanceolate, acuminate;
abaxial stamens perfect, with a Z-shaped filament;
roots of vivid red or orange colour . . . . . . . . . . . . .
...................... 4. Craterostigma s.str.
Leaves small, ovoid to lanceolate, obtuse; abaxial
stamina usually reduced to staminodes, roots whitish
........................ 4. Craterostigma s.l.
11. Plants from tropical Africa usually confined to rock
outcrops ................................ 12
Plants from Africa, Asia and South America on open
soil in savanna or rainforest, or as weeds . . . . . . 13
12. Robust plants, sometimes small subshrubs; stems
usually elongate with distant leaves, abaxial stamens
fertile (except C. niamniamense, C. pusillum with
abaxial staminodes) . . . . . . . . 4. Craterostigma s.l.
Small and delicate plants, usually annuals; basal
leaves either in small inconspicuous rosettes or dis-
tant; abaxial stamens usually reduced to staminodes,
rarely fertile (in L. bolusii, L. boutiqueana, L. brevi-
dens and L. cerastioides) ....... 10. Linderniella
13. Leaf venation palmate . . . . . . . . . . . . . . . . . . . . . 14
Leaf venation pinnate . . . . . . . . . . . . . . . . . . . . . . 15
14. Inflorescence capitate, enclosed by closely overlap-
ping large involucral leaves with semiorbicular base
and long acuminate apex, diering from linear-lan-
ceolate stem leaves . . . . . . . . . . . 12. Pierranthus
Inflorescence a lax raceme, rarely capitate (C. crassi-
folium), but then involucral leaves not diering from
stem leaves . . . . . . . . . . . . . . 4. Craterostigma s.l.
15. The 2 abaxial stamens reduced to staminodes . . . 16
All 4 stamens perfect . . . . . . . . . . . . . . . . . . . . . . 17
16. Leaves serrate or pinnatifid with aristate teeth, gener-
ally only midvein clearly visible; corolla with long (c.
15mm) weakly curved tube; staminodes mammiform
or obtuse at apex . . . . . . . . . . . . 14. Scolophyllum
Leaves entire or denticulate, rarely serrate, generally
lateral veins clearly visible; corolla with short tube
(less than 10mm, often less than 5mm); staminodes
clavate ......................... 2. Bonnaya
220 Fischer & al.: The phylogeny of Linderniaceae
17. Plants large; stem generally more than 50cm tall; co-
rolla more than 2 cm long, abaxial stamens with 2
boss-like geniculations . . . . . . . . . . . 1. Artanema
Plants much smaller, stem generally not more than
30 cm tall, if stem decumbent then occasionally to
50 cm long; corolla generally <2 cm long (but see
Torenia); abaxial stamens with 1 curved or spur-like
geniculation ............................ 18
18. Calyx suborbicular, with 3 broad wings between
ridges, auriculate at base, 3-dentate at apex . . . . . .
............................... 8. Legazpia
Calyx tubular, winged or wingless, if winged then
with 5 wings, 5-lobed . . . . . . . . . . . . . . . . . . . . . 19
19. Dehiscence of capsule poricidal, opening on both
sides of septum at same time as dehiscence of calyx;
ovary with apical and internal hairs . . . . . . . . . . . .
........................... 16. Torenia s.str.
Dehiscence of capsule septicidal; ovary without api-
cal hairs or hairs within locule . . . . . . . . . . . . . . 20
20. Fruit usually not exceeding length of calyx; leaves
usually ovate with distinctly pinnate venation . . . . .
........... 16. Torenia s.l. (T. crustacea group)
Fruit usually distinctly exceeding length of calyx;
leaves either ovate with distinctly pinnate venation or
linear to lanceolate, then often only midvein visible
.............................. 17. Vandellia
1. Artanema D. Don in Sweet, Brit. Fl. Gard. 6: t. 234.
1834, nom cons. Type: A. fimbriatum (Hook. ex Gra-
ham) D. Don.
1. Artanema angustifolium Benth., Scroph. Ind.: 39.
1835.
Distribution — India.
2. Artanema bantamense Backer in Bull. Jard. Bot. Bui-
tenzorg, ser. 2, 12: 37. 1913.
Distribution — Indonesia (Java).
3. Artanema fimbriatum (Hook. ex Graham) D. Don in
Sweet, Brit. Fl. Gard. 6: t. 234. 1834 Torenia fimbriata
Hook. ex Graham in Edinburgh New Philos. J. 1831: 379.
1831.
DistributionAustralia.
4. Artanema longifolium (L.) Vatke in Linnaea 43: 307.
1882 Columnea longifolia L., Syst. Nat., ed. 12, 2: 427;
Mant. Pl. 1: 90. 1767.
= Artanema sesamoides Benth., Scroph. Ind.: 39. 1835.
= Artanema cabrae De Wild. & T. Durand in Bull. Soc.
Roy. Bot. Belgique 38(1, Compt. Rend.): 131. 1899.
Distribution — Côte d’Ivoire, Ghana, Togo, Nigeria,
Cameroon, Gabon, Congo-Brazzaville, Central African
Republic, Congo-Kinshasa, Uganda, Tanzania, Sri Lan-
ka, India, Thailand, Cambodia, Vietnam, Malaysia, Indo-
nesia (Java, Sumatra), Borneo, Philippines.
2. Bonnaya Link & Otto, Icon. Pl. Select.: 25. 1821
Lindernia subg. Bonnaya (Link & Otto) T. Yamaz. –
Type: B. brachiata Link & Otto [= B. ciliata (Colsm.)
Spreng. subsp. ciliata].
1. Bonnaya aculeata (Bonati) Eb. Fisch., Schäferh. &
Kai Müll., comb. nov. Ilysanthes aculeata Bonati in
Bull. Soc. Bot. France 55: 541. 1908 Lindernia aculea-
ta (Bonati) T. Yamaz. in J. Jap. Bot. 53: 2. 1978.
Distribution — Cambodia, Laos.
2. Bonnaya antipoda (L.) Druce in Rep. Bot. Exch.
Club. Brit. Isles 3: 415. 1914 Ruellia antipoda L., Sp.
Pl. 2: 635. 1753 Ilysanthes antipoda (L.) Merr., Interpr.
Rumph. Herb. Amboin.: 467. 1917 Lindernia antipoda
(L.) Alston in Trimen, Handb. Fl. Ceyl. 6(Suppl., 2): 214.
1931 Vandellia antipoda (L.) T. Yamaz. in J. Jap. Bot.
30: 177. 1955.
= Lindernia verbenifolia (Colsm.) Pennell in Acad. Nat.
Sci. Philadelphia Mongr. 5: 31. 1943 Gratiola verbeni-
folia Colsm., Prodr. Descr. Gratiol.: 9. 1793.
Distribution Sri Lanka and India to China, Ja-
pan, Malaysia, Australia, New Guinea, Micronesia and
Polynesia; introduced in U.S.A. (Louisiana), Mexico and
Venezuela.
3. Bonnaya cephalantha (T. Yamaz.) Eb. Fisch., Schäf-
erh. & Kai Müll., comb. nov. Lindernia cephalantha T.
Yamaz. in J. Jap. Bot. 53: 1. 1978.
Distribution — Thailand.
4. Bonnaya ciliata (Colsm.) Spreng., Syst. Veg. 1: 41.
1824 Gratiola ciliata Colsm., Prodr. Descr. Gratiol.:
14. 1793 Lindernia ciliata (Colsm.) Pennell in Brittonia
2: 182. 1936 Ilysanthes ciliata (Colsm.) Kuntze, Revis.
Gen. Pl. 2: 461. 1891.
4.1. Bonnaya ciliata (Colsm.) Spreng. subsp. ciliata
= Gratiola serrata Roxb., Fl. Ind. 1: 140. 1820 Ilysan-
thes serrata (Roxb.) Urb. in Ber. Deutsch. Bot. Ges. 2:
436. 1884.
= Bonnaya brachiata Link & Otto, Icon. Pl. Select.: 25.
1821.
= Bonnaya bracteoides Blatt. & Hallb. in J. Bombay Nat.
Hist. Soc. 25: 416. 1918 Lindernia bracteoides (Blatt.
& Hallb.) Mukerjee in J. Ind. Bot. Soc. 24: 133. 1945.
Distribution — India, Nepal, Myanmar, S China,
Cambodia, Laos, Vietnam, Malaysia, Philippines, New
Guinea, Australia; introduced in U.S.A. (Florida).
4.2. Bonnaya ciliata subsp. sivarajanii (Tandyekk. & N.
Mohanan) Eb. Fisch., Schäferh. & Kai Müll., comb. nov.
Lindernia ciliata subsp. sivarajanii Tandyekk. & N.
Mohanan in Nordic. J. Bot. 28: 202. 2010.
Distribution — India.
5. Bonnaya cyrtotricha (P. C. Tsoong & T. C. Ku) Eb.
Fisch., Schäferh. & Kai Müll., comb. nov. Lindernia
cyrtotricha P. C. Tsoong & T. C. Ku in P. C. Tsoong &
H. P. Yang, Fl. Reipubl. Popularis Sin. 67(2): 398. 1979.
Distribution — China.
6. Bonnaya estaminodiosa Blatt. & Hallb. in J. Bombay
Nat. Hist. Soc. 25: 416. 1918 Lindernia estaminodiosa
(Blatt. & Hallb.) Mukerjee in J. Ind. Bot. Soc. 24: 133.
1945.
Distribution — India.
7. Bonnaya multiflora Bonati in Bull. Soc. Bot. France
55: 542. 1908 Vandellia multiflora (Bonati) T. Yamaz.
221Willdenowia 43 – 2013
in J. Jap. Bot. 30: 177. 1955, nom. illeg. [non Vandellia
multiflora (Roxb.) G. Don, Gen. Hist. 4: 549. 1837 1838]
Vandellia bonatii T. Yamaz. in Bull. Natl. Sci. Mus.,
Tokyo, n.s., 6: 371. 1963 Lindernia bonatii (T. Yamaz.)
Philcox in Taxon 19: 649. 1970 [non Lindernia multiflora
(Roxb.) Mukerjee in J. Ind. Bot. Soc. 24: 131. 1945].
= Ilysanthes viatica Kerr ex Barnett in Kew Bull. 16:
489. 1963 Lindernia viatica (Kerr ex Barnett) Philcox
in Taxon 19: 649. 1970.
Distribution Taiwan, Thailand, Cambodia, Vietnam.
8. Bonnaya oppositifolia (Retz.) Spreng., Syst. Veg. 1:
41. 1824 Gratiola oppositifolia Retz., Observ. Bot. 4:
8. 1786 Ilysanthes oppositifolia (Retz.) Urb. in Ber.
Deutsch. Bot. Ges. 2: 435. 1884 Vandellia oppositifolia
(Retz.) Haines, Bot. Bihar Orissa 4: 634. 1922 Linder-
nia oppositifolia (Retz.) Mukherjee in J. Ind. Bot. Soc.
24: 134. 1945.
Distribution — India.
9. Bonnaya ruellioides (Colsm.) Spreng., Syst. Veg. 1:
41. 1824 Gratiola ruellioides Colsm., Prodr. Descr.
Gratiol.: 12. 1793 Lindernia ruellioides (Colsm.) Pen-
nell in Brittonia 2: 182. 1936 Ilysanthes ruellioides
(Colsm.) Kuntze, Revis. Gen. Pl. 2: 462. 1891.
= Gratiola reptans Roxb., Fl. Ind. 1: 140. 1820 Bon-
naya reptans (Roxb.) Spreng., Syst. Veg. 1: 41. 1824
Lindernia reptans (Roxb.) F. Muell., Descr. Notes Pa-
puan Pl. 2(7): 31. 1886.
Distribution — India, Myanmar, S China, Cambodia,
Laos, Vietnam, Malaysia, Philippines, New Guinea.
10. Bonnaya succosa (Kerr ex Barnett) Eb. Fisch.,
Schäferh. & Kai Müll., comb. nov. Ilysanthes succosa
Kerr ex Barnett in Kew Bull. 16: 489. 1963 Lindernia
succosa (Kerr ex Barnett) Philcox in Taxon 19: 649. 1970.
Distribution — Thailand, Laos.
11. Bonnaya tenuifolia (Colsm.) Spreng., Syst. Veg.
1: 42. 1824 Gratiola tenuifolia Colsm., Prodr. Descr.
Gratiol.: 8. 1793 Vandellia tenuifolia (Colsm.) Haines,
Bot. Bihar Orissa 4: 634. 1922 Ilysanthes tenuifolia
(Colsm.) Urb. in Ber. Deutsch. Bot. Ges. 2: 435. 1884.
Distribution — Sri Lanka, India, S China, Cambodia,
Vietnam, Malaysia.
12. Bonnaya zanzibarica (Eb. Fisch. & Hepper) Eb.
Fisch., Schäferh. & Kai Müll., comb. nov. Lindernia
zan zibarica Eb. Fisch. & Hepper in Kew Bull. 46: 529.
1991.
Distribution Kenya, Tanzania (including Zanzi-
bar).
3. Chamaegigas Dinter, Ber. Tätigk. Naturw. Ges. Isis
1913 1915: 38. 1916 – Type: C. intrepidus Dinter.
1. Chamaegigas intrepidus Dinter, Ber. Tätigk. Naturw.
Ges. Isis 1913 1915: 38. 1916.
Distribution — Namibia.
4. Craterostigma Hochst. in Flora 24: 668. 1841. – Type
(designated by Phillips, Gen. S. Afr. Fl. Pl., ed. 2: 673.
1951): C. plantagineum Hochst.
= Strigina Engl. in Bot. Jahrb. Syst. 23: 516. 1897. –
Type: S. pusilla Engl. Craterostigma pusillum (Engl.)
Eb. Fisch., Schäferh. & Kai Müll.
Craterostigma plantagineum group
Mainly tropical African with two species in Arabia ex-
tending to India.
1. Craterostigma alatum Hepper in Kew Bull. 42: 945.
1987.
Distribution — Kenya, Tanzania.
2. Craterostigma hirsutum S. Moore in J. Bot. 38: 461.
1900.
Distribution Rwanda, Uganda, Kenya, Tanzania,
Malawi, Zimbabwe, Mozambique.
3. Craterostigma lanceolatum (Engl.) Skan in Oliver &
al., Fl. Trop. Afr. 4(2): 331. 1906 Craterostigma nanum
var. lanceo latum Engl., Pflanzenw. Ost-Afrikas C: 357.
1895.
= Craterostigma ndassekerense Engl. in Bot. Jahrb. Syst.
57: 611. 1922.
Distribution Congo-Kinshasa, Rwanda, Burundi,
Uganda, Kenya, Tanzania, Malawi, Zambia, Zimbabwe.
4. Craterostigma longicarpum Hepper in Kew Bull. 42:
945. 1987.
Distribution — Ethiopa, Uganda, Kenya, Tanzania.
5. Craterostigma plantagineum Hochst. in Flora 24:
669. 1841 Torenia plantaginea (Hochst.) Benth. in
Candolle, Prodr. 10: 411. 1846.
Distribution — Niger, Chad, Sudan, Ethiopia, Soma-
lia, Congo-Kinshasa, Rwanda, Burundi, Uganda, Kenya,
Tanzania, Angola, Zambia, Zimbabwe, Botswana, Na-
mibia, South Africa, Yemen, India.
6. Craterostigma pumilum Hochst. in Flora 24: 670.
1841 Torenia pumila (Hochst.) Benth. in Candolle,
Prodr. 10: 411. 1846.
Distribution — Ethiopia, Sudan, Somalia, Uganda,
Kenya, Tanzania, Zambia, Botswana, Yemen (including
Socotra), Saudi Arabia.
7. Craterostigma purpureum Lebrun & L. Touss. in
Bull. Jard. Bot. État Bruxelles 17: 83. 1943 Lindernia
purpurea (Lebrun & L. Touss.) R. Germ. in Bull. Jard.
Bot. État Bruxelles 22: 75. 1952.
Distribution — Congo-Kinshasa.
8. Craterostigma smithii S. Moore in J. Bot. 38: 461.
1900.
= Craterostigma boranense Chiov., Missione Biol. Bo-
rana 4: 206. 1939.
Distribution — Ethiopia, Kenya, Tanzania.
9. Craterostigma wilmsii Engl. ex Diels in Bot. Jahrb.
Syst. 26: 122. 1898.
Distribution — South Africa.
Craterostigma abyssinicum group
The following species appear within the Craterostigma
clade and are included here in a broadly defined genus
Craterostigma.
10. Craterostigma abyssinicum (Engl.) Eb. Fisch.,
Schäferh. & Kai Müll., comb. nov. Lindernia abyssin-
ica Engl. in Bot. Jahrb. Syst. 23: 503. 1897.
222 Fischer & al.: The phylogeny of Linderniaceae
Distribution — Nigeria, Cameroon, Sudan, Ethiopia,
Uganda, Kenya, Tanzania.
11. Craterostigma angolense (Skan) Eb. Fisch.,
Schäferh. & Kai Müll., comb. nov. Torenia angolen-
sis Skan in Oliver & al., Fl. Trop. Afr. 4(2): 335. 1906
Lindernia angolensis (Skan) Eb. Fisch. in Trop. Subtrop.
Pflanzenwelt 81: 254. 1992.
Distribution — Angola.
12. Craterostigma crassifolium Engl. in Bot. Jahrb. Syst.
23: 500. 1897 Lindernia crassifolia (Engl.) Eb. Fisch.
in Trop. Subtrop. Pflanzenwelt 81: 256. 1992.
Distribution — Angola.
13. Craterostigma engleri Eb. Fisch., Schäferh. & Kai
Müll., nom. nov. Ilysanthes welwitschii Engl. in Bot.
Jahrb. Syst. 23: 504. 1897 [non Craterostigma wel witschii
Engl. in Bot. Jahrb. Syst. 23: 501. 1897] Lindernia wel-
witschii (Engl.) Eb. Fisch. in Trop. Subtrop. Pflanzenwelt
81: 283. 1992.
Distribution Angola.
A replacement name is required because the name
Craterostigma welwitschii already exists.
14. Craterostigma gossweileri (S. Moore) Eb. Fisch.,
Schäferh. & Kai Müll., comb. nov. Ilysanthes goss-
weileri S. Moore in J. Bot. 57: 215. 1919 Lindernia
scapoidea Eb. Fisch. in Trop. Subtrop. Pflanzenwelt 81:
285. 1992 [non Lindernia gossweileri S. Moore in J. Bot.
45: 87. 1907].
Distribution Angola.
The original epithet gossweileri, which when com-
bined under Lindernia would form a homo nym of L. goss-
weileri and therefore required the replacement name L.
scapoidea, can now be maintained in Craterostigma.
15. Craterostigma kigomense (Eb. Fisch.) Eb. Fisch.,
Schäferh. & Kai Müll., comb. nov. Lindernia kigomen-
sis Eb. Fisch. in Bull. Jard. Bot. Natl. Belg. 67: 362. 1999.
Lindernia “species alpha” Philcox in Ghazanfar & al.,
Fl. Trop. E. Africa, Scrophular.: 82. 2008.
Distribution — Tanzania.
16. Craterostigma lindernioides E. A. Bruce in Bull.
Misc. Inform. Kew 1933: 474. 1933 Lindernia lindern-
ioides (E. A. Bruce ) Philcox in Ghazanfar & al., Fl. Trop.
E. Africa, Scrophular.: 76. 2008.
Distribution — Uganda, Tanzania.
17. Craterostigma newtonii (Engl.) Eb. Fisch., Schäferh.
& Kai Müll., comb. nov. Lindernia newtonii Engl. in
Bot. Jahrb. Syst. 23: 503. 1897.
= Vandellia lobelioides Oliv. in Trans. Linn. Soc. London
29: 120. 1875, nom. illeg. [non Vandellia lobelioides F.
Muell. in Trans. & Proc. Philos. Inst. Victoria 3: 61. 1859]
Lindernia lobelioides Wettst. in Engler & Prantl, Nat.
Pflanzenfam. 4(3b): 80. 1891, nom. illeg. [non Lindernia
lobelioides (F. Muell.) F. Muell., Syst. Census Austral.
Pl.: 97. 1882] Lindernia oliveriana Dandy in Andrews,
Fl. Pl. Sudan 3: 139. 1956.
DistributionTogo, Nigeria, Central African Repub-
lic, Congo-Kinshasa, Rwanda, Sudan, Ethiopia, Uganda,
Kenya, Tanzania, Zambia, Zimbabwe, Malawi, Angola.
The new combination is based on the earliest legitimate
name for the species, L. newtonii. Vandellia lobelioides
Oliv. and Lindernia lobelioides Wettst. are earlier names,
but both are later homonyms and therefore illegitimate.
18. Craterostigma niamniamense (Eb. Fisch. & Hepper)
Eb. Fisch., Schäferh. & Kai Müll., comb. nov. Linder-
nia niamniamensis Eb. Fisch. & Hepper in Kew Bull. 46:
534. 1991.
Distribution — Sudan, Uganda.
19. Craterostigma nummulariifolium (D. Don) Eb.
Fisch., Schäferh. & Kai Müll., comb. nov. Vandellia
num mulariifolia D. Don, Prodr. Fl. Nepal.: 86. 1825
Lindernia nummulariifolia (D. Don) Wettst. in Engler &
Prantl, Nat. Pflanzenfam. 4(3b): 79. 1891 Pyxidaria
nummulariifolia (D. Don) Kuntze, Revis. Gen. Pl. 2: 464.
1891.
= Vandellia chinensis T. Yamaz. in J. Jap. Bot. 30: 178.
1955 Lindernia chinensis (T. Yamaz.) Philcox in Taxon
19: 649. 1970.
= Mitranthus latifolius Hochst. in Flora 27: 108. 1844.
= Vandellia corymbosa Baker in J. Bot. 20: 221. 1882.
= Bryodes perrieri Bonati, Bull. Soc. Bot. Genève, ser.
2, 15: 104. 1924.
Distribution — Sierra Leone, Côte d’Ivoire, Nigeria,
Cameroon, Gabon, Sudan, Ethiopia, Congo-Kinshasa,
Rwanda, Burundi, Uganda, Kenya, Tanzania, Angola,
Zambia, Malawi, Zimbabwe, Madagascar; India, Nepal,
Myanmar, China, Thailand, Vietnam.
20. Craterostigma pusillum (Engl.) Eb. Fisch., Schäf erh.
& Kai Müll., comb. nov. Strigina pusilla Engl. in Bot.
Jahrb. Syst. 23: 516. 1897 Lindernia acicularis Eb.
Fisch. in Bull. Jard. Bot. Natl. Belg. 59: 446. 1989 [non
Lindernia pusilla (Willd.) Bold., Zakfl. Java: 165. 1916].
Distribution — Rwanda, Burundi, Tanzania.
Strigina was described by Engler as a new genus
closely related to Striga. However, it represents a close
relative of Lindernia and was subsequently transferred to
that genus (Fischer 1989; Bidgood 1992).
21. Craterostigma stuhlmannii (Engl.) Eb. Fisch.,
Schäferh. & Kai Müll., comb. nov. Lindernia stuhl-
mannii Engl., Pflanzenw. Ost-Afrikas C: 357. 1895.
Distribution — Tanzania.
22. Craterostigma sudanicum (Eb. Fisch. & Hepper)
Eb. Fisch., Schäferh. & Kai Müll., comb. nov. Linder-
nia sudanica Eb. Fisch. & Hepper in Kew Bull. 46: 530.
1991.
Distribution — Sudan, Uganda.
23. Craterostigma syncerus (Seine, Eb. Fisch. &
Barth lott) Eb. Fisch., Schäferh. & Kai Müll., comb. nov.
Lindernia syncerus Seine, Eb. Fisch. & Barthlott in
Feddes Repert. 106: 8. 1995.
Distribution — Zimbabwe.
24. Craterostigma tanzanicum Eb. Fisch., Schäferh. &
Kai Müll., nom. nov. Lindernia longicarpa Eb. Fisch.
& Hepper in Kew Bull. 46: 534. 1991 [non Craterostig-
ma longicarpum Hepper in Kew Bull. 42: 945. 1987].
Distribution — Tanzania.
223Willdenowia 43 – 2013
A replacement name is required because the name
Craterostigma longicarpum Hepper already exists.
25. Craterostigma yaundense (S. Moore) Eb. Fisch.,
Schäferh. & Kai Müll., comb. nov. Ilysanthes yaun-
densis S. Moore in J. Bot. 57: 216. 1919 Lindernia
yaundensis (S. Moore) Eb. Fisch. in Trop. Subtrop. Pflan-
zenwelt 81: 286. 1992.
Distribution — Cameroon.
5. Crepidorhopalon Eb. Fisch. in Feddes Repert. 100:
443. 1989. – Type: C. schweinfurthii (Oliv.) Eb. Fisch.
1. Crepidorhopalon anis (De Wild.) Eb. Fisch. ex Go-
vaerts, World Checkl. Seed Pl. 3(1): 20. 1999 Torenia
anis De Wild. in Ann. Mus. Congo Belge, Bot., ser. 4,
1: 122. 1903.
Distribution — Congo-Kinshasa.
2. Crepidorhopalon alatocalycinus Eb. Fisch. in Bull.
Jard. Bot. Natl. Belg. 59: 454. 1989.
Distribution — Congo-Kinshasa, Zambia.
3. Crepidorhopalon bifolius (Skan) Eb. Fisch. in Bull.
Jard. Bot. Natl. Belg. 60: 413. 1990 Lindernia bifolia
Skan in Oliver & al., Fl. Trop. Afr. 4(2): 343. 1906.
Distribution — Congo-Kinshasa, Tanzania, Zambia.
4. Crepidorhopalon chironioides (S. Moore) Eb. Fisch.
in Feddes Repert. 100: 444. 1989 Craterostigma chiro-
nioides S. Moore in J. Bot. 57: 215. 1919.
= Craterostigma kundalungense Engl. in Bot. Jahrb. Syst.
57: 612. 1922.
Distribution — Congo-Kinshasa.
5. Crepidorhopalon debilis (Skan) Eb. Fisch. in Feddes
Repert. 106: 8. 1995 Lindernia debilis Skan in Oliver
& al., Fl. Trop. Afr. 4(2): 344. 1906.
= Lindernia minima R. G. N. Young in Candollea 14: 9.
1952, nom. illeg. [non Lindernia minima (Benth.) Muk-
erjee in J. Ind. Bot. Soc. 24: 132. 1945] Lindernia fugax
R. G. N. Young in Candollea 15: 125. 1956.
Distribution — Senegal, Côte d’Ivoire, Ghana, Ni-
geria, Cameroon, Sudan, Congo-Kinshasa, Uganda, Tan-
zania, Zimbabwe, South Africa.
6. Crepidorhopalon goetzei (Engl.) Eb. Fisch. in Feddes
Repert. 100: 444. 1989 Craterostigma goetzei Engl. in
Bot. Jahrb. Syst. 28: 477. 1900 Torenia goetzei (Engl.)
Hepper in Bol. Soc. Brot., ser. 2, 60: 271. 1987.
= Torenia brevifolia Engl. & Pilg. in Bot. Jahrb. Syst. 45:
214. 1910.
Distribution Congo-Kinshasa, Burundi, Tanzania,
Zam bia, Malawi.
7. Crepidorhopalon gracilis (Pilg.) Eb. Fisch. in Fed-
des Repert. 100: 443. 1989 Craterostigma gracile Pilg.
in Bot Jahrb. Syst. 45: 213. 1910 Torenia ledermannii
Hepper in Bol. Soc. Brot., ser. 2, 60: 271. 1987.
= Craterostigma guineense Hepper in Kew Bull. 14: 407.
1960.
Distribution Senegal, Mali, Guinea-Bissau, Gui-
nea, Cameroon, Congo-Kinshasa, Tanzania, Zambia.
8. Crepidorhopalon hartlii Eb. Fisch. in Bull. Jard. Bot.
Natl. Belg. 59: 453. 1989.
Distribution — Congo-Kinshasa.
9. Crepidorhopalon hepperi Eb. Fisch. in Bull. Jard.
Bot. Natl. Belg. 60: 410. 1990 Lindernia hepperi (Eb.
Fisch.) Philcox in Ghazanfar & al., Fl. Trop. E. Africa,
Scrophular.: 72. 2008.
Distribution — Kenya, Tanzania.
10. Crepidorhopalon involucratus (Philcox) Eb. Fisch. in
Trop. Subtrop. Pflanzenwelt 81: 197. 1992 Torenia in-
volucrata Philcox in Bol. Soc. Brot., ser. 2, 60: 267. 1987.
Distribution — Zambia.
11. Crepidorhopalon latibracteatus (Skan) Eb. Fisch. in
Feddes Repert. 100: 444. 1989 Craterostigma latibrac-
teatum Skan in Oliver & al., Fl. Trop. Afr. 4(2): 333. 1906
Torenia latibracteata (Skan) Hepper in Bol. Soc. Brot.,
ser. 2, 60: 271. 1987.
Distribution — Congo-Brazzaville, Congo-Kinshasa.
12. Crepidorhopalon laxiflorus Eb. Fisch. in Bull. Jard.
Bot. Natl. Belg. 60: 410. 1990.
Distribution — Congo-Kinshasa.
13. Crepidorhopalon malaissei Eb. Fisch. in Bull. Jard.
Bot. Natl. Belg. 59: 450. 1989.
Distribution — Congo-Kinshasa.
14. Crepidorhopalon manganicolus Eb. Fisch. in Bull.
Jard. Bot. Natl. Belg. 59: 455. 1989.
Distribution — Congo-Kinshasa, Angola, Zambia.
15. Crepidorhopalon membranocalycinus Eb. Fisch. in
Feddes Repert. 100: 444. 1989.
Distribution — Congo-Kinshasa.
16. Crepidorhopalon microcarpaeoides (Bonati) Eb.
Fisch. in Bull. Mus. Natl. Hist. Nat., B, Adansonia, ser.
4, 17: 252. 1995 Lindernia microcarpaeoides Bonati in
Bull. Soc. Bot. Genève, ser. 2, 15: 101. 1924.
Distribution — Madagascar.
17. Crepidorhopalon parviflorus (Philcox) Eb. Fisch.
in Bull. Jard. Bot. Natl. Belg. 67: 378. 1999 Torenia
latibracteata subsp. parviflora Philcox in Bol. Soc. Brot.,
ser. 2, 60: 267. 1987 Crepidorhopalon latibracteatus
subsp. parviflorus (Philcox) Eb. Fisch. in Trop. Subtrop.
Pflanzenwelt 81: 155. 1992.
Distribution Burundi, Kenya, Tanzania, Zambia,
Mozambique.
18. Crepidorhopalon perennis (P. A. Duvign.) Eb. Fisch.
in Bull. Jard. Bot. Natl. Belg. 60: 413. 1990 Lindernia
perennis P. A. Duvign. in Bull. Soc. Roy. Bot. Belgique
90: 256. 1958.
Distribution — Congo-Kinshasa.
19. Crepidorhopalon robynsii Eb. Fisch. in Bull. Jard.
Bot. Natl. Belg. 67: 371. 1999.
Distribution — Congo-Kinshasa.
20. Crepidorhopalon rupestris (Engl.) Eb. Fisch. in Bull.
Jard. Bot. Natl. Belg. 67: 376. 1999 Lindernia rupestris
Engl. in Bot. Jahrb. Syst. 30: 402. 1901.
= Crepidorhopalon insularis (Skan) Eb. Fisch. in Bull.
Jard. Bot. Natl. Belg. 59: 457. 1989 Lindernia insularis
Skan in Oliver & al., Fl. Trop. Afr. 4(2): 342. 1906.
= Lindernia subscaposa Mildbr. in Notizbl. Bot. Gart.
Berlin-Dahlem 8: 233. 1922.
224 Fischer & al.: The phylogeny of Linderniaceae
Distribution Congo-Kinshasa, Burundi, Uganda,
Kenya, Tanzania, Zambia, Malawi, Zimbabwe.
21. Crepidorhopalon scaettae (Staner) Eb. Fisch. in
Bull. Jard. Bot. Natl. Belg. 66: 78. 1997 Stemodiopsis
scaettae Staner in Rev. Zool. Bot. Africaines 24: 220.
1933.
Distribution — Congo-Kinshasa.
22. Crepidorhopalon schweinfurthii (Oliv.) Eb. Fisch. in
Trop. Subtrop. Pflanzenwelt 81: 130. 1992 Torenia sch-
weinfurthii Oliv. in Hooker’s Icon. Pl. 13: t. 1256. 1878
Craterostigma schweinfurthii (Oliv.) Engl. in Bot. Jahrb.
Syst. 23: 501. 1897.
Distribution — Mali, Côte d’Ivoire, Ghana, Togo,
Benin, Nigeria, Cameroon, Central African Republic,
Sudan, Congo-Kinshasa, Uganda, Tanzania, Angola,
Zambia, Mozambique.
23. Crepidorhopalon spicatus (Engl.) Eb. Fisch. in Fed-
des Repert. 100: 444. 1989 Torenia spicata Engl. in
Bot. Jahrb. Syst. 23: 502. 1897.
= Torenia inaequalifolia Engl. in Bot. Jahrb. Syst. 23:
502. 1897.
= Lindernia dinteri Schinz in Mém. Herb. Boissier 20:
27. 1900.
Distribution — Senegal, Mali, Guinea, Côte d’Ivoire,
Burkina Faso, Ghana, Togo, Nigeria, Cameroon, Chad,
Sudan, Congo-Kinshasa, Tanzania, Angola, Zambia, Ma-
lawi, Zimbabwe, Botswana, Namibie, South Africa.
24. Crepidorhopalon symoensii Eb. Fisch. in Bull. Jard.
Bot. Natl. Belg. 59: 451. 1989.
Distribution — Congo-Kinshasa.
25. Crepidorhopalon tanzanicus Eb. Fisch. in Bull. Jard.
Bot. Natl. Belg. 67: 374. 1999.
Distribution — Tanzania.
26. Crepidorhopalon tenuifolius (Philcox) Eb. Fisch. in
Trop. Subtrop. Pflanzenwelt 81: 141. 1992 Torenia ten-
uifolia Philcox in Bol. Soc. Brot., ser. 2, 60: 267. 1987.
Distribution — Congo-Kinshasa, Burundi, Zambia.
27. Crepidorhopalon tenuis (S. Moore) Eb. Fisch. in
Bull. Jard. Bot. Natl. Belg. 59: 457. 1989 Lindernia
tenuis S. Moore in J. Bot. 56: 10. 1918.
= Lindernia damblonii P. A. Duvign., Bull. Soc. Roy. Bot.
Belgique 90: 256. 1958 Crepidorhopalon damblonii (P.
A. Duvign.) Eb. Fisch. in Feddes Repert. 100: 444. 1989.
Distribution — Congo-Kinshasa, Burundi, Tanzania,
Zambia.
28. Crepidorhopalon uvens (Hiern) Eb. Fisch. in Bull.
Jard. Bot. Natl. Belg. 60: 413. 1990 Lindernia uvens
Hiern, Cat. Afr. Pl. 1: 763. 1898.
Distribution — Cameroon, Congo-Kinshasa, Angola,
Zambia.
29. Crepidorhopalon welwitschii (Engl.) Eb. Fisch. in
Feddes Repert. 100: 443. 1989 Craterostigma wel-
witschii Engl. in Bot. Jahrb. Syst. 23: 501. 1897.
DistributionAngola.
30. Crepidorhopalon whytei (Skan) Eb. Fisch. in Feddes
Repert. 100: 444. 1989 Lindernia whytei Skan in Oliver
& al., Fl. Trop. Afr. 4(2): 340. 1906.
= Lindernia gossweileri S. Moore in J. Bot. 45: 87. 1907.
= Torenia mildbraedii Pilg. in Mildbraed, Wiss. Erg.
Deut. Zentr.-Afr. Exped., Bot. 2: 285. 1911.
Distribution Ethiopia, Congo-Kinshasa, Rwanda,
Burundi, Sudan, Uganda, Kenya, Tanzania, Zimbabwe,
Angola.
6. Hartliella Eb. Fisch. in Trop. Subtrop. Pflanzenwelt
81: 204. 1992. – Type: H. suruticosa (Lisowski & Miel-
carek) Eb. Fisch.
The genus has aulacospermous seeds and is probably
closely related to Crepidorhopalon.
1. Hartliella bampsii (Eb. Fisch.) Eb. Fisch. in Trop.
Subtrop. Pflanzenwelt 81: 209. 1992 Lindernia bamp-
sii Eb. Fisch. in Bull. Jard. Bot. Natl. Belg. 59: 449.
1989.
DistributionCongo-Kinshasa.
2. Hartliella capitata (Eb. Fisch.) Eb. Fisch. in Trop.
Subtrop. Pflanzenwelt 81: 211. 1992 Lindernia capita-
ta Eb. Fisch. in Bull. Jard. Bot. Natl. Belg. 59: 448. 1989.
DistributionCongo-Kinshasa, Zambia.
3. Hartliella cupricola Eb. Fisch. in Trop. Subtrop. Pflan-
zenwelt 81: 207. 1992.
DistributionCongo-Kinshasa.
4. Hartliella suffruticosa (Lisowski & Mielcarek) Eb.
Fisch. in Trop. Subtrop. Pflanzenwelt 81: 204. 1992
Lindernia suffruticosa Lisowski & Mielcarek, Bull. Jard.
Bot. Natl. Belg. 54: 127. 1984.
DistributionCongo-Kinshasa.
7. Hemiarrhena Benth., Fl. Austral. 4: 518. 1868. –
Type: H. plantaginea (F. Muell.) Benth.
The genus has bothrospermous seeds and shows rela-
tionships to Bonnaya. The abaxial stamens are reduced to
staminodes and the leaves show a pinnate venation.
1. Hemiarrhena plantaginea (F. Muell.) Benth., Fl. Aus-
tral. 4: 518. 1868 Vandellia plantaginea F. Muell. in
Trans. & Proc. Philos. Inst. Victoria 3: 62. 1859 Linder-
nia plantaginea (F. Muell.) F. Muell., Fragm. 6: 102. 1868.
Distribution — Tropical NW Australia.
8. Legazpia Blanco, Fl. Filip., ed. 2: 338. 1845. – Type: L.
triptera Blanco [= L. polygonoides (Benth.) T. Yamaz.].
1. Legazpia polygonoides (Benth.) T. Yamaz. in J. Jap.
Bot. 30: 359. 1955 Torenia polygonoides Benth., Scro-
ph. Ind.: 39. 1835.
= Legazpia mucronulata (Benth.) T. Yamaz. in J. Jap.
Bot. 30: 359. 1955 Torenia mucronulata Benth. in Can-
dolle, Prodr. 10: 409. 1846.
= Legazpia triptera Blanco, Fl. Filip., ed. 2: 339. 1845.
Distribution — Cambodia, Laos, Vietnam, Thailand,
Micronesia, New Guinea.
Legazpia polygonoides has bothrospermous seeds. It
is closely related to Torenia, but diers in the suborbicu-
lar calyx with 3 broad wings between the ridges, auricu-
late at base and 3-dentate at apex (Fischer 2004). Torenia
has a calyx with 5 wings.
225Willdenowia 43 – 2013
9. Lindernia All. in Mélanges Philos. Math. Soc. Roy.
Turin 3(1): 178. 1766 Pyxidaria Kuntze, Revis. Gen.
Pl. 2: 464. 1891. – Type: L. palustris Hartmann [= L.
procumbens (Krock.) Philcox].
= Ilysanthes Raf., Ann. Nat.: 13. 1820. – Type: I. riparia
Raf. [= Lindernia dubia (L.) Pennell].
= Bryodes Benth. in Candolle, Prodr. 10: 433. 1846. –
Type: B. micrantha Benth. Lindernia benthamii Eb.
Fisch., Schäferh. & Kai Müll.
= Psammetes Hepper in Hooker’s Icon. Pl. 36: t. 3582.
1962. – Type: P. nigerica Hepper [= Lindernia madagas-
cariensis (Bonati) Eb. Fisch., Schäferh. & Kai Müll.].
1. Lindernia alsinoides R. Br., Prodr. Fl. Nov. Holland.:
441. 1810 Tittmannia alsinoides (R. Br.) Spreng., Syst.
Veget. 2. 1825 Ilyogeton alsinoides (R. Br.) Endl. in
Repert. Bot. Syst. 3: 297. 1844 Vandellia alsinoides
(R. Br.) Benth., Prodr. (DC) 10: 415. 1846 Pyxidaria
alsinoides (R. Br.) Kuntze, Rev. Gen. Pl. 2: 464. 1891.
= Ilyogeton subulatum Endl. in Repert. Bot. Syst. 3: 297.
1844.
DistributionAustralia.
2. Lindernia alterniflora (C. Wright) Alain in Contr.
Ocas. Mus. Hist. Nat. Colegio “De La Salle” 15: 13.
1956 Bonnaya alterniflora C. Wright in Anales Acad.
Ci. Med. Habana 7: 40. 1870 Ilysanthes alterniflora
(C. Wright) Urb. in Ber. Deutsch. Bot. Ges. 2: 436. 1884.
Distribution — Cuba.
3. Lindernia benthamii Eb. Fisch., Schäferh. & Kai
Müll., nom. nov. Bryodes micrantha Benth. in Can-
dolle, Prodr. 10: 433. 1846 [non Lindernia micrantha D.
Don, Prodr. Fl. Nepal.: 85. 1825].
Distribution — Madagascar.
The monotypic genus Bryodes (Fischer & Hepper
1997) is nested within Lindernia. A replacement name is
required because the name Lindernia micrantha already
exists.
4. Lindernia brachyphylla Pennell in Field iana, Bot. 28:
519. 1953.
= Lindernia barrosorum L. B. Sm. in Los Angeles Coun-
ty Mus. Contr. Sci. 23: 5. 1958.
Distribution — Venezuela, Guyana, Brazil.
5. Lindernia bryoides Eb. Fisch. in Bull. Mus. Natl. Hist.
Nat., B, Adansonia, ser. 4, 17: 248. 1995.
Distribution — Madagascar.
6. Lindernia capensis Thunb., Prodr. Pl. Cap.: 104. 1800
Ilysanthes capensis (Thunb.) Benth. in Candolle, Prodr.
10: 419. 1846.
Distribution — South Africa.
7. Lindernia conferta (Hiern) Philcox, Bol. Soc. Brot.,
ser. 2, 60: 268. 1987 Ilysanthes conferta Hiern in Har-
vey, Fl. Cap. 4(2): 365. 1904.
= Ilysanthes plantaginella S. Moore in J. Bot. 43: 49. 1905.
Distribution Congo-Kinshasa, Zimbabwe, South
Africa.
8. Lindernia congesta (A. Raynal) Eb. Fisch. in Trop.
Subtrop. Pflanzenwelt 81: 317. 1992 Ilysanthes con-
gesta A. Raynal in Adansonia, n.s, 7: 348. 1967.
Distribution — Senegal (Cap-Vert), Tanzania.
9. Lindernia dubia (L.) Pennell in Acad. Nat. Sci. Phila-
delphia Monogr. 1: 141. 1935 Gratiola dubia L., Sp.
Pl. 1: 17. 1753 Capraria gratioloides L., Syst. Nat.,
ed. 10, 2: 1117. 1759, nom. illeg. Lindernia pyxidaria
L., Mant. Pl. 2: 252. 1771, nom. illeg. Ilysanthes grati-
oloides Benth. in Candolle, Prodr. 10: 419. 1846, nom.
illeg. Lindernia gratioloides J. Lloyd & Foucaud, Fl.
Ouest France, ed. 4: 246. 1886, nom. illeg. Limnophi-
la dubia (L.) M. R. Almeida, Fl. Maharashtra 3B: 393.
2001, nom. illeg. [non Limnophila dubia Bonati in Bull.
Soc. Bot. France 55: 511. 1908].
= Lindernia dubia var. riparia (Raf.) Fernald in Rhodora
44: 444. 1942 Ilysanthes riparia Raf. in Ann. Nat.: 13.
1820.
= Lindernia anagallidea (Michx.) Pennell in Acad. Nat.
Sci. Philadelphia Monogr. 1: 152. 1935 Gratiola ana-
gallidea Michx., Fl. Bor.-Amer. (Michaux) 1: 6. 1803.
= Lindernia dilatata Muhl. ex Elliott, Sketch Bot. S.
Carolina 1: 16. 1816.
Distribution Canada, U.S.A., Cuba, Dominican
Republic, Puerto Rico, Mexico, Belize, Guatemala, Hon-
duras, Nicaragua, Panama, Argentina, Bolivia, Brazil,
Chile, Colombia, Ecuador, Paraguay, Peru, Venezuela;
introduced in Europe and E Asia.
10. Lindernia grandiflora Nutt., Gen. N. Amer. Pl. 2: 43.
1818 Ilysanthes grandiflora (Nutt.) Benth. in Candolle,
Prodr. 10: 418. 1846.
= Bazina nudiflora Raf., Autik. Bot.: 45. 1840.
Distribution — U.S.A. (Florida).
11. Lindernia hyssopioides (L.) Haines, Bot. Bihar Oris-
sa 4: 635. 1922 Gratiola hyssopioides L., Mant. Pl. 2:
174. 1771 Ilysanthes hyssopioides (L.) Benth. in Can-
dolle, Prodr. 10: 419. 1846.
Distribution Sri Lanka, India, S China, Vietnam,
Indonesia (Java, Sumatra).
12. Lindernia jiuhuanica X. H. Guo & X. L. Liu in Acta
Phytotax. Sin. 26: 153. 1988.
Distribution — China.
13. Lindernia lemuriana Eb. Fisch., Schäferh. & Kai
Müll., nom. nov. Ilysanthes micrantha Bonati in Bull.
Soc. Bot. Genève, ser. 2, 18: 10. 1926, nom. illeg. [non
Ilysanthes micrantha S. Moore in J. Bot. 49: 158. 1911,
nec Lindernia micrantha D. Don, Prodr. Fl. Nepal.: 85.
1825] Lindernia bonatii Eb. Fisch. in Bull. Mus. Natl.
Hist. Nat., B, Adansonia, ser. 4, 17: 242. 1995, nom. il-
leg. [non Lindernia bonatii (T. Yamaz.) Philcox in Taxon
19: 649. 1970].
Distribution — Madagascar.
Lindernia bonatii was published as a replacement
name because Ilysanthes micrantha is a later homonym,
and therefore illegitimate, and the name L. micrantha al-
ready exists. However, because L. bonatii is also a later
homonym, a new replacement name is required.
14. Lindernia linearifolia (Engl.) Eb. Fisch. in Trop.
Subtrop. Pflanzenwelt 81: 319. 1992 Craterostigma
linearifolium Engl. in Bot. Jahrb. Syst. 23: 501. 1897.
226 Fischer & al.: The phylogeny of Linderniaceae
Distribution — Angola.
15. Lindernia madagascariensis (Bonati) Eb. Fisch.,
Schäferh. & Kai Müll., comb. nov. Bryodes madagas-
cariensis Bonati in Bull. Soc. Bot. Genève, ser. 2, 15:
103. 1924 Psammetes madagascariensis (Bonati) Eb.
Fischer & Hepper in Kew Bull. 52: 750. 1997.
= Psammetes nigerica Hepper in Hooker’s Icon. Pl. 36:
t. 3582. 1962.
Distribution — Nigeria, Madagascar.
The monotypic Psammetes is nested within Lindernia.
16. Lindernia madayiparensis Rateesh, Sunil & Nanda-
kumar in Int. J. Pl. Anim. Environ. Sci. 2(3): 59. 2012.
Distribution — India.
17. Lindernia manilaliana Sivar. in Kew Bull. 31: 151.
1976.
Distribution — India.
18. Lindernia microcalyx Pennell & Stehlé in Stehlé &
al., Fl. Guadeloupe 2(1): 217. 1938.
Distribution — Guadeloupe, Jamaica, Martinique.
Lewis (2000) placed this species into the synonymy
of Lindernia rotundifolia. However, our analysis shows
that it diers from typical L. rotundifolia (Fig. 4, 5) and
it is therefore reinstated here. Probably L. rotundifolia s.l.
comprises several taxa, and a critical revision of the com-
plex is required.
19. Lindernia minima (Benth.) Mukerjee in J. Ind. Bot.
Soc. 24: 132. 1945 Ilysanthes minima Benth. in Can-
dolle, Prodr. 10: 420. 1846.
Distribution — India.
20. Lindernia monroi (S. Moore) Eb. Fisch. in Trop. Sub-
trop. Pflanzenwelt 81: 322. 1992 Craterostigma monroi
S. Moore in J. Bot. 57: 214. 1919 Torenia monroi (S.
Moore) Philcox in Bol. Soc. Brot., ser. 2, 60: 267. 1987.
= Lindernia dongolensis E. A. Bruce in Bothalia 6: 235.
1951.
Distribution — Zimbabwe, Angola, South Africa.
21. Lindernia monticola Nutt., Gen. N. Amer. Pl. 2: Ad-
dit. [1]. 1818 Ilysanthes monticola (Nutt.) Raf., Autik.
Bot.: 45. 1840.
= Lindernia refracta Elliott, Sketch Bot. S. Carolina 1:
579. 1821 Ilysanthes refracta (Elliott) Raf., Autik. Bot.:
46. 1840.
= Lindernia saxicola M. A. Curtis in Amer. J. Sci. 44:
83. 1843 Ilysanthes saxicola (M. A. Curtis) Chapm.,
Fl. South. U.S.: 294. 1860 Ilysanthes refracta var. saxi-
cola (M. A. Curtis) A. Gray, Syn. Fl. N. Amer. 2(1): 283.
1878.
Distribution — SE U.S.A.
22. Lindernia natans Eb. Fisch. in Bull. Mus. Natl. Hist.
Nat., B, Adansonia, ser. 4, 17: 250. 1995.
Distribution — Madagascar.
23. Lindernia paludosa (Bonati) Eb. Fisch. in Bull. Mus.
Natl. Hist. Nat., B, Adansonia, ser. 4, 17: 245. 1995 Ily-
santhes paludosa Bonati in Bull. Soc. Bot. Genève, ser.
2, 18: 5. 1926.
= Ilysanthes longipes Bonati in Bull. Soc. Bot. Genève,
ser. 2, 18: 7. 1926.
= Ilysanthes macrantha Bonati in Bull. Soc. Bot. Genève,
ser. 2, 18: 6. 1926.
= Ilysanthes macrophylla Bonati in Bull. Soc. Bot.
Genève, ser. 2, 18: 8. 1926.
= Ilysanthes perrieri Bonati in Bull. Soc. Bot. Genève,
ser. 2, 18: 11. 1926.
Distribution — Madagascar.
24. Lindernia parviflora (Roxb.) Haines, Bot. Bihar
Orissa 4: 635. 1922 Gratiola parviflora Roxb., Pl.
Coromandel 3: 3. 1811 Ilysanthes parviflora (Roxb.)
Benth. in Candolle, Prodr. 10: 419. 1846.
= Ilysanthes radicans Pilg. in Mildbraed, Wiss. Erg.
Deut. Zentr.-Afr. Exped., Bot. 2: 286. 1911.
Distribution — Senegal, Mali, Ghana, Nigeria, Gabon,
Chad, Sudan, Ethiopia, Somalia, Congo-Kinshasa, Rwan-
da, Burundi, Uganda, Kenya, Tanzania, Zambia, Malawi,
Zimbabwe, Mozambique, Botswana, Namibia, South Af-
rica, Madagascar, Mauritius, India, Sri Lanka, Vietnam.
25. Lindernia procumbens (Krock.) Philcox in Taxon
14: 30. 1965 Anagalloides procumbens Krock., Fl.
Siles. 2(1): 398. 1790 Pyxidaria procumbens (Krock.)
Borbás, Békésvármegye Fl.: 80. 1881.
= Lindernia palustris Hartmann, Primae Lin. Inst. Bot.,
ed. 2: 77. 1767, nom. utique rej.
= Vandellia erecta Benth., Scroph. Ind.: 36. 1835.
Distribution — Europe, E to SE Asia.
26. Lindernia rotundata (Pilg.) Eb. Fisch. in Bull. Jard.
Bot. Natl. Belg. 67: 366. 1999 Ilysanthes rotundata
Pilg. in Bot. Jahrb. Syst. 45: 214. 1910.
DistributionNigeria, Cameroon, Ethiopia, Congo-
Kinshasa, Rwanda, Uganda, Tanzania, Malawi.
27. Lindernia rotundifolia (L.) Alston in Trimen, Handb.
Fl. Ceyl. 6(Suppl., 2): 214. 1931 Gratiola rotundifolia
L., Mant. Pl. 2: 174. 1771 Ilysanthes rotundifolia (L.)
Benth. in Candolle, Prodr. 10: 420. 1846.
= Ilysanthes oblongifolia Baker in J. Bot. 20: 221. 1882.
= Ilysanthes hypericifolia Bonati in Bull. Soc. Bot.
Genève, ser. 2, 18: 10. 1926.
= Ilysanthes madagascariensis Bonati in Bull. Soc. Bot.
Genève, ser. 2, 18: 12. 1926.
Distribution Belize, Guatemala, Honduras, Nica-
ragua, Brazil, Colombia, French Guiana, Paraguay, Peru,
E Africa, Madagascar, Sri Lanka, India, China.
28. Lindernia srilankana L. H. Cramer & Philcox in
Ceylon J. Sci., Biol. Sci. 11: 35. 1974.
Distribution — Sri Lanka.
29. Lindernia tridentata (Small) D. Q. Lewis in Casta-
nea 65: 105. 2000 Ilysanthes tridentata Small in Bull.
Torrey Bot. Club 23: 297. 1896.
Distribution — Mexico.
30. Lindernia viguieri (Bonati) Eb. Fisch. in Bull. Mus.
Natl. Hist. Nat., B, Adansonia, ser. 4, 17: 240. 1995
Ilysanthes viguieri Bonati in Bull. Soc. Bot. Genève, ser.
2. 18: 3. 1926.
= Ilysanthes pseudoviguieri Bonati in Bull. Soc. Bot.
Genève, ser. 2. 18: 4. 1926.
Distribution — Madagascar.
227Willdenowia 43 – 2013
10. Linderniella Eb. Fisch., Schäferh. & Kai Müll., gen.
nov.
Plantae rosulatae vel caulibus elongatis instructae, folia
opposita nervibus palmatis, flores staminibus fertilibus
2 et staminodiis geniculatis abaxialibus instructi, raro
staminibus fertilibus 4 (solo in L. boutiqueana, L. brevi-
dente et L. cerastioidi), semina bothrosperma. – Typus:
L. pygmaea (Bonati) Eb. Fisch., Schäferh. & Kai Müll.
The new genus covers mainly rosulate rupicolous
species; some of them, however, also have elongated
stems. Except for Linderniella boutiqueana, L. brevidens
and L. cerastioides the abaxial stamens are reduced to
staminodes or bear only minute, reduced anthers (in L.
horombensis and L. pygmaea). The seeds are bothrosper-
mous. Linderniella brevidens is the only species that is
not restricted to rock outcrops.
1. Linderniella andongensis (Hiern) Eb. Fisch., Schäferh.
& Kai Müll., comb. nov. Ilysanthes andongensis Hiern,
Cat. Afr. Pl. 1: 765. 1898 Lindernia andongensis (Hi-
ern) Eb. Fisch. in Trop. Subtrop. Pflanzenwelt 81: 304.
1992, comb. inval.
DistributionAngola.
2. Linderniella bolusii (Hiern) Eb. Fisch., Schäferh.
& Kai Müll., comb. nov. Ilysanthes bolusii Hiern in
Harvey, Fl. Cap. 4(2): 367. 1904 Lindernia bolusii
(Hiern) Eb. Fisch. in Trop. Subtrop. Pflanzenwelt 81:
261. 1992.
DistributionEthiopia, Uganda, Kenya, South Af-
rica.
3. Linderniella boutiqueana (Germain) Eb. Fisch.,
Schäferh. & Kai Müll., comb. nov. Lindernia bou-
tiqueana R. Germ. in Bull. Jard. Bot. État. Bruxelles 22:
74. 1952.
Distribution — Burundi.
4. Linderniella brevidens (Skan) Eb. Fisch., Schäferh. &
Kai Müll., comb. nov. Lindernia brevidens Skan in Ol-
iver & al., Fl. Trop. Afr. 4(2): 339. 1906.
Distribution — Kenya, Tanzania.
5. Linderniella cerastioides (Bonati) Eb. Fisch., Schäf-
erh. & Kai Müll., comb. nov. Craterostigma cerastio-
ides Bonati in Bull. Soc. Bot. Genève, ser. 2, 15: 106.
1924 Lindernia andringitrae Eb. Fisch. in Bull. Mus.
Natl. Hist. Nat., B, Adansonia, ser. 4, 17: 236. 1995 [non
Lindernia cerastioides T. Yamaz., J. Jap. Bot. 53: 97.
1978].
Distribution — Madagascar.
6. Linderniella gracilis (Skan) Eb. Fisch., Schäferh. &
Kai Müll., comb. nov. Ilysanthes gracilis Skan in Ol-
iver & al., Fl. Trop. Afr. 4(2): 349. 1906 Lindernia exi-
lis Philcox in Bol. Soc. Brot., ser. 2, 60: 269. 1987 [non
Lindernia gracilis (Bonati) Bonati, Fl. Gén. Indoch. 4:
416. 1927].
Distribution Côte d’Ivoire, Sierra Leone, Ghana,
Nigeria, Central African Republic, Sudan, Congo-Kin-
shasa, Rwanda, Burundi.
7. Linderniella hartlii (Eb. Fisch. & Hepper) Eb. Fisch.,
Schäferh. & Kai Müll., comb. nov. Lindernia hartlii
Eb. Fisch. & Hepper in Bull. Jard. Bot. Natl. Belg. 59:
447. 1989.
Distribution — Congo-Kinshasa, Tanzania.
8. Linderniella horombensis (Eb. Fisch.) Eb. Fisch.,
Schäferh. & Kai Müll., comb. nov. Lindernia ho rom-
bensis Eb. Fisch. in Bull. Mus. Natl. Hist. Nat., B, Adan-
sonia, ser. 4, 17: 238. 1995.
Distribution — Madagascar.
9. Linderniella mbalaensis (Eb. Fisch.) Eb. Fisch.,
Schäferh. & Kai Müll., comb. nov. Lindernia mbalaen-
sis Eb. Fisch. in Bull. Jard. Bot. Natl. Belg. 67: 361. 1999.
Distribution — Zambia.
10. Linderniella nana (Engl.) Eb. Fisch., Schäferh. & Kai
Müll., comb. nov. Ilysanthes nana Engl. in Bot. Jahrb.
Syst. 23: 505. 1897 Lindernia nana (Engl.) Roessler in
Mitt. Bot. Staatssamml. München 5: 691. 1965.
= Ilysanthes schlechteri Hiern in Harvey, Fl. Cap. 4(2):
365. 1904.
Distribution Ethiopia, Congo-Kinshasa, Rwanda,
Uganda, Kenya, Tanzania, Angola, Zambia, Zimbabwe,
Namibia, South Africa.
11. Linderniella pulchella (Skan) Eb. Fisch., Schäf erh.
& Kai Müll., comb. nov. Ilysanthes pulchella Skan in
Oliver & al., Fl. Trop. Afr. 4(2): 348. 1906 Lindernia
pulchella (Skan) Philcox in Bol. Soc. Brot., ser. 2, 60:
268. 1987.
= Ilysanthes purpurascens Hutch., Botanist S. Africa:
461. 1946.
= Ilysanthes saxatilis Norl. in Bot. Not. 1951: 115. 1951.
= Ilysanthes pulchella subsp. rhodesiana Norl. in Bot.
Not. 1951: 113. 1951.
Distribution — Sudan, Ethiopia, Congo-Kinshasa,
Uganda, Tanzania, Angola, Zambia, Malawi, Zimbabwe,
Mozambique, South Africa.
12. Linderniella pusilla Eb. Fisch., Schäferh. & Kai
Müll., nom. nov. Bonnaya pusilla Oliv. in Trans. Linn.
Soc. London 29: 121. 1875, nom. illeg. [non Bonnaya pu-
silla Benth., Scroph. Ind.: 33. 1835, nec Bonnaya pusilla
Gri., Not. Pl. Asiat. 4: 107. 1854] Ilysanthes pusilla
Urb. in Ber. Deutsch. Bot. Ges. 2: 435. 1884 Linder-
nia philcoxii Eb. Fisch. in Trop. Subtrop. Pflanzenwelt
81: 295. 1992, nom. illeg. Lindernia serpens Philcox
in Ghazanfar & al., Fl. Trop. E. Africa, Scrophular.: 89.
2008 [non Lindernia pusilla (Willd.) Bold., Zakfl. Java:
165. 1916].
= Ilysanthes rupicola Lanza, Missione Biol. Borana 4:
208. 1939.
Distribution Ethiopia, Congo-Kinshasa, Rwanda,
Burundi, Uganda, Kenya, Tanzania.
The taxonomic history of this taxon is confusing.
Described as Bonnaya pusilla Oliv., which is a later
homonym and therefore illegitimate, it was transferred
to Ilysanthes by Urban (1884), whose I. pusilla was a re-
placement name. Fischer (1992) transferred it to Linder-
nia, proposing L. philcoxii as a replacement name,
because the name L. pusilla already existed, but I. ugan-
densis was cited as a synonym, which made the name
228 Fischer & al.: The phylogeny of Linderniaceae
L. philcoxii nomenclaturally superfluous and therefore
illegitimate. Careful study of the type of I. ugandensis
showed that it is dierent from I. pusilla. Thus Phil-
cox (2008) proposed L. serpens as another replacement
name. The replacement name published here in Lindern-
iella reuses the original epithet of B. pusilla, which is
also that of the earliest legitimate name for the species,
I. pusilla.
13. Linderniella pygmaea (Bonati) Eb. Fisch., Schäf-
erh. & Kai Müll., comb. nov. Craterostigma pygmae-
um Bonati in Bull. Soc. Bot. Genève, ser. 2, 15: 107.
1924 Lindernia pygmaea (Bonati) Eb. Fisch. in Bull.
Mus. Natl. Hist. Nat., B, Adansonia, ser. 4, 17: 234.
1995.
Distribution — Madagascar.
14. Linderniella trichotoma (Oliv.) Eb. Fisch., Schäf erh.
& Kai Müll., comb. nov. Bonnaya trichotoma Oliv. in
Trans. Linn. Soc. London 29: 121. 1875 Ilysanthes tri-
chotoma (Oliv.) Urb. in Ber. Deutsch. Bot. Ges. 4: 435.
1884 Lindernia madiensis Dandy in Andrews, Fl. Pl.
Sudan 3: 139. 1956 [non Lindernia trichotoma Schltr. in
Bot. Jahrb. Syst. 59: 107. 1924, nom. illeg.].
= Ilysanthes schweinfurthii Engl. in Bot. Jahrb. Syst. 23:
504. 1897 Lindernia schweinfurthii (Engl.) Dandy in
Andrews, Fl. Pl. Sudan 3: 139. 1956.
= Ilysanthes barteri Skan in Oliver & al., Fl. Trop. Afr.
4(2): 350. 1906.
= Ilysanthes albertina S. Moore in J. Bot. 45: 331. 1907.
Distribution Senegal, Guinea-Bissau, Guinea, Si-
erra Leone, Liberia, Côte d’Ivoire, Burkina Faso, Ghana,
Togo, Nigeria, Cameroon, Central African Republic, Su-
dan, Ethiopia, Congo-Kinshasa, Rwanda, Burundi, Ugan-
da, Kenya, Tanzania, Zambia.
15. Linderniella ugandensis (Skan) Eb. Fisch., Schäf erh.
& Kai Müll., comb. nov. Ilysanthes ugandensis Skan
in Oliver & al., Fl. Trop. Afr. 4(2): 348. 1906 Lindernia
ugandensis (Skan) Philcox in Ghazanfar & al., Fl. Trop.
E. Africa, Scrophular.: 86. 2008.
Distribution — Uganda.
16. Linderniella wilmsii (Engl. ex Diels) Eb. Fisch.,
Schäferh. & Kai Müll., comb. nov. Ilysanthes wilm-
sii Engl. ex Diels in Bot. Jahrb. Syst. 26: 123. 1898
Lindernia wilmsii (Engl. ex Diels) Philcox in Bol. Soc.
Brot., ser. 2, 60: 268. 1987.
= Ilysanthes muddii Hiern in Harvey, Fl. Cap. 4(2): 366.
1904.
Distribution — Congo-Kinshasa, Rwanda, Burundi,
Ugan da, Kenya, Tanzania, Angola, Zambia, Zimbabwe,
Mozambique, South Africa.
11. Micranthemum Michx., Fl. Bor.-Amer. 1: 10. 1803,
nom. cons. Globifera J. F. Gmel., Syst. Nat. 2: 32.
1791. – Type: M. orbiculatum Michx., nom. illeg. M.
umbrosum (J. F. Gmel.) S. F. Blake.
= Hemianthus Nutt. in J. Acad. Nat. Sci. Philadelphia 1:
119. 1817. – Type: H. micranthemoides Nutt. Micran-
themum micranthemoides (Nutt.) Wettst.
= Amphiolanthus Griseb., Cat. Pl. Cub.: 186. 1866. – Type
(designated by Pennell in Proc. Acad. Nat. Sci. Philadel-
phia 75: 15. 1923): A. bryoides Griseb. Micranthemum
bryoides (Griseb.) M. Gómez.
The genus Micranthemum is in need of a taxonomic
revision because species delimitations are sometimes
weakly defined.
1. Micranthemum arenarioides (Griseb.) M. Gómez,
Anal. Hist. Nat. 23: 277. 1894 Amphiolanthus arenari-
oides Griseb., Cat. Pl. Cub.: 187. 1866 Globifera are-
narioides (Griseb.) Kuntze, Revis. Gen. Pl. 2: 461. 1891.
Distribution — Cuba.
2. Micranthemum bryoides (Griseb.) M. Gómez, Anal.
Hist. Nat. 23: 277. 1894 Amphiolanthus bryoides
Griseb., Cat. Pl. Cub.: 186. 1866 Globifera bryoides
(Griseb.) Kuntze, Revis. Gen. Pl. 2: 461. 1891.
Distribution — Cuba.
3. Micranthemum callitrichoides (Griseb.) C. Wright in
Anales Acad. Ci. Med. Habana 7: 40. 1870 Hemian-
thus callitrichoides Griseb., Pl. Wright. 2: 522. 1862
Globifera callitrichoides (Griseb.) Kuntze, Revis. Gen.
Pl. 2: 461. 1891.
= Micranthemum adenandrum (C. Wright ex Griseb.)
C. Wright in Anales Acad. Ci. Med. Habana 7: 41. 1870
Hemianthus adenander C. Wright ex Griseb., Cat. Pl.
Cub.: 185. 1866 Globifera adenandra (C. Wright ex
Griseb.) Kuntze, Revis. Gen. Pl. 2: 461. 1891.
= Micranthemum ciliolatum C. Wright in Anales Acad.
Ci. Med. Habana 7: 41. 1870 Globifera ciliolata (C.
Wright) Kuntze, Revis. Gen. Pl. 2: 461. 1891.
= Micranthemum elatinoides (Griseb.) C. Wright in
Anales Acad. Ci. Med. Habana 7: 40. 1870 Hemian-
thus elatinoides Griseb., Cat. Pl. Cub.: 185. 1866 Glob-
ifera elatinoides (Griseb.) Kuntze, Revis. Gen. Pl. 2: 461.
1891.
= Micranthemum multiflorum (C. Wright ex Griseb.) C.
Wright in Anales Acad. Ci. Med. Habana 7: 40. 1870
Hemianthus multiflorus C. Wright ex Griseb., Cat. Pl.
Cub.: 185. 1866.
= Micranthemum punctatum (C. Wright ex Griseb.) C.
Wright in Anales Acad. Ci. Med. Habana 7: 40. 1870
Hemianthus punctatus C. Wright ex Griseb., Cat. Pl. Cub.:
184. 1866 Globifera punctata (C. Wright ex Griseb.)
Kuntze, Revis. Gen. Pl. 2: 461. 1891.
Distribution — Cuba.
4. Micranthemum erosum (C. Wright ex Griseb.) Eb.
Fisch., Schäf erh. & Kai Müll., comb. nov. Hemianthus
erosus C. Wright ex Griseb., Cat. Pl. Cub.: 184. 1866
Globifera erosa (C. Wright ex Griseb.) Kuntze, Revis.
Gen. Pl. 2: 461. 1891.
Distribution — Cuba.
5. Micranthemum glomeratum (Chapm.) Shinners in
Sida 1: 252. 1964 M. nuttallii var. glomeratum Chapm.,
Fl. South. U.S., ed. 2, Suppl. 2: 690. 1892 Hemianthus
glomeratus (Chapm.) Pennell in Proc. Acad. Nat. Sci.
Philadelphia 71: 248. 1920.
Distribution — U.S.A. (Florida).
229Willdenowia 43 – 2013
6. Micranthemum micranthemoides (Nutt.) Wettst. in
Engler & Prantl, Nat. Pflanzenfam. 4(3b): 77. 1891
Hemianthus micranthemoides Nutt. in J. Acad. Nat. Sci.
Philadelphia 1: 119. 1817 Micranthemum nuttallii A.
Gray, Manual, ed. 5: 331. 1867, nom. illeg. Globifera
micranthemodes (Nutt.) Kuntze, Revis. Gen. Pl. 2: 461.
1891.
Distribution — U.S.A., Cuba.
7. Micranthemum pilosum Ernst in Flora 57: 215. 1874.
Distribution — Venezuela.
8. Micranthemum procerorum L. O. Williams in Fieldi-
ana, Bot. 34: 123. 1972.
Distribution — Mexico.
9. Micranthemum reflexum (C. Wright ex Griseb.) C.
Wright in Anales Acad. Ci. Med. Habana 7: 42. 1870
Hemianthus reflexus C. Wright ex Griseb., Cat. Pl. Cub.:
186. 1866 Globifera reflexa (C. Wright ex Griseb.)
Kuntze, Revis. Gen. Pl. 2: 461. 1891.
Distribution — Cuba.
10. Micranthemum rotundatum C. Wright ex Griseb.,
Cat. Pl. Cub.: 184. 1866 Globifera rotundata (C.
Wright ex Griseb.) Kuntze, Revis. Gen. Pl. 2: 461. 1891.
Distribution — Cuba.
11. Micranthemum standleyi L. O. Williams in Fieldi-
ana, Bot. 34: 124. 1972.
Distribution — Mexico.
12. Micranthemum tetrandrum C. Wright in Anales
Acad. Ci. Med. Habana 7: 41. 1870 Globifera tetran-
dra (C. Wright) Kuntze, Revis. Gen. Pl. 2: 461. 1891
Hemianthus tetrandrus (C. Wright) Pennell in Proc.
Acad. Nat. Sci. Philadelphia 75: 16. 1923.
Distribution — Cuba.
13. Micranthemum trisetosum (C. Wright ex Griseb.)
C. Wright in Anales Acad. Ci. Med. Habana 7: 40. 1870
Hemianthus trisetosus C. Wright ex Griseb., Cat. Pl.
Cub.: 185. 1866 Globifera trisetosa (C. Wright ex
Griseb.) Kuntze, Revis. Gen. Pl. 2: 461. 1891.
Distribution — Cuba.
14. Micranthemum umbrosum (J. F. Gmel.) S. F. Blake
in Rhodora 17: 131. 1915 Globifera umbrosum J. F.
Gmel., Syst. Nat. 2: 32. 1791 Micranthemum orbicula-
tum Michx., Fl. Bor.-Amer. 1: 10. 1803, nom. illeg.
= Micranthemum emarginatum Elliott, Sketch Bot. S.
Carolina 1: 18. 1816 Micranthemum orbiculatum var.
emarginatum (Elliott) C. Mohr in Bull. Torrey Bot. Club
24: 26. 1897.
Anonymos umbrosus Walter, Fl. Carol.: 63. 1788, nom.
inval.
DistributionSE U.S.A. to South America, Puerto
Rico.
12. Pierranthus Bonati in Bull. Soc. Bot. Genève, ser. 2,
4: 254. 30 Nov 1912 Delpya Pierre ex Bonati in Bull.
Soc. Bot. Genève, ser. 2, 4: 238. 30 Sep 1912, nom. illeg.
[non Delpya Pierre ex Radlkofer in Notul. Syst. (Paris) 1:
304. 1910]. – Type: P. capitatus (Bonati) Bonati.
The genus has bothrospermous seeds, but diers from
Vandellia in its overall morphology (capitate inflores-
cence, shape of leaves, bracts, calyx and corolla).
1. Pierranthus capitatus (Bonati) Bonati in Bull. Soc.
Bot. Genève, ser. 2, 4: 254. 1912 Vandellia capitata
Bonati in Bull. Soc. Bot. France 55: 514. 1908.
Distribution — Thailand, Cambodia, Vietnam.
13. Schizotorenia T. Yamaz. in J. Jap. Bot. 53: 101. 1978
– Type: S. finetiana (Bonati) T. Yamaz.
The genus resembles Torenia, but diers in the shape
of the calyx and the aulacospermous seeds.
1. Schizotorenia atropurpurea (Ridl.) T. Yamaz. in J.
Jap. Bot. 53: 102. 1978 Torenia atropurpurea Ridl. in J.
Straits Branch Roy. Asiatic Soc. 49: 24. 1907.
Distribution — Thailand, Peninsular Malaysia.
2. Schizotorenia finetiana (Bonati) T. Yamaz. in J. Jap.
Bot. 53: 101. 1978 Torenia finetiana Bonati in Bull.
Soc. Bot. Genève, ser. 2, 5: 312. 1914 Artanema fine-
tianum (Bonati) T. Yamaz. in J. Jap. Bot. 28: 34. 1953.
= Torenia evrardii Bonati in Bull. Soc. Bot. France 71:
1097. 1924 Artanema evrardii (Bonati) T. Yamaz. in J.
Jap. Bot. 28: 36. 1953.
Distribution — Vietnam.
14. Scolophyllum T. Yamaz. in J. Jap. Bot. 53: 98. 1978.
– Type: S. ilicfolium (Bonati) T. Yamaz.
The genus has bothrospermous seeds and is probably
closely related to Bonnaya.
1. Scolophyllum ilicifolium (Bonati) T. Yamaz. in J.
Jap. Bot. 53: 100. 1978 Ilysanthes ilicfolia Bonati in
Bull. Soc. Bot. France 55: 541. 1908 Bonnaya ilicifolia
(Bonati) Bonati in Lecomte & al., Fl. Indo-Chine 4: 435.
1927 Lindernia ilicifolia (Bonati) Philcox in Taxon 19:
649. 1970.
Distribution — Thailand, Cambodia, Vietnam.
2. Scolophyllum longitubum T. Yamaz. & Chuakul in J.
Jap. Bot. 74: 82. 1999.
Distribution — Thailand.
3. Scolophyllum spinifidum (Kerr ex Barnett) T. Yamaz.,
J. Jap. Bot. 53: 101. 1978 Ilysanthes spinifida Kerr ex
Barnett in Kew Bull. 16: 489. 1963 Lindernia spinifida
(Kerr ex Barnett) Philcox in Taxon 19: 649. 1970.
Distribution — Thailand.
15. Stemodiopsis Engl. in Annuario Reale Ist. Bot. Roma
7: 25. 1898. – Type: S. rivae Engl.
1. Stemodiopsis buchananii Skan in Oliver & al., Fl.
Trop. Afr. 4(2): 315. 1906.
= Sutera cymbalariifolia Chiov., Res. Sci. Somalia Ital.
1: 129. 1916.
Distribution — Sudan, Somalia, Kenya, Tanzania,
Zambia, Zimbabwe, Malawi, Mozambique.
2. Stemodiopsis eylesii S. Moore in J. Bot. 46: 71. 1908.
Distribution — Zimbabwe.
3. Stemodiopsis glandulosa Philcox, Fl. Zambes. 8(2):
42. 1990.
Distribution — Zambia.
230 Fischer & al.: The phylogeny of Linderniaceae
4. Stemodiopsis linearis S. Moore in J. Bot. 49: 157. 1911.
Distribution — Congo-Kinshasa.
5. Stemodiopsis rivae Engl. in Annuario Reale Ist. Bot.
Roma 7: 25. 1898.
= Stemodiopsis humilis Skan in Oliver & al., Fl. Trop.
Afr. 4(2): 316. 1906.
DistributionCameroon, Sudan, Ethiopia, Burundi,
Uganda, Kenya, Tanzania, Zambia, Zimbabwe, Mozam-
bique, South Africa.
6. Stemodiopsis ruandensis Eb. Fisch. in Bot. Jahrb.
Syst. 119: 319. 1997.
Distribution — Rwanda.
16. Torenia L., Sp. Pl. 2: 619. 1753. – Type: T. asiatica L.
1. Torenia asiatica L., Sp. Pl. 2: 619. 1753.
= Torenia glabra Osbeck, Dagbok Ostind. Resa.: 210.
1757.
DistributionIndia, Nepal, China; widely natural-
ized, e.g. in Hawaii.
2. Torenia benthamiana Hance in Ann. Sci. Nat., Bot.,
ser. 4, 18: 226. 1862.
= Torenia nantoensis Hayata, Icon. Pl. Formosan. 9: 81.
1920.
= Torenia tonkinensis Bonati in Lecomte & al., Fl. Indo-
Chine 4: 399. 1927.
Distribution — China, Taiwan, Vietnam.
3. Torenia biniflora T. L. Chin & D. Y. Hong in P. C.
Tsoong & H. P. Yang, Fl. Reipubl. Popularis Sin. 67(2):
399. 1979.
Distribution — China.
4. Torenia chevalieri Bonati in Bull. Soc. Bot. France 71:
1095. 1925.
Distribution — Cambodia, Vietnam.
5. Torenia concolor Lindl. in Edwards’s Bot. Reg. 32: t.
62. 1846.
= Torenia albomarginata Bonati, Bull. Soc. Bot. France
71: 1096. 1925.
= Torenia alboviolacea Bonati in Bull. Soc. Bot. France
71: 1094. 1925.
= Torenia annamitica Bonati in Bull. Soc. Bot. France
71: 1094. 1925.
Distribution — China, Laos, Vietnam.
6. Torenia cordata (Gri.) N. M. Dutta, Bull. Bot. Soc.
Bengal 19: 25. 1965 Treisteria cordata Gri., Not. Pl.
Asiat. 4: 109. 1854.
Distribution — Myanmar.
7. Torenia cordifolia Roxb., Pl. Coromandel 2: 32. 1802.
Distribution — India, Nepal, Myanmar, China, Thai-
land.
8. Torenia courtallensis Gamble, Fl. Madras 2: 956.
1922.
Distribution — India.
9. Torenia cyrtandriflora B. L. Burtt in Rheedea 1: 3.
1991.
Distribution — Malaysia (Sarawak).
10. Torenia daubyi Eb. Fisch. & O. Lachenaud in Phyto-
taxa 125: 42. 2013.
Distribution — Gabon.
11. Torenia diusa D. Don, Prodr. Fl. Nepal.: 86. 1825.
= Torenia vagans Roxb., Fl. Ind., ed. 1832, 3: 96. 1832.
Distribution — Nepal, Bhutan, India (Assam, Sik-
kim).
12. Torenia dinklagei Engl. in Bot. Jahrb. Syst. 57: 610.
1922.
DistributionSierra Leone, Liberia, Côte d’Ivoire,
Ghana, Togo, Nigeria, Equatorial Guinea (Bioko), Cam-
eroon, Gabon, Congo-Kinshasa.
13. Torenia flava Buch.-Ham. ex Benth., Scroph. Ind.:
38. 1835.
= Torenia hokutensis Hayata, Icon. Pl. Formosan. 9: 80.
1920.
Distribution — India, China, Taiwan, Thailand, Laos,
Cambodia, Vietnam, Myanmar, Malaysia, Indonesia.
14. Torenia fordii Hook. f. in Bot. Mag. 111: t. 6797B.
1885.
Distribution — China.
15. Torenia fournieri Linden ex E. Fourn. in Ill. Hort.
23: 129. 1876.
Distribution — Thailand, Cambodia, Laos, Vietnam;
introduced (escaped from cultivation) in Mexico, Central
America and South America.
16. Torenia hayatae Bonati in Bull. Soc. Bot. France 71:
1096. 1925.
Distribution — Vietnam.
17. Torenia hirsutissima Bonati in Bull. Soc. Bot. France
55: 512. 1908.
Distribution — Thailand, Cambodia, Laos.
18. Torenia indica C. J. Saldanha in Bull. Bot. Surv. In-
dia 8: 127. 1967.
Distribution — India.
19. Torenia laotica Bonati in Bull. Soc. Bot. France 55:
512. 1908.
Distribution — Cambodia, Laos, Vietnam.
20. Torenia mannii Skan in Oliver & al., Fl. Trop. Afr.
4(2): 336. 1906.
Distribution — Gabon, Equatorial Guinea.
21. Torenia pierreana Bonati in Bull. Soc. Bot. France
55: 513. 1908.
= Torenia reptans Kerr ex Barnett in Kew Bull. 16: 487.
1963.
Distribution — Thailand, Cambodia.
22. Torenia poilanei Bonati in Bull. Soc. Bot. France 71:
1093. 1925.
= Torenia rupestris Bonati in Lecomte & al., Fl. Indo-
Chine 4: 402. 1927.
Distribution — Cambodia, Vietnam.
23. Torenia ranongensis T. Yamaz. in J. Jap. Bot. 58: 14.
1983.
Distribution — Thailand.
24. Torenia scandens Bonati in Bull. Soc. Bot. France
71: 1095. 1925.
Distribution — Vietnam.
25. Torenia siamensis T. Yamaz. in J. Jap. Bot. 53: 103.
1978.
231Willdenowia 43 – 2013
Distribution — Thailand.
26. Torenia silvicola A. Raynal in Adansonia, n.s., 5:
522. 1965.
Distribution — Cameroon, Gabon, Congo-Brazza-
ville.
27. Torenia stolonifera Bojer ex Benth. in Candolle,
Prodr. 10: 410. 1846.
Distribution — Madagascar.
28. Torenia thailandica T. Yamaz. in J. Jap. Bot. 65: 265.
1990.
Distribution — Thailand.
29. Torenia thorelii Bonati in Bull. Soc. Bot. France 55:
514. 1908.
Distribution — Cambodia, Laos, Vietnam.
30. Torenia thouarsii (Cham. & Schltdl.) Kuntze, Re-
vis. Gen. Pl. 2: 468. 1891 Nortenia thouarsii Cham.
& Schltdl. in Linnaea 3: 18. 1828 Torenia parviflora
Buch.-Ham. ex Benth., Scroph. Ind.: 39. 1835, nom.
illeg.
= Torenia ramosissima Vatke in Oesterr. Bot. Z. 25: 10.
1875.
= Torenia chamaedrys Bonati in Bull. Soc. Bot. Genève,
ser. 2, 18: 30. 1926.
= Torenia nana Bonati in Bull. Soc. Bot. Genève, ser. 2,
18: 32. 1926, nom. illeg. [non Torenia nana Benth. in
Candolle, Prodr. 10: 412. 1846].
= Torenia pedunculariformis Bonati in Bull. Soc. Bot.
Genève, ser. 2, 18: 33. 1926.
= Torenia viguieri Bonati in Bull. Soc. Bot. Genève, ser.
2, 18: 30. 1926.
Distribution — Senegal, Mali, Guinea, Sierra Leone,
Liberia, Côte d’Ivoire, Ghana, Togo, Nigeria, São Tomé
and Príncipe (Príncipe), Cameroon, Gabon, Congo-Braz-
zaville, Central African Republic, Sudan, Ethiopia, Con-
go-Kinshasa, Rwanda, Burundi, Uganda, Kenya, Tanza-
nia, Angola, Zambia, Malawi, Zimbabwe, Mozambique,
Botswana, South Africa, Madagascar, Mauritius, India,
Myanmar; introduced in Central America and South
America.
31. Torenia travancorica Gamble, Fl. Madras 2: 957.
1922.
Distribution — India.
32. Torenia vientianica T. Yamaz. in J. Jap. Bot. 55: 328.
1980.
Distribution — Vietnam.
33. Torenia violacea (Blanco) Pennell in J. Arnold Arbor.
24: 255. 1943 Mimulus violaceus Blanco, Fl. Filip., ed.
2: 357. 1845.
= Torenia exappendiculata Regel in Gartenflora 26: 34.
1877.
= Torenia peduncularis Benth. ex Hook. f., Fl. Brit. India
4: 276. 1884.
= Torenia edentula Gri. ex Benth. in Candolle, Prodr.
10: 410. 1846.
Distribution Bhutan, India, China, Taiwan, Thai-
land, Laos, Cambodia, Vietnam, Malaysia, Philippines,
Indonesia (Java).
Torenia crustacea group
Lindernia sect. Torenioides (Benth.) Pennell in Acad.
Nat. Sci. Philadelphia Monogr. 1: 139. 1935 Vandel-
lia sect. Torenioides Benth. in Candolle, Prodr. 10: 413.
1846.
The following species are closely related to Torenia
crustacea and have been assigned to Lindernia in most
recent treatments (e.g. Philcox 1968). They are here in-
cluded in Torenia.
34. Torenia blancoi Merr., Sp. Blancoan.: 346. 1918
Lindernia blancoi (Merr.) Philcox in Kew Bull. 22: 23.
1968.
Distribution — Philippines; introduced in Mexico.
35. Torenia bonatii Eb. Fisch., Schäferh. & Kai Müll.,
nom. nov. Vandellia pierreana Bonati in Bull. Soc.
Bot. France 55: 538. 1908 Lindernia pierreana (Bona-
ti) Bonati in Lecomte & al., Fl. Indo-Chine 4: 415. 1927
[non Torenia pierreana Bonati in Bull. Soc. Bot. France
55: 513. 1908].
= Lindernia maculata Bonati in Lecomte & al., Fl. Indo-
Chine 4: 414. 1927.
= Vandellia ligulata T. Yamaz. in J. Jap. Bot. 28: 37. 1953
Lindernia ligulata (T. Yamaz.) Philcox in Taxon 19:
649. 1970.
Distribution — Vietnam.
A replacement name is required because the name
Torenia pierreana already exists.
36. Torenia cambodgiana Bonati in Bull. Soc. Bot.
France 55: 513. 1908 Lindernia cerastioides T. Yamaz.
in J. Jap. Bot. 53: 97. 1978 [non Lindernia cambodgiana
(Bonati) Philcox in Taxon 19: 649. 1970].
Distribution — Cambodia.
37. Torenia crenata (Pennell) Pennell in J. Arnold Arbor.
24: 254. 1943 Lindernia crenata Pennell in J. Arnold
Arbor. 20: 79. 1939.
Distribution — Indonesia (Sulawesi), New Guinea.
38. Torenia crustacea (L.) Cham. & Schltdl. in Lin naea
2: 570. 1827 Capraria crustacea L., Syst. Nat., ed. 12,
2: 419; Mant. Pl. 1: 87. 1767 Lindernia crustacea (L.)
F. Muell., Syst. Census Austral. Pl.: 97. 1882 1883
Vandellia crustacea (L.) Benth., Scroph. Ind.: 35. 1835.
= Vandellia racemosa Bonati in Bull. Soc. Bot. France
55: 537. 1908, nom. illeg. [non Vandellia racemosa
Spreng., Neue Entd. 1: 262. 1820].
= Vandellia gracilis Bonati in Bull. Soc. Bot. France 55:
539. 1908 Lindernia gracilis (Bonati) Bonati in Le-
comte & al., Fl. Indo-Chine 4: 416. 1927.
DistributionSenegal, Guinea-Bissau, Sierra Leo-
ne, Côte d’Ivoire, Ghana, Nigeria, Cameroon, Equato-
rial Guinea (Annobón), Central African Republic, Ga-
bon, Congo-Brazzaville, Congo-Kinshasa, Burundi,
Madagascar, India, Nepal, Sri Lanka, China, Thailand,
Laos, Vietnam, Cambodia, Japan, Philippines, Austral-
ia, New Guinea, Micronesia, Polynesia; introduced in
S U.S.A., Mexico, Cuba, Dominica, Grenada, Guad-
eloupe, Jamaica, Martinique, Puerto Rico, Saint Lucia,
Trinidad and Tobago, Belize, Costa Rica, Guatemala,
232 Fischer & al.: The phylogeny of Linderniaceae
Honduras, Nicaragua, Panama, Bolivia, Brazil, Colom-
bia, Ecuador, French Guiana, Guyana, Peru, Suriname
and Venezuela.
39. Torenia davidii Eb. Fisch., Schäferh. & Kai Müll.,
nom. nov. Lindernia celebica Philcox in Kew Bull. 22:
15. 1968 [non Torenia celebica T. Yamaz. in J. Jap. Bot.
65: 262. 1990].
Distribution — Indonesia (Sulawesi).
A replacement name is required because the name
Torenia celebica already exists.
40. Torenia dictyophora (P. C. Tsoong) Eb. Fisch.,
Schäferh. & Kai Müll., comb. nov. Lindernia dictyo-
phora P. C. Tsoong in P. C. Tsoong & H. P. Yang, Fl.
Reipubl. Popularis Sin. 67(2): 397. 1979.
Distribution — Thailand, China (Yunnan).
41. Torenia grandiflora (Merr.) Eb. Fisch., Schäferh. &
Kai Müll., comb. nov. Vandellia grandiflora Merr. in
Philipp. J. Sci. 1(Suppl. 3): 237. 1906 Lindernia gran-
diflora (Merr.) Merr., Enum. Philipp. Fl. Pl. 3: 438. 1923,
nom. illeg. [non Lindernia grandiflora Nutt., Gen. N.
Amer. Pl. 2: 43. 1818] Lindernia philippinensis Phil-
cox in Kew Bull. 22: 13. 1968.
Distribution — Philippines, New Guinea.
42. Torenia molluginoides (Benth.) Eb. Fisch., Schäf-
erh. & Kai Müll., comb. nov. Vandellia molluginoides
Benth., Scroph. Ind.: 35. 1835 Lindernia molluginoides
(Benth.) Wettst. in Engler & Prantl, Nat. Pflanzenfam.
4(3b): 80. 1891.
Distribution — Thailand, Myanmar.
43. Torenia oblonga (Benth.) Steud., Nomencl. Bot., ed.
2, 2: 692. 1841 Vandellia oblonga Benth., Scroph. Ind.:
35. 1835 Lindernia oblonga (Benth.) Merr. & Chun in
Sunyatsenia 5: 180. 1940.
= Lindernia subcrenulata (Miq.) Merr. in Sunyatsenia 1:
33. 1930 Vandellia subcrenulata Miq. in J. Bot. Néerl.
1: 113. 1861.
Distribution — S China, Vietnam.
44. Torenia perennans (T. Yamaz.) Eb. Fisch., Schäf-
erh. & Kai Müll., comb. nov. Vandellia perennans T.
Yamaz. in J. Jap. Bot. 28: 39. 1953 Lindernia peren-
nans (T. Yamaz.) Philcox in Taxon 19: 649. 1970.
Distribution — Laos, Vietnam.
45. Torenia philcoxii Eb. Fisch., Schäferh. & Kai Müll.,
nom. nov. Lindernia glabra Philcox in Kew Bull. 22:
23. 1968 [non Torenia glabra Osbeck, Dagbok Ostind.
Resa.: 210. 1757].
Distribution — New Guinea.
A replacement names is required because the name
Torenia glabra already exists.
46. Torenia pierreanoides (T. Yamaz.) Eb. Fisch.,
Schäferh. & Kai Müll., comb. nov. Lindernia pierrea-
noides T. Yamaz. in J. Jap. Bot. 53: 2. 1978.
Distribution — Thailand, Cambodia.
47. Torenia pterogona (T. Yamaz.) Eb. Fisch., Schäferh.
& Kai Müll., comb. nov. Lindernia pterogona T.
Yamaz. in J. Jap. Bot. 55: 328. 1980.
Distribution — Laos.
48. Torenia spathacea (Bonati) Eb. Fisch., Schäferh. &
Kai Müll., comb. nov. Vandellia spathacea Bonati in
Notul. Syst. (Paris) 1: 333. 1911 Lindernia spathacea
(Bonati) Bonati in Lecomte & al., Fl. Indo-Chine 4: 414.
1927.
Distribution — Cambodia, Laos, Vietnam.
49. Torenia subconnivens (Philcox) Eb. Fisch., Schäferh.
& Kai Müll., comb. nov. Lindernia subconnivens Phil-
cox in Kew Bull. 22: 13. 1968.
Distribution — Philippines.
50. Torenia udawnensis (T. Yamaz.) Eb. Fisch., Schäferh.
& Kai Müll., comb. nov. Lindernia udawnensis T.
Yamaz. in J. Jap. Bot. 53: 9. 1978.
Distribution — Thailand.
51. Torenia umbellata (T. Yamaz.) Eb. Fisch., Schäferh.
& Kai Müll., comb. nov. Lindernia umbellata T.
Yamaz. in J. Jap. Bot. 53: 10. 1978.
Distribution — Cambodia.
17. Vandellia L., Syst. Nat., ed. 12, 2: 384, 422; Mant. Pl.
1: 12. 1767. – Type: V. diusa L.
= Tittmannia Rchb., Iconogr. Bot. Exot. 1: 26. 1824, nom.
rej. [non Tittmannia Brongn. in Ann. Sci. Nat. (Paris) 8:
385. 1826, nom. cons., Bruniaceae] Type: T. viscosa
(Hornem.) Rchb. Vandellia viscosa (Hornem.) Merr.
= Georaya Bonati in Notul. Syst. (Paris) 1: 334. 1911.
– Type (designated by Yamazaki in Leroy, Fl. Cambodge,
Laos Viêt-nam 21: 117. 1985): G. junciformis Bonati
Vandellia junciformis (Bonati) Eb. Fisch., Schäferh. &
Kai Müll.
= Trichotaenia T. Yamaz. in J. Jap. Bot. 28: 40. 1953
Type (designated by Yamazaki in Leroy, Fl. Cambodge,
Laos Viêt-nam 21: 118. 1985): Trichotaenia saginiformis
(Bonati) T. Yamaz. Vandellia saginiformis Bonati.
= Lindernia subg. Didymadenia W. R. Barker in J. Ad-
elaide Bot. Gard. 13: 79. 1990. – Type: L. chrysoplectra
W. R. Barker Vandellia chrysoplectra (W. R. Barker)
Eb. Fisch., Schäferh. & Kai Müll.
1. Vandellia anagallis (Burm. f.) T. Yamaz. in J. Jap. Bot.
30: 176. 1955 Ruellia anagallis Burm. f., Fl. Indica:
135. 1768 Lindernia anagallis (Burm. f.) Pennell in J.
Arnold Arbor. 24: 252. 1943.
= Gratiola cordifolia Colsm., Prodr. Descr. Gratiol.: 15.
1793 Lindernia cordifolia (Colsm.) Merr., Bibl. Enum.
Born. Pl.: 524. 1921.
= Vandellia pedunculata Benth., Scroph. Ind.: 37. 1835
Lindernia pedunculata (Benth.) Wettst. in Engler &
Prantl, Nat. Pflanzenfam 4(3b): 79. 1891.
= Vandellia laotica T. Yamaz. in J. Jap. Bot. 37: 268.
1962, nom. illeg. [non Vandellia laotica Bonati, Bull.
Soc. Bot. Genève, ser. 2, 5: 140. 1913].
= Vandellia emarginata T. Yamaz. in Bull. Natl. Sci.
Mus., Tokyo, n.s., 6: 370. 1963 Lindernia emarginata
(T. Yamaz.) Philcox in Taxon 19: 649. 1970.
Distribution Madagascar, India, S China, Thai-
land, Laos, Vietnam, Malaysia, Australia, Philippines,
New Guinea.
233Willdenowia 43 – 2013
2. Vandellia annamensis (T. Yamaz.) Eb. Fisch.,
Schäferh. & Kai Müll., comb. nov. Lindernia anna-
mensis T. Yamaz. in J. Fac. Sci. Univ. Tokyo, Sect. 3, Bot.
13: 48. 1981.
Distribution — Vietnam.
3. Vandellia aplectra (W. R. Barker) Eb. Fisch., Schäferh.
& Kai Müll., comb. nov. Lindernia aplectra W. R.
Barker in J. Adelaide Bot. Gard. 13: 80. 1990.
DistributionAustralia.
4. Vandellia brevipedunculata (Migo) T. Yamaz. in J.
Jap. Bot. 30: 175. 1955 Lindernia brevipedunculata
Migo in J. Shanghai Sci. Inst., Sect. 3, 4: 160. 1939.
Distribution — China.
5. Vandellia brucei (R. A. Howard) Eb. Fisch., Schäferh.
& Kai Müll., comb. nov. Lindernia brucei R. A.
Howard in J. Arnold Arbor. 56: 454. 1975.
Distribution — Saint Vincent and the Grenadines
(Saint Vincent).
6. Vandellia cambodgiana (Bonati) Eb. Fisch., Schäferh.
& Kai Müll., comb. nov. Ilysanthes cambodgi-
ana Bonati in Bull. Soc. Bot. France 55: 542. 1908
Lindernia cambodgiana (Bonati) Philcox in Taxon 19:
649. 1970.
Distribution — Thailand, Cambodia, Laos, Vietnam.
7. Vandellia chrysoplectra (W. R. Barker) Eb. Fisch.,
Schäferh. & Kai Müll., comb. nov. Lindernia chryso-
plectra W. R. Barker in J. Adelaide Bot. Gard. 13: 81.
1990.
DistributionAustralia.
8. Vandellia cleistandra (W. R. Barker) Eb. Fisch.,
Schäferh. & Kai Müll., comb. nov. Lindernia cleistan-
dra W. R. Barker in J. Adelaide Bot. Gard. 13: 82. 1990.
DistributionAustralia.
9. Vandellia cowiei (W. R. Barker) Eb. Fisch., Schäferh.
& Kai Müll., comb. nov. Lindernia cowiei W. R. Bark-
er in J. Adelaide Bot. Gard. 18: 162. 1998.
DistributionAustralia.
10. Vandellia delicatula (P. C. Tsoong & T. C. Ku) Eb.
Fisch., Schäferh. & Kai Müll., comb. nov. Lindernia
delicatula P. C. Tsoong & T. C. Ku in P. C. Tsoong &
H. P. Yang, Fl. Reipubl. Popularis Sin. 67(2): 398. 1979.
Distribution — China.
11. Vandellia diusa L., Syst. Nat., ed. 12, 2: 422; Mant.
Pl. 1: 89. 1767 Lindernia diusa (L.) Wettst. in Engler
& Prantl, Nat. Pflanzenfam. 4(3b): 79. 1891 Pyxidaria
diusa (L.) Kuntze, Revis. Gen. Pl. 2: 464. 1891.
DistributionSenegal, Sierra Leone, Liberia, Côte
d’Ivoire, Burkina Faso, Ghana, Togo, Nigeria, Cam-
eroon, Equatorial Guinea (Annobón, Bioko), São Tomé
and Príncipe, Gabon, Central African Republic, Congo-
Brazzaville, Congo-Kinshasa, Burundi, Uganda, Tanza-
nia, Madagascar; introduced in Mexico, Belize, Costa
Rica, Guatemala, Honduras, Panama, Cuba, Dominica,
Dominican Republic, Grenada, Guadeloupe, Jamaica,
Martinique, Puerto Rico, Saint Vincent and the Gren-
adines, Trinidad and Tobago, Bolivia, Brazil, Colombia,
Ecuador, Guyana, Peru, Suriname and Venezuela.
12. Vandellia eberhardtii (Bonati) Eb. Fisch., Schäferh.
& Kai Müll., comb. nov. Lindernia eberhardtii Bonati
in Bull. Soc. Bot. Genève, ser. 2, 15: 111. 1924.
Distribution — Vietnam.
13. Vandellia elata Benth., Scroph. Ind.: 36. 1835
Lindernia elata (Benth.) Wettst. in Engler & Prantl, Nat.
Pflanzenfam. 4(3b): 79. 1891.
= Vandellia urticifolia Hance in J. Bot. 6: 301. 1868
Lindernia urticifolia (Hance) Bonati in Lecomte & al.,
Fl. Indo-Chine 4: 421. 1927.
= Lindernia noei Kerr ex Barnett in Kew Bull. 16: 488.
1963.
Distribution — Myanmar, S China, Thailand, Cam-
bodia, Vietnam, Malaysia.
14. Vandellia eremophiloides (W. R. Barker) Eb. Fisch.,
Schäferh. & Kai Müll., comb. nov. Lindernia eremophi-
loides W. R. Barker in J. Adelaide Bot. Gard. 13: 84. 1990.
Distribution — Australia.
15. Vandellia harmandii (Bonati) Eb. Fisch., Schäf erh.
& Kai Müll., comb. nov. Vandellia elata var. harmandii
Bonati in Bull. Soc. Bot. France 55: 539. 1908 Linder-
nia elata var. harmandii (Bonati) Bonati in Lecomte &
al., Fl. Indo-Chine 4: 418. 1927 Lindernia harmandii
(Bonati) T. Yamaz. in J. Fac. Sci. Univ. Tokyo, Sect. 3,
Bot. 13: 21. 1981.
Distribution — Cambodia, Vietnam.
16. Vandellia hookeri C. B. Clarke ex Hook. f., Fl. Brit.
India 4: 280. 1884 Lindernia hookeri (C. B. Clarke ex
Hook.f.) Wettst. in Engler & Prantl, Nat. Pflanzenfam.
4(3b): 80. 1891.
Distribution — India, Nepal, Myanmar, China, Thai-
land, Vietnam.
17. Vandellia humilis (Bonati) Eb. Fisch., Schäferh. &
Kai Müll., comb. nov. Lindernia humilis Bonati in
Bull. Soc. Bot. Genève, ser. 2, 15: 100. 1924.
= Lindernia subreniformis Philcox in Bol. Soc. Brot., ser.
2, 60: 268. 1987.
Distribution Kenya, Tanzania (including Zanzi-
bar), Mozambique, Madagascar.
18. Vandellia hypandra (W. R. Barker) Eb. Fisch.,
Schäferh. & Kai Müll., comb. nov. Lindernia hypan-
dra W. R. Barker in J. Adelaide Bot. Gard. 13: 85. 1990.
DistributionAustralia.
18. Vandellia junciformis (Bonati) Eb. Fisch., Schäf-
erh. & Kai Müll., comb. nov. Georaya junciformis
Bonati in Notul. Syst. (Paris) 1: 334. 1911 Lindernia
junciformis (Bonati) T. Yamaz. in J. Jap. Bot. 53: 97.
1978.
= Georaya cuspidata Bonati in Notul. Syst. (Paris) 1:
334. 1911.
Distribution — Thailand, Vietnam.
19. Vandellia khaoyaiensis (T. Yamaz.) Eb. Fisch., Schäf-
erh. & Kai Müll., comb. nov. Lindernia khaoyaiensis
T. Yamaz. in J. Jap. Bot. 55: 331. 1980.
Distribution — Thailand.
20. Vandellia kiangsiensis (P. C. Tsoong) Eb. Fisch.,
Schäferh. & Kai Müll., comb. nov. Lindernia kiang-
234 Fischer & al.: The phylogeny of Linderniaceae
siensis P. C. Tsoong in P. C. Tsoong & H. P. Yang, Fl.
Reipubl. Popularis Sin. 67(2): 396. 1979.
Distribution — China.
21. Vandellia laotica Bonati in Bull. Soc. Bot. Genève,
ser. 2, 5: 140. 1913 Lindernia laotica (Bonati) Bonati
in Lecomte & al., Fl. Indo-Chine 4: 423. 1927.
Distribution — Thailand, Laos.
22. Vandellia latifolia (Blume) Benth. in Candolle, Pro-
dr. 10: 415. 1846 Diceros latifolius Blume, Bijdr. Fl.
Ned. Ind. 14: 752. 1826 Lindernia latifolia (Blume)
Koord., Exkurs.-Fl. Java 3: 179. 1912.
= Vandellia foliosa Bonati in Bull. Soc. Bot. France 56:
468. 1909 Lindernia foliosa (Bonati) Bonati in Le-
comte & al., Fl. Indo-Chine 4: 418. 1927.
Distribution — Myanmar, Thailand, Laos, Vietnam,
Indonesia (Java, Sumatra), Borneo.
23. Vandellia longituba T. Yamaz. in Bull. Natl. Sci.
Mus., Tokyo, n.s., 6: 370. 1963 Lindernia longituba (T.
Yamaz.) Philcox in Taxon 19: 649. 1970.
Distribution — Vietnam.
24. Vandellia macrobotrys (P. C. Tsoong) Eb. Fisch.,
Schäferh. & Kai Müll., comb. nov. Lindernia macro-
botrys P. C. Tsoong in P. C. Tsoong & H. P. Yang, Fl.
Reipubl. Popularis Sin. 67(2): 396. 1979.
Distribution — China.
25. Vandellia macrosiphonia (F. Muell.) Eb. Fisch.,
Schäferh. & Kai Müll., comb. nov. Rhamphicarpa mac-
rosiphonia F. Muell. in Proc. Linn. Soc. New South Wales,
ser. 2, 6: 473. 1892 Lindernia macrosiphonia (F. Muell.)
W. R. Barker in J. Adelaide Bot. Gard. 13: 86. 1990.
DistributionAustralia.
26. Vandellia maxwellii (T. Yamaz.) Eb. Fisch., Schäf-
erh. & Kai Müll., comb. nov. Lindernia maxwellii T.
Yamaz. in J. Jap. Bot. 53: 2. 1978.
Distribution — Thailand.
27. Vandellia megaphylla (P. C. Tsoong) Eb. Fisch.,
Schäferh. & Kai Müll., comb. nov. Lindernia mega-
phylla P. C. Tsoong in Nordic J. Bot. 26: 41. 2008.
Distribution — China.
28. Vandellia micrantha (D. Don) Eb. Fisch., Schäf erh.
& Kai Müll., comb. nov. Lindernia micrantha D. Don,
Prodr. Fl. Nepal.: 85. 1825.
= Vandellia angustifolia Benth., Scroph. Ind.: 37. 1835
Lindernia angustifolia (Benth.) Wettst. in Engler &
Prantl, Nat. Pflanzenfam. 4(3b): 79. 1891.
Distribution — Sri Lanka, India, Nepal, Myanmar,
China, Japan, Cambodia, Laos, Vietnam, Malaysia, New
Guinea.
29. Vandellia montana (Blume) Benth. in Candolle,
Prodr. 10: 415. 1846 Diceros montanus Blume, Bi-
jdr. Fl. Ned. Ind. 14: 752. 1826 Lindernia montana
(Blume) Koord., Exkurs.-Fl. Java 3: 178. 1912, nom. il-
leg. [non Lindernia montana Hiern, Cat. Afr. Pl. 1: 764.
1898].
= Vandellia mollis Benth., Scroph. Ind.: 37. 1835
Lindernia mollis (Benth.) Wettst. in Engler & Prantl, Nat.
Pflanzenfam. 4(3b): 79. 1891.
= Vandellia sericea Bonati in Bull. Soc. Bot. France 55:
540. 1908.
= Vandellia nutans T. Yamaz. in J. Jap. Bot. 28: 36. 1953
Lindernia nutans (T. Yamaz.) Philcox in Taxon 19: 649.
1970.
Distribution Pakistan, India, Myanmar, S China,
Cambodia, Laos, Vietnam, Malaysia, Indonesia (Java,
Sumatra).
30. Vandellia multiflora (Roxb.) G. Don, Gen. Hist. 4:
549. 1837 1838 Torenia multiflora Roxb., Fl. Ind., ed.
1832, 3: 96. 1832 Pyxidaria trichotoma Kuntze, Revis.
Gen. Pl. 2: 464. 1891, nom. illeg. Lindernia trichotoma
Schltr. in Bot. Jahrb. Syst. 59: 107. 1924, nom. illeg.
Lindernia multiflora (Roxb.) Mukerjee in J. Ind. Bot.
Soc. 24: 131. 1945.
= Lindernia papuana Pennell in J. Arnold Arbor. 24: 251.
1943.
Tittmannia trichotoma Benth. in Wallich, Numer. List:
no. 3943. 1831, nom. nud.
DistributionIndia, Malaysia, Indonesia (Java, Mo-
luccas), Philippines, New Guinea.
31. Vandellia purpurea (Kerr ex Barnett) Eb. Fisch.,
Schäferh. & Kai Müll., comb. nov. Georaya purpurea
Kerr ex Barnett in Kew Bull. 16: 486. 1963 Lindernia
kerrii T. Yamaz. in J. Jap. Bot. 53: 98. 1978 [non Lindern-
ia purpurea (Lebrun & L. Touss.) R. Germ. in Bull. Jard.
Bot. État Bruxelles 22: 75. 1952].
Distribution — Thailand, Laos.
32. Vandellia pusilla (Willd.) Merr. in Philipp. J. Sci., C
7: 246. 1912 Gratiola pusilla Willd., Sp. Pl. 1: 105.
1797 Lindernia pusilla (Willd.) Bold., Zakfl. Java: 165.
1916.
= Diceros caespitosus Blume, Bijdr. Fl. Ned. Ind. 14:
753. 1826 Lindernia caespitosa (Blume) Panigrahi in
Taxon 33: 320. 1984.
= Torenia hirta Cham. & Schtdl. in Linnaea 2: 571.
1827.
=Vandellia scabra Benth., Scroph. Ind.: 36. 1835 Lin-
dernia scabra (Benth.) Wettst. in Engler & Prantl, Nat.
Pflanzenfam. 4(3b): 79. 1891.
= Lindernia stellariifolia Hayata, Icon. Pl. Formosan. 9:
77. 1920.
Distribution — Sri Lanka, India, Myanmar, S China,
Taiwan, Thailand, Cambodia, Laos, Philippines, New
Guinea.
33. Vandellia rivularis (Kerr ex Barnett) Eb. Fisch.,
Schäferh. & Kai Müll., comb. nov. Lindernia rivularis
Kerr ex Barnett in Kew Bull. 16: 488. 1963.
Distribution — Thailand.
34. Vandellia saginiformis Bonati in Bull. Soc. Bot.
Genève, ser. 2, 4: 240. 1912 Lindernia saginiformis
(Bonati) Bonati in Lecomte & al., Fl. Indo-Chine 4: 412.
1927 Trichotaenia saginiformis (Bonati) T. Yamaz. in
J. Jap. Bot. 28: 41. 1953.
= Lindernia fasciculata Bonati in Bull. Soc. Bot. Genève,
ser. 2, 15: 112. 1924 Trichotaenia fas ciculata (Bonati)
T. Yamaz. in J. Jap. Bot. 28: 42. 1953.
235Willdenowia 43 – 2013
Distribution — Cambodia, Vietnam.
35. Vandellia satakei (T. Yamaz.) Eb. Fisch., Schäferh. &
Kai Müll., comb. nov. Lindernia satakei T. Yamaz. in
J. Jap. Bot. 55: 331. 1980.
Distribution — Thailand.
36. Vandellia scapigera (R. Br.) Benth. in Candolle, Pro-
dr. 10: 415. 1846 Lindernia scapigera R. Br., Prodr.:
441. 1810.
Distribution — New Guinea, Australia.
37. Vandellia scutellariiformis (T. Yamaz.) T. Yamaz. in
J. Jap. Bot. 30: 174. 1955 Lindernia scutellariiformis T.
Yamaz. in J. Jap. Bot. 27: 67. 1952.
DistributionTaiwan.
38. Vandellia senegalensis Benth. in Candolle, Prodr. 10:
416. 1846 Lindernia senegalensis (Benth.) Hiern, Cat.
Afr. Pl. 1: 763. 1898 Pyxidaria senegalensis (Benth.)
Kuntze, Revis. Gen. Pl. 2: 465. 1891.
DistributionSenegal, Guinea-Bissau, Sierra Leo-
ne, Liberia, Côte d’Ivoire, Ghana, Nigeria, Cameroon,
Gabon, Central African Republic, Congo-Kinshasa.
39. Vandellia setulosa (Maxim.) T. Yamaz. in J. Jap. Bot.
30: 174. 1955 Torenia setulosa Maxim. in Bull. Acad.
Imp. Sci. Saint-Pétersbourg 31: 72. 1887 Lindernia
setulosa (Maxim.) Tuyama ex H. Hara in J. Jap. Bot. 19:
207. 1943.
= Vandellia cavaleriei H. Lév. in Bull. Soc. Agric. Sarthe,
ser. 2, 31: 324. 1904.
Distribution — China, Japan.
40. Vandellia stemodioides Miq., Fl. Ned. Ind., Eerste
Bijv. 3: 563. 1861 Lindernia stemodioides (Miq.) Kerr
in Craib, Fl. Siam. 3(2): 79. 1954.
Distribution — Indonesia (Bangka, Sulawesi).
41. Vandellia stictantha (Hiern) Eb. Fisch., Schäferh. &
Kai Müll., comb. nov. Ilysanthes stictantha Hiern, Cat.
Afr. Pl. 1: 765. 1898 Lindernia stictantha (Hiern) Skan
in Oliver & al., Fl. Trop. Afr. 4(2): 339. 1906.
Distribution São Tomé and Príncipe (São Tomé),
Zambia, Malawi, Angola.
42. Vandellia stolonifera (T. Yamaz.) Eb. Fisch., Schäferh.
& Kai Müll., comb. nov. Lindernia stolonifera T.
Yamaz. in J. Jap. Bot. 53: 6. 1978.
Distribution — Vietnam.
43. Vandellia stricta (P. C. Tsoong & T. C. Ku) Eb. Fisch.,
Schäferh. & Kai Müll., comb. nov. Lindernia stricta P.
C. Tsoong & T. C. Ku in P. C. Tsoong & H. P. Yang, Fl.
Reipubl. Popularis Sin. 67(2): 397. 1979.
Distribution — China.
44. Vandellia subracemosa (De Wild.) Eb. Fisch.,
Schäferh. & Kai Müll., comb. nov. Lindernia subrace-
mosa De Wild., Pl. Bequaert. 5: 425. 1932.
Distribution Ethiopia, Congo-Kinshasa, Rwanda,
Burundi, Uganda, Tanzania.
45. Vandellia subulata (R. Br.) Benth. in Candolle, Pro-
dr. 10: 415. 1846 Lindernia subulata R. Br., Prodr.:
441. 1810.
Distribution — New Guinea, Australia.
46. Vandellia taishanensis (F. Z. Li) Eb. Fisch., Schäferh.
& Kai Müll., comb. nov. Lindernia taishanensis F. Z.
Li in Bull. Bot. Res., Harbin. 6(1): 169. 1986.
Distribution — China.
47. Vandellia tectanthera (W. R. Barker) Eb. Fisch.,
Schäferh. & Kai Müll., comb. nov. Lindernia tect-
anthera W. R. Barker in J. Adelaide Bot. Gard. 13: 86.
1990.
Distribution — Australia.
48. Vandellia thorelii Bonati in Bull. Soc. Bot. France
55: 539. 1908 Lindernia thorelii (Bonati) Bonati in Le-
comte & al., Fl. Indo-Chine 4: 423. 1927.
Distribution — Thailand, Laos, Vietnam.
49. Vandellia tonkinensis Bonati in Bull. Soc. Bot.
France 55: 540. 1908 Lindernia tonkinensis (Bonati)
Bonati in Lecomte & al., Fl. Indo-Chine 4: 422. 1927.
Distribution — Vietnam.
50. Vandellia viscosa (Hornem.) Merr. in Philipp. J. Sci.,
C 7: 246. 1912 Gratiola viscosa Hornem., Enum. Pl.
Hort. Hafn., rev. ed.: 19. 1807 Hornemannia viscosa
(Hornem.) Willd., Enum. Pl. 2: 654. 1809 Tittmannia
viscosa (Hornem.) Rchb., Iconogr. Bot. Exot. 1: 26. 1824
Lindernia viscosa (Hornem.) Bold., Zakfl. Java: 165. 1916
Vandellia hirsuta Buch.-Ham. ex Benth., Scroph. Ind.:
36. 1835, nom. illeg. Lindernia hirsuta Wettst. in Engler
& Prantl, Nat. Pflanzenfam., 4(3b): 79. 1891, nom. illeg.
= Lindernia elata var. floribunda Bonati in Lecomte &
al., Fl. Indo-Chine 4: 419. 1927.
= Lindernia aprica Kerr ex Barnett in Kew Bull. 16: 487.
1963.
Distribution — India, Nepal, Myanmar, S China,
Thailand, Cambodia, Laos, Vietnam, Malaysia, Philip-
pines; introduced in Brazil.
51. Vandellia vogelii (Skan) Eb. Fisch., Schäferh. & Kai
Müll., comb. nov. Lindernia vogelii Skan in Oliver &
al., Fl. Trop. Afr. 4(2): 339. 1906.
Distribution — Côte d’Ivoire, Nigeria, Equatorial
Guinea (Bioko), Gabon.
52. Vandellia yaoshanensis (P. C. Tsoong) Eb. Fisch.,
Schäferh. & Kai Müll., comb. nov. Lindernia yaoshan-
ensis P. C. Tsoong in P. C. Tsoong & H. P. Yang, Fl.
Reipubl. Popularis Sin. 67(2): 396. 1979.
Distribution — China.
Excluded or unassigned species
Several names described in, or transferred to, Lindernia
have to be excluded from this genus or could not be as-
signed to one of the genera listed above:
Bonnaya quinqueloba Blatt. & Hallb. in J. Bombay Nat.
Hist. Soc. 25: 417. 1918 Lindernia quinqueloba
(Blatt. & Hallb.) Mukerjee in J. Ind. Bot. Soc. 24: 133.
1945. – According to Sivarajan & Mathew (1983),
this taxon is probably an aberrant form of L. bracte-
oides (Blatt. & Hallb.) Mukerjee (= Bonnaya ciliata
236 Fischer & al.: The phylogeny of Linderniaceae
(Colsm.) Spreng. subsp. ciliata). However, more ma-
terial is needed to evaluate the taxon.
Conobea vandellioides Benth. in Candolle, Prodr. 10:
391. 1846 Lindernia vandellioides (Benth.) Pen-
nell ex G. M. Barroso in Rodriguésia 27: 44. 1952
Stemodia vandellioides (Benth.) V. C. Souza, Fl.
Fanerog. Estado São Paulo 3: 316. 2003. – Originally
described from Brazil, it was transferred to Lindernia
by Barroso (1952), who compared it to L. diusa. The
generic description of Conobea in Bentham (1846)
states “Semina numerosa ovoidea striata”, indicating
that the endosperm is probably not alveolate, and the
species may therefore represent a member of Linder-
nias.str.
Ellobum montanum Blume, Bijdr. Fl. Ned. Ind. 14: 747.
1826 Vandellia ellobum Benth. in Candolle, Prodr .
10: 417. 1846 [non Vandellia montana (Blume)
Benth., l.c.: 415. 1846] Lindernia ellobum (Benth.)
Koord., Exkurs.-Fl. Java 3: 179. 1912 [non Linder-
nia montana Hiern, Cat. Afr. Pl. 1: 764. 1898, nec
Lindernia montana (Blume) Koord., l.c.: 178. 1912,
nom. illeg.] Didissandra elongata subsp. montana
(Blume) A. Weber & B. L. Burtt in Beitr. Biol. Pflan-
zen 70: 213. 1998 [Gesneriaceae].
Lindernia dianthera Sw., Prodr.: 92. 1788 Mecardonia
dianthera (Sw.) Pennell in Proc. Acad. Nat. Sci. Phil-
adelphia 98: 87. 1946 [Plantaginaceae].
Lindernia grossidentata O. Schwarz in Repert. Spec.
Nov. Regni Veg. 24: 95. 1927.
Lindernia montevidensis Spreng., Syst. Veg. 2: 769.
1825.
Lindernia neocaledonica S. Moore in J. Linn. Soc., Bot.
45: 372. 1921. – The holotype specimen (New Cal-
edonia, Tonine, 30 Sep 1914, R. H. Compton 1932,
BM), although very scanty, very much resembles L.
dubia, and it seems quite reasonable to tentatively
place L. neocaledonica as a synonym of that species.
It was probably introduced to New Caledonia as a
neophytic weed.
Lindernia sesamoides Spreng., Neue Entdeck. Pflanzenk.
1: 261. 1820.
Vandellia clausa F. Muell. in Trans. Philos. Inst. Victoria
3: 60. 1859 Lindernia clausa (F. Muell.) F. Muell.,
Fragm. 6: 102. 1868 Bonnaya clausa (F. Muell.)
Benth., Fl. Austral. 4: 499. 1868 Ilysanthes clausa
(F. Muell.) Urb. in Ber. Deutsch. Bot. Ges. 2: 436.
1884.
Vandellia pubescens Benth. in Candolle, Prodr. 10: 415.
1846 Lindernia pubescens (Benth.) F. Muell., Syst.
Census Austral. Pl.: 97. 1882 Pyxidaria pubescens
(Benth.) Kuntze, Revis. Gen. Pl. 2: 465. 1891.
Vandellia punctata Prain in J. Asiat. Soc. Bengal, Pt.
2, Nat. Hist. 72(2): 19. 1903 Lindernia punctata
(Prain) Mukerjee in J. Ind. Bot. Soc. 24: 131. 1945.
Vriesea nervosa Hassk. in Flora 25(2, Beibl.): 28. 1842
Lindernia nervosa (Hassk.) Koord., Exkurs.-Fl. Java
3: 179. 1912.
Vriesea sphaerocarpa Hassk. in Flora 25(2, Beibl.): 27.
1842 Lindernia sphaerocarpa (Hassk.) Koord.,
Exkurs.-Fl. Java 3: 179. 1912.
Acknowledgements
The first author (EF) would like to thank the Akademie
der Wissenschaften und Literatur Mainz for financial
support of the field trips to Rwanda and Madagascar. He
would like to thank the Rwanda Development Board,
Dept. of Conservation and Tourism for collecting and
export permissions. We are indebted to the sta of the
Parc Botanique et Zoologique Tsimbaza in Madagascar
for facilitating permissions to collect and export plant
material for scientific study. Our gratitude goes to the
General Commissioner of the CENAREST in Gabon,
Prof. Dr Franck Idiata for obtaining research and export
permits in Gabon. We are grateful to Dr Hugues Eyi
Ndong from CENAREST/Libreville for help in support
and organization of field work in Gabon. We would like
to thank the curators of the following herbaria for the
loan of specimens: BR, K, P, WAG. The Director and the
sta of the Bonn University Botanical Gardens provided
living material and opportunities to cultivate species of
Linderniaceae for further studies. Additional financial
support by DFG grant “Carnivory in Lamiales: under-
standing character evolution, substitution rate plasticity,
and genome miniaturization”, MU2875/2, to K.F.M. is
kindly acknowledged. We also thank two anonymous
reviewers for their comments on an earlier draft of this
paper.
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  • ... (Pennell 1935(Pennell , 1943Philcox 1968;Rahmanzadeh et al. 2004). But Fischer et al. (2013) reinstated the genera Bonnaya and Vandellia with one new genus Linderniella Eb. Fisch., Schäferh. ...
    ... based on phylogenetic study of the family. According to Fischer et al. (2013), the family comprises 17 genera and 253 species. In India, it is represented by 50 species belonging to five genera, Bonnaya, Craterostigma Hochst., Lindernia, Torenia L. and Vandellia (Sivarajan and Mathew 1983;Ratheesh Narayanan et al. 2012;Fischer et al. 2013;Sunil et al. 2014;Wang 2014, 2016;Prasad et al. 2014Prasad et al. , 2017Prasad and Sunojkumar 2014a, 2014b, 2015Sardesai et al. 2019;Shimpale et al. 2019). ...
    ... According to Fischer et al. (2013), the family comprises 17 genera and 253 species. In India, it is represented by 50 species belonging to five genera, Bonnaya, Craterostigma Hochst., Lindernia, Torenia L. and Vandellia (Sivarajan and Mathew 1983;Ratheesh Narayanan et al. 2012;Fischer et al. 2013;Sunil et al. 2014;Wang 2014, 2016;Prasad et al. 2014Prasad et al. , 2017Prasad and Sunojkumar 2014a, 2014b, 2015Sardesai et al. 2019;Shimpale et al. 2019). ...
    Article
    The article deals with the information on lectotypification of five names, Bonnaya bracteoides Blatt. & Hallb., B. micrantha Blatt. & Hallb., Torenia diffusa D.Don, Vandellia hookeri C.B. Clarke ex Hook.f., V. molluginoides Benth. and second-step lectotypification of Bonnaya pusilla Benth.
  • ... Philcox 1968Philcox , 2008Sivarajan & Mathew 1983;Yamazaki 1985Yamazaki , 1990Fischer 1992Fischer , 1995Fischer , 2004Lewis 2000). Based on DNA sequence data, Rahmanzadeh et al., (2005) transferred Bonnaya and 12 genera to the monophyletic family Linderniaceae which was also supported by Fischer et al., (2013). As currently circumscribed, Bonnaya is characterized by pinnate leaf venation, deeply five lobed calyx, two clavate staminodes without appendages and a liner cylindrical capsule (Fischer et al., 2013;Liang & Wang 2014). ...
    ... Based on DNA sequence data, Rahmanzadeh et al., (2005) transferred Bonnaya and 12 genera to the monophyletic family Linderniaceae which was also supported by Fischer et al., (2013). As currently circumscribed, Bonnaya is characterized by pinnate leaf venation, deeply five lobed calyx, two clavate staminodes without appendages and a liner cylindrical capsule (Fischer et al., 2013;Liang & Wang 2014). The twelve species currently accepted (Mabberley 2018) have a subcosmopolitan (B. ...
    ... The twelve species currently accepted (Mabberley 2018) have a subcosmopolitan (B. antipoda Druce 1914: 415) or pantropical distribution (Fischer 1992;Lewis 2000;Liang et al., 2012;Fischer et al., 2013). ...
    Article
    Bonnaya milindii (Linderniaceae), a new species from the Western Ghats of India is described and illustrated. The species is similar to Bonnaya sanpabloensis but differs in its petaloid staminodes and glabrous corolla.
  • ... Samenvatting -Schijngenadekruid (Lindernia dubia (L.) Pennell; Fig. 1, 2) is een kleine eenjarige plant, behorend tot een eigen familie (Fischer et al. 2013), de Linderniaceae, die nauw verwant is aan de Gratiolaceae en Plantaginaceae en vroeger werd geplaatst in de Scrophulariaceae s.l. De plant is haarloos en vormt geen rozet, groeit liggend tot opstijgend, met op doorsnede vierkante stengels tot 20 cm lang. ...
    ... It occurs in wet ditches, wet meadows, borders of ponds and lakes, depressions and along streams (Lewis 2000). Its current distribution has expanded to the entire North American continent, Central America and a large part of South America (Pennell 1935, Lewis 2000, Fischer et al. 2013. Nowadays it is a cosmopolitan weed; it is more or less naturalised in Japan (Yamazaki 1993), China and Taiwan (Shu 1998) and has been discovered in India recently (Krishnasamy & Arumugam 2015). ...
    ... However, the genus Lindernia All. itself, with approximately 100 species, turned out to be polyphyletic and has been split up by Fischer et al. (2013). The remaining genus Lindernia s. str., to which Lindernia dubia belongs, contains 30 species, the majority of them occurs in (sub-)tropical Africa and Southeast Asia; a few species are known from southeastern North America, the Carribean and Eurasia. ...
  • ... Vandellia L. and Craterostigma Hochst. Fischer et al. (2013), however, provided a satisfactory circumscription of Torenia by including some former members of Lindernia All. With this modern circumscription, Torenia contains about 50 species distributed mainly in Asia and Africa. ...
    ... Torenia godefroyi has long been treated as a variety of L. crustacea. However, Fischer et al. (2013) proposed a revised circumscription of the family Linderniaceae and placed L. crustacea under Torenia, within a separate group called the T. crustacea group. However, as shown here, T. godefroyi is morphologically distinct from T. crustacea by its distinctly winged calyx and the fruit which is completely included in the fruiting calyx. ...
    Article
    Torenia godefroyi, an Asian species overlooked since first described, has been considered as a variety of Lindernia crustacea (now treated as Torenia crustacea). However, detailed examination of fresh specimens, type and additional material clearly indicates that T. godefroyi is distinct from T. crustacea. Therefore, T. godefroyi is re-established here as a distinct species. Furthermore, the taxonomic identity of another Asian species, Torenia lindernioides, described from India is discussed. We observe that morphological characters of T. lindernioides overlap with T. godefroyi and suggest to synonymise it under T. godefroyi. An amended description, photographs, and notes about the distribution and habitat of T. godefroyi are provided.
  • ... & Kai Müll. In the 'Flora of Tropical East Africa' account of the Scrophulariaceae , Crepidorhopalon was not upheld, and species that had previously been transferred to that genus were treated in the genera Lindernia All. and Torenia L. However, molecular studies (Rahmanzadeh et al. 2005, Fischer et al. 2013) support the morphological evidence for recognition of Crepidorhopalon as a distinct genus, and it is now widely accepted (e.g., African Plant Database, continuously updated, Stevens 2001 onwards). ...
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    Full-text available
    The Crepidorhopalon whytei ( Linderniaceae ) species complex is revised using morphological analyses. Based primarily on variations in indumentum, floral morphology, corolla colour and seed morphology, four species are formally recognised within this group in eastern Africa. Crepidorhopalon whytei s.str. is widespread in the highlands of eastern Africa, extending from South Sudan and Ethiopia in the north through to western Tanzania in the south. A new combination in Crepidorhopalon is made for Lindernia flava (= C. flavus ), which is confined to the Manica Highlands of the Mozambique-Zimbabwe border. Two new species are described, C. namuliensis which is known only from Mt Namuli in northern Mozambique and C. kwaleensis which is known only from the coastal lowlands of southeast Kenya. A fifth, imperfectly known species is documented from the Nguru Mountains of Tanzania where it is so far known from a single collection. Three names are lectotypified. The habitat requirements and distribution are documented and the extinction risk is assessed for each species. Crepidorhopalon flavus is assessed as globally Vulnerable and C. kwaleensis as globally Endangered, while C. namuliensis and C. whytei are currently considered to be of Least Concern, although the latter is declining markedly in parts of its range. The botanical importance of the key sites for the newly recognised taxa is discussed.
  • ... The generic concept of Lindernia Allioni (1766: 178) sensu lato has been revised and updated by Fischer et al. (2013) as a result of their molecular phylogenetic studies on the family Linderniaceae. Their study proposed several taxonomic changes within the family including (1) the re-establishment of Bonnaya Link & Otto (1820: 25) and Vandellia P. Browne ex Linnaeus (1767: 384) which were earlier been treated under Lindernia s. l., (2) inclusion of some taxa into Craterostigma Hochstetter (1841: 668) and Torenia Linnaeus (1753: 619) (T. ...
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    The taxonomic identity of two Indian species of Lindernia s. str. is revised and updated. Thus, the following names at the species rank are reduced to synonymy of what follows them in parenthesis viz. L. calemeriana (L. srilankana) and L. parviflora (L. dubia). Lectotype of L. srilankana is also designated for the first time.
  • Technical Report
    Full-text available
    A product of the Bishop Museum’s Hawaii Biological Survey, this special edition of the Hawaiian Naturalized Vascular Plants Checklist is an authority file of names of accepted naturalized angiosperm, gymnosperm, and fern/lycophyte taxa in the Hawaiian Islands, updated through February 2019. The file is maintained as a Microsoft Access database in the Department of Natural Sciences/Botany, Bernice P. Bishop Museum. The foundation for the dicot and monocot sections of the checklist is the Manual of the Flowering Plants of Hawai‘i (Wagner et al. 1990, 1999, hereafter referred to as the Manual), while that of the fern/lycophyte section is Hawai‘i’s Ferns & Fern Allies (Palmer 2003). Naturalized records of gymnosperms have been included since being first reported in 2002 (Oppenheimer 2002). This checklist is an offshoot of the Hawaiian Native and Naturalized Vascular Plants Checklist (Imada 2012), which is periodically updated to incorporate newly published native and naturalized island and state records, nomenclatural and taxonomic changes, and newly described native taxa. Many of the additions are garnered from the annual Bishop Museum publication series, Records of the Hawaii Biological Survey (1994 and continuing). Additions of new island or state records are based on published sources supported by voucher specimens; not included are citations based on anecdotal reports or casual observations, or on unpublished checklists or manuscripts. This checklist tracks and attempts to maintain conformity with the nomenclature and taxonomy of the Smithsonian Institution’s National Museum of Natural History Flora of the Hawaiian Islands website (https://naturalhistory2.si.edu/botany/hawaiianflora/), which periodically publishes online supplements that update names and distributions since publication of the Manual in 1990. At this writing, the latest available supplement is dated 12 April 2012 (version 1.3, http://botany.si.edu/pacificislandbiodiversity/hawaiianflora/Hawaiian_vas cular_plant_updates_1.3.pdf). Pending nomenclatural or taxonomic changes or in-press taxa are generally excluded until they are published, although exceptions are made. Proposed additions or changes to the checklist are reviewed and evaluated by staff botanists, often with outside consultation, before acceptance. While the formal checklist lists only currently accepted names, Appendix A allows the user to track down the current disposition of scientific names no longer used in this checklist. It is arranged alphabetically by the old scientific name. Appendix B extracts all of the newly naturalized taxa published since the Manual was released in 1990. Appendix C similarly extracts all of the questionably naturalized taxa included in this checklist. An index to accepted families and genera is included to assist the user in navigating the checklist. Pagination is provided for the first page on which an accepted family or genus is listed in the main body of checklist (Sections A and B). Also integrated into the index is a finding list of families and genera no longer treated in Hawai‘i’s naturalized flora, with links to their present dispositions. A complete bibliography is included for all citations included in the text.
  • Article
    Full-text available
    During the exceptionally low water level of the Danube River in the 2018 vegetation season the third locality of Lindernia dubia was found in Slovakia. This alien species, native to North America, grew east of the Štúrovo town, in the confluence area of the rivers Danube and Hron. Its population counted ca. 50 plants which grew sporadically on the exposed, muddy shores in the vegetation of the association Cyperetum micheliani, class Isoëto-Nanojuncetea. Except the characteristic taxa of this class, the stands were enriched with several nitrophilous species of the class Bidentetea. Lindernia dubia was most frequently found in the same vegetation type in the surrounding countries of Central Europe.
  • Article
    The new species Craterostigma loitense from the Loita Hills in Kenya is described and illustrated. It differs from Craterostigma purpureum in the two abaxial staminodes, the basal tufts of hairs on the anthers, and the subentire to serrulate leaf margin.
  • Article
    Full-text available
    Three new species from the genus Lindernia are described: L. stantonii Wannan, L. beasleyi Wannan from northern Queensland (Cape York Peninsula), and L. barkeri Wannan from northern Western Australia (Kimberley). Illustrations of flowers, fruits, seeds, leaves, stem anatomy, type specimen and a distribution map are provided for all species. Notes on habitat and conservation status are also provided for each. A key to Queensland species is provided.
  • Article
    The genus Lindernia All., as treated here, is represented in the New World by fourteen species. Six species are members of the Section Brachycarpae, five of which are postulated to be native in the New World. One native and one introduced species are in each of the sections Lindernia and Tittmannia. Old World introductions also include two species in Section Torenioides and one species each in sections Numularia and Bonnaya. Keys are provided to the fourteen species and four varieties that occur in the New World. Descriptions and distribution maps are included for the native taxa and most of the introduced species, and most are also illustrated. Nomenclatural changes include a new variety and new combinations for a species and a variety. Lindernia viscosa is reported for the first time in the New World.
  • Article
    Craterostigma alatum Hepper and C. longicarpum Hepper are described as new species from East Africa.
  • Article
    While checking some names in Scrophulariaceae in connection with preparation of the list of genera accepted at Kew, my attention was drawn to Strigina Engl., a monotypic genus from NW Tanzania whose sole species, S. pusilla Engl., was described from a single collection (Stuhlmann 3367). No other collections have since been identified with this species. The holotype was destroyed in Berlin, but an isotype is present at Kew. I was struck by the similarity between this plant and Lindernia acicularis E. Fischer, a species recently described from the same area. Closer examination has shown conclusively that Strigina pusilla is indeed the same species as Lindernia acicularis. The name Lindernmia pusilla (Willd.) Boldingh is validly published for a different species that comes from Asia, so that this combination is not available for the present plant; its correct name is Lindernia acicularis. According to the original description of Engler (1897, p. 516), Strigina should be placed next to Striga, differing only in having the stamens inserted in the throat of the corolla (inserted in the corolla-tube in Striga) and in having two of the stamens reduced to staminodes (having four fertile stamens in the tribe). It seems odd that Engler should have placed Strigina in the tribe Gerardicae whilst at the same time distinguishing it by these two characters which separate it from all other African genera within Gerardieae. These two are among the characters normally used to distinguish the tribe Gratioleae from Gerardieae. Engler's judgement was, however, accepted by Hemsley & Skan (1906, p. 266), who kept Strigina in the tribe Gerardieae as the only genus in that tribe with only two stamens. It also seems that this position for Strigina has never been questioned by any subsequent botanists, despite the fact that it appears to be a completely typical Lindernia.