Willdenowia 43 – 2013 209 EBerhard Fischer1*, Bastian Schäferhoff2 & Kai Müller2 The phylogeny of Linderniaceae – The new genus Linderniella, and new combinations within Bonnaya, Craterostigma, Lindernia, Micranthemum, Torenia and Vandellia Abstract Fischer E., Schäferhoff 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 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. Additional key words: trnK, matK, Lamiales, poikilohydric plants, desiccation tolerance Introduction Research interest in Lindernia and related genera has increased 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 Nanocyperion communities in ponds) (Šumberová & al. 2012). Another focus has been the research on poikilohydric 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 circumscribed as Linderniaceae was first considered by Bentham (1846) to represent a subtribe within the Gratio­leae. Based on Lindernia, the circumscription of genera was, however, controversially discussed resulting in a complex taxonomic history with proposals to unite all species into a broadly defined genus Lindernia or splitting them into 13 different 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 Arta­nema, 1 Institut für Integrierte Naturwissenschaften – Biologie, Universität Koblenz-Landau, Universitätsstr. 1, 56070 Koblenz, Germany; *e-mail: efischer@uni-koblenz.de (author for correspondence). 2 Institut für Evolution and Biodiversität, Universität Münster, Hüfferstr. 1, 48149 Münster, Germany; e-mail: schaeferhoff@unimuenster.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 horombensis, Madagascar; E: Linderniella gracilis, Rwanda; F: Linderniella brevidens, Kenya; G: Chamaegigas intrepidus, Namibia; H: Vandellia diffusa, Gabon; I, K: Torenia stolonifera, Madagascar; L: Torenia thouarsii, Gabon; M: Torenia crustacea, Gabon. – All photographs by E. Fischer except G by N. Jürgens. Willdenowia 43 – 2013 211 Fig. 2. Linderniaceae – A: Lindernia viguieri, Madagascar; B: Lindernia rotundifolia, Madagascar; C: Lindernia benthamii (= Bryodes micrantha), Madagascar; D: Lindernia madagascariensis (= Psammetes madagascariensis), Madagascar; E: Crepidorhopalon microcarpaeoides, Madagascar; F: Crepidorhopalon tenuis, Congo-Kinshasa; G, H: Crepidorhopalon whytei, Rwanda; I: Hartliella capitata, Congo-Kinshasa; K: Stemodiopsis ruandensis, Rwanda. – All photographs by E. Fischer except I by M. Schaijes. 212 Fischer & al.: The phylogeny of Linderniaceae 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. Bonnaya, Curanga Juss. (= Picria Lour.), Hemianthus, Ilysanthes, Lindernia, Micranthemum, Peplidium Delile, Torenia (Fig. 1I, K) and Vandellia as genera within subtribe 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 Ilysanthes (palmate venation). Urban (1884) lumped Bonnaya and Ilysanthes together with Bazina Raf. into Ilysanthes with 2 fertile stamens and Ilyogeton, Tittmannia and Vandellia with 4 fertile stamens into Lindernia. This concept of separating Lindernia and Ilysanthes, was also accepted 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 Africa. 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 apparently 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 Ilysanthes and Lindernia into a broadly circumscribed genus Lindernia. This was accepted by the majority of authors (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 monophyletic. Yamazaki (1954a – b, 1955) already proposed a division into Vandellia with pinnate leaf venation, serrate leaf margin, a 1-celled chalazar haustorium, and alveolate 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) Willdenowia 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 Crepidorhopalon (Fischer 1992). This highly unnatural concept of Crate­rostigma 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), nonrosulate habit and anatomically highly complex clavate hairs on the lower corolla-lip comprising a distinct multicellular base into the new genus Crepidorhopalon. Also plants formerly placed in Lindernia, Torenia or even Stemodiopsis (Fig. 2K) showed the above mentioned characters and were subsequently transferred to Crepidorhopalon (Fischer 1992, 1995, 1997a, 1999a – b). A further new genus of suffruticose metallophytes from Central Africa (Katanga), Hartliella Eb. Fisch. (Fig. 2I), was described and based on former species of Lindernia (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 including members of the former tribe Lindernieae (Rahmanzadeh & al. 2005) revealed that they represented a new family, Linderniaceae as a further lineage distinct from Scrophula­riaceae and Plantaginaceae. It also became evident that Lindernia sensu Pennell is not monophyletic, and that Crepidorhopalon and Torenia represent different clades. Linderniaceae as a separate lineage was confirmed by Albach & al. (2005), who included also Micranthemum, by Oxelman & al. (2005), who additionally included Stemodiopsis, and by Schäferhoff & 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 different genera of Linderniaceae as we could obtain. Also several species per genus were included 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 phylogeny (Schäferhoff & al. 2010). Within Linderniaceae, the genera Stemodiopsis and Micranthemum were included to show their exact position. Voucher information and GenBank accession numbers of taxa included in this study are listed in Table 1. 213 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 markers, 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, Schäferhoff & al. 2010); for the 3´part the primers ACmatK500F (TTCTTCTTTGCATT TATTACG, Müller & Borsch 2005), LindmatK1580F (TCAATTCATTCAACWTTTCCC, Schäferhoff & al. 2010), and trnK2R (AACTAGTCGGATGGAGTAG, Johnson & Soltis 1995) were used. Reactions were performed 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 carried out on a Biometra T3 thermocycler using the following 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 random 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 (Swofford 1998). Tree evaluation was performed with 10 000 bootstrap 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- Linderniaceae Linderniaceae Linderniaceae Linderniaceae Linderniaceae Linderniaceae Linderniaceae Linderniaceae Crepidorhopalon tenuis Crepidorhopalon whytei Lindernia benthamii (Bryodes micrantha) Lindernia bryoides Lindernia dubia Lindernia madagascariensis (Psammetes madagascariensis) Lindernia procumbens Lindernia rotundata Artanema angustifolium Artanema fimbriatum Bonnaya ciliata (Lindernia ciliata) Bonnaya multiflora (Lindernia bonatii) Bonnaya ruellioides (Lindernia ruellioides) Chamaegigas intrepidus Craterostigma crassifolium (Lindernia crassifolia) Craterostigma engleri (Lindernia welwitschii) Craterostigma hirsutum Craterostigma newtonii (Lindernia oliveriana) Craterostigma nummulariifolium (Lindernia nummulariifolia) Craterostigma plantagineum Craterostigma pumilum Craterostigma pusillum (Lindernia acicularis) Craterostigma sp. nov. Craterostigma yaundense (Lindernia yaundensis) Crepidorhopalon bifolius Crepidorhopalon goetzei Crepidorhopalon microcarpaeoides Crepidorhopalon perennis Linderniaceae Linderniaceae Linderniaceae Linderniaceae Linderniaceae Linderniaceae Linderniaceae Linderniaceae Linderniaceae Linderniaceae Linderniaceae Linderniaceae Linderniaceae Linderniaceae Linderniaceae Linderniaceae Linderniaceae Linderniaceae Linderniaceae Linderniaceae Species Family Portugal, Baixo Alentejo, H. Kalheber 99-1990 (BONN) Rwanda, Musanze, E. Fischer 7320 (KOBL) Rwanda, Nyamata, E. Fischer s.n. (BG Bonn 19988) Kenya, Nairobi, E. Fischer 8759 (KOBL) Rwanda, Nyarubuye, E. Fischer 4230 (KOBL) Angola, Huilla, T. Leyens 401 (BONN) Cameroon, Yaunde, S. Porembski 3858 (KOBL) Congo-Kinshasa, Katanga, Lubumbashi, M. Faucon s.n. (BRLU, KOBL) Congo-Kinshasa, Katanga, Lubumbashi, M. Faucon s.n. (BRLU, KOBL) Madagascar, Ambalavao, E. Fischer 10296 (KOBL) Congo-Kinshasa, Katanga, Lubumbashi, M. Faucon s.n., E. Fischer 16 (BRLU, KOBL) Congo-Kinshasa, Katanga, M. Faucon s.n., E. Fischer 3 (BRLU, KOBL) Rwanda, Butare, E. Fischer 7657 (KOBL) Madagascar, Antsirabé, E. Fischer 10258 (KOBL) Madagascar, Ambositra, E. Fischer 10240 (KOBL) Germany, F. Müller s.n. (BONN) Madagascar, Ambositra, E. Fischer 10234 (KOBL) Singapore, Perak, Kampong Permatang Serai, Dindings, J. Sinclair 9876 (M) Germany, Botanical Gardens Bonn 0-BONN-15545, T. Borsch 3790 (BONN) Thailand, Koh Samui, E. Fischer EF Th 677 (KOBL) Madagascar, Ambositra, E. Fischer 10236 (KOBL) Thailand, Koh-Samui, E. Fischer EF Th 690 (KOBL) Südwestafrika [Namibia], Distr. Omaruru, Ohere-Oos, H. Merxmüller 1580 (M) Angola, Huilla, T. Leyens 347 (BONN) Angola, Huilla, Lobin & al. s.n. (BONN) Rwanda, Nyarubuye, 9003 (KOBL) Rwanda, Lac Ihema, Akagera National Park, E. Fischer s.n. (KOBL) Nyungwe National Park, Cyamundongo, E. Fischer 10256 (KOBL) Voucher Table 1. Species, voucher information, and GenBank accession numbers of the sequences used in the present study. FR728409 AY667469 FR728413 AY667463 FN773532 FR728404 FR728411 FR728410 FR728387 AY667462 AY667465 FR728389 FR728388 FR728417 FR728419 FR728418 FR728415 FR728403 AY667460 FR728401 FR728408 FR728402 FR728398 FR728390 FR728391 AF531776 FR728393 FR728392 GenBank accession number continued on next page this study Rahmanzadeh & al. 2005 this study Rahmanzadeh & al. 2005 Schäferhoff & al. 2010 this study this study this study this study Rahmanzadeh & al 2005 Rahmanzadeh & al 2005 this study this study this study this study this study this study this study Rahmanzadeh & al. 2005 this study this study this study this study this study this study Rahmanzadeh & al. 2005 this study this study Citation 214 Fischer & al.: The phylogeny of Linderniaceae Species Lindernia rotundifolia Lindernia microcalyx Lindernia viguieri Linderniella brevidens (Lindernia brevidens) Linderniella gracilis (Lindernia exilis) Linderniella horombensis (Lindernia horombensis) Linderniella pusilla (Lindernia philcoxii) Linderniella pygmaea (Lindernia pygmaea) Micranthemum umbrosum Stemodiopsis ruandensis Torenia crustacea (Lindernia crustacea) Torenia diffusa (Torenia vagans) Torenia stolonifera Vandellia diffusa (Lindernia diffusa) Vandellia pusilla (Lindernia pusilla) Vandellia senegalensis (Lindernia senegalensis) Vandellia subracemosa (Lindernia subracemosa) Byblis gigantea Lindl. Byblis lamellata Conran & Lowrie Byblis liniflora Salisb. Family Linderniaceae Linderniaceae Linderniaceae Linderniaceae Linderniaceae Linderniaceae Linderniaceae Linderniaceae Linderniaceae Linderniaceae Linderniaceae Linderniaceae Linderniaceae Linderniaceae Linderniaceae Linderniaceae Linderniaceae Byblidaceae Byblidaceae Byblidaceae Madagascar, Ambositra, E. Fischer 10241 (KOBL) Germany, Botanical Garden Mainz, E. Fischer s.n. (BONN) Madagascar, Antsirabe, E. Fischer 10267 (KOBL) Kenya, Teita Hills, E. Fischer 8022 (KOBL) Rwanda, Nyarubuye, E. Fischer 12225 (KOBL) Madagascar, Ambalavao, E. Fischer 10298 (KOBL) Rwanda, Nyarubuye, E. Fischer 553/87 (KOBL) Madagascar, Ambalavao, E. Fischer s.n. (KOBL) Germany, Botanical Gardens Bonn, Schäferhoff 43 (BONN) Rwanda, Rugarama, E. Fischer 10352 (KOBL) Thailand, Koh-Samui, E. Fischer EF Th 692 (KOBL) U.K., Royal Botanical Garden Edinburgh, B. L. Burtt s.n. (BONN) Madagascar, Ranomafana, E. Fischer 10249 (KOBL) Dominican Republic, T. Borsch 3837 (B) Thailand, Koh-Samui, E. Fischer, EF Th 691 (KOBL) Gabon, Makokou, E. Fischer s.n. (KOBL) Rwanda, Uwinka, E. Fischer 1350 (BG Bonn 19990-2) (KOBL) Germany, Botanical Gardens Bonn 0-BONN-25514, K. Müller 733 (MSUN) Germany, Botanical Gardens Bonn 0-BONN-25513, K. Müller s.n. (MSUN) Germany, Botanical Gardens Bonn 0-BONN-16008, K. Müller s.n. (MSUN) Voucher FR728406 AY667468 FR728407 FN773545 FR728396 FR728394 AY667466 FR728395 FR773548 FN773559 FR728405 AF531812 FN773561 FR728399 FR728397 FR822526 FR728400 AF531774 FR773534 FR773535 GenBank accession number this study Rahmanzadeh & al. 2005 this study Schäferhoff & al. 2010 this study this study Rahmanzadeh & al. 2005 this study Schäferhoff & al. 2010 Schäferhoff & al. 2010 this study Rahmanzadeh & al 2005 Schäferhoff & al. 2010 this study this study this study this study Müller & al. 2004 Schäferhoff & al. 2010 Schäferhoff & al. 2010 Citation Willdenowia 43 – 2013 215 216 portion of invariable sites, and a uniform topological prior. Using the formula provided by Brown & al. (2010), we calculated an exponential rate parameter for the branchlength prior based on an average branch length estimate from an initial neighbor joining analysis in PAUP. Four categories were used to approximate the gamma distribution. 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 conservatively 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 total alignment comprised 2742 characters; 92 characters were excluded from phylogenetic analyses due to uncertain 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 sequences 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 phylogeny is given in Fig. 4, together with posterior probabilities (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, Micranthemum and Stemodiopsis (Schäferhoff & 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 Linderniaceae. 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 Fischer & al.: The phylogeny of Linderniaceae 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-supported 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. diffusa 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) com­prises L. oliveriana 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 supported (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 Stemodiopsis and other taxa from Linderniaceae. The abaxial filaments are curved or twisted and thus match the general staminal morphology of Linderniaceae. Stemodiopsis 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 aulacospermous 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. However, this is a feature that regularly occurs in Lindernia and Willdenowia 43 – 2013 217 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, numbers 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. also Vandellia. The type species of Lindernia, L. procumbens usually bears small cleistogamous flowers, and the form with chasmogamous flowers occurring mainly in the tropics has even been described as a different 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 glandular 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 studied, Micranthemum is not included in Lindernia. Also several morphological differences ­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 bothrospermous seeds with alveolate endosperm of the Torenia type (Hartl 1959; Fischer 1992). Torenia is maintained here, and L. crustacea, suggested to have affinities to Torenia by some previous authors, is shown to be indeed a member 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 ciliata, L. ruellioides and Artanema form a moderately supported 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 filament 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 218 1990) it was used as a subgenus of Lindernia. The species of Bonnaya share abaxial staminodes, pinnate leaf venation and bothrospermous seeds. The well-supported clade around Lindernia diffusa can be called Vandellia as L. diffusa is the type species of that genus. Yamazaki (1954a – b, 1955) already proposed a division into Vandellia with pinnate leaf venation, serrate leaf margin, a 1-celled chalazar haustorium, and alveolate endosperm (bothrospermous seeds), and Lindernia 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 morphological 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 morphologically very distinct plant with seeds with non-alveolate 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, Plantago-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 yelloworange 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. oliveriana, 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 aulaco­spermous seeds comprises four species endemic to the copper belt of Katanga province in Congo-Kinshasa and N Zambia (Fischer 1999b, 2004). Schizotorenia with two species from Indo-Malesia also has aulacospermous seeds (Fischer 2004). The monotypic Hemiarrhena plantaginea from NW Australia, the monotypic Legazpia polygonoides from SE Asia, Micronesia and New Guinea, the monotypic Pierranthus capitatus from SE Asia and Scolophyllum, with three species from Thailand to Cambodia and Vietnam, have bothrospermous seeds (Fischer Fischer & al.: The phylogeny of Linderniaceae 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. 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 presence 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, Crate­rostigma, Lindernia, Micranthemum, Torenia and Vandellia. The species not represented in the molecular analysis are assigned to genera according to their morphology. Willdenowia 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 corolla 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 Ca­ribbean . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 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 furrows (aulacospermous), endosperm star-shaped in transverse section . . . . . . . . . . . . . . . . . . . . . . . . . 5 4. Aquatic plants with basal rosette, with 2-lobed complex 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 flowers 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); sometimes 2 stamens reduced to staminodes . . . . . . . . . 8 6. Plants procumbent, creeping; leaf venation pinnate; flo­wers 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 occasionally 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 basal 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 219 weakly curved with a small basal knee-like appendage present or absent; perennial plants with large subterraneous woody rhizomes on heavy metal soil . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6. Hartliella 8. Rosulate herbs; fertile stamens 2, one pair reduced to filiform staminodes, anthers with 2 unequal mucronate 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 often 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 distant; abaxial stamens usually reduced to staminodes, rarely fertile (in L. bolusii, L. boutiqueana, L. brevidens and L. cerastioides) . . . . . . . . 10. Linderniella 13. Leaf venation palmate . . . . . . . . . . . . . . . . . . . . . . 14 – Leaf venation pinnate . . . . . . . . . . . . . . . . . . . . . . . 15 14. Inflorescence capitate, enclosed by closely overlapping large involucral leaves with semiorbicular base and long acuminate apex, differing from linear-lanceolate stem leaves . . . . . . . . . . . . 12. Pierranthus – Inflorescence a lax raceme, rarely capitate (C. crassifolium), but then involucral leaves not differing 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, generally 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 17. Plants large; stem generally more than 50 cm tall; corolla 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 apical 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 Graham) 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. Distribution — Australia. 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 Lanka, India, Thailand, Cambodia, Vietnam, Malaysia, Indonesia (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]. Fischer & al.: The phylogeny of Linderniaceae 1. Bonnaya aculeata (Bonati) Eb. Fisch., Schäferh. & Kai Müll., comb. nov. ≡ Ilysanthes aculeata Bonati in Bull. Soc. Bot. France 55: 541. 1908 ≡ Lindernia aculeata (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 verbenifolia Colsm., Prodr. Descr. Gratiol.: 9. 1793. Distribution — Sri Lanka and India to China, Japan, 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 ≡ Ilysanthes 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. Willdenowia 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 ≡ Lindernia 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.) Pennell in Brittonia 2: 182. 1936 ≡ Ilysanthes ruellioides (Colsm.) Kuntze, Revis. Gen. Pl. 2: 462. 1891. = Gratiola reptans Roxb., Fl. Ind. 1: 140. 1820 ≡ Bonnaya reptans (Roxb.) Spreng., Syst. Veg. 1: 41. 1824 ≡ Lindernia reptans (Roxb.) F. Muell., Descr. Notes Papuan 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 zan­zibarica Eb. Fisch. & Hepper in Kew Bull. 46: 529. 1991. Distribution — Kenya, Tanzania (including Zanzibar). 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. 221 = 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 extending 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. lanceo­latum 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, Somalia, Congo-Kinshasa, Rwanda, Burundi, Uganda, Kenya, Tanzania, Angola, Zambia, Zimbabwe, Botswana, Namibia, 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. Borana 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 Distribution — Nigeria, Cameroon, Sudan, Ethiopia, Uganda, Kenya, Tanzania. 11. Craterostigma angolense (Skan) Eb. Fisch., Schäferh. & Kai Müll., comb. nov. ≡ Torenia angolensis 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 wel­witschii Engl. in Bot. Jahrb. Syst. 23: 501. 1897] ≡ Lindernia welwitschii (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 gossweileri 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 combined under Lindernia would form a homo­nym 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 kigomensis 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 lindernioides (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. Distribution — Togo, Nigeria, Central African Republic, Congo-Kinshasa, Rwanda, Sudan, Ethiopia, Uganda, Kenya, Tanzania, Zambia, Zimbabwe, Malawi, Angola. Fischer & al.: The phylogeny of Linderniaceae 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. ≡ Lindernia 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 num­mulariifolia 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., Schäf­erh. & 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 stuhlmannii Engl., Pflanzenw. Ost-Afrikas C: 357. 1895. Distribution — Tanzania. 22. Craterostigma sudanicum (Eb. Fisch. & Hepper) Eb. Fisch., Schäferh. & Kai Müll., comb. nov. ≡ Lindernia sudanica Eb. Fisch. & Hepper in Kew Bull. 46: 530. 1991. Distribution — Sudan, Uganda. 23. Craterostigma syncerus (Seine, Eb. Fisch. & ­Barth­lott) 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 Craterostigma longicarpum Hepper in Kew Bull. 42: 945. 1987]. Distribution — Tanzania. Willdenowia 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 yaundensis S. Moore in J. Bot. 57: 216. 1919 ≡ Lindernia yaundensis (S. Moore) Eb. Fisch. in Trop. Subtrop. Pflanzenwelt 81: 286. 1992. Distribution — Cameroon. 5. Crepidorhopalon Eb. Fisch. in Feddes Repert. 100: 443. 1989. – Type: C. schweinfurthii (Oliv.) Eb. Fisch. 1. Crepidorhopalon affinis (De Wild.) Eb. Fisch. ex Govaerts, World Checkl. Seed Pl. 3(1): 20. 1999 ≡ Torenia affinis 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 chironioides 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.) Mukerjee 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, Nigeria, Cameroon, Sudan, Congo-Kinshasa, Uganda, Tanzania, 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, Zam­bia, Malawi. 7. Crepidorhopalon gracilis (Pilg.) Eb. Fisch. in Feddes 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. 223 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 involucrata 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 latibracteatum 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 Distribution — Congo-Kinshasa, Burundi, Uganda, ­ enya, Tanzania, Zambia, Malawi, Zimbabwe. K 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 schweinfurthii 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 Feddes 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, Malawi, 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 tenuifolia 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. Distribution — Angola. 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. Fischer & al.: The phylogeny of Linderniaceae = 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. suffruticosa (Lisowski & Mielcarek) 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 bampsii Eb. Fisch. in Bull. Jard. Bot. Natl. Belg. 59: 449. 1989. Distribution — Congo-Kinshasa. 2. Hartliella capitata (Eb. Fisch.) Eb. Fisch. in Trop. Subtrop. Pflanzenwelt 81: 211. 1992 ≡ Lindernia capitata Eb. Fisch. in Bull. Jard. Bot. Natl. Belg. 59: 448. 1989. Distribution — Congo-Kinshasa, Zambia. 3. Hartliella cupricola Eb. Fisch. in Trop. Subtrop. Pflanzenwelt 81: 207. 1992. Distribution — Congo-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. Distribution — Congo-Kinshasa. 7. Hemiarrhena Benth., Fl. Austral. 4: 518. 1868. – Type: H. plantaginea (F. Muell.) Benth. The genus has bothrospermous seeds and shows relationships to Bonnaya. The abaxial stamens are reduced to staminodes and the leaves show a pinnate venation. 1. Hemiarrhena plantaginea (F. Muell.) Benth., Fl. Austral. 4: 518. 1868 ≡ Vandellia plantaginea F. Muell. in Trans. & Proc. Philos. Inst. Victoria 3: 62. 1859 ≡ Lindernia 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., Scroph. Ind.: 39. 1835. = Legazpia mucronulata (Benth.) T. Yamaz. in J. Jap. Bot. 30: 359. 1955 ≡ Torenia mucronulata Benth. in Candolle, 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 differs in the suborbicular calyx with 3 broad wings between the ridges, auriculate at base and 3-dentate at apex (Fischer 2004). Torenia has a calyx with 5 wings. Willdenowia 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 madagascariensis (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. Distribution — Australia. 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 Candolle, 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 Field­iana, Bot. 28: 519. 1953. = Lindernia barrosorum L. B. Sm. in Los Angeles County 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 Harvey, 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 congesta A. Raynal in Adansonia, n.s, 7: 348. 1967. 225 Distribution — Senegal (Cap-Vert), Tanzania. 9. Lindernia dubia (L.) Pennell in Acad. Nat. Sci. Philadelphia 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 gratioloides Benth. in Candolle, Prodr. 10: 419. 1846, nom. illeg. ≡ Lindernia gratioloides J. Lloyd & Foucaud, Fl. Ouest France, ed. 4: 246. 1886, nom. illeg. ≡ Limnophila 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 anagallidea 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, Honduras, 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 Orissa 4: 635. 1922 ≡ Gratiola hyssopioides L., Mant. Pl. 2: 174. 1771 ≡ Ilysanthes hyssopioides (L.) Benth. in Candolle, 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. illeg. [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 already 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 Distribution — Angola. 15. Lindernia madagascariensis (Bonati) Eb. Fisch., Schäferh. & Kai Müll., comb. nov. ≡ Bryodes madagascariensis 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 & Nandakumar 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 differs 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 complex is required. 19. Lindernia minima (Benth.) Mukerjee in J. Ind. Bot. Soc. 24: 132. 1945 ≡ Ilysanthes minima Benth. in Candolle, Prodr. 10: 420. 1846. Distribution — India. 20. Lindernia monroi (S. Moore) Eb. Fisch. in Trop. Subtrop. 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: Addit. [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. saxicola (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 ≡ Ilysanthes 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. Fischer & al.: The phylogeny of Linderniaceae = 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, Rwanda, Burundi, Uganda, Kenya, Tanzania, Zambia, Malawi, Zimbabwe, Mozambique, Botswana, Namibia, South Africa, 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. Distribution — Nigeria, Cameroon, Ethiopia, CongoKinshasa, 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, Nicaragua, 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 Castanea 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. Willdenowia 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. brevidente 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 bothrospermous. 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 (Hiern) Eb. Fisch. in Trop. Subtrop. Pflanzenwelt 81: 304. 1992, comb. inval. Distribution — Angola. 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. Distribution — Ethiopia, Uganda, Kenya, South Africa. 3. Linderniella boutiqueana (Germain) Eb. Fisch., Schäferh. & Kai Müll., comb. nov. ≡ Lindernia boutiqueana 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 Oliver & al., Fl. Trop. Afr. 4(2): 339. 1906. Distribution — Kenya, Tanzania. 5. Linderniella cerastioides (Bonati) Eb. Fisch., Schäf­ erh. & Kai Müll., comb. nov. ≡ Craterostigma cerastioides 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 Oliver & al., Fl. Trop. Afr. 4(2): 349. 1906 ≡ Lindernia exilis 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-Kinshasa, Rwanda, Burundi. 7. Linderniella hartlii (Eb. Fisch. & Hepper) Eb. Fisch., Schäferh. & Kai Müll., comb. nov. ≡ Lindernia hartlii 227 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 ho­rom­ bensis Eb. Fisch. in Bull. Mus. Natl. Hist. Nat., B, Adansonia, ser. 4, 17: 238. 1995. Distribution — Madagascar. 9. Linderniella mbalaensis (Eb. Fisch.) Eb. Fisch., Schäferh. & Kai Müll., comb. nov. ≡ Lindernia mbalaensis 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., Schäf­erh. & 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 pusilla Benth., Scroph. Ind.: 33. 1835, nec Bonnaya pusilla Griff., Not. Pl. Asiat. 4: 107. 1854] ≡ Ilysanthes pusilla Urb. in Ber. Deutsch. Bot. Ges. 2: 435. 1884 ≡ Lindernia 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 replacement name. Fischer (1992) transferred it to Lindernia, proposing L. philcoxii as a replacement name, because the name L. pusilla already existed, but I. ugandensis was cited as a synonym, which made the name 228 L. philcoxii nomenclaturally superfluous and therefore illegitimate. Careful study of the type of I. ugandensis showed that it is different from I. pusilla. Thus Philcox (2008) proposed L. serpens as another replacement name. The replacement name published here in Linderniella 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 pygmaeum 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., Schäf­erh. & Kai Müll., comb. nov. ≡ Bonnaya trichotoma Oliv. in Trans. Linn. Soc. London 29: 121. 1875 ≡ Ilysanthes trichotoma (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, Sierra Leone, Liberia, Côte d’Ivoire, Burkina Faso, Ghana, Togo, Nigeria, Cameroon, Central African Republic, Su­ dan, Ethiopia, Congo-Kinshasa, Rwanda, Burundi, Uganda, Kenya, Tanzania, Zambia. 15. Linderniella ugandensis (Skan) Eb. Fisch., Schäf­erh. & 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 wilmsii 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, Ugan­da, 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. ≡ Micranthemum micranthemoides (Nutt.) Wettst. Fischer & al.: The phylogeny of Linderniaceae = Amphiolanthus Griseb., Cat. Pl. Cub.: 186. 1866. – Type (designated by Pennell in Proc. Acad. Nat. Sci. Philadelphia 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 arenarioides Griseb., Cat. Pl. Cub.: 187. 1866 ≡ Globifera arenarioides (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 ≡ Hemianthus 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 ≡ Hemianthus elatinoides Griseb., Cat. Pl. Cub.: 185. 1866 ≡ Globifera 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., Schäf­erh. & 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). Willdenowia 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 Fieldiana, 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 Fieldiana, Bot. 34: 124. 1972. Distribution — Mexico. 12. Micranthemum tetrandrum C. Wright in Anales Acad. Ci. Med. Habana 7: 41. 1870 ≡ Globifera tetrandra (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 orbiculatum 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. Distribution — SE 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 differs from 229 Vandellia in its overall morphology (capitate inflorescence, 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 differs 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 finetianum (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 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. Distribution — Cameroon, Sudan, Ethiopia, Burundi, Uganda, Kenya, Tanzania, Zambia, Zimbabwe, Mozambique, 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. Distribution — India, Nepal, China; widely naturalized, 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. IndoChine 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 (Griff.) N. M. Dutta, Bull. Bot. Soc. Bengal 19: 25. 1965 ≡ Treisteria cordata Griff., Not. Pl. Asiat. 4: 109. 1854. Distribution — Myanmar. 7. Torenia cordifolia Roxb., Pl. Coromandel 2: 32. 1802. Distribution — India, Nepal, Myanmar, China, Thailand. 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 Phytotaxa 125: 42. 2013. Fischer & al.: The phylogeny of Linderniaceae Distribution — Gabon. 11. Torenia diffusa D. Don, Prodr. Fl. Nepal.: 86. 1825. = Torenia vagans Roxb., Fl. Ind., ed. 1832, 3: 96. 1832. Distribution — Nepal, Bhutan, India (Assam, Sikkim). 12. Torenia dinklagei Engl. in Bot. Jahrb. Syst. 57: 610. 1922. Distribution — Sierra Leone, Liberia, Côte d’Ivoire, Ghana, Togo, Nigeria, Equatorial Guinea (Bioko), Cameroon, 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. India 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. IndoChine 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. Willdenowia 43 – 2013 Distribution — Thailand. 26. Torenia silvicola A. Raynal in Adansonia, n.s., 5: 522. 1965. Distribution — Cameroon, Gabon, Congo-Brazzaville. 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, Revis. 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-Brazzaville, Central African Republic, Sudan, Ethiopia, Congo-Kinshasa, Rwanda, Burundi, Uganda, Kenya, Tanzania, 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 Griff. ex Benth. in Candolle, Prodr. 10: 410. 1846. Distribution — Bhutan, India, China, Taiwan, Thailand, Laos, Cambodia, Vietnam, Malaysia, Philippines, Indonesia (Java). 231 Torenia crustacea group Lindernia sect. Torenioides (Benth.) Pennell in Acad. Nat. Sci. Philadelphia Monogr. 1: 139. 1935 ≡ Vandellia 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 included 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 (Bonati) 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. IndoChine 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 Lin­naea 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 Lecomte & al., Fl. Indo-Chine 4: 416. 1927. Distribution — Senegal, Guinea-Bissau, Sierra Leone, Côte d’Ivoire, Ghana, Nigeria, Cameroon, Equatorial Guinea (Annobón), Central African Republic, Gabon, Congo-Brazzaville, Congo-Kinshasa, Burundi, Madagascar, India, Nepal, Sri Lanka, China, Thailand, Laos, Vietnam, Cambodia, Japan, Philippines, Australia, New Guinea, Micronesia, Polynesia; introduced in S U.S.A., Mexico, Cuba, Dominica, Grenada, Guadeloupe, Jamaica, Martinique, Puerto Rico, Saint Lucia, Trinidad and Tobago, Belize, Costa Rica, Guatemala, 232 Honduras, Nicaragua, Panama, Bolivia, Brazil, Colombia, 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 dictyophora 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 grandiflora (Merr.) Merr., Enum. Philipp. Fl. Pl. 3: 438. 1923, nom. illeg. [non Lindernia grandiflora Nutt., Gen. N. Amer. Pl. 2: 43. 1818] ≡ Lindernia philippinensis Philcox 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 perennans (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 pierreanoides 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. Fischer & al.: The phylogeny of Linderniaceae 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 Philcox 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. diffusa 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. = Geoffraya 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. Adelaide 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, Thailand, Laos, Vietnam, Malaysia, Australia, Philippines, New Guinea. Willdenowia 43 – 2013 2. Vandellia annamensis (T. Yamaz.) Eb. Fisch., Schäferh. & Kai Müll., comb. nov. ≡ Lindernia annamensis 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. Distribution — Australia. 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 chrysoplectra W. R. Barker in J. Adelaide Bot. Gard. 13: 81. 1990. Distribution — Australia. 8. Vandellia cleistandra (W. R. Barker) Eb. Fisch., Schäferh. & Kai Müll., comb. nov. ≡ Lindernia cleistandra W. R. Barker in J. Adelaide Bot. Gard. 13: 82. 1990. Distribution — Australia. 9. Vandellia cowiei (W. R. Barker) Eb. Fisch., Schäferh. & Kai Müll., comb. nov. ≡ Lindernia cowiei W. R. Barker in J. Adelaide Bot. Gard. 18: 162. 1998. Distribution — Australia. 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 diffusa L., Syst. Nat., ed. 12, 2: 422; Mant. Pl. 1: 89. 1767 ≡ Lindernia diffusa (L.) Wettst. in Engler & Prantl, Nat. Pflanzenfam. 4(3b): 79. 1891 ≡ Pyxidaria diffusa (L.) Kuntze, Revis. Gen. Pl. 2: 464. 1891. Distribution — Senegal, 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, CongoBrazzaville, Congo-Kinshasa, Burundi, Uganda, Tanzania, Madagascar; introduced in Mexico, Belize, Costa Rica, Guatemala, Honduras, Panama, Cuba, Dominica, Dominican Republic, Grenada, Guadeloupe, Jamaica, Martinique, Puerto Rico, Saint Vincent and the Grenadines, Trinidad and Tobago, Bolivia, Brazil, Colombia, Ecuador, Guyana, Peru, Suriname and Venezuela. 233 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, Cambodia, Vietnam, Malaysia. 14. Vandellia eremophiloides (W. R. Barker) Eb. Fisch., Schäferh. & Kai Müll., comb. nov. ≡ Lindernia eremophiloides W. R. Barker in J. Adelaide Bot. Gard. 13: 84. 1990. Distribution — Australia. 15. Vandellia harmandii (Bonati) Eb. Fisch., Schäf­erh. & Kai Müll., comb. nov. ≡ Vandellia elata var. harmandii Bonati in Bull. Soc. Bot. France 55: 539. 1908 ≡ Lindernia 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, Thailand, 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 Zanzibar), Mozambique, Madagascar. 18. Vandellia hypandra (W. R. Barker) Eb. Fisch., Schäferh. & Kai Müll., comb. nov. ≡ Lindernia hypandra W. R. Barker in J. Adelaide Bot. Gard. 13: 85. 1990. Distribution — Australia. 18. Vandellia junciformis (Bonati) Eb. Fisch., Schäf­ erh. & Kai Müll., comb. nov. ≡ Geoffraya junciformis Bonati in Notul. Syst. (Paris) 1: 334. 1911 ≡ Lindernia junciformis (Bonati) T. Yamaz. in J. Jap. Bot. 53: 97. 1978. = Geoffraya 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 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, Prodr. 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 macrobotrys 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 macrosiphonia 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. Distribution — Australia. 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 megaphylla P. C. Tsoong in Nordic J. Bot. 26: 41. 2008. Distribution — China. 28. Vandellia micrantha (D. Don) Eb. Fisch., Schäf­erh. & 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, Bijdr. Fl. Ned. Ind. 14: 752. 1826 ≡ Lindernia montana (Blume) Koord., Exkurs.-Fl. Java 3: 178. 1912, nom. illeg. [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. Fischer & al.: The phylogeny of Linderniaceae = 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. Distribution — India, Malaysia, Indonesia (Java, Mo­ luccas), Philippines, New Guinea. 31. Vandellia purpurea (Kerr ex Barnett) Eb. Fisch., Schäferh. & Kai Müll., comb. nov. ≡ Geoffraya purpurea Kerr ex Barnett in Kew Bull. 16: 486. 1963 ≡ Lindernia kerrii T. Yamaz. in J. Jap. Bot. 53: 98. 1978 [non Lindernia 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 fas­ciculata (Bonati) T. Yamaz. in J. Jap. Bot. 28: 42. 1953. Willdenowia 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, Prodr. 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. Distribution —  Taiwan. 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. Distribution — Senegal, Guinea-Bissau, Sierra Leone, 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 subracemosa De Wild., Pl. Bequaert. 5: 425. 1932. Distribution — Ethiopia, Congo-Kinshasa, Rwanda, Burundi, Uganda, Tanzania. 45. Vandellia subulata (R. Br.) Benth. in Candolle, Prodr. 10: 415. 1846 ≡ Lindernia subulata R. Br., Prodr.: 441. 1810. Distribution — New Guinea, Australia. 235 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 tectanthera 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 Lecomte & 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, Philippines; 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 yaoshanensis 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 assigned 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. bracteoides (Blatt. & Hallb.) Mukerjee (= Bonnaya ciliata 236 (­ Colsm.) Spreng. subsp. ciliata). However, more material is needed to evaluate the taxon. Conobea vandellioides Benth. in Candolle, Prodr. 10: 391. 1846 ≡ Lindernia vandellioides (Benth.) Pennell 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. diffusa. 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 Lindernia 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 Lindernia 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. Pflanzen 70: 213. 1998 [Gesneriaceae]. Lindernia dianthera Sw., Prodr.: 92. 1788 ≡ Mecardonia dianthera (Sw.) Pennell in Proc. Acad. Nat. Sci. Philadelphia 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 Caledonia, 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. Fischer & al.: The phylogeny of Linderniaceae 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 staff 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 staff 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: understanding character evolution, substitution rate plasticity, and genome miniaturization”, MU2875/2, to K.F.M. is kindly acknowledged. 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