TAXON 60 (3) • June 201 1 : 767-790 Weber & al. • Remodelling of Chirita Molecular systematics and remodelling of Chirit genera (Gesneriaceae) Anton Weber,1 David J. Middleton,2 Alan Forrest,2 Ruth Kiew,3 Chung Lu Sontag,1 Pramote Triboun,4 Yi-Gang Wei,5 Tze Leong Yao3 & Michael Molle 1 University of Vienna, Faculty Centre of Biodiversity, Department of Structural and Function 1030 Vienna, Austria 2 Royal Botanic Garden Edinburgh, 20 A Inverleith Row, Edinburgh, EH3 5LR, Scotland, U.K 3 Forest Research Institute Malaysia, 52109 Kepong, Selangor, Malaysia 4 Bangkok Herbarium, Princess Sirindhorn Herbarium Building, Plant Variety Protection Div Chatuchak, Bangkok 10900, Thailand 5 Guangxi Institute of Botany, Guangxi Zhuang Autonomous Region and the Chinese Academ Author for correspondence: David J. Middleton, d.middleton@rbge.ac.uk Abstract The polyphyletic genus Chirita is remodelled after an extensive molecular phylog and to other associated genera. Most of Chirita sect. Chirita and the monotypic Hemibo sect. Henckelia , resulting in a very differently circumscribed genus Henckelia and the sy ing species of Chirita sect. Chirita are accommodated in the revived genus Damrongia. the genus Liebigia. Chirita sect. Microchirita is recognised as the genus Microchirita. C with Chiritopsis and Wentsaiboea , included in the originally monotypic and now enorm necessary combinations are made and a general list showing the present accommodatio under Chirita , Chiritopsis , Hemiboeopsis , Primulina and Wentsaiboea is provided. Keywords Bayesian inference analysis; Chirita ; Chiritopsis ; Damrongia ; Hemiboeopsis maximum parsimony; Microchirita ; molecular phylogeny; Primulina ; taxonomy; trnL- Supplementary Material Diagnostics of the Bayesian inference analysis are available in and Figures SI and S2 and Table SI are available in Electronic Supplement 2 to the onlin (http://www.ingentaconnect.com/content/iapt/tax). ■ INTRODUCTION 1998; Li & Wang, 2004) been divided into three sections: Chirita sect. Chirita , Chirita sect. Gibbosaccus and Chirita sect. Micro- In its traditional sense, the genus Chirita Buch.-Ham. chiritaex C.B. Clarke. Wang (1985a, b) also recognised subsections D. Don, established formally by Don (1822) and revised by and Canseries in the genus and Hilliard (2004) resurrected Chirita dolle (1845), Clarke (1883) and Wood (1974), is a large genus of sect. Liebigia (Endl.) C.B. Clarke from synonymy of Chirita sect. Chirita to create a fourth section. perennial or annual herbs or subshrubs ranging from Sri Lanka, the Himalayas and southern China through continental SouthIn this paper we set out to investigate the relationships of Chirita to the other didymocarpoid genera (see Weber, 2004; (1974) recognised 77 species in three sections but the genus hasMoller & al., 2009, 201 1) and to establish a stable framework that grown rapidly in recent years, particularly through the addition can be used as a structure for investigating species delimitation. of many new species from China (see Wang & al., 1998; Li & There have been several molecular phylogenetic studies that have east Asia and into Malesia as far as Sumatra and Java. Wood Moller, 2009). Wang (1985a, b), Wang & al. (1998) and Wang &included species of Chirita , beginning with Mayer & al. (2003), al. (2011) estimated that there were about 140 species and Li &and continuing with Li & Wang (2007), Moller & al. (2009), Moller (2009) 150 species, two-thirds of which belong to ChiritaWang & al. (2011) and Moller & al. (2011). The focus of the pasect. Gibbosaccus C.B. Clarke and most of which are endemic to per by Li & Wang (2007) was to study the relationship between China. Weber (2004) gave a very large range of 80-140 species,Chirita sect. Gibbosaccus and Chiritopsis W.T. Wang. They did marked the genus as highly heterogeneous and noted there werenot include representatives of other genera except as outgroups. problems in the delimitation of the genus and the subdivisions Of the three sections recognised by Wood (1974) and Wang & within it. The great variance in the estimate of the number ofal. (1998), Chirita sect. Chirita , Chirita sect. Microchirita and species is also testament to the considerable problems of specificChirita sect. Gibbosaccus , Li & Wang (2007) found the first delimitation in Chirita but this problem will need to be addressed two to be monophyletic and the last to be paraphyletic due to the in subsequent revisions of the species rather than in this work.inclusion of Chiritopsis. However, this result was more or less The genus has mostly (Wood, 1974; Wang, 1985a, b; Wang & al.,inevitable given the lack of sampling of other didymocarpoid 767 This content downloaded from 142.51.238.106 on Fri, 09 Jun 2017 20:39:35 UTC All use subject to http://about.jstor.org/terms Weber & al. • Remodelling of Chirita TAXON 60 (3) • June 201 1: 767-790 the genus is polyphyletic? Are the three (Wood, 1974) or genera. In Moller & al. (2009, 2011), with more limited (2) samfour sections only of Chirita each monophyletic? pling in Chirita but considerably more in(Hilliard, other2004) genera, (3) What are the relationships of any possible new groupings Chirita sect. Microchirita remained monophyletic, albeit with of Chirita Chirita species otherconcept didymocarpoid genera, particularly low sampling in all sections. Chirita sect. (intothe to Henckelia , Primulinain andMoller Chiritopsis ? of Wood, 1974) was represented only by five species In thisOne paper the for the various aspects of the & al. (2009) and these did not group together. ofresponsibility these was papertype are as follows: D.J. Middleton Chirita asperifolia (Blume) B.L. Burtt, the of Chirita sect.and A. Weber initiated the need to(2004) further investigate generic delimitation in Chirita and Liebigia (Endl.) C.B. Clarke which Hilliard resurrected. have(Hook.f.) compiled the B.L. taxonomic sections (with contributions by The other species were Chirita lacunosa Burtt, in Microchirita R. Kiew, C.L. Lim, D.J. MiddleC. walkerae Gardner (as C. walkeri ), A.R. C. Rafidah pumila D. Don,); and ton, M.type Moller, of A.R. Chirita. Rafidah, R Triboun, Y.-G. Wei and T.L. Yao C. urticifolia Buch.-Ham. ex D. Don, the In collected sect. and identified most of the new material sequenced; Moller & al. (2009) four species of Chirita Gibbosaccus Forrest, C.L. Lim,due M. Moller and T.L. Yao generated the sewere included but these were found to beA. paraphyletic to the quence data; helped to compile the formal taxonomy inclusion of the monotypic Primulina Hance andS.aSontag single species and procured literature required; and M. Moller oversaw all of Chiritopsis. These results were confirmed in athe larger analysis aspects of the molecular work and analysed the molecular data. of didymocarpoid Gesneriaceae (Moller & al., 2011). Li & Wang (2007) did not include Primulina in their study. The largest published study to date is by Wang & al. (2011), a ■ MATERIAL AND paper submitted for publication after we submitted ours toMETHODS Taxon but which was reviewed and published in just a few weeks.1 Plant - The samples included in the present In their paper they included a larger sample ofmaterial. Chiritopsis and study are with those used in a recent study (Moller & al., 201 1), plus Chirita sect. Gibbosaccus species, combined sequences 41 Chinese samples from additional species of Chirita , Chiritopsis and from Moller & al. (2009) for the crucial species Primulina tabacum Hance and Petrocodon dealbatus Hance and for Henckelia. Some sequences came from GenBank, others from some non-Chinese taxa. They confirmed the basic topology of recent field collections (Table SI in Electronic Suppl. 2). In total the previous findings by Moller & al. (2009) and proceeded to 232 samples have been included covering 66 out of the 80 genraise Chirita sect. Microchirita and Chirita sect. Liebigia to ge- era classified in the Old World didymocarpoid Gesneriaceae nus rank and to reduce Chirita sect. Gibbosaccus , Chiritopsis (Weber & Skog, 2007 onwards; Wei & al., 2010), or 58 out of 61 and Wentsaiboea D. Fang & D.H. Qin to synonymy of Primulina. of the advanced Asiatic and Malesian genera to which Chirita They did not address the problem of Chirita lacunosa nor the re- belongs (Moller & al., 2011). Details of the sample coverage, lationship between species of Chirita sect. Chirita and Henckelia ingroup and outgroup selection and rooting are given in Moller & al. (2011). Here we focus on the genus Chirita. Sampling within the highly polyphyletic genus Chirita has Chirita is morphologically quite variable (Wood, 1974; Weber, 2004). Clarke (1883, 1884) commented on how Chirita been expanded from 15 to 42 samples (representing 33 species "is with difficulty distinguished from Didymocarpus ". Wood and three undescribed species), with nine species (plus one un(1974: 126) remarked that the difference between the apically described species) from section Chirita (out of 40 described), 14 fused anthers in Chirita sect. Microchirita , in contrast to face- species from section Gibbosaccus (out of 117 described), nine to-face fusion in the other sections, "together with the possi- species (plus two undescribed species) from section Microfrom Sri Lanka which were highlighted by Moller & al. (2009). bility that the syndrome of bilobed stigma and geniculate fila- chirita (out of 19 described) and one species from section Liebiments has originated several times independently, suggests that gia (out of 12 species described, all of which were previously Chirita may be polyphyletic." Clarke's and Wood's comments included in synonymy of a variable Chirita asperifolia). The genus Chiritopsis was earlier shown to be nested in may have been pertinent indeed and an investigation of Chirita clearly needs to take into account any possible relationships the Chirita sect. Gibbosaccus (Li & Wang, 2007; Moller & al., species may have with other didymocarpoid genera. Clarke 2009, 2011). To investigate closer the relationships of Chiri(1883, 1884) was largely unaware of the large number of species topsis with Chirita , five out of the nine described species have of Chirita sect. Gibbosaccus that were yet to be collected in also been included. The polyphyletic Henckelia is represented by 15 species China and his comments mainly reflected his understanding of the species in Chirita sect. Chirita. Chirita sect. Gibbosaccus (out of ca. 180). We have increased the sampling in Henckelia has since become by far the largest of the sections in Chirita because in Moller & al. (2009, 2011), with very low sampling, some species of Henckelia were nested in Chirita sect. Chirita. (Wang & al., 1998; Li & Wang, 2004). From the comments in the older literature and from the These 15 samples have been selected from a very much larger preliminary findings of Moller & al. (2009, 2011), the ques- sample of Henckelia species, the results of which will be pretions that need to be addressed for a better understanding ofsented in follow-up papers concentrating on that genus (results Chirita are: (1) Will a better sampling of Chirita confirm that of the larger Henckelia study confirm the results presented with the smaller 15 species sample included in this paper). 1 An invitation from us to combine the two papers before publication remained unanswered. DNA extraction, PCR, and direct sequencing. - Molecular methods and protocols basically followed Moller & al. 768 This content downloaded from 142.51.238.106 on Fri, 09 Jun 2017 20:39:35 UTC All use subject to http://about.jstor.org/terms TAXON 60 (3) • June 201 1: 767-790 Weber & al. • Remodelling of Chirita Phylogenetic analyses. - The MP analysis on the com(2009, 201 1), acquiring sequences of the plastid trnL-F intronbined data resulted intransribed 1 17,344 most parsimonious trees of 671 1 spacer (i trnL-F ) and the nuclear ribosomal internal steps length, a consistency index of 0.2845 and retention index spacer (ITS) region. of 0.6648. majority-rule tree was highly resolved Phylogenetic analysis. - For this study 27 The trnL-F and consensus ITS sequences were newly generated and an and was additional similar in topology 14 trnL-F to the previous phylogeny of Moller and ITS sequences were obtained from&GenBank (both sources al. (2011) where the clades received high branch support (Fig. given in bold in Appendix 2 and distinguished inSuppl. the 2). last 51 in Electronic The two BI tree recovered was basically columns). Both sequence regions were congruent available for to the MP all tree,samples in areas where the branches were well included (Appendix 2). The newly acquired GenBank sesupportedand (Fig. S2 in Electronic Suppl. 2). A detailed report quences were added to existing matrices (Moller al., 2011). and discussion of& the interand intra-generic relationships of The trnL-F alignment was unproblematic, onlydidymocarpoid 66 characters the advanced Gesneriaceae is given in Moller had to be excluded at the 5' end due to the poor quality of se& al. (2011) and we here refrain from repeating these. Instead quences, particularly those retrieved we from GenBank. focus on the positionsFor of thethe Chirita samples and associated ITS matrix, initially aligned automatically and byBI analyses are summarised genera. Thethen resultsadjusted of the MP and eye (see Moller & al., 2009), 155 characters had to be excluded in a simplified tree (Fig. 1) and subtrees (Figs. 2-6), but given due to alignment ambiguities. in full as Figs. SI and S2 (in Electronic Suppl. 2). The phylogenetic analysis basically followed Moller & different clades. Several nodes Chirita samples fell in six al. (2009, 2011). In brief, two analyses, maximum parsimony separating these clades received medium to high support values (MP) implemented in PAUP* v.4.0bl0 (Fig. (Swofford, 2002), and the separate phylogenetic 1, grey squares), supporting Bayesian inference (BI) in MrBayes v.3.1.2 (Huelsenbeck & positions of the six respective Chirita positions; i.e., they repRonquist, 2001, 2007) were carried out. The trnL-F and ITS resent separate evolutionary lineages. The branches leading sequences were analysed together after to their the sixcombinability clades with Chirita was samples (Fig. 1, grey circles) all checked by the incongruence length difference (ILD; P = 0.07) received high support values; these clades represent well detest implemented in PAUP*. fined separate genetic entities. This can also be inferred from For the MP analysis characters were and untheunordered long branches leading to these clades in the BI tree (Fig. weighted with alignment gaps treated52as data. in missing Electronic Suppl. 2). StartChirita (Nixon, elata Ridl. was1999), nested inside ing trees were found by parsimony ratchet in the Malesian HenckPAUPRat (Sikes & Lewis, 2001) and elia PAUP*, and saved clade, as sister to the Henckelia nana A. Weber (BS = 100%, trees further optimised in PAUP*, with TBR and Multrees PP both = 1.0) (Fig. 1; Figs. SI and S2 in Electronic Suppl. 2). Chirita on. Statistical branch support was obtained from 10,000 heulacunosa , C. purpureolineata (Kerr ex Craib) D. Wood and an ristic bootstrap replicates (Moller & al., 2009, Chirita 2011)sample in PAUP*. undescribed formed a clade (BS = 100%, PP = For the BI analysis models and parameter settings 1.0) sister toprior a highly supported clade of Paraboea (C.B. Clarke) were obtained independently for the trnL-F ' the ITS spacers Ridl., Trisepalum C.B. Clarke and Phylloboea Benth. samples and 5.8S sequences using MrModeltest(BS v.2.3 2004), = 98%,(Nylander, PP = 1.0), but with no statistical support for this and were GTR+G, GTR+I+G and SYM+I+G, respectively, sister relationship (Figs. 1 and 2).as suggested by the Akaike Information Criterion (AIC; Akaike, samples (not including The 1 1 Chirita sect. Microchirita 1974). Five million generations were run Chiritain elatatwo which independent is discussed above), representing eight speanalyses each with four Markov chain Carlo (MCMC) cies Monte (plus two undescribed), were placed in a highly supported chains (Electronic Supplement 1 A-D). One was clade (BS = tree 100%, PP = 1.0)sampled (Fig. 3) forming the basal branch every 250 generations (= 20,000 trees), and first 100,000 among thethe straight-fruited advanced Asiatic and Malesian gengenerations (the first 400 trees) discarded asFig. burn-in, deterera (Fig. 1; SI in Electronic Suppl. 2), though this position mined after plotting the generations against the Log(n) likelireceived no branch support, and was not present in the BI tree hoods (Electronic Supplement 1E-F). The trees (Fig.remaining S2 in Electronic Suppl. 2). were Within the Microchirita clade used to reconstruct a majority rule consensus tree in PAUP*, the two samples of Chirita hamosa R. Br. were sister (BS = 100%, The PP = 1.0). and posterior probabilities (PP) obtained. PP branch support values showed a high correlation between the of two parallel Ten samples seven species of Chirita sect. Chirita Bayesian runs (Electronic Supplementformed 1G). a strongly supported clade (BS = 92%, PP = 1.0) that also included samples of the monotypic Hemiboeopsis and two Henckelia samples, Henckelia floccosa (Thwaites) A. Weber ■ RESULTS & B.L. Burtt and H. incana (Vahl) Spreng. (Fig. 4). The relationships between Hemiboeopsis and Chirita urticifolia and Matrix characteristics. - After the exclusion of ambigC. walkerae received no branch support. The two Henckelia uous characters, the combined matrix had a length of samples 1966 were sister (BS = 100%, PP = 1.0), and this pair was characters, 1132 for trnL-F and 834 for ITS. In the combined sister to five Chirita species (BS = 83%, PP = 1.0). analysis 906 of the remaining 1966 characters were constant, The grouping of Chirita asperifolia with Allocheilos 309 variable but parsimony-uninformative, and 751 (38.2%) W.T. Wang, Gyrocheilos W.T. Wang and Didymocarpus Wall, parsimony-informative. was highly supported (BS = 99%, PP = 1.0) (Figs. 1 and 5), 769 This content downloaded from 142.51.238.106 on Fri, 09 Jun 2017 20:39:35 UTC All use subject to http://about.jstor.org/terms Weber & al. • Remodelling of Chirita TAXON 60 (3) • June 201 1 : 767-790 ^ I I Boea nyqrometrica 1 00 1 00; 1 00/1.0 I 8068 P™'Pf>ensis 100 I ^ 100 Chirita sect. Chirita 1 LasT 1.0 1-° 100 100 821 98/i I inn o<::~~~~- *1* ioo boo. */0.81 ^ *1* QC ^ i-^ Henckelia sect. Loxocarpus - Q5 f <T | 100 | 53/10 100 | 100 *r " ' | r^r- Aeschynanthus 1*°° 10offe^=== */* ' Metapetrocosmea peltata */* ' */* 100 1 oo 1001 1 o"==r===-~- ~ ioo 83/1-0 I 100 - I | Tjoo, 88/10 ioo - I 1^°° ' Lagarosolen- dominated RooT 100 ' 73 10 rao/iTo Chirita sect. Glbbosaccus */0.55 | r 70 99 LoxostigmalPetrocosmea- dom. ^ HemiboealLysionotus-dom. Fig.1. Simplified maximum parsimony majority-rule consensus tree based on combined trnL-F and ITS by names of genera dominating these clades. Chirita clades are highlighted in grey. Numbers above br numbers below branches are bootstrap values (bold italics) and Bayesian posterior probabilities (bold). bootstrap or <0.5 posterior probability values. Numbers in grey circles indicate branches supporting the m Numbers in grey squares indicate branches supporting the independent origins and relationships of the dif 100 Fig. 2. Cladogram of the majority-rule consensus i 100 too/i.o 1 from the full analysis on combined trnL-F and ITS 1001 1.0 100 I - ioo 1 P9H 00/1.0 n 100 sequences of 232 didymocarpoid Gesneriaceae sam i depicting the Chirita sect. Chirita clade 1 (- ► Dam 1 gia). For further explanation see Fig. 1. <100 | 1 nn ioon.o | 1 - OA/in 100 i 100 52/0.96 1 60/0.98 100 i 84i 1.0 ' 770 This content downloaded from 142.51.238.106 on Fri, 09 Jun 2017 20:39:35 UTC All use subject to http://about.jstor.org/terms TAXON 60 (3) • June 201 1: 767-790 Weber & al. • Remodelling of Chirita Fig. 3. Cladogram of the majority-rule consen LJ°o inn 95/10 1 I 1001 from the full analysis on combined trnL-F and sequences of 232 didymocarpoid Gesneriaceae depicting the Chirita sect. Microchirita clade (- 1 chirita ). For further explanation see Fig. 1. ioo */* 100 */* 100 gg/i.o 100 93/1 o 100 i 1001 1.01 Fig. 4. Cladogram of the majority-rule con tree from the full analysis on combined tr - 7jr- ioo ITS sequences of 232 didymocarpoid Ges 1f)n 10QI i o| ioo i l n u -i oo | iooi 1 83/1 Jffjn 83/1 0 samples, depicting the Chirita sect. Chirit (- ► Henckelia). For further explanation see 1.0 1 0 100 72/0.85 fls/i n 10o ' 63/0.9 a 100 1 79/1 .01 Fig. 5. Cladogram of the majority consensus tree from the full analy 100 gg/1 0 loon.o 100 | 52/0 "100pS | 1Qo 10011.0 100 | t 58/1.0' | on combined trnL-F and ITS sequ es of 232 didymocarpoid Gesneri ceae samples, depicting the Chiri sect. Liebigia clade (- ► Liebigia). further explanation see Fig. 1. 100 | ' 1Qo 1001 1.0 ' 1001 1.0 |g iMTo 100 100/1.0 100 i 68/0.79' Fig. 6. Cladogram of the majority-rul consensus tree from the full analysis 100 1001 100/10 combined trnL-F and ITS sequences 232 didymocarpoid Gesneriaceae sam depicting the Chirita sect. Gibbosac ' 1.0 - ioo - __ - 10011.0 - 10Q| * 10.52 100 clade (- ► Primulina). For further exp tion see Fig. 55/0.87 100 1 Chirita sinensis 1. 100 aWjffiJ 100 ^5 100 95/ro1 10o i w/0.93 ' 100 Wis 99/1.0 100 100 1 YoWml 100 100 */* 1 t 10o i W 1 100| 100 85/1. o' T | ^ | 100 1 aoTTTo] 100 1 iooITW 111 This content downloaded from 142.51.238.106 on Fri, 09 Jun 2017 20:39:35 UTC All use subject to http://about.jstor.org/terms Weber & al. • Remodelling of Chirita TAXON 60 (3) • June 201 1: 767-790 though the position among these genera is unclear. In both theHenckelia MP synonymy. Given that and Chirita were large tree (Fig. SI in Electronic Suppl. 2) Chirita is far-reaching sister genera asperifolia this conclusion has implications, not only to all three genera, though the branch supporting this relationbecause Chirita now disappears altogether, but also due to the ship received no bootstrap support. Inradical the change BI tree S2 in in the(Fig. delimitation of Henckelia. Henckelia now Electronic Suppl. 2) it was in a sister position to the Didymoincludes only the species of Henckelia from South India and carpus samples (BS=100%, PP=0.99). Sri Lanka plus most of the species of Chirita sect. Chirita and The clade including the samples of Chirita excludes allsect. species Gibbosacformerly placed in Henckelia from Thailand cus formed a strongly supported sister relationship to a clade and Malesia. Unfortunately the Thai and Malesian species of dominated by Lagarosolen W.T. Wang samples (BS 100%, PPmajority of species. ManuHenckelia made up=the very large = 1.0). The Chirita sect. Gibbosaccus clade received high scriptsitself on the fate of these species, along with further molecular branch support values (BS = 100%, PP = 1.0). Aswork, well the 14 Wang & al. (201 1) did not phylogenetic are as in preparation. species of Chritia sect. Gibbosaccus it include included six of species any species Henckeliaof in their study and concluded Chiritopsis , two species of Wentsaiboeathat and thisthe clademonotypic corresponded to geChirita sect. Chirita as defined nus Primulina (Fig. 6). The topology within by Woodthe (1974)clade except differed for Liebigia (see below). Chiritato sect. Microchirita clade. - Chirita sect. Microbetween the MP and BI analysis mainly due the changing position of Primulina tabacum Hance and Wentsaiboea renifochirita is a morphologically well defined group that, surprislia D. Fang & D.H. Qin (Figs. SI and S2 ingly, in Electronic had never been Suppl. recognised2). at generic rank before being The Chiritopsis samples appeared in three raised places, by Wang &in al. strongly (2011). We agree that generic rank is supported relationships with Chirita samples Wentsaiboea appropriate or for Microchirita. Although the sampling within luochengensis Yan Liu & W.B. Xu (Fig. the6). section is relatively good, the two subclades found do not have clear morphological distinction (both include species with crested inflorescences), but do reflect the two base chromo■ DISCUSSION some numbers in the section (C. involucrata Craib, C. caligi- nosa C.B. Clarke and C. rupestris Ridl. belong to the group This work is an extension of the recent most comprehenwith x = 9, while C. lavandulacea Stapf and C. hamosa belong sive study on the "didymocarpoid Gesneriaceae" to (Moller & having x = 17 - see Kiehn & al., 1998) as well as the group al., 2011), with an additional 27 Chirita , four Chiritopsis and the geographical distribution (all diploids with x = 9 occur in 10 Henckelia samples added. The present study focuses on the Peninsular Malaysia and Southern Thailand, while the species highly polyphyletic genus Chirita which appears in with six differx = 17, possibly dys-tetraploids, derived from 2n = 18, ent locations in the resulting phylogenies. Despite have the differa more northerly distribution in Central and Northern Thailand, and Southern China (Wood's, 1974 records ences in topology between the MP and BI analysis, whichVietnam is of C. hamosa for Southern Thailand and Northern Peninsular mainly the result of a poorly supported backbone structure, the separate positions of the Chirita samples were highlyMalaysia supported are erroneous due to confusion with C. viola Ridl.). and are not likely artefacts. Although the intergeneric clade Theand species (including the undescribed species) which have notthe had high their chromosomes counted also divide into the same relationships cannot be elucidated from these results, support values for the individual clades containing Chirita northernspeand southern clades. The group of species highlighted cies means that we can with some confidence propose byredefined Wood (1974) as having the unusual feature of the presence genera for these different clades, in some cases agreeing with of hairs with swollen tips on the upper surface of the corolla the very recent changes proposed by Wang & al. (201 1) and throat, C. in caliginosa , C. mollis sima Ridl. and C. viola , do come some cases disagreeing. out as a group sister to C. rupestris. Chirita sp. nov. 3, which is very close to C. sericea , is also in this group. sect. Chirita 1 in Fig. 1 contains Chirita purpureolineata, C. la- Chirita elata. - The generic position of Chirita elata has been problematic since it was first described (see Rafidah & al., cunosa and an undescribed species. Chirita purpureolineata was originally described as Damrongia purpureolineata Kerr 201 1). Our results show it nested firmly amongst the Malesian Chirita sect. Chirita 1 clade. - The clade labelled Chirita species of Henckelia (in the sense of Weber & Burtt, 1998), ex Craib and is the type species of the genus Damrongia. This genus name is available for this clade. This clade was not found which is in agreement with the diandrous flowers. by Wang & al. (201 1), nor did they discuss the isolated position Chirita sect. Liebigia clade. - Chirita sect. Liebigia was of Chirita lacunosa in Moller & al. (2009). relatively recently resurrected as a distinct section within Chirita by Hilliard (2004), having been included in Chirita sect. Chirita 2 in Fig. 1 contains the type species of the genera sect. Chirita by Wood (1974). Wood (1974) recognised only one Chirita (C. urticifolia), Henckelia (H. incana) and Hemiboeopsis widespread and variable species, Chirita asperifolia , but HillChirita sect. Chirita 2 clade. - The clade labelled Chirita (H. longisepala (H.W. Li) W.T. Wang, the only species). The other iard (2004) recognised 12 species, 10 of them based on material species of Henckelia included, H. floccosa , is also a memberalso of available to Woods (who freely admitted that "the account Henckelia sect. Henckelia which is a group of 14 rather similar of Chirita asperifolia is probably the least satisfactory in this species in South India and Sri Lanka. The other taxa in this clade revision"). We have sampled only material identified by Hill- iard as Chirita asperifolia , the type species of the section, and are all species of Chirita sect. Chirita. This clade can be recog- nised as the genus Henckelia with Chirita and Hemiboeopsisitinis quite distinct from the other parts of Chirita sect. Chirita 772 This content downloaded from 142.51.238.106 on Fri, 09 Jun 2017 20:39:35 UTC All use subject to http://about.jstor.org/terms TAXON 60 (3) • June 201 1: 767-790 Weber & al. • Remodelling of Chirita and from other didymocarpoid genera. Liebigia , at rank, accounted for in genus the new classification framework and hope is already available for this group. Chirita sect. Gibbosaccus clade. - Chirita sect. Gibbosac- their status can be better resolved in the future. One new combination is made in the genus Codonoboea cus is by far the largest section within Chirita. Our results show for Chirita humilis Miq. Chirita humilis was previously also in Henckelia , the Malesian members of which must all species in a well-supported clade but with the species included of Chiritopsis , Wentsaiboea renifolia , W. luochengensis and Primnow be excluded from the genus. Manuscripts focusing on this ulina nested within. In this group the genus name Primulina, group are in preparation but for the sake of completeness this previously monotypic but very recently expanded to include new combination is proposed in Appendix 1. these groups by Wang & al. (2011), has priority. Henckelia Spreng., Anleit. Kenntn. Gew., ed. 2, 2(1): 402. 1817 = Henckelia sect. Henckelia Weber & Burtt in Beitr. Biol. ■ TAXONOMIC IMPLICATIONS Pflanzen 70: 334. 1998 ('1997') = Didymocarpus sect. Orthoboea Benth. in Benthem & Hooker, Gen. PI. 2(2): Each of the six clades identified above has already been 1022. 1876 - Type: Henckelia incana (Vahl) Spreng. highlighted in previous taxonomic work as morphologically = Chirita Buch.-Ham. ex D. Don, Edinburgh Philos. J. 7: 83. 1822 = Chirita distinct. The molecular work presented here lends weight to the sect. Euchirita C.B. Clarke in Candolle & Canrecognition of each of the groups as distinct taxa (or in adolle, differMonogr. Phan. 5(1): 111. 1883 = Didymocarpus sect. ent genus in the case of Chirita elata). However, theseEuchirita groups (C.B. Clarke) Chun in Sunyatsenia 6: 294. 1946 = are not always easily distinguished from each other and not sect. Euchirita (C.B. Clarke) Fritsch in Engler & Roettlera easily distinguished from other didymocarpoid Gesneriaceae, Prantl,Nat. Pflanzenfam. IV/3b: 148. 1895- Type (lectotype a not uncommon problem in the family. Burtt, 1954): Chirita urticifolia Buch.-Ham. ex D. Don (= The single character which was used to unite the species Henckelia urticifolia (Buch.-Ham. ex D. Don.) A. Dietr.). and sections of Chirita into one genus and distinguish it from Wall., Numer. List: 800-806. 1829. All species = Calosacme other genera was the lack of development of the upper lobe ofestablished in Calosacme by Wallich (1829) are names nomina nuda. the stigma and the usually bifid development of the lower lobe (henceforth referred to as the chiritoid stigma). It was alreadyDC. ex Meisn., PI. Vase. Gen. 1: tab. diag. 302, = Babactes recognised that this character, although useful, had its probComm. 211. 1840 - Type: Babactes oblongifolia (Roxb.) lems and led to confusion over the correct placement of DC.some (= Henckelia oblongifolia (Roxb.) D.J. Middleton & species. Below we discuss the morphological basis of Mich. each Moller). of the genera we now recognise but acknowledge that further = Gonatostemon Regel in Gartenflora 15: 353. 1866 - Type: research is necessary in the didymocarpoid Gesneriaceae to Gonatostemon boucheanum Regel (= Henckelia urticifolia better understand the morphological as well as molecular evo(Buch.-Ham. ex D. Don.) A. Dietr.). lution in the group. = Ceratoscyphus Chun in Sunyatsenia 6: 276. 1946 - Type: Below we list all of the taxa we now recognise within each Ceratoscyphus caerulea Chun (= Henckelia ceratoscyphus genus, including the many necessary new combinations. (B.L.We Burtt) D.J. Middleton & Mich. Moller). list heterotypic synonyms when the synonym was= published Hemiboeopsis W.T. Wang in Acta Bot. Yunnan. 6: 397. 1984 in Chirita , but we must emphasise that we have not critically - Type: Hemiboeopsis longisepala (H.W. Li) W.T. Wang (= checked the status of these synonyms. Instead we haveHenckelia gener- longisepala (H.W. Li) D.J. Middleton & Mich. ally adopted heterotypic synonyms for names in Chirita Moller). as they have appeared in recent revisions, especially by WoodPerennial (1974) or annual herbs, sometimes woody at base, and Wang & al. (1998). Further work may show that some caulescent or acaulescent, rarely creeping. Leaves opposite, names presently found in synonymy must be re-established alternate or whorled, sometimes clustered at the stem apex, or and transferred to one of the present genera. In Appendix 1 one or two leaves, petiolate, lamina variable, reduced to just we give a complete list of all names ever publishedsometimes in Chirita pinnately lobed. Inflorescences axillary (or someand the current name for each of them. times appearing terminal by reduction of vegetative shoot), It should be borne in mind that we have moved all curflowers 1-15; bracts paired or in whorls, free or connate at rently recognised taxa out of Chirita even when we cannot bebase, orbicular to linear, narrowly ovate or narrowly triangular, sure we have made the correct decision. Chirita as a genus issometimes early caducous. Calyx of 5 free lobes or fused into a a synonym of Henckelia and no currently recognised speciestube for part of length, when fused often more deeply divided can bear this generic name. To leave species in Chirita due toon dorsal side, segments triangular or narrowly triangular. Cosome uncertainties would result in competing and confusingrolla infundibuliform, more rarely tubular, tube often slightly classification frameworks for this group. Instead we have madepouched, sometimes rather constricted in throat; bilabiate with new combinations for all taxa based on our best knowledge of2-lobed upper lip and 3-lobed lower lip; colouration variable, where each taxon now belongs. We freely admit that in someoften with yellow markings in throat. Stamens 2; filaments cases we may be mistaken and have highlighted these taxa in geniculate or straight; anthers fused face to face, glabrous or the Appendices. These taxa, however, were already previouslypubescent. Disc a simple annular ring or 5-lobed, often very poorly understood and we have merely ensured that they aresmall. Ovary shortly stipitate or not; stigma chiritoid, often 773 This content downloaded from 142.51.238.106 on Fri, 09 Jun 2017 20:39:35 UTC All use subject to http://about.jstor.org/terms Weber & al. • Remodelling of Chirita TAXON 60 (3) • June 201 1 : 767-790 weakly so and rarely almost truncate, lower lobe bifid or not. the newly circumscribed Henckelia do not occu Fruit a capsule splitting into 2 valves or but opening only the rather on along sandstone, granite and other non-li dorsal side, stipitate or not, plagiocarpic or not, persistent This is calyx in marked contrast to Damrongia , Mic or not in fruit. Seeds minute, ellipsoid. Primulina which are almost exclusively found Distribution: Sri Lanka, Southern and Northeastern Although theIndia, name Hemiboeopsis suggests a cl Nepal, Bhutan, Southern China, Northern Vietnam, Northern to Hemiboea, it relates only to the similarity Laos, Northern Thailand. - As far as is known Henckelia (1984) himself stressed a possible relationship w diagnosis ("fortasse species are found on soils and rocks butthe not on limestone. Ap- affinis Chiritae D. Don"). proximately 56 species. the stigma shape (though with short upper lobe) Clarke (1883, 1884) noted the difficulty ofcylindrical distinguishing long, and orthocarpic capsules are n Didymocarpus and Chirita but it is likely that he was, in rather particuHemiboea but confirm Wang's suggestion lar, referring to those species which were later Burtt, ship to (Weber Chirita.&The molecular data show that it sect. Chirita clade. 1998) removed from Didymocarpus to a Chirita resurrected Henckelia. Henckelia was distinguished by having a fruit which dehisces Wang & al. (2011) included several species of Chirita sect. only along the dorsal side, whilst Didymocarpus hasanalysis fruits Chirita in their butwhich as they did not include any species dehisce into two valves. It was noted by Weberthey & incorrectly Burtt (1998) of Henckelia maintained the genus Chirita to acthat the South Indian and Sri Lankan commodate speciesthese were distinct species. They also did not include Hemiboeopenough to form a section, Henckelia sect. the sis inHenckelia their analysis so, did not type reduce this genus to synonymy. section of the genus. Its morphology is described by these of authors There are a number species which we include in Henck- in some detail under "South Indian Didymocarpus (Weber & in the molecular analyses elia but which have"not been included and and for which their inclusion could be debated. These include Burtt, 1998: 316-317). Study of specimens evaluation of the older literature and illustrations have revealed Henckelia forrestii that stigma (J. Anthony) shape D.J. Middleton & Mich. Moller, H. smitinandii (B.L. Burtt) Middleton & Mich. Moller. is variable in this group, ranging from truncate-capitate to D.J. definitely chiritoid with a short or undeveloped upper stigma and H. tibetica (Franch.) D.J.lobe Middleton & Mich. Moller. Only and a large lower lobe. In one individual of Henckelia incana it a more focused study on these species can resolve this issue has even been observed that the lower lobe is bifid, theis same but until contraryand evidence found they should be included in is suggested in the description of the stigma ofalong Henckelia repens Henckelia with most other species in Chirita sect. Chirita. by Clarke (1883, as Didymocarpus repens). Therefore, no fundamentally from the In its new sense,there Henckeliaisdiffers sharp boundary between Henckelia sect. Henckelia andconcept Chirita (purely morphological) of Weber & Burtt (1998): it is with regard to the crucial stigma character. A chiritoid stigma, an amalgamation of the type sections of Henckelia and Chirita however, has never been observed in the Malesian species placed (excluding the species placed in Damrongia ), but excludes all in Henckelia by Weber & Burtt (1998). In other addition the taxa fruit charinfrageneric referred to Henckelia by these authors acter used to characterise Henckelia , dehiscence only along the, Heteroboea and Glossad(sects. Loxocarpus , Didymanthus dorsal side, has also been observed in Chirita smitinandii enia). The fate of the species inB.L. these sections will be discussed Burtt and C. lachenensis C.B. Clarke and may occur in forthcoming papers.in other Chirita species in fresher material (drying of specimens often makes this character difficult to observe, particularly those D.J. Middleton & Mich. Henckelia adenocalyx in (Chatterjee) species with very long and thin fruits such as Chirita Alladenocalyx Chatterjee in Moller, comb.pumila). nov. = Chirita Kews.str. Bull. 1948: 1948. - Distr.: India (Arunachal but one species of Henckelia sect. Henckelia are63. acaulesPradesh), Northern Burma, China (Yunnan). cent. Most species of Chirita sect. Chirita (without Damrongia) Henckelia anachoreta (Hance) D.J. Middleton & Mich. Moller, are caulescent but acaulescent species of Chirita sect. Chirita comb. nov. Chirita anachoreta are known such as C. briggsioides W.T. Wang, C. = dielsii (Borza) Hance in Ann. Sci. Nat., Bot., ser. 5, 5: Clarke. 231. 1866 = It Didymocarpus anachoretus B.L. Burtt, C. lachenensis and C. primulacea C.B. should also be noted that both caulescent (Hance) and acaulescent H. Lev. in Compt.speRend. Assoc. Fran?. 34: 427. 1906 ('1905') = Chirita minutiserrulata Hayata, Icon. PI. cies are known from other Gesneriaceae genera (e.g., Paraboea). Recorded chromosome numbers are n = 4,Formosan. 9, 10, 5: 14, 27 India (Sikkim), Burma, 133.16, 1915.17, - Distr.: Northern Thailand, (Guangdong, Guangxi, Hunan, and ±45 (see "Webcyte", Moller & al., 2002 onwards). TheChina newly defined genus Henckelia is variable but noYunnan, more so than other Xizang), Taiwan, Laos, Northern Vietnam. large didymocarpoid genera. It occurs Henckelia in South India and Sri angusta (C.B. Clarke) D.J. Middleton & Mich. Moller, comb. nov. = Chirita zeylanica var. angusta C.B. Clarke Lanka, the Himalayas and continental Southeast Asia, excluding in Candolle Candolle, Monogr. Phan. 5(1): 113. 1883 = Peninsular Thailand and Malaysia. It does not get & into Malesia. Chirita angusta Clarke) W.L. Theob. & Grupe in CeySome of the species we include in Henckelia are(C.B. known J. Sci.,only Biol. Sci. 10(1): 70. 1972. - Distr.: Sri Lanka. to us only from herbarium specimens and lon a few from the Henckelia auriculata (J.M. Li & S.X. Zhu) D.J. Middleton literature. Habitat preference is something not well documented & Mich. Moller, comb. nov. = on specimens or in the literature with most species recorded asChirita auriculata J.M. Li & S.X. Zhu inthose Ann. Bot. Fenn. 47: 222. 2010. - Distr.: occurring on rocks. We have the impression from species (Yunnan). that we do know from the field that most,China if not all, species in 774 This content downloaded from 142.51.238.106 on Fri, 09 Jun 2017 20:39:35 UTC All use subject to http://about.jstor.org/terms TAXON 60 (3) • June 201 1: 767-790 Weber & al. • Remodelling of Chirita Henckelia bifolia (D. Don) A. Dietr., Sp. Biol.PL, Pflanzen ed. 90: 6, 344. 1: 574. 1998 ('1997') = Didymocarpus 1831 = Chirita bifolia D. Don, Prodr. Fl. Nepal.: 90. 1825 fischeri Gamble in Bull. Misc. Inform. Kew 1923: 117. - Distr.: = Chirita amplectens Steud., Nomencl.1923. Bot., ed.South 2, India. 1: 351. 1840, nom. nud. = Chirita scabra Royle ex floccosa C.B. Clarke Henckelia (Thwaites) in A. Weber & B.L. Burtt in Beitr. Candolle & Candolle, Monogr. Phan. 5(1): 1883, nom. Biol.113. Pflanzen 90: 344. 1998 ('1997') = Didymocarpus illeg. pro syn. - Distr.: India (Himachal Pradesh, Uttar floccosus Thwaites, Enum. PI. Zeyl.: 207. 1860. - Distr.: Sri Lanka. Pradesh), Nepal, Bhutan. Henckelia briggsioides (W.T. Wang) D.J. Middleton &Anthony) Mich. D.J. Middleton & Mich. Henckelia forrestii (J. Moller, comb. nov. = Chirita briggsioides W.T. Wang Moller, comb. nov. = Chiritain forrestii J. Anthony in Notes Bull. Bot. Res., Harbin 2(2): 138. 1982. - Distr.: China Roy. Bot. Gard. Edinburgh 18: 192. 1934 = Didymocarpus (Hubei). forrestii (J. Anthony) Hand.-Mazz., Symb. Sin. 7: 882. Henckelia burttii D.J. Middleton & Mich. Moller, nom. nov. 1936 = Chirita forrestii var. acutidentata W.T. Wang in = Chirita reptans B.L. Burtt & Panigrahi in Notes Roy. Acta Phytotax. Sin. 13(3): 103. 1975. - Distr.: China Bot. Gard. Edinburgh 26: 265. 1965, non Henckelia reptans (NW Yunnan). - Chirita forrestii was referred to Chirita (Jack) Spreng. (1827). - Distr.: Northern Burma. sect. Chirita by Wood (1974), but to Chirita sect. GibHenckelia calva (C.B. Clarke) D.J. Middleton & Mich. Moller, bosaccus by Wang & al. (1998) and Li & Wang (2004) comb. nov. = Chirita calva C.B. Clarke in Candolle & which would result in it now being transferred to PrimCandolle, Monogr. Phan. 5(1): 116. 1883. - Distr.: Indiaulina. We conclude that it is a Henckelia but further study (Assam, Sikkim), Northern Burma. is clearly necessary to resolve the issue of its correct Henckelia ceratoscyphus (B.L. Burtt) D.J. Middleton & Mich. placement. Moller, comb. nov. = Chirita ceratoscyphus B.L. Burtt in fruticola (H.W. Li) D.J. Middleton & Mich. Moller, Henckelia Notes Roy. Bot. Gard. Edinburgh 21: 212. 1954 = Ceratocomb. nov. = Chirita fruticola H.W. Li in Bull. Bot. Res., scyphus caeruleus Chun in Sunyatsenia 6: 278. 1946, non Harbin 3(2): 37. 1983. - Distr.: China (Yunnan) Chirita caerulea R. Br. (1840), non Henckelia caerulea Henckelia gambleana (C.E.C. Fisch.) A. Weber & B.L. Burtt (Ridl.) A. Weber (1998) = Chirita corniculata Pellegr. in in Beitr. Biol. Pflanzen 90: 345. 1998 ('1997') = DidymoBull. Soc. Bot. France 73: 420. 1926, non Henckelia corcarpus gambleanus C.E.C. Fisch. in Bull. Misc. Inform. Kew 1938: 36. 1938. - Distr.: South India. niculata (Jack) Spreng. (1827). - Distr.: China (Guangxi), Northern Vietnam. Henckelia grandifolia A. Dietr., Sp. PI., ed. 6, 1: 576. 1831 = Henckelia communis (Gardner) D.J. Middleton & Mich. Chirita macrophylla Wall., PL Asiat. Rar. 1: 56, t. 72. 1830), Moller, comb. nov. = Chirita communis Gardner in Cal- non Henckelia macrophylla (D. Don) Spreng. = Chirita cutta J. Nat. Hist. 4: 481. 1846 = Chirita zeylanica Hook, macrophylla Wall. var. tirapensis Panigrahi in Bull. Bot. in Bot. Mag. 71: t. 4182. 1845, non Henckelia zeylanica Soc. Bengal. 21: 32. 1967. - Distr.: Nepal, India (Assam, (R. Br.) A. Weber & B.L. Burtt (1998) = Chirita vulgaris Sikkim, West Bengal), Bhutan, Northern Burma, China Morren in Belgique Hort. 3: 238, t. 36. 1853, nom. nud. (Yunnan), Northern Thailand. - Distr.: Sri Lanka. Henckelia heterostigma (B.L. Burtt) D.J. Middleton & Mich. Henckelia dibangensis (B.L. Burtt, S.K. Srivastava & Mehro-Moller, comb. nov. = Chirita heterostigma B.L. Burtt in tra) D.J. Middleton & Mich. Moller, comb. nov. = ChiritaNotes Roy. Bot. Gard. Edinburgh 26: 262. 1965. - Distr.: Burma. dibangensis B.L. Burtt, S.K. Srivastava & Mehrotra in Notes Roy. Bot. Gard. Edinburgh 45(3): 469. 1989 ('1988'). Henckelia hookeri (C.B. Clarke) D.J. Middleton & Mich. - Distr.: India (Arunachal Pradesh). Moller, comb. nov. = Chirita hookeri C.B. Clarke, Henckelia dielsii (Borza) D.J. Middleton & Mich. Moller,Commelyn. Cyrtandr. Bengal.: t. 70. 1874. - Distr.: India comb. nov. = Chirita dielsii (Borza) B.L. Burtt in Notes (Assam). Roy. Bot. Gard. Edinburgh 23: 97. 1960 = Chirita orbicuHenckelia humboldtiana (Gardner) A. Weber & B.L. Burtt in laris W.W. Sm. in Notes Roy. Bot. Gard. Edinburgh 9: 94.Beitr. Biol. Pflanzen 90: 346. 1998 ('1997') = Didymocar1916. - Distr.: China (Yunnan, Sichuan). pus humboldtianus Gardner in Calcutta J. Nat. Hist. 6: 477. 1846. - Distr.: Sri Lanka. Henckelia dimidiata (Wall, ex C.B. Clarke) D.J. Middleton & Mich. Moller, comb. nov. = Chirita dimidiata Wall, Henckelia incana (Vahl) Spreng., Syst. Veg. 1: 38. 1824 = ex C.B. Clarke in Candolle & Candolle, Monogr. Phan.Rottlera incana Vahl, Enum. PL 1: 88. 1804. - Distr.: South India. 5(1): 115. 1883 = Chirita polyneura Miq. var. amabilis C.B. Clarke, Commelyn. Cyrtandr. Bengal.: t. 76. 1874 = Henckelia infundibuliformis (W.T. Wang) D.J. Middleton Chirita dimidiata Steud., Nomencl. Bot., ed. 2, 1: 351. 1840,& Mich. Moller, comb. nov. = Chirita infundibuliformis nom. nud. - Distr.: India (Assam, Sikkim). W.T. Wang in Acta Phytotax. Sin. 17(1): 111. 1979. Henckelia fasciculiflora (W.T. Wang), D.J. Middleton & Mich.Distr.: China (Xizang). Moller, comb. nov. = Chirita fasciculiflora W.T. Wang in Henckelia innominata (B.L. Burtt) A. Weber & B.L. Burtt Bull. Bot. Res., Harbin 2(2): 139. 1982. - Distr.: Chinain Beitr. Biol. Pflanzen 90: 347. 1998 ('1997') = Didymo(Yunnan). carpus innominatus B.L. Burtt in Notes Roy. Bot. Gard. Henckelia fischeri (Gamble) A. Weber & B.L. Burtt in Beitr. Edinburgh 21: 201. 1954. - Distr.: South India. 775 This content downloaded from 142.51.238.106 on Fri, 09 Jun 2017 20:39:35 UTC All use subject to http://about.jstor.org/terms Weber & al. • Remodelling of Chirita TAXON 60 (3) • June 201 1: 767-790 Henckelia moonii Middleton & Mich. Moller, Henckelia insignis (C.B. Clarke) D.J. Middleton & (Gardner) Mich.D.J. Moller, comb. nov. = Chirita moonii Gardner, comb. nov. = Chirita insignis C.B. Clarke in Candolle & Calcutta J. Nat. Hist. 6: 479. 1846. - Distr.: Sri Lanka. Candolle, Monogr. Phan. 5(1): 121. 1883. - Distr.: Burma. Henckelia lacei (W.W. Sm.) D.J. Middleton &oblongifolia Mich. Moller, Henckelia (Roxb.) D.J. Middleton & Mich. Moller, comb. nov. = Incarvillea oblongifolia Roxb., Fl. Ind., ed. comb. nov. = Didymocarpus lacei W.W. Sm. in Rec. Bot. Surv. India 6: 41. 1913 = Chirita 2, lacei Sm.)oblongifolia (Roxb.) J. Sinclair in 3: 113.(W.W. 1832 = Chirita Bull. Bot. Soc. Bengal 9: 102. 1957 = Chirita oblongifoB.L. Burtt in Notes Roy. Bot. Gard. Edinburgh 21: 187. lia (Roxb.) B.L. Burtt in Notes Roy. Bot. Gard. Edinburgh 1954. - Distr.: Burma. - Wood (1974: 161-162) placed 307. 1958, ,comb, the species tentatively in Chirita sect.22:Chirita butsuperfl. ear- = Chirita acuminata Wall, ex R. Br.,whose Cyrtandreae: 117. 1839, nom. nud. = Chirita acumimarked it as "another anomalous" species, position "remains uncertain". We make the transfer to Henckelia nata Steud., Nomencl. Bot., ed. 2, 1: 351. 1840, nom. nud. - Distr.: India (Assam), Northern Burma, China (Yunnan). but note that further study is required. Henckelia lachenensis (C.B. Clarke) D.J. Middleton & Mich. Henckelia ovalifolia (Wight) A. Weber & B.L. Burtt in Beitr. Moller, comb. nov. = Chirita lachenensis C.B. Clarke in Biol. Pflanzen 90: 351. 1998 ('1997') = Didymocarpus Candolle & Candolle, Monogr. Phan. 5(1): 118. 1883 = Chirita clarkei Hook.f., Fl. Brit. India 4: 359. 1884, nom. ovalifolius Wight, Icon. PI. Ind. Orient. 4(2): 10, t. 1351. 1848. - Distr.: South India. illeg. = Chirita umbricola W.W. Sm. in Notes Roy. Bot.Henckelia peduncularis (B.L. Burtt) D.J. Middleton & Mich. Moller, comb. nov. = Chirita peduncularis B.L. Burtt in Gard. Edinburgh 10: 172. 1918 = Chirita stolonifera C.E.C. Fisch. & Kaul in Kew Bull. Misc. Inform. 1940: 196. Notes Roy. Bot. Gard. Edinburgh 26: 265. 1965. - Distr.: Northern Burma. 1940. - Distr.: Bhutan, India (Assam, Sikkim), Northern Burma, China (Yunnan). Henckelia primulacea (C.B. Clarke) D.J. Middleton & Mich. comb. nov. = Chirita primulacea C.B. Clarke, Henckelia longipedicellata (B.L. Burtt) D.J. MiddletonMoller, & Commelyn. Cyrtandr. Bengal.: t. 82. 1874. - Distr.: India Mich. Moller, comb. nov. = Chirita longipedicellata (Sikkim), Nepal. B.L. Burtt in Notes Roy. Bot. Gard. Edinburgh 26: 264. 1965. - Distr.: Burma. Henckelia puerensis (Y.Y. Qian) D.J. Middleton & Mich. Henckelia longisepala (H.W. Li) D.J. Middleton & Mich.Moller, comb. nov. = Chirita puerensis Y.Y. Qian in Acta Moller, comb. nov. = Lysionotus longisepalus H.W. Li, Phytotax. Sin. 34(2): 216. 1996. - Distr.: China (Yunnan). Henckelia pumila (D. Don) A. Dietr., Sp. PL, ed. 6, 1: 574. Bull. Bot. Res., Harbin 3(2): 1. 1983 = Hemiboeopsis longisepala (H.W. Li) W.T. Wang in Acta Bot. Yunnan. 6(4): 397. 1831 = Chirita pumila D. Don, Prodr. Fl. Nepal.: 90. 1825 1984. - Distr.: China (Yunnan), Laos. = Chirita edgeworthii A. DC., Prodr. 9: 269. 1845 = Chirita Henckelia lyrata (Wight) A. Weber & B.L. Burtt in Beitr. Biol.polyneura Miq. var. thomsonii C.B. Clarke, Commelyn. Pflanzen 90: 349. 1998 ('1997') = Didymocarpus lyratus Cyrtandr. Bengal.: t. 75. 1874 = Chirita sphagnicola Wight, Icon. PI. Ind. Orient. 42: 10, t. 1350. 1848. - Distr.: H. Lev. & Vaniot in Bull. Soc. Bot. France 53: 550. 1906 = Chirita diaphana Royle, 111. Bot. Himal. Mts.: 294. 1836, South India. Henckelia macrostachya (E. Barnes) A. Weber & B.L. Burtt nom. nud. = Chirita flava Wall, ex R. Br., Cyrtandreae: 1 17. in Beitr. Biol. Pflanzen 90: 349. 1998 ('1997') = Didymo- 1839, nom. superfl. = Chirita flava Steud., Nomencl. Bot., carpus macrostachyus E. Barnes in Bull. Misc. Inform. ed. 2, 1: 351. 1840, nom. nud. - Distr.: Northern India Kew 1938: 37. 1938. - Distr.: South India. (Himachal Pradesh to Assam), Nepal, Bhutan, Northern Burma, Thailand, Southern China (Yunnan, Guizhou), Henckelia meeboldii (W.W. Smith & Ramaswami) A. Weber & B.L. Burtt in Beitr. Biol. Pflanzen 90: 349. 1998 ('1997') Northern Vietnam. Henckelia pycnantha (W.T. Wang) D.J. Middleton & Mich. = Didymocarpus meeboldii W.W. Smith & Ramaswami in Rec. Bot. Surv. India 6: 43. 1913. - Distr.: South India. Moller, comb. nov. = Chirita pycnantha W.T. Wang in Novon 7(4): 425. 1998 ('1997'). - Distr.: China (Yunnan). Henckelia mishmiensis (Debbarm. ex Biswas) D.J. Middleton & Mich. Moller, comb. nov. = Chirita mishmiensis Deb-Henckelia repens (Bedd.) A. Weber & B.L. Burtt in Beitr. Biol. barm. ex Biswas in Assam Forest Rec., Bot. 1: 21. 1934. Pflanzen 90: 354. 1998 ('1997') = Didymocarpus repens Bedd., Icon. PI. Ind. Orient. 1: 24, t. 120. 1874. - Distr.: - Distr.: India (Assam). South India. Henckelia missionis (Wall, ex R. Br.) A. Weber & B.L. Burtt in Beitr. Biol. Pflanzen 90: 350. 1998 ('1997') = Didymo- Henckelia rotundata (Barnett) D.J. Middleton & Mich. Moller, comb. nov. = Chirita rotundata Barnett in Kew Bull. 15: carpus missionis Wall, ex R. Br., Cyrtandreae: 119. 1839. - Distr.: South India. 254. 1961. - Distr.: Northern Thailand. Henckelia shuii (Z. Yu Li) D.J. Middleton & Mich. Moller, Henckelia monantha (W.T. Wang) D.J. Middleton & Mich. Moller, comb. nov. = Chirita monantha W.T. Wang in Bull. comb. nov. = Chirita shuii Z. Yu Li in Novon 7(4): 427. Bot. Res., Harbin 2(4): 53. 1982. - Distr.: China (Hunan). 1998 ('1997'). - Distr.: China (Yunnan). Henckelia smitinandii (B.L. Burtt) D.J. Middleton & Mich. Henckelia monophylla (C.B. Clarke) D.J. Middleton & Mich. Moller, comb. nov. = Chirita monophylla C.B. Clarke inMoller, comb. nov. = Chirita smitinandii B.L. Burtt in Thai J.D. Hooker, Fl. Brit. India 4: 360. 1884. - Distr.: India Forest Bull., Bot. 29: 89. 2001. - Distr.: Eastern Thailand (Khao Yai N.P.). - Burtt (2001) did not give the sectional (Assam). 776 This content downloaded from 142.51.238.106 on Fri, 09 Jun 2017 20:39:35 UTC All use subject to http://about.jstor.org/terms TAXON 60 (3) • June 201 1 : 767-790 Weber & al. • Remodelling of Chirita margin, to ovate. Calyx of 5 free lobes or fused into a tube placement of the species but suggested itlinear was related to forplace part of length, segments triangular or narrowly triangular, Chirita poilanei Pellegr., a species we in Primulina. However, in its distribution, thin leaves, dehissometimesdorsally reduced to 3 lobes by fusion of upper 3 lobes. Corolla bilabiate with cent capsule and habitat preference infundibuliform; (not on limestone) it2-lobed upper lip and 3-lobed lower lip; white to deep purple,in sometimes with yellow markings would appear to be more similar to species now placed Henckelia. Further study is necessary. in throat. Stamens 2; filaments straight or curved; anthers fused Henckelia speciosa (Kurz) D.J. Middleton & glabrous. Mich.Stigma Moller, face to face, chiritoid, lower lobe bifid or not. Fruit capsule11: splitting 2 valves. Seeds minute, ellipsoid. comb. nov. = Chirita speciosa Kurz in J. aBot. 195.into 1873 = Chirita brevipes C.B. Clarke in Candolle & Candolle, Distribution: Thailand and Peninsular Malaysia. - Found Monogr. Phan. 5(1): 120. 1883 = Chirita trailliana Forrest on limestone rocks, usually in shade. Approximately 6 species. & W.W. Sm. in Notes Roy. Bot. Gard. Damrongia Edinburgh 9: 95. was coined by Kerr but published by Craib 1916. - Distr.: Southern China (Yunnan). (1918) for the accommodation of a single species, D. purHenckelia tibetica (Franch.) D.J. Middleton & Mich. Moller, pureolineata and was later transferred to Chirita by Wood comb. nov. = Roettlera tibetica Franch. in Bull. Mus. (1972). Wood did not provide any argument for his decision to Hist. Nat. (Paris) 5: 251. 1899 = Didymocarpus tibeticus reduce the genus to a synonym of Chirita . A second species, (Franch.) Hand.-Mazz., Symb. Sin. 7: 881. 1936 = Chirita D. kerrii , was incorrectly attributed to the genus by Pellegrin tibetica (Franch.) B.L. Burtt in Notes Roy. Bot. Gard. Edin(1930) and is now recognised in Petrocosmea. The genus name burgh 23: 99. 1960. - Distr.: China (Yunnan, Sichuan). honours Damrong Raja Nubhap (1862-1943) of Thailand who "has done so much for the advancement of education in his - Burtt (1960: 99) discussed the transfer of Hemiboea subacaulis Hand.-Mazz. and Roettlera tibetica Franch. to country" (Craib, 1918: 365). Chirita and he regarded the two species as closely allied. Damrongia purpureolineata belongs to a small group of Wang & al. (1998) did not accept the transfer of Hemiboea similar species restricted to Thailand and Peninsular Malaysia. subacaulis to Chirita , and thus the two species are curThe most-widespread species is perhaps Damrongia lacunosa , rently placed in different genera. The placement of the originally described as Didymocarpus lacunosus. It was transspecies thus is most problematic and requires further study. ferred to Chirita , along with Didymocarpus cyaneus , by Burtt Given that to us it has closer affinities with Chirita sect. (1965). Wood (1974) combined the two species under Chirita Chirita than to Chirita sect. Gibbosaccus we make the lacunosa and suggested it was closely related to Chirita fulva , transfer to Henckelia pending these studies. C. Integra and C. trisepala , distinguishable "from the other Henckelia urticifolia (D. Don.) A. Dietr., Sp. PL, ed. 6, 1: 574. species in the section [sect. Chirita] by an acaulescent habit and 1831 = Chirita urticifolia Buch.-Ham. ex D. Don, Prodr. a scapose inflorescence". Three species of the group have been Fl. Nepal.: 90. 1825 = Chirita grandiflora Wall., PL Asiat. included in the molecular analysis under their names in Chirita : Rar. 1: 43. 1830. - Distr.: India (Assam, Sikkim), Nepal, Chirita purpureolineata (the type species of Damrongia ), Bhutan, Northern Burma, Southern China (Yunnan). Chirita lacunosa and an as yet undescribed species, Chirita Henckelia walkerae (Gardner) D.J. Middleton & Mich. Moller, sp. nov. 2. The clade, with all species possessing straight fruits, comb. nov. = Chirita walkerae Gardner in Calcutta J. Nat. is nevertheless nested amongst the twisted-fruited genera. At Hist. 6: 480. 1846 (' walker /') = Chirita walkerae vax.parvi- present, only the chromosome number of Damrongia lacunosa flora C.B. Clarke in Candolle & Candolle, Monogr. Phan. is known with n = 9 (Kiehn & al., 1998). 5(1): 112. 1883 = Chirita walkerae Gardner subsp. parviWang & al. (2011) included several Chirita samples from flora (C.B. Clarke) W.L. Theob. & Grupe in Ceylon J. Sci., Moller & al. (2009) in their analysis but, curiously, did not Biol. Sci. 10(1): 70. 1972. - Distr.: Sri Lanka. - The spe- include Chirita lacunosa despite its isolated position in Moller cific epithet has previously almost always been spelt 'walk- & al. (2009). They, therefore, incorrectly included Damrongia ed' but Gardner makes very clear that his plant is named in synonymy of their maintained genus Chirita. after a Mrs. Walker so must be corrected to 'walkerae' under Art. 60.11 of the ICBN (McNeill & al., 2006). Damrongia cyanea (Ridl.) D.J. Middleton & A. Weber, comb, Henckelia wightii (C.B. Clarke) A. Weber & B.L. Burtt in nov. = Didymocarpus cyaneus Ridl. in J. Bot. 38: 68. Beitr. Biol. Pflanzen 90: 359. 1998 ('1997') = Didymocarpus 1900 = Chirita cyanea (Ridl.) B.L. Burtt in Notes Roy. rottlerianus var. wightii C.B. Clarke in Candolle & CanBot. Gard. Edinburgh 26: 267. 1965. - Distr.: Southern dolle, Monogr. Phan. 5(1): 101. 1883. - Distr.: South India. Thailand (Surat Thani). Henckelia zeylanica (R. Br.) A. Weber & B.L. Burtt in Beitr.Damrongia fulva (Barnett) D.J. Middleton & A. Weber, comb, Biol. Pflanzen 90: 359. 1998 ('1991') = Didymocarpus zeylanicus R. Br., Cyrtandreae: 1 19. 1839. - Distr.: Sri Lanka. nov. = Chirita fulva Barnett in Nat. Hist. Bull. Siam Soc. 20: 15. 1961. - Distr.: Southern Thailand (Nakhon Si Thammarat, Surat Thani). Damrongia Kerr ex Craib in Bull. Misc. Inform. Kew 1918: 364. Damrongia integra (Barnett) D.J. Middleton & A. Weber, comb, 1918 - Type: Damrongia purpureolineata Kerr ex Craib. nov. = Chirita integra Barnett in Nat. Hist. Bull. Siam Soc. Perennial herbs, acaulescent. Leaves whorled, petiolate, 20: 16. 1961. - Distr.: Southern Thailand (Krabi). lamina ovate or elliptic. Inflorescences axillary, scapose, Damrongia lacunosa (Hook, f.) D.J. Middleton & A. Weber, flowers 1-10; bracts paired or ternate, free or fused along one comb. nov. = Didymocarpus lacunosus Hook. f. in Bot. Ill This content downloaded from 142.51.238.106 on Fri, 09 Jun 2017 20:39:35 UTC All use subject to http://about.jstor.org/terms Weber & al. • Remodelling of Chirita TAXON 60 (3) • June 201 1: 767-790 Microchirita barbata (Sprague) A. Weber & D.J. Middleton, Mag. 1892: t. 7236. 1892 = Chirita lacunosa (Hook.f.) comb. nov. 26: = Chirita barbata Sprague in Bot. Mag. 134: B.L. Burtt in Notes Roy. Bot. Gard. Edinburgh 267. t. 8200. 1908. - Distr.: Only known from cultivated mate1965. - Distr.: Peninsular Malaysia (Pahang, Perak), rial but probably from Indochina. Southern Thailand (Krabi, Satun). Microchirita bimaculata Wood) A. Weber & D.J. MiddleDamrongia purpureolineata Kerr ex Craib in Bull. Misc. (D. Inton, comb. nov. = Chirita bimaculata D. Wood in Notes form. Kew 1918: 364. 1918 = Chirita purpureolineata (Kerr Roy. Bot. Gard. Edinburgh 31(3): 368. 1972. - Distr.: ex Craib) D. Wood in Notes Roy. Bot. Gard. Edinburgh Thailand (Chiang Mai). 31(3): 371. 1972. - Distr.: Northern Thailand (Lamphun). Microchirita caerulea (R. Br.) Y.Z. Wang in J. Syst. Evol. 49: Damrongia trisepala (Barnett) D.J. Middleton & A. Weber, 201 1 Hist. = ChiritaBull. caerulea R. Br., Cyrtandreae: 1 17. 1839 comb. nov. = Chirita trisepala Barnett in60. Nat. Siam Soc. 20: 18. 1961; et in Kew Bull =15: 254. 1961. - Clarke in Candolle & Candolle, Chirita zollingeri C.B. Monogr. Phan. 5(1): 125. 1883. - Distr.: Indonesia (Bali, Distr.: Central Thailand (Chanthaburi, Nakhon Nayok). Java, Sumba). Microchirita caliginosa (C.B.59. Clarke) Y.Z. Wang in J. Syst. Microchirita (C.B. Clarke) Y.Z. Wang in J. Syst. Evol. 49: 2011 = Chirita sect. Microchirita C.B. Clarke in Candolle Evol. 49: 60. 201 1 = Chirita caliginosa C.B. Clarke in Candolle & Candolle, Monogr. Phan. 5(1) 122. 1883 = Chirita & Candolle, Monogr. Phan. 5: 127. 1883 =Roettlera sect. fusca C.B. Clarke in Candolle & Candolle, Monogr. Phan. Microchirita (C.B. Clarke) Fritsch in Engler & Prantl, 5(1) 129. 1883 = Chirita parvula Ridl. in J. Fed. Malay Nat. Pflanzenfam. IV/3b: 148. 1895 = Didymocarpus sect. States Mus. 10: 149. 1920. - Distr.: Peninsular Malaysia Microchirita (C.B. Clarke) Chun in Sunyatsenia 6: 290. (Perak, Pahang, Selangor), Sarawak. 1946 - Type (lectotype Burtt, 1954): Microchirita hamosa Microchirita elphinstonia (Craib) A. Weber & D.J. Middleton, (R. Br.) Y.Z. Wang. comb. nov. = Chirita elphinstonia Craib in Bull. Misc. Annual or longer lived (flowering and fruiting for many Inform. Kew 1932: 149. 1932. - Distr.: Thailand (Prachin years in a ± aseasonal climate, dying down to a rootstock in Buri). the dry season of more seasonal climates), caulescent, stems often fleshy or juicy. Leaves in distant pairs, except lowermost Microchirita hamosa (R. Br.) Y.Z. Wang in J. Syst. Evol. 49: 60. 2011 = Chirita hamosa R. Br., Cyrtandreae: 117. 1839 = leaf solitary (macrocotyledon). Inflorescences usually two or Chirita hamosa vax.finlaysonia C.B. Clarke in Candolle & several in a leaf axil, often displaced onto the petiole, often Candolle, Monogr. Phan. 5(1) 128. 1883 = Chirita hamosa consisting of a short-stalked serial flower pair only, but this R. Br. var. unifolia C.B. Clarke in Candolle & Candolle, repeated several times (serial arrangement, "inflorescence Monogr. Phan. 5(1) 129. 1883 = Chirita cristata (Dalzell) crest"). Calyx lobes free to base, narrowly triangular to narB.L. Burtt in Notes Roy. Bot. Gard. Edinburgh 23: 96. 1960. rowly ovate, appressed to fruit. Corolla usually infundibuli- Distr.: Northern and Northeastern India, China (Guangxi, form, more rarely tubular or funnel-shaped; limb bilabiate, Yunnan), Burma, Thailand, Laos, Northern Vietnam. upper lip 2-lobed, lower lip 3-lobed, lobes rounded; colour Microchirita involucrata (Craib) Y.Z. Wang in J. Syst. Evol. white, bluish, blue, purple, yellow, orange or reddish. Stamens 49: 60. 201 1 (" involucrate ") = Chirita involucrata Craib in 2, anthers usually joined by an apical ligature (projections on Gard. Chron., ser. 3, 83: 140. 1928 = Chirita involucellata the connectives). Ovary not stipitate, stigma chiritoid. Fruit Craib in Gard. Chron., ser. 3, 83: 140. 1928 = Chirita capitis an elongate, straight or curved capsule, usually splitting into Craib in Bull. Misc. Inform. Kew 1930: 173. 1930 = Chirita 2 valves. Seeds minute, ellipsoid; (always?) with conspicuous knobs on the testa cells. geoffrayi Pellegr. in Lecomte & al., Fl. Indo-Chine 4: 529. 1930. - Distr.: Central and southern Thailand, Cambodia, Distribution: Ranging from the Western Ghats of India, the Peninsular Malaysia (Kedah, Kelantan, Penang, Perak, foothills of the Himalayas, through continental Southeast Asia Pahang). to Borneo, Sumatra and Java. - Rock plants, on limestone, growing in forest or on exposed rocks in the open or in light Microchirita lavandulacea (Stapf) Y.Z. Wang in J. Syst. Evol. 49: 60. 2011 = Chirita lavandulacea Stapf in Bot. Mag. shade. Approximately 18 species. 150: t. 9047. 1925. - Distr.: China, Northern Vietnam. Microchirita is perhaps the most readily recognisable of the genera to arise from Chirita. The highest concentration Microchirita marcanii (Craib) A. Weber & D.J. Middleton, comb. nov. = Chirita marcanii Craib in Bull. Misc. Inform. of species is in Thailand. Chromosome numbers of n = 9, 17, Kew 1926: 171. 1926. - Distr.: Thailand (Saraburi). 18 have been recorded (see 'Webcyte', Moller & al., 2002 onwards). All species currently known are found only on karst Microchirita micromusa (B.L. Burtt) A. Weber & D.J. Middlelimestone rocks, being sun-tolerant to some degree. Wang & al. ton, comb. nov. = Chirita micromusa B.L. Burtt in J. Roy. Hort. Soc. 85: 28. 1960. - Distr.: Thailand (Nakhon Nayok). (201 1), using sequence data from Li & Wang (2007) and Moller Microchirita mollissima (Ridl.) A. Weber & D.J. Middleton, & al. (2009), raised this group to genus level. comb. nov. = Chirita mollissima Ridl. in J. Linn. Soc., Bot. 32: 517. 1896. - Distr.: Thailand (Phangnga, Surat Thani). Microchirita aratriformis (D. Wood) A. Weber & D.J. MiddleMicrochirita oculata (Craib) A. Weber & D.J. Middleton, ton, comb. nov. = Chirita aratriformis D. Wood in Notes comb. nov. = Chirita oculata Craib in Bull. Misc. Inform. Roy. Bot. Gard. Edinburgh 31(3): 367. 1972. - Distr.: Northern Vietnam. Kew 1930: 174. 1930. - Distr.: Thailand (Prachin Buri). 778 This content downloaded from 142.51.238.106 on Fri, 09 Jun 2017 20:39:35 UTC All use subject to http://about.jstor.org/terms TAXON 60 (3) • June 201 1 : 767-790 Weber & al. • Remodelling of Chirita Liebigia is morphologically and biogeographically well Microchirita rupestris (Ridl.) A. Weber & Rafidah, comb. nov. defined. The speciesAsiat. representSoc. large, coarse perennial herbs, = Chirita rupestris Ridl. in J. Straits Branch Roy. elongate stems (to Branch 2 m long) and scabrid leaves in distant 44: 59. 1905 = Chirita glasgovii Ridl. with in J. Straits For a detailed description of the morphology see Hilliard Roy. Asiat. Soc. 44: 60. 1905. = Chirita pairs. kerrii Craib in Bull. Misc. Inform. Kew 1914: 129. 1914. - Distr.: Southern (2004) and the generic description above. Chromosome num- Thailand, Peninsular Malaysia (Perak, Perlis, Kedah). bers have been reported as n = 14 (Lee, 1962, as C. blumei) and 16 (Ratter & Prentice, 1967). Microchirita sahyadriensis (Punekar & Lakshmin.) A. Weber & D.J. Middleton, comb. nov. = Chirita sahyadriensis Punekar & Lakshmin. in Fol. Malaysiana 10: 18. 2009. Liebigia adenonema (Hilliard) Mich. Moller & A. Weber, comb. nov. = Chirita adenonema Hilliard in Edinburgh Distr.: India (Western Ghats). Microchirita sericea (Ridl.) A. Weber & Rafidah, comb. nov. J. Bot. 60: 381. 2004 ('2003'). - Distr.: Indonesia: Sumatra (North Sumatra). = Chirita sericea Ridl. in J. Straits Branch Roy. Asiat. Soc. 44: 59. 1905 = Chirita sericea var. scortechinii Ridl. Liebigia dissimilis (Hilliard) Mich. Moller & A. Weber, comb, nov. = Chirita dissimilis Hilliard in Edinburgh J. Bot. 60: in J. Straits Branch Roy. Asiat. Soc. 43: 59. 1905. - Distr.: 372. 2004 ('2003'). - Distr.: Indonesia: Sumatra (West Peninsular Malaysia (Kedah, Perak, Kelantan). Sumatra). Microchirita tubulosa (Craib) A. Weber & D.J. Middleton, comb. nov. = Chirita tubulosa Craib in Bull. Misc. Inform. Liebigia glabra (Miq.) Mich. Moller & A. Weber, comb. nov. Kew 1922: 173. 1922. - Distr.: Southern Thailand. = Chirita glabra Miq., Fl. Ind. Bat. 2: 729. 1858. - Distr.: Indonesia: Sumatra (West Sumatra, Jambi). Microchirita viola (Ridl.) A. Weber & Rafidah, comb. nov. = Chirita viola Ridl., J. Linn. Soc., Bot. 32: 516. 1896. Liebigia horsfieldii (R. Br.) Mich. Moller & A. Weber, comb. nov. = Chirita horsfieldii R. Br., Cyrtandreae: Distr.: S Thailand, Peninsular Malaysia (Kedah: Langkawi 117. 1839 = Chirita scaberrima R. Br., Cyrtandreae: 117. Islands). 1839 = Chirita horsfieldii R. Br. var. scaberrima (R. Br.) C.B. Clarke in Candolle & Candolle, Monogr. Phan. 5(1): Liebigia Endl., Gen. PI.: 1407. Feb-Mar. 1841 = Tromsdorf124. 1883 = Liebigia bracteosa Zoll. & Moritzi, Natuurfia Blume, Bijdr. Fl. Ned. Ind.: 762. 1826, non Bernhardi (1800), nec Martius (1826) = Morstdorffia Steud., No- Geneesk. Arch. Ned.-Indie 2: 574. 1845 = Chirita brac- mencl. Bot., ed. 2, 2: 162. Mar. 1841 = Chirita sect. Liebi- teosa (Zoll. & Moritzi) Miq., Fl. Ned. Ind. 2: 728. 1858. gia (Endl.) C.B. Clarke in Candolle & Candolle, Monogr. Phan. 5(1): 122. 1883; Hilliard in Edinburgh J. Bot. 60(3): - Distr.: Indonesia: Java (West Java), Bali. 364. 2004 ('2003') - Type: Liegibia speciosa (Blume) Endl. Liebigia leuserensis (Hilliard) Mich. Moller & A. Weber, comb. nov. = Chirita leuserensis Hilliard in Edinburgh J. Bot. 60: 383. 2004 ('2003'). - Distr.: Indonesia: Su- = Hypopteron Hassk., Cat. Hort. Bot. Bogor.: 154. 1844, nom. inval. = Bilabium Miq., Fl. Ned. Ind. 2: 730. 1858 - Type: Bilabium limans Miq. (= Liebigia limans (Miq.) Mich. Moller & A. Weber). Coarse perennial herbs or subshrubs, caulescent, stems often to 2 m tall. Leaves opposite, petiolate, lamina usually broadly elliptic to ovate, upper surface more or less asperous from harsh acute hairs. Inflorescences axillary, sometimes arising from small brachyblasts, bracts paired, either linearlanceolate and free from one another or broadly lanceolate to suborbicular and margins contiguous and partly connate. Calyx funnel-shaped or tubular, subequally 5-lobed or spathaceous (split open on one side). Corolla broad-tubed, ventricose, white or pinkish, limb bilabiate, upper lip 2-lobed, lower lip 3-lobed, lobes rounded. Stamens 2; anthers globose or heart-shaped with a distinct apiculus, anther surface often with conspicuous beard; anthers cohering face to face at tips. Ovary shortly stipi- tate, stigma chiritoid, lower lobe deeply bilobed (fish-tailed). Fruit an elongate capsule splitting into 2 valves; seeds minute, ellipsoid. matra (North Sumatra). Liebigia limans (Miq.) Mich. Moller & A. Weber, comb. nov. = Bilabium limans Miq., Fl. Ned. Ind. 2: 730. 1858 = Chirita bilabium C.B. Clarke in Candolle & Candolle, Monogr. Phan. 5(1): 127. 1883, nom. illeg. = Chirita limans (Miq.) B.L. Burtt in Notes Roy. Bot. Gard. Edinburgh 21: 196. 1954. - Distr.: Indonesia: Sumatra (West Sumatra). Liebigia neoforbesii (Hilliard) Mich. Moller & A. Weber, comb. nov. = Chirita neoforbesii Hilliard in Edinburgh J. Bot. 60: 368. 2004 ('2003') = Chirita blumei C.B. Clarke var. cordifolia S. Moore in J. Bot. 63 (Suppl.): 74. 1925. Distr.: Indonesia: Sumatra (South Sumatra). Liebigia polyneura (Miq.) Mich. Moller & A. Weber, comb, nov. = Chirita polyneura Miq., Fl. Ned. Bat. 2: 728. 1858 = Chirita polyneura Miq. var. albiflora Miq., Fl. Ned. Ind. 2: 728. 1858 = Chirita spectabilis Var. 13? pedicellata C.B. Clarke in Candolle & Candolle, Monogr. Phan. 5(1): 125. 1883 = Chirita alba Ridl. in J. Fed. Malay States Mus. 8(4): 68. 1917 = IChirita glabra Miq. var. intermedia in disturbed forest, open places and forest margins, river banks Ridl. in J. Fed. Malay States Mus. 8(4): 68. 1917. - Distr.: Indonesia: Sumatra (West Sumatra). Liebigia praeterita (Hilliard) Mich. Moller & A. Weber, comb, nov. = Chirita praeterita Hilliard in Edinburgh J. Bot. 60: etc.; probably growing in acid soil (limestone not recorded, but ecological information generally scanty). Twelve species in Hilliard (2004). Liebigia speciosa (Blume) Endl., Gen. PL, Suppl. I: 1407. 1841 = Tromsdorffia speciosa Blume, Bijdr. Fl. Ned. Ind.: 763. Distribution: Sumatra, Java and Bali. - Forest plants, also 370. 2004 ('2003'). - Distr.: Java. 779 This content downloaded from 142.51.238.106 on Fri, 09 Jun 2017 20:39:35 UTC All use subject to http://about.jstor.org/terms Weber & al. • Remodelling of Chirita TAXON 60 (3) • June 201 1: 767-790 1826 = Chirita blumei C.B. Clarke in Candolle & CanPrimulina , now comprising Chiritopsis , Wentsaiboea (except W. tiandengensis Yan Liu & B. Pan, a species found to be dolle, Monogr. Phan. 5(1): 122. 1883 = Chirita asperifolia more closely related to Lagarosolen [Moller & al., 2011] and (Blume) B.L. Burtt in Notes Roy. Bot. Gard. Edinburgh will be dealt with in a forthcoming paper) and the large 24: 41. 1962 = Chirita spectabilis Miq., Fl. Ned. Ind. which 3: 564. 1861 = Chirita forbesii S. Moore in J. Bot. 37: number 175. of species described in Chirita sect. Gibbosaccus , is a genus of perennial, acaulescent rosette plants, with no excep1899 = Chirita forbesii Ridl. in J. Straits Branch Roy. tions known so far. The inflorescences are axillary and scapose. Asiat. Soc. 44: 60. 1905, nom. illeg. = Chirita ridleyana Phyllotaxis is primarily opposite (easily seen in the type species S. Moore, J. Bot. 63 (Suppl.): 74. 1925. - Distr.: Indonesia: Sumatra (Benkulu, South Sumatra), Java, Krakatau Primulina tabacum and P. dryas = Chirita sinensis , the type of island. Chirita sect. Gibbosaccus - see photos of many other species in Li & Wang, 2004), or in whorls of three, but this may become Liebigia tenuipes (Hilliard) Mich. Moller & A. Weber, comb, nov. = Chirita tenuipes Hilliard in Edinburgh J. Bot.blurred 60: in many-leaved rosettes or phyllotaxy may change from 384. 2004 ('2003'). - Distr.: Indonesia: Sumatra (North opposite to alternate in the course of development. The flowSumatra). ers are usually infundibuliform or tubular, with a slightly to Liebigia tobaensis (Hilliard) Mich. Moller & A. Weber, comb, distinctly bilabiate limb. Only Primulina tabacum has salverform flowers with an almost actinomorphic limb, apparently nov. = Chirita tobaensis Hilliard in Edinburgh J. Bot. 60: 379. 2004 ('2003'). - Distr.: Indonesia : Sumatra (North reflecting a different pollination syndrome (see below). The Sumatra). distribution is essentially in the southern half of China, with about a dozen species spreading into or being restricted to Vietnam. There is some geographical overlap with Henckelia Primulina Hance in J. Bot. 21: 169. 1883 - Type: Primulina tabacum Hance. (in the present sense), especially in western and southern China, = Chirita sect. Gibbosaccus C.B. Clarke in Candolle & Can- but the species of the two genera seem to be ecologically well those of Primulina growing in limestone areas, those dolle, Monogr. Phan. 5(1): 130. 1883 - Type: Chirita separated, sinensis Lindl. (= Primulina dryas (Dunn) Mich. Moller of & Henckelia not. There is no overlap with Damrongia and A. Weber). Liebigia. Only one species of Microchirita , the widespread M. hamosa , is found within the range of Primulina. = Deltocheilos W.T. Wang in Bull. Bot. Res., Harbin, 1(3): 39. Available information suggests that Primulina is essen1981 - Type: Deltocheilos tenuitubum W.T. Wang (= Primtially a calciphilous genus with typical habitats being the karst ulina tenuituba (W.T. Wang) Y.Z. Wang). limestone towers in southern China. Guangxi province is par= Chiritopsis W.T. Wang in Bull. Bot. Res., Harbin, 1: 21. ticularly rich in species with about half of the species described 1981 - Type: Chiritopsis repanda W.T. Wang (= Primulina occurring there, either as local endemics or also occurring in repanda (W.T. Wang) Y.Z. Wang). other provinces such as Guizhou, Yunnan, Hunan, etc. = Wentsaiboea D. Fang & D.H. Qin in Acta Phytotax. Sin. It may be that some of the species transferred to Primulina 42(6): 534. 2004 - Type: Wentsaiboea renifolia D. Fang here may prove misplaced in future studies. Targeted studies & D.H. Qin (= Primulina renifolia (D. Fang & D.H. Qin) are necessary to identify problematic species and eventually Y.Z. Wang). Perennial rhizomatous herbs, acaulescent, rarely with a to find a better place for them. It is also our impression that too many species have been described with numerous pairs or short woody stem, rhizome usually stout. Leaves in a basal Inflorescences axillary, scapose, flowers in pair-flowered small groups of species growing in adjacent areas and differing only in slight, quantitative characters. We accept that after a critical revision of the genus many of the species described in Chirita sect. Gibbosaccus , and here transferred to Primulina , cymes, often umbel-like; bracts usually present, small to con- will have been sunk into synonymy. rosette, opposite (at least in young plants; later phyllotaxy often becoming blurred or possibly changing to alternate), petiolate or sessile, lamina form variable (including pinnately lobed). spicuous. Calyx lobes free to base or rarely fused at base to form a short tube, segments narrowly triangular to narrowly ovate, sometimes toothed or fimbriate. Corolla usually infundibuliform, more rarely tubular or salverform; limb bilabiate with 2-lobed upper lip and 3-lobed lower lip, rarely almost actinomorphic, lobes rounded, colouration variable, often with two raised yellow bars on lower lip. Stamens 2, anthers fused face to face; ovary not stipitate, sometimes one carpel sterile, stigma chiritoid, rarely with short upper lobe, lower lobe usually bilobed. Fruit a capsule splitting into 2 valves, sometimes (esp. in species with only one fertile carpel) probably opening only on dorsal side and plagiocarpic. Seeds minute, ellipsoid. Distribution: Western and Southern China, Vietnam. Plants growing (as far as is known) on limestone. Approximately 131 species (but see comments below). The relationship of Chiritopsis to Chirita (sect. Gibbosaccus) is evident from the name. The main difference is in the small flowers and the short ovoid capsule. The stigma is chiritoid in the enlarged and often bilobed condition of the lower stigma lobe. Recently described species further blur the boundaries between the two genera (e.g., Chirita macrodonta , with Chiritopsis- like foliage - Fang & al., 1994; Chiritopsis lingchuanensis with typical Chirita characteristics, such as an ovary shorter than the style, and a calyx length longer than the capsule - Liu & al., 2006). It is, therefore, no great surprise to see Chiritopsis embedded in Chirita sect. Gibbosaccus (as was already found by Li & Wang, 2007, Moller & al., 2009 and Wang & al., 2011) and Primulina , respectively. Though Primulina tabacum , previously the sole representative of Primulina , looks quite different from the species 780 This content downloaded from 142.51.238.106 on Fri, 09 Jun 2017 20:39:35 UTC All use subject to http://about.jstor.org/terms TAXON 60 (3) • June 201 1: 767-790 Weber & al. • Remodelling of Chirita W.T.the Wang &narrow-tubed D.Y. Chen in Bull. Bot. Res., Harbin 5(3): of Chirita sect. Gibbosaccus , especially in 54. 1985. - Distr.: China (Guizhou). flowers with an almost actinomorphic patent limb (a salverform Primulina brachytricha var. magnibracteata flower), the relationship is uncontroversial. The species has (W.T. Wang & D.Y.habit Chen) Mich. Moller opposite & A. Weber, comb. nov. = Chirita a typical chiritoid stigma and the rosette with brachytricha var. magnibracteata W.T. leaves also suggests a close relationship. In the treatments of Wang & D.Y. Chen in Bull. Bot.Primulina Res., Harbin 5(3): 55. Wang & al. (1990, 1998) and Li & Wang (2004) is1985. - Distr.: China (Guizhou). interpretation directly associated with Chirita. The functional of the different flower form seems rather easy. Primulina Primulina brassicoides (W.T. Wang)tabaMich. Moller & A. We- ber, comb. nov.to = Chirita brassicoides W.T. Wang in Bull. cum has apparently switched from bee-pollination butterfly- pollination. In their form and vivid colouration the flowers Bot. Res., Harbin 1(4): 63. 1981. - Distr.: China (Guangxi). exhibit the typical syndrome of psychophilous flowers. Field Primulina brunnea (W.T. Wang) Mich. Moller & A. Weber, studies are urgently needed to confirm or reject hypothesis. comb. nov.this = Chirita brunnea W.T. Wang in Bull. Bot. Res., Harbin 3(4):said 48. 1983. - Distr.: The recently described genus Wentsaiboea was to be Vietnam. similar to Dolicholoma and ChiritopsisPrimulina by Fang & Qin (2004). carnosifolia (C.Y. Wu ex H.W. Li) Y.Z. Wang in The differences to the latter, now in Primulina in 49: the palmately J. Syst.,Evol. 60. 201 1 = Chirita carnosifolia C.Y. Wu ex H.W. and Li in Bull. Bot. "obliquely Res., Harbin 3(2): 36. 1983. - Distr.: nerved leaves, the campanulate corolla tube the Chinathe (Yunnan). hippocrepiform" stigma would appear, from molecular results, not to be very significant. Its position within Primulina Primulina colaniae (Pellegr.) Mich. Moller & A. Weber, comb, thus does not pose a great problem. nov. = Chirita colaniae Pellegr. in Bull. Soc. Bot. France 73: 418. 1926. - Distr.: Despite the large number of species described, the Northern chro- Vietnam. mosome number of only one species, Chirita sinensis (which Primulina confertiflora (W.T. Wang) Mich. Moller & A. Weis the type species of Chirita sect. Gibbosaccus is confertiflora W.T. Wang ber, comb.and nov. =which Chiritopsis in Bull. Bot. Res., Harbin 1(3): 25. 1981. - Distr.: China now Primulina dryas : n = 18) has so far been published. Further counts, confirming that number, have been made by F. Christie (Guangdong). Primulina cordata Mich. Moller & A. Weber, nom. nov. = at the Royal Botanic Garden Edinburgh (unpub.). Chirita cordifolia W.T. Wang in Bull. Bot. Res., Harbin Primulina annamensis (Pellegr.) Mich. Moller & A. Weber, 1(4): 51. 1981. - Distr.: China (Guangxi). comb. nov. = Chirita annamensis Pellegr. Lecomte Primulina in cordifolia (D. Fang&& W.T. Wanga) Y.Z. Wang in al., Fl. Indo-Chine 4: 530. 1930. - Distr.: Vietnam. J. Central Syst. Evol. 49: 61. 2011 = Chiritopsis cordifolia D. Fang Primulina atroglandulosa (W.T. Wang) Mich. & A. & W.T.Moller Wang in Bull. Bot.WeRes., Harbin 2(4): 54. 1982, non ber, comb. nov. = Chirita atroglandulosa W.T. Wang in Chirita cordifolia W.T. Wang. - Distr.: China (Guangxi). Novon 7(4): 423. 1998 ('1997'). - Distr.: China (Guangxi). Primulina crassituba (W.T. Wang) Mich. Moller & A. Weber, Primulina atropurpurea (W.T. Wang) Mich. & A. We- W.T. Wang in Bull. Bot. comb. Moller nov. = Chirita crassituba ber, comb. nov. = Chirita atropurpurea W.T. Wang Res., Harbin 9(4):in 24.Bull. 1989. - Distr.: China (Hunan). Bot. Res., Harbin 1(4): 60. 1981. - Distr.: China (Guangxi). Primulina cruciformis (Chun) Mich. Moller & A. Weber, Primulina baishouensis (Y.G. Wei, H.Q. Wen & S.H. Zhong)cruciformis Chun in Sunyatcomb. nov. = Didymocarpus Y.Z. Wang in J. Syst. Evol. 49: 60. 2011 = 6: Chirita baishosenia 296. 1946 = Chirita cruciformis (Chun) W.T. Wang uensis Y.G. Wei, H.Q. Wen & S.H. Zhong Phytotax. in in Bull.Acta Bot. Res., Harbin 1(4): 45. 1981. - Distr.: China Sin. 38(3): 299. 2000. - Distr.: China (Hunan). (Guangxi). Primulina balansae (Drake) Mich. Moller & A. Weber, comb, Primulina cycnostyla (B.L. Burtt) Mich. Moller & A. Weber, nov. = Chirita balansae Drake in Bull. Annuel Soc. Philom. comb. nov. = Chirita cycnostyla B.L. Burtt in Notes Roy. Paris, ser. 8, 2: 138. 1890. - Distr.: Northern Vietnam. Bot. Gard. Edinburgh 23: 96. 1960. - Distr.: Central Viet- nam. Primulina bicolor (W.T. Wang) Mich. Moller & A. Weber, comb. nov. = Chirita bicolor W.T. Wang in Bull. Bot. Res., Primulina cyrtocarpa (D. Fang & L. Zeng) Mich. Moll Harbin 5(3): 59. 1985. - Distr.: China (Guangdong). A. Weber, comb. nov. = Chirita cyrtocarpa D. Fang Primulina bipinnatifida (W.T. Wang) Y.Z. J.Phytotax. Syst. L. Wang Zeng in in Acta Sin. 31(5): 468. 1993. - Dist Evol. 49: 60. 2011 = Chiritopsis bipinnatifida W.T. Wang China (Guangxi). in Bull. Bot. Res., Harbin 1(3): 26. 1981. - Distr.: China Primulina demissa (Hance) Mich. Moller & A. Weber, com (Guangxi). nov. = Didymocarpus demissus Hance in J. Bot. 21: 1 Primulina bogneriana (B.L. Burtt) Mich. Moller & A. Weber, 1883 = Chirita demissa (Hance) W.T Wang in Novon 7( comb. nov. = Chirita bogneriana B.L. Burtt in Gard. Bull. 424. 1998 ('1997'). - Distr.: China (Guangdong). - F Singapore 54: 239. 2002. - Distr.: Vietnam. ther studies are required to confirm its generic placeme Primulina brachystigma (W.T. Wang) Mich. Moller & A. Weber, Primulina depressa (Hook, f.) Mich. Moller & A. Weber, co comb. nov. = Chirita brachystigma W.T Wang in Bull. Bot. nov. = Chirita depressa Hook. f. in Bot. Mag. t. 7213. 189 Res., Harbin 4(1): 22. 1984. - Distr.: China (Guangxi). Primulina brachytricha (W.T. Wang & D.Y. Chen) Y.Z. Wang - Distr.: China (Guangdong). Primulina drakei (B.L. Burtt) Mich. Moller & A. Web in J. Syst. Evol. 49: 60. 2011. = Chirita brachytricha comb. nov. = Chirita bracteosa Drake in Bull. Annuel 781 This content downloaded from 142.51.238.106 on Fri, 09 Jun 2017 20:39:35 UTC All use subject to http://about.jstor.org/terms Weber & al. • Remodelling of Chirita TAXON 60 (3) • June 201 1: 767-790 Soc. Philom. Paris, ser. 8, 2: 39. 1890,2011 nom. = Chiritailleg., gemella D. non Wood in Notes Roy. Bot. Gard. Edinburgh 31(3): 1972. - Distr.: Vietnam (Quang Chirita bracteosa (Zoll. & Moritzi) Miq. (1858) =370. Chirita Ninh). Edinburgh 23: drakei B.L. Burtt in Notes Roy. Bot. Gard. 98. 1960. - Distr.: Northern Vietnam. Primulina glabrescens (W.T. Wang & D.Y. Chen) Mich. Moller Primulina dryas (Dunn) Mich. Moller & A. Weber, comb, & A. Weber, comb. nov. = Chirita glabrescens W.T. Wang nov. = Chirita dryas Dunn in Bull. Misc. Inform. Kew, & D.Y. Chen in Bull. Bot. Res., Harbin 5(3): 57. 1985. Addit. Ser. 10: 195. 1912 = Chirita sinensis Lindl. in EdDistr.: China (Guizhou). wards's Bot. Reg. 30: t. 59. 1844. = Didymocarpus sinensis Primulina glandulosa (D. Fang, L. Zeng & D.H. Qin) Y.Z. Wang in J. Syst. Evol. 49: 61. 2011 = Chiritopsis glandulosa (Lindl.) H. Lev. in Compt. Rend. Assoc. Fran?. Avancem. Sci. 34: 427. 1906 = Primulina sinensis (Lindl.) Y.Z. Wang D. Fang, L. Zeng & D.H. Qin in Acta Phytotax. Sin. 31(5): 470. 1993. - Distr.: China (Guangxi). in J. Syst. Evol. 49: 62. 2011, nom. illeg., won Primulina sinensis Hook. f. in Bot. Mag. 1 16: pl. 71 17. 1890, nom. illeg. = Primulina glandulosa var. yangshuoensis (F. Wen, Yue Wang & Q.X. Zhang) Mich. Moller & A. Weber, comb. nov. = Chirita sinensis var. angustifolia Dunn in J. Bot. (London) 45: 403. 1907 = Chirita sinensis var. bodinieri H. Lev. in Chiritopsis glandulosa D. Fang, L. Zeng & D.H. Qin var. yangshuoensis F. Wen, Yue Wang & Q.X. Zhang in GuiFeddes Repert. 9: 328. 1911. - Distr.: China (Guangdong). haia 28(3): 291. 2008. - Distr.: China (Guangxi). Primulina eberhardtii (Pellegr.) Mich. Moller & A. Weber, comb. nov. = Chirita eberhardtii Pellegr. in Bull. Soc. Bot. Primulina grandibracteata (J.M. Li & Mich. Moller) Mich. France 73: 418. 1926. - Distr.: Central Vietnam. Moller & A. Weber, comb. nov. = Chirita grandibracteata Primulina eburnea (Hance) Y.Z. Wang in J. Syst. Evol. 49: 61. 2011 = Chirita eburnea Hance in J. Bot. 21: 168. 1883 = J.M. Li & Mich. Moller in Ann. Bot. Fenn. 46(2): 126. 2009. - Distr.: China (Yunnan). Chirita fauriei Franch. in Bull. Mens. Soc. Linn. Paris 1: Primulina gueilinensis (W.T. Wang) Y.Z. Wang in J. Syst. Evol. 49: 61. 2011 = Chirita gueilinensis W.T. Wang in 45. 1885. - Distr.: China (Guangdong, Guangxi, Guizhou, Bull. Bot. Res., Harbin 1(4): 43. 1981. - Distr.: China Hubei, Hunan, Sichuan). Primulina fangii (W.T. Wang) Mich. Moller & A. Weber, (Guangdong, Guangxi). comb. nov. = Chirita fangii W.T. Wang in Bull. Bot. Res.,Primulina guihaiensis (Y.G. Wei, B. Pan & W.X. Tang) Mich. Harbin 2(4): 51. 1982. - Distr.: China (Sichuan). Primulina fimbrisepala (Hand.-Mazz.) Y.Z. Wang in J. Syst. Moller & A. Weber, comb. nov. = Chirita guihaiensis Bull. Bot. Res., Harbin 5(3): 40. 1985. - Distr.: China 61. 201 1 = Chirita heterotricha Merr. in Lingnan Sci. J. 13: Y.G. Wei, B. Pan & W.X. Tang in Nordic J. Bot. 25(5-6): 296. 2008 ('2007'). Distribution: China (Guangxi). Evol. 49: 61. 2011 = Chirita fimbrisepala Hand.-Mazz. in Anz. Akad. Wiss. Wien, Math.-Naturwiss. Kl. 62: 65. Primulina halongensis (Kiew & T.H. Nguyen) Mich. Moller & A. Weber, comb. nov. = Chirita halongensis Kiew & 1925. - Distr.: China (Fujian, Guangdong, Guangxi, T.H. Nguyen in Gard. Bull. Singapore 52(1): 189. 2000. Guizhou, Hunan, Jiangxi). Distr.: Vietnam (Quang Ninh). Primulina fimbrisepala var. mollis (W.T. Wang) Mich. Moller & A. Weber, comb. nov. = Chirita fimbrisepala Hand.- Primulina hedyotidea (Chun) Y.Z. Wang in J. Syst. Evol. 49: 61. 201 1 = Didymocarpus hedyotideus Chun in Sunyatsenia 6: Mazz. var. mollis W.T. Wang in Bull. Bot. Res., Harbin 291. 1946 = Chirita hedyotidea (Chun) W.T. Wang in Bull. 2(4): 50. 1982. - Distr.: China (Guangxi). Bot. Res., Harbin 1(4): 65. 1981. - Distr.: China (Guangxi). Primulina flavimaculata (W.T. Wang) Mich. Moller & A. Weber, comb. nov. = Chirita flavimaculata W.T. Wang in Primulina heterotricha (Merr.) Y.Z. Wang in J. Syst. Evol. 49: 71 (1934). - Distr.: China (Hainan). (probably Guangxi). Primulina floribunda (W.T. Wang) Mich. Moller & A. Weber,Primulina hiepii (Kiew) Mich. Moller & A. Weber, comb. nov. = Chirita hiepii Kiew in Gard. Bull. Singapore 52(1): 192. comb. nov. = Chirita floribunda W.T. Wang in Bull. Bot. 2000. - Distr.: Vietnam (Quang Ninh). Res., Harbin 4(1): 23. 1984. - Distr.: China (probably Primulina hochiensis (C.C. Huang & X.X. Chen) Mich. Moller Guangxi). & A. Weber, comb. nov. = Chirita hochiensis C.C. Huang Primulina fordii (Hemsl.) Y.Z. Wang in J. Syst. Evol. 49: 61. 2011 = Didymocarpus fordii Hemsl. in J. Linn. Soc., Bot. 26: 229. 1890 = Chirita fordii (Hemsl.) D. Wood in Notes Roy. Bot. Gard. Edinburgh 31(3): 371. 1972 = Chirita longi styla W.T. Wang in Bull. Bot. Res., Harbin 1(4): 48. 1981. - Distr.: China (Guangdong, Guangxi, Guizhou, Hunan, Sichuan). Primulina fordii var. dolichotricha (W.T. Wang) Mich. Moller & A. Weber, comb. nov. = Chirita gueilinensis W.T. Wang var. dolichotricha W.T. Wang in Bull. Bot. Res., Harbin 2(4): 50. 1982 = Chirita fordii (Hemsl.) D. Wood var. dolichotricha (W.T. Wang) W.T. Wang in Guihaia 12(4): 296. 1992. - Distr.: China (Guangdong). Primulina gemella (D. Wood) Y.Z. Wang in J. Syst. Evol. 49: 61. & X.X. Chen in Bot. J. S. China 1: 14 (1992). - Distr.: China (Guangxi). Primulina jiuwanshanica (W.T. Wang) Y.Z. Wang in J. Syst. Evol. 49: 61. 2011 = Chirita jiuwanshanica W.T. Wang in Guihaia 12(4): 294 (1992). - Distr.: China (Guangxi). Primulina juliae (Hance) Mich. Moller & A. Weber, comb, nov. = Chirita juliae Hance in J. Bot. 21: 168. 1883 = Chirita mangshanensis W.T. Wang in Bull. Bot. Res., Harbin 1(4): 49. 1981 = Chirita mangshanensis W.T. Wang var. lasiandra W.T. Wang in Bull. Bot. Res., Harbin 4(1): 15. 1984 = Chirita gueilinensis W.T. Wang var. brachycarpa W.T. Wang in Bull. Bot. Res., Harbin 4(1): 15. 1984. Distr.: China (Fujian, Guangdong, Hunan, Jiangxi). 782 This content downloaded from 142.51.238.106 on Fri, 09 Jun 2017 20:39:35 UTC All use subject to http://about.jstor.org/terms TAXON 60 (3) • June 201 1: 767-790 Weber & al. • Remodelling of Chirita Primulina langshanica (W.T. Wang) Primulina Y.Z. Wanglunglinensis in J. Syst. (W.T. Wang) Mich. Mo Evol. 49: 61. 2011 = Chirita langshanica W.T. Wang in= Chirita lunglinensis W.T W ber, comb. nov. Guihaia 12(4): 293 (1992). - Distr.: China (Hunan). Bot. Res., Harbin 1(4): 53. 1981. - Distr.: Ch Primulina latinervis (W.T. Wang) Mich. Guizhou). Moller & A. Weber, comb. nov. = Chirita latinervis W.T. Wang in lunglinensis Bull. Bot. Primulina var. amblyosepala (W. Res., Harbin 9(4): 21. 1989. - Distr.: China (Hunnan). Moller & A. Weber, comb. nov. = Chirita lun Primulina laxiflora (W.T. Wang) Y.Z. Wang in J. Syst.W.T. Evol. amblyosepala Wang in Bull. Bot. Res., 49: 61. 2011 = Chirita laxiflora W.T. Wang in Bull. Bot. China (Guangxi). 48. 1982. - Distr.: Res., Harbin 4(1): 21. 1984. - Distr.:Primulina China (Guangxi). lungzhouensis (W.T. Wang) Mic Primulina leeii (F. Wen, Yue Wang & A. Q.X. Zhang) Mich. Weber, comb. nov. = Chirita lungzhouen Moller & A. Weber, comb. nov. = Chirita leeiiBot. F. Wen, in Bull. Res., Harbin 1(4): 56. 1981. - D Yue Wang & Q.X. Zhang in Guihaia (Guangxi). 29(6): 719. 2009. Distr.: China (Guangxi). Primulina luochengensis (Yan Liu & W.B. Xu) Mich. Moller Primulina leiophylla (W.T. Wang) Y.Z. Wang in comb. J. Syst. Evol. luochengensis Yan & A. Weber, nov. = Wentsaiboea & W.B. Xu Nordic J. Bot. 28: 743. 2010. - Distr.: 49: 61. 2011 = Chirita leiophylla W.T. Liu Wang ininGuihaia 6(3): 159 (1986). - Distr.: China (Guangxi). China (Guangxi). Primulina liboensis (W.T. Wang & D.Y. Chen) Mich. Primulina lutea (Yan LiuMoller & Y.G. Wei) Mich. Moller & A. Weber, comb. nov.W.T. = Chirita lutea Yan Liu & Y.G. Wei, J. Wuhan & A. Weber, comb. nov. = Chirita liboensis Wang & D.Y Chen in Bull. Bot. Res., Harbin 56. 1985. Bot. 5(3): Res. 22(5): 391. 2004. - Distr.: China (Guangxi). Distr.: China (Guizhou). Primulina luzhaiensis (Yan Liu, Y.S. Huang & W.B. Xu) Mich. Moller & A. Weber, nov. = Chirita luzhaiensis Yan Primulina lienxienensis (W.T. Wang) Mich. Moller & A.comb. Weber, comb. nov. = Chirita lienxienensis W.T.Liu, Wang in Bull. Bot. Y.S. Huang & W.B. Xu in J. Trop. Subtrop. Bot. 18(2): Res., Harbin 2(4): 48. 1982. - Distr.: China (Guangdong). 137. 2010. - Distr. China (Guangxi). Primulina liguliformis (W.T. Wang) Mich. Moller & A.(D. Weber, Primulina macrodonta Fang & D.H. Qin) Mich. Moller & comb. nov. = Chirita liguliformis W.T. A. Wang Bull. Bot. Weber, in comb. nov. = Chirita macrodonta D. Fang & Res., Harbin 4(1): 20. 1984. - Distr.: China D.H. Qin(Guizhou). in Acta Phytotax. Sin. 32(6): 566. 1994. - Distr.: Primulina linearifolia (W.T. Wang) Y.Z. Wang in J. Syst. Evol. China (Guangxi). 49: 61. 2011 = Chirita linearifolia W.T. Wang in Bull. Bot. Primulina macrorhiza (D. Fang & D.H. Qin) Mich. Moller & Res., Harbin 2(2): 136. 1982. - Distr.: China A. Weber,(Guangxi). comb. nov. = Chirita macrorhiza D. Fang & Primulina lingchuanensis (Yan Liu & Y.G. Moller D.H.Wei) Qin in Mich. Acta Phytotax. Sin. 32(6): 568. 1994. - Distr.: & A. Weber, comb. nov. = Chiritopsis China lingchuanensis Yan (Guangxi). Liu & Y.G. Wei in Acta Phytotax. Sin. 44(3): 340. 2006. Primulina maguanensis (Z. Yu Li, H. Jiang & H. Xu) Mich. - Distr.: China (Guangxi). Moller & A. Weber, comb. nov. = Chirita maguanensis Primulina linglingensis (W.T. Wang) Mich. &&A. WeZ. Yu Moller Li, H. Jiang H. Xu in Bot. J. Linn. Soc. 158(2): ber, comb. nov. = Chirita linglingensis W.T. Wang in China Bull. (Yunnan). 269. 2008. - Distr.: Bot. Res., Harbin 1(4): 42. 1981. - Distr.: PrimulinaChina medica (D.(Hunan). Fang) Y.Z. Wang in J. Syst. Evol. 49: 61. Primulina liujiangensis (D. Fang & D.H. Qin) Y.Z. Wang in 2011 = Chirita medica D. Fang in Bull. Bot. Res., Harbin J. Syst. Evol. 49: 61. 2011 = Chirita liujiangensis Fang & (Guangxi). 1(4): 64. 1981. -D. Distr.: China D.H. Qin in Acta Phytotax. Sin. 32(6): 569. minutihamata 1994. - Distr.: Primulina (D. Wood) Mich. Moller & A. Weber, China (Guangxi). comb. nov. = Chirita minutihamata D. Wood in Notes Roy. Primulina lobulata (W.T. Wang) Mich. Moller &Edinburgh A. Weber, Bot. Gard. 31(3): 370. 1972 = Chirita cicatricomb. nov. = Chiritopsis lobulata W.T. cosa Wang in Bull. Bot. W.T. Wang in Bull. Bot. Res., Harbin 1(4): 69. 1981. Res., Harbin 2(4): 55. 1982. - Distr.: China - Distr.:(Guangdong). China (Guangxi), Central Vietnam. Primulina longgangensis (W.T. Wang)Primulina Y.Z. Wang in J. (D. Syst. minutimaculata Fang & W.T. Wang) Y.Z. Wang Evol. 49: 61. 2011 = Chirita longgangensis W.T in = Chirita minutimaculata in J. Syst. Evol.Wang 49: 61. 2011 Guihaia 2(4): 171. 1982. - Distr.: ChinaD.(Guangxi). Fang & W.T. Wang in Bull. Bot. Res., Harbin 1(4): 55. Primulina longgangensis var. hongyao1981. (S.Z. Huang) Mich. - Distr.: China (Guangxi). Moller & A. Weber, comb. nov. = Chirita Primulinalonggangensis modesta (Kiew & T.H. Nguyen) Mich. Moller A. Weber, comb. nov. = -Chirita modesta Kiew & var. hongyao S.Z. Huang in Guihaia& 2(4): 173. 1982. Distr.: China (Guangxi). T.H. Nguyen in Gard. Bull. Singapore 52(1): 194. 2000. - Distr.: Vietnam. Primulina longicalyx (J.M. Li & Y.Z. Wang) Mich. Moller & A. Weber, comb. nov. = Chirita longicalyx J.M.(D. LiFang & & W.T. Wang) Y.Z. Wang in Primulina mollifolia Y.Z. Wang in Ann. Bot. Fenn. 45(3): 212. 2008. - Distr.: J. Syst. Evol. 49: 61. 2011 = Chiritopsis mollifolia D. Fang China (Guangxi). & W.T. Wang in Guihaia 6(1-2): 6. 1986. - Distr.: China Primulina longii (Z. Yu Li) Y.Z. Wang in J. Syst. Evol. 49: 61. (Guangxi). 2011 = Chirita longii Z. Yu Li in Novon 12(4): 492. 2002. Primulina nandanensis (S.X. Huang, Y.G. Wei & W.H. Luo) Mich. Moller & A. Weber, comb. nov. = Chirita nandanen- Distr.: China (Guangxi). 783 This content downloaded from 142.51.238.106 on Fri, 09 Jun 2017 20:39:35 UTC All use subject to http://about.jstor.org/terms Weber & al. • Remodelling of Chirita TAXON 60 (3) • June 201 1 : 767-790 sis S.X. Huang, Y.G. Wei & W.H. Luo in Ann. Bot. Evol. 49: 62. 2011 Fenn. = Wentsaiboea renifolia D. Fang & 47: 139. 2010. - Distr.: China (Guangxi). D.H. Qin in Acta Phytotax. Sin. 42(6): 534. 2004. - Distr.: Primulina napoensis (Z. Yu Li) Mich. Moller & A. Weber, China (Guangxi). comb. nov. = Chirita napoensis Z. Yu Li in repanda Novon 7(4): Primulina (W.T. Wang) Y.Z. Wang in J. Syst. Evol. 424. 1998 ('1997'). - Distr.: China (Guangxi). 49: 62. 201 1 = Chiritopsis repanda W.T. Wang in Bull. Bot. Primulina obtusidentata( W.T. Wang) Mich. Res., Moller & 1(3): A. 23. Weber, Harbin 1981. - Distr.: China (Guangxi). comb. nov. = Chirita obtusidentata Primulina W.T. Wang repanda in var.Bull. guilinensis (W.T. Wang) Mich. Moller Bot. Res., Harbin 1(4): 57. 1981 = Chirita laifengensis & A. Weber, comb. nov. = Chiritopsis repanda var. guilin- W.T. Wang in Bull. Bot. Res., Harbin 4(1): 1984. ensis W.T.17. Wang in Guihaia 12(4): 299. 1992. - Distr.: Distr.: China (Guizhou, Hubei). China (Guangxi). Primulina obtusidentata var. mollipes (W.T. Wang) (D. Mich. Primulina ronganensis Fang & Y.G. Wei) Mich. Moller Moller & A. Weber, comb. nov. = Chirita obtusidentata & A. Weber, comb. nov. = Chirita ronganensis D. Fang & var. mollipes W.T. Wang in W.T. WangY.G. & Wei al.,inKeys Vase. Sin. 39(5): 467. 2001. - Distr.: Acta Phytotax. PI. Wuling Mts.: 581. 1995. - Distr.: China (Hunan). China (Guangxi). Primulina ophiopogoides (D. Fang & W.T. Wang) Y.Z. Wang Primulina roseoalba (W.T. Wang) Mich. Moller & A. Weber, in J. Syst. Evol. 49: 62. 2011 = Chirita comb.ophiopogoides nov. = Chirita roseoalba W.T. Wang in Bull. Bot. Res.,Harbin Harbin 1(4): 44. 1981. D. Fang & W.T. Wang in Bull. Bot. Res., 1(4): 68. - Distr.: China (Hunan). 1981. - Distr.: China (Guangxi). Primulina rotundifolia (Hemsl.) Mich. Moller & A. Weber, Primulina orthandra (W.T. Wang) Mich. Moller & =A. Weber, comb. nov. Didymocarpus rotundifolius Hemsl. in comb. nov. = Chirita orthandra W.T. Wang Bull. Bot. J. Linn.in Soc., Bot. 26: 230. 1890. = Chirita rotundifolia Res., Harbin 5(3): 42. 1985. - Distr.: China (Hemsl.)(Guangdong). D. Wood in Notes Roy. Bot. Gard. Edinburgh 31: Primulina parvifolia (W.T. Wang) Y.Z. Wang in J. Syst. Evol. 371. 1972. - Distr.: China (Guangdong). 49: 62. 2011 = Chirita parvifolia W.T. Wang in Bull. Bot. Primulina sclerophylla (W.T. Wang) Y.Z. Wang in J. Syst. Evol. Res., Harbin 1(4): 50. 1981. - Distr.: China (Guangxi). 49: 62. 201 1 = Chirita sclerophylla W.T. Wang in Bull. Bot. Primulina pinnata (W.T. Wang) Y.Z. Wang Res., in J. Syst. 49: Harbin 1(4): Evol. 46. 1981. - Distr.: China (Guangxi). 62. 2011 = Chirita pinnata W.T. Wang in Bull. Bot. Res., Primulina secundiflora (Chun) Mich. Moller & A. Weber, Harbin 4(1): 25. 1984. - Distr.: China (Guangxi). comb. nov. = Didymocarpus secundiflorus Chun in Primulina pinnatifida (Hand.-Mazz.) Y.Z. Sunyatsenia Wang in 6: J. 293.Syst. 1946 = Chirita secundiflora (Chun) Evol. 49: 62. 2011 = Didymocarpus pinnatifidus Hand.W.T. Wang in Bull. Bot. Res., Harbin 2(4): 52, as 4 Chirta' 1982. - Distr.: China (Guizhou). Mazz. in Sinensia 5: 8. 1934 = Chirita pinnatifida (Hand.Mazz.) B.L. Burtt in Notes Roy. Bot. Primulina Gard. Edinburgh 23: semicontorta (Pellegr.) Mich. Moller & A. Weber, 99. 1960 = Chirita quercifolia D. Wood in Notes comb. nov. = Roy. Chirita Bot. semicontorta Pellegr. in Lecomte & Gard. Edinburgh31: 369. 1972. - Distr.: (Fujian, al., Fl.China Indo-Chine 4: 531. 1930. - Distr.: Vietnam (Quang Guangdong, Guangxi, Hunan, Jiangxi,Ninh). Zhejiang). Primulina poilanei (Pellegr.) Mich. Moller & shouchengensis A. Weber, comb, Primulina (Z. Yu Li) Y.Z. Wang in J. Syst. nov. = Chirita poilanei Pellegr. in Bull. Soc. France Evol. 49: 62.Bot. 2011 = Chirita shouchengensis Z.Y. Li in 73: 419. 1926. - Distr.: Central Vietnam. Novon 7(4): 426. 1998 ('1997'). - Distr.: China (Guangxi). Primulina polycephala (Chun) Mich. Moller & A. Weber, Primulina sichuanensis (W.T. Wang) Mich. Moller & A. Wecomb. nov. = Didymocarpus polycephalus Chun in ber, comb. nov. = Chirita sichuanensis W.T. Wang in Bull. Bot. Res., Harbin 4(1): 16. 1984. - Distr.: China (Sichuan). Sunyatsenia 6: 299. 1946 = Chirita polycephala (Chun) W.T. Wang in Bull. Bot. Res., Harbin 1(4): 58. 1981. Primulina skogiana (Z. Yu Li) Mich. Moller & A. Weber, comb. nov. = Chirita skogiana Z. Yu Li in Novon 7(4): Primulina pseudoeburnea (D. Fang & W.T. Wang) Mich. 427. 1998 ('1997'). - Distr.: China (Gansu). Distr.: China (Guangdong). Moller & A. Weber, comb. nov. = Chirita pseudoeburnea Primulina spadiciformis (W.T. Wang) Mich. Moller & A. Weber, D. Fang & W.T. Wang in Bull. Bot. Res., Harbin 1(4): 54. comb. nov. = Chirita spadiciformis W.T. Wang in Bull. Bot. Res., Harbin 5(3): 49. 1985. - Distr.: China (prob1981. - Distr.: China (Guangxi). Primulina pseudoheterotricha (T.J. Zhou, B. Pan & W.B. Xu)ably Guangxi). Primulina speluncae (Hand.-Mazz.) Mich. Moller & A. Weber, Mich. Moller & A. Weber, comb. nov. = Chirita pseudoheterotricha T.J. Zhou, B. Pan & W.B. Xu in J. Trop. comb. nov. = Didymocarpus speluncae Hand.-Mazz., Subtrop. Bot. 17(6): 596. 2009. - Distr.: China (Guangxi). Symb. Sin. 7: 880. 1936 = Chirita speluncae (Hand.-Mazz.) Primulina pteropoda (W.T. Wang) Y.Z. Wang in J. Syst. Evol. D. Wood in Notes Roy. Bot. Gard. Edinburgh 31(3): 370. 49: 62. 201 1 = Chirita pteropoda W.T. Wang in Bull. Bot. 1972. - Distr.: China (Yunnan). Primulina spinulosa (D. Fang & W.T. Wang) Y.Z. Wang in Res., Harbin 5(3): 51. 1985. - Distr.: China (Guangdong). Primulina pungentisepala (W.T. Wang) Mich. Moller & A. We-J. Syst. Evol. 49: 62. 2011 = Chirita spinulosa D. Fang & ber, comb. nov. = Chirita pungentisepala W.T. Wang in W.T. Wang in Bull. Bot. Res., Harbin 1(4): 67. 1981. Novon 7(4): 425. 1998 ('1997'). - Distr.: China (Guangxi). Distr.: China (Guangxi). Primulina subrhomboidea (W.T. Wang) Y.Z. Wang in J. Syst. Primulina renifolia (D. Fang & D.H. Qin) Y.Z. Wang in J. Syst. 784 This content downloaded from 142.51.238.106 on Fri, 09 Jun 2017 20:39:35 UTC All use subject to http://about.jstor.org/terms TAXON 60 (3) • June 201 1 : 767-790 Weber & al. • Remodelling of Chirita Evol. 49: 62. 2011 = Chirita subrhomboidea W.T. Wang Primulina villosissima (W.T. Wang) Mich. Molle in Bull. Bot. Res., Harbin 1(4): 47. 1981. - Distr.: comb. nov.China = Chirita villosissima W.T. Wan (Guangxi). 6(3): 161. 1986. - Distr.: China (Guangdong). Primulina subulata (W.T. Wang) Mich. Primulina Moller & A. Weber, wangiana (Z. Yu Li) Mich. Moller comb. nov. = Chiritopsis subulata W.T. Wangnov. in Guihaia comb. = Chirita wangiana Z. Yu Li in N 6(1-2): 8. 1986. - Distr.: China (Guangdong). 428. 1998 01997'). - Distr.: China (Guangx Primulina weii Mich. Mich. Moller & A. Weber, nom. Primulina subulata var. yangchunensis (W.T. Wang) Moller & A. Weber, comb. nov. = Chiritopsis subulata var. Y.G. Wei & J. Murata in Ac mollifolia D. Fang, yangchunensis W.T. Wang in Guihaia 12(4): 299. 592. 1992. Sin. 37(6): 1999, non Chiritopsis mollifo Distr.: China (Guangdong). W.T. Wang (1986). - Distr.: China (Guangxi) Primulina subulatisepala (W.T. Wang) Mich. Moller & A. WePrimulina wentsaii (D. Fang & L. Zeng) Y.Z. Wa ber, comb. nov. = Chirita subulatisepala W.T. in = Chirita wentsaii D. Fan Evol. 49:Wang 62. 2011 Bull. Bot. Res., Harbin 4(1): 18. 1984. Distr.: China Sin. 31(5): 466. 1993. - Di in-Acta Phytotax. (Hubei, Sichuan). (Guangxi). Primulina swinglei (Merr.) Mich. Moller & A. Weber, comb, (W.T. Wang) Mich. Moll Primulina xinningensis nov. = Didymocarpus swinglei Merr. in comb. Philipp. J. = Sci. 13: nov. Chirita xinningensis W.T. Wang 156. 1918 = Chirita swinglei (Merr.) W.T. Wang in 9(4): Bull. 22. 1989. - Distr.: China ( Res., Harbin Bot. Res., Harbin 1(4): 62. 1981 = Chirita pellegriniana Primulina xiuningensis (X.L. Liu & X.H. Guo) B.L. Burtt in Notes Roy. Bot. Gard. Edinburgh 23: 98. 1960 & A. Weber, comb. nov. = Chiritopsis xiuni = Didymocarpus balansae Pellegr. in Bull. Soc. Bot. Guo France Liu & X.H. in Bull. Bot. Res., Harbin 9( 73: 415. 1926, non Chirita balansae Drake (1890). China - Distr.: - Distr.: (Anhui). China (Guangdong, Guangxi), Northern Vietnam. Primulina yungfuensis (W.T. Wang) Mich. Moll Primulina tabacum Hance, J. Bot. 21: 169. 1883nov. = Primulina comb. = Chirita yungfuensis W.T. Wan sinensis Hook.f., Bot. Mag. 116: t. 7117.7(2): 1890, nom. illeg. 125 (1987). - Distr.: China (Guangxi). - Distr.: China (Guangdong). Primulina tamiana (B.L. Burtt) Mich. Moller & A. Weber, ■ ACKNOWLEDGMENTS comb. nov. = Chirita tamiana B.L. Burtt, in Gloxinian 49(4): 20. 1999. - Distr.: Vietnam (Vinh Phu). Primulina tenuifolia (W.T. Wang) Y.Z. Wang in J. The authors areSyst. gratefulEvol. to the many people who have contributed 49: 62. 2011 = Chirita tenuifolia W.T. Wang Bull.here. Bot. to the work in presented For providing DNA samples we thank, particular, Bhaskar Adhikari and Paul Smith. Thanks also to the Res., Harbin 1(4): 61. 1981. - Distr.: inChina (Guangxi). Primulina tenuituba (W.T. Wang) Y.Z. Wang in J. Evol. horticulturists at Syst. the Royal Botanic Garden Edinburgh (RBGE), Steve 49: 62. 2011 = Deltocheilos tenuitubum W.T. Wang in Scott and Sadie Barber, for expertly maintaining and curating the liv- Bull. Bot. Res., Harbin 1(3): 40. 1981 =Gesneriaceae Chirita collections tenuituba ing at RBGE. The work was, in part, carried (W.T. Wang) W.T. Wang, Fl. Reipubl. 69: out Popularis as a co-operationSin. between the University of Vienna (supported by Austrian Science Fonds, FWF-proj. No. P-13107-Bio), the Forest 388. 1990 = Chirita shennungjiaensisthe W.T. Wang in Bull. Bot. Res., Harbin 1(4): 66. 1981. - Distr.: China (Guizhou, Research Institute Malaysia (FRIM) and RBGE. RBGE is supported Hubei, Hunan, Sichuan). by the Scottish Government Rural and Environment Research and Analysis Directorate (RERAD). The contribution of the FRIM staff Primulina tribracteata (W.T. Wang) Mich. Moller & A. Weber, comb. nov. = Chirita tribracteata W.T. Wang by inthe Bull. is supported Flora Bot. of Peninsular Malaysia Project (01-04-01Res., Harbin 1(4): 58. 1981. - Distr.: 0000 China Khas) (Guangxi). funded by the Ministry of Science, Technology and InPrimulina tribracteata var. zhuana (Z. Yu Li, (MOSTI). Q. Xing Yuan of Michael Moller was supported novation The & fieldwork B. Li) Mich. Moller & A. Weber, comb. nov. = Chirita by the Davis Expedition Fund of the University of Edinburgh and the Oleg Polunin The fieldwork of David Middleton was tribracteata var. zhuana Z. Yu Li, Q. Xing &Memorial Yuan Fund. B. Li in Acta Phytotax. Sin. 44(6): 649. 2006. - Distr.: supported by RBGE, China the Biodiversity Research and Training Program (Guangxi). (BRT) of Thailand and FRIM. Yi-Gang Wei benefitted from support Primulina varicolor (D. Fang & D.H. Qin) Y.Z. &Wang in from the Science Technology Innovation Program of the Guangxi of Sciences Fund. & J. Syst. Evol. 49: 62. 2011 = ChiritaAcademy varicolor D. Fang D.H. Qin in Acta Phytotax. Sin. 37(6): 591. 1999. - Distr.: China (Guangxi). LITERATURE CITED Primulina verecunda (Chun) Mich. Moller■& A. Weber, comb, nov. = Didymocarpus verecundus Chun, Sunyatsenia 6: Akaike, H. 1974. A new look at the statistical model identification. 297. 1946 = Chirita verecunda (Chun) W.T. Wang in Bull. IEEE Trans. Automat. Control 19: 716-723. Bot. Res., Harbin 1(4): 50. 1981. - Distr.: China (Guangxi). Burtt, B.L. 1954. Studies in the Gesneriaceae of the Old World II. Types Primulina vestita (D. Wood) Mich. Moller & A. 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The families and genera of vascular plants, vol. 7, China. Acta Phytotax. Sin. 44: 340-344. Mayer, V., Moller, M., Perret, M. & Weber, A. 2003. Phylogenetic po-Flowering plants: Dicodyledons; Lamiales (except Acanthaceae sition and generic differentiation of Epithemateae (Gesneriaceae)including Avicenniaceae). Berlin & Heidelberg: Springer. Weber, A. & Burtt, B.L. 1998 ("1997"). Remodelling of Didymocarinferred from cpDNA sequence data. Amer. J. Bot. 90: 321-329. McNeill, J., Barrie, F.R., Burdet, H.M., Demoulin, V., Hawksworth, pus and associated genera (Gesneriaceae). Beitr. Biol. Pflanzen 70: 293-363. D.L., Marhold, K., Nicolson, D.H., Prado, J., Silva, P.C., Skog, J.E., Wiersema, J.H. & Turland, N.J. (eds.). 2006. International Weber, A. & Skog, L.E. 2007 onwards. The genera of Gesneriaceae. Basic information with illustration of selected species, ed. 2. http :// code of botanical nomenclature (Vienna Code). Regnum Vegetawww.genera-gesneriaceae.at. ble 146. Ruggell: Gantner. Wei, Y.G., Wen, F., Moller, M., Monro, A., Zhang, Q., Gao, Q., Moller, M., Forrest, A., Wei, Y.G. & Weber, A. 2011. A molecuMou, H.F., Zhong, S.H. & Cui, C. 2010. Gesneriaceae of South lar phylogenetic assessment of the advanced Asian and Malesian China. Yanshan, Guilin, Guangxi: Guangxi Science and Technoldidymocarpoid Gesneriaceae with focus on non-monophyletic and ogy Publishing House. monotypic genera. PL Syst. Evol. 292: 223-248. Moller, M., Perez-Espona, S., Pullan, M., Kiehn, M. & Skog,Wood, L.E. D. 1972. Studies in the Gesneriaceae of the Old World XXXV. New species and combinations in Chirita. Notes Roy. Bot. Gard. 2002 onwards. RBGE WebCyte - Gesneriaceae cytology database. Edinburgh 31: 367-371. http://elmer.rbge.org.uk/webcyte/webcyteintro.php. Wood, Moller, M., Pfosser, M., Jang, C.G., Mayer, V., Clark, A., Hol- D. 1974. A revision of Chirita (Gesneriaceae). Notes Roy. Bot. Gard. Edinburgh 33: 123-205. lingsworth, M.L., Barfuss, M.H.J., Wang, Y.Z., Kiehn, M. & 786 This content downloaded from 142.51.238.106 on Fri, 09 Jun 2017 20:39:35 UTC All use subject to http://about.jstor.org/terms TAXON 60 (3) • June 201 1 : 767-790 Weber & al. • Remodelling of Chirita Appendix 1. Present placement of species hitherto referred to Chirita , Chiritopsis, Hemiboeopsis, Primu epithets, irrespective of generic name). Chirita B.L. calva C.B. Clarke = Henckelia calva (C.B. Clarke) D.J. Middleton Chirita acaulis Merr. = Opithandra acaulis (Merr.) Burtt (see Wood, & Mich. Moller 1974; Wang & al., 1998). Chirita acuminata Wall, ex R. Br., nom. nud. = Henckelia oblongifolia Chirita capitis Craib = Microchirita involucrata (Craib) Y.Z. Wang (Roxb.) D.J. Middleton & Mich. Moller Chirita carnosifolia C.Y. Wu ex H.W. Li = Primulina carnosifolia (C.Y. Wu ex H.W. Li) Y.Z. Chirita acuminata sensu A.DC. = Didymocarpus acuminatus R.Wang Br. (see Clarke, 1874; Burtt, 1958) Chirita ceratoscyphus B.L. Burtt = Henckelia ceratoscyphus (B.L. Burtt) D.J. Middleton & Mich. Moller Chirita adenocalyx Chatterjee = Henckelia adenocalyx (Chatterjee) D.J.(see Wang & al., 1998) Middleton & Mich. Moller Chirita chanetii H. Lev. = Rehmannia glutinosa (Gaertn.) Libosch. ex Fisch. & C.A. Mey. (see Burtt, 1960; Wood, 1974) Chirita adenonema Hilliard = Liebigia adenonema (Hilliard) Mich. Moller & A. Weber Chirita chlamydata W.W. Sm. = Raph wear pus begoniifolius (H. Lev.) Chirita alba Ridl. = Liebigia polyneura (Miq.) Mich. Moller & A. Weber B.L. Burtt (see Weber & Burtt, 1998) Chirita cicatricosa W.T. Wang = Primulina minutihamata (D. Wood) Chirita amplectens Steud., nom. nud. = Henckelia bifolia (D. Don) A. Dietr. Mich. Moller & A. Weber Chirita anachoreta Hance = Henckelia anachoreta (Hance) D.J. Chirita clarkei Hook, f., nom. illeg. = Henckelia lachenensis (C.B. Middleton & Mich. Moller Clarke) D.J. Middleton & Mich. Moller Chirita angusta (C.B. Clarke) Theobald & Grupe = Henckelia angusta Chirita colaniae Pellegr. = Primulina colaniae (Pellegr.) Mich. Moller & A. Weber (C.B. Clarke) D.J. Middleton & Mich. Moller Chirita annamensis Pellegr. = Primulina annamensis (Pellegr.) Mich. Chirita communis Gardner = Henckelia communis (Gardner) D.J. Moller & A. Weber Middleton & Mich. Moller Chirita aratriformis D. Wood = Microchirita aratriformis (D. Wood) Chiritopsis confertiflora W.T. Wang = Primulina confertiflora (W.T. A. Weber & D.J. Middleton Wang) Mich. Moller & A. Weber Chirita asperifolia (Blume) B.L. Burtt = Liebigia speciosa (Blume) Endl.Chiritopsis cordifolia D. Fang & W.T. Wang = Primulina cordifolia Chirita atroglandulosa W.T. Wang = Primulina atroglandulosa (W.T. (D. Fang & W.T. Wang) Y.Z. Wang Wang) Mich. Moller & A. Weber Chirita cordifolia W.T. Wang = Primulina cordata (W.T. Wang) Mich. Moller & A. Weber Chirita atropurpurea W.T. Wang = Primulina atropurpurea (W.T. Wang) Mich. Moller & A. Weber Chirita corniculata Pellegr. = Henckelia ceratoscyphus (B.L. Burtt) D.J. Chirita auriculata J.M. Li & S.X. Zhu = Henckelia auriculata (J.M. Li & S.X. Zhu) D.J. Middleton & Mich. Moller Chirita baishouensis Y.G. Wei, H.Q. Wen & S.H. Zhong = Primulina baishouensis (Y.G. Wei, H.Q. Wen & S.H. Zhong) Y.Z. Wang Chirita balansae Drake = Primulina balansae (Drake) Mich. Moller & A. Weber Chirita barbata Sprague = Microchirita barbata (Sprague) A. Weber & D.J. Middleton Middleton & Mich. Moller Chirita cortusifolia Hance = Didymocarpus cortusifolius (Hance) W.T. Wang. - Wood (1974) retained the species in Chirita , while Wang (1990) transferred it to Didymocarpus , a position retained in Wang & al. (1998). According to Moller & al. (201 1) the species is in a clade with Gyrocheilos mdAllocheilos. Chirita crassituba W.T. Wang = Primulina crassituba (W.T. Wang) Mich. Moller & A. Weber Chirita bicolor W.T. Wang = Primulina bicolor (W.T. Wang) Mich. Moller Chirita cristata (Dalzell) B.L. Burtt = Microchirita hamosa (R. Br.) Y.Z. & A. Weber Wang Chirita bicornuta Hayata = Hemiboea bicornuta (Hayata) Ohwi (see Chirita cruciformis (Chun) W.T. Wang = Primulina cruciformis (Chun) Mich. Moller & A. Weber Wood, 1974; Wang & al., 1998) Chirita bifolia D. Don = Henckelia bifolia (D. Don) A. Dietr. Chirita cyanea (Ridl.) B.L. Burtt = Damrongia cyanea (Ridl.) D.J. Middleton & A. Weber Chirita bilabium C.B. Clarke, nom. illeg. = Liebigia limans (Miq.) Mich. Moller & A. Weber Chirita eyenostyla B.L. Burtt = Primulina eyenostyla (B.L. Burtt) Mich. Moller & A. Weber Chirita bimaculata D. Wood = Microchirita bimaculata (D. Wood) A. Weber & D.J. Middleton Chirita cyrtocarpa D. Fang & L. Zeng = Primulina cyrtocarpa (D. Fang Chiritopsis bipinnatifida W.T. Wang = Primulina bipinnatifida (W.T. & L. Zeng) Mich. Moller & A. Weber Wang) Y.Z. Wang Chirita dalzielii W.W. Sm. = Opithandra dalzielii (W.W. Sm.) B.L. Burtt Chirita blumei C.B. Clarke = Liebigia speciosa (Blume) Endl. (see Wood, 1974; Wang & al., 1998) Chirita blumei C.B. Clarke var. cordifolia S. Moore = Liebigia neoforbesii Chirita demissa (Hance) W.T. Wang = Primulina demissa (Hance) Mich. Moller & A. Weber (Hilliard) Mich. Moller & A. Weber Chirita bogneriana B.L. Burtt = Primulina bogneriana (B.L. Burtt) Mich. Chirita depressa Hook. f. = Primulina depressa (Hook, f.) Mich. Moller Moller & A. Weber & A. Weber Chirita brachystigma W.T. Wang = Primulina brachystigma (W.T. Wang) Chirita diaphana Royle, nom. nud. = Henckelia pumila (D. Don) A. Dietr. Mich. Moller & A. Weber Chirita dibangensis B.L. Burtt, S.K. Srivastava & Mehrotra = Henckelia dibangensis (B.L. Burtt, S.K. Srivastava & Mehrotra) D.J. Middleton Chirita brachytricha W.T. Wang & D. Y. Chen = Primulina brachytricha & Mich. Moller (W.T. Wang & D.Y. Chen) Y.Z. Wang Chirita brachytricha W.T. Wang & D.Y. Chen var. magnibracteata W.T.Chirita dielsii (Borza) B.L. Burtt = Henckelia dielsii (Borza) D.J. Middleton & Mich. Moller Wang & D.Y. Chen = Primulina brachytricha var. magnibracteata (W.T. Wang & D.Y. Chen) Mich. Moller & A. Weber Chirita dimidiata Wall, ex C.B. Clarke = Henckelia dimidiata (Wall, ex Chirita bracteosa Drake, nom. illeg. = Primulina drakei (B.L. Burtt) C.B. Clarke) D.J. Middleton & Mich. Moller Mich. Moller & A. Weber Chirita dissimilis Hilliard = Liebigia dissimilis (Hilliard) Mich. Moller &A. Weber Chirita bracteosa (Zoll. & Moritzi) Miq. = Liebigia horsfieldii (R. Br.) Mich. Moller & A. Weber Chirita drakei B.L. Burtt = Primulina drakei (B.L. Burtt) Mich. Moller & A. Weber Chirita brassicoides W.T. Wang = Primulina brassicoides (W.T. Wang) Mich. Moller & A. Weber Chirita dryas Dunn = Primulina dryas (Dunn) Mich. Moller & A. Weber Chirita brevipes C.B. Clarke = Henckelia speciosa (Kurz) D.J. Middleton Chirita eberhardtii Pellegr. = Primulina eberhardtii (Pellegr.) Mich. & Mich. Moller Moller & A. Weber Chirita briggsioides W.T. Wang = Henckelia briggsioides (W.T. Wang)Chirita eburnea Hance = Primulina eburnea (Hance) Y.Z. Wang D.J. Middleton & Mich. Moller Chirita edgeworthii A. DC. = Henckelia pumila (D. Don) A. Dietr. Chirita brunnea W.T. Wang = Primulina brunnea (W.T. Wang) Mich. Chirita elata Ridl. = Codonoboea elata (Ridl.) Rafidah Moller & A. Weber Chirita elphinstonia Craib = Microchirita elphinstonia (Craib) A. Weber & D.J. Middleton Chirita caerulea R. Br. = Microchirita caerulea (R. Br.) Y.Z. Wang Chirita caliginosa C.B. Clarke = Microchirita caliginosa (C.B. Clarke) Chirita fangii W.T. Wang = Primulina fangii (W.T. Wang) Mich. Moller & A. Weber Y.Z. Wang 787 This content downloaded from 142.51.238.106 on Fri, 09 Jun 2017 20:39:35 UTC All use subject to http://about.jstor.org/terms Weber & al. • Remodelling of Chirita TAXON 60 (3) • June 201 1: 767-790 Appendix 1. Continued. heterotricha Merr. = Primulina heterotricha (Merr.) Y.Z. Wang Chirita fasciculiflora W.T. Wang = Henckelia Chirita fasciculiflora (W.T. Wang) D.J. Middleton & Mich. Moller Chirita heucherifolia (Hand.-Mazz.) D. Wood = Didymocarpus heucherifolius Hand.-Mazz. - Wood (1972, 1974) placed the Chirita fauriei Franch. = Primulina eburnea (Hance) Y.Z. Wang species in Chirita , while Wang & al. (1998) left it in Didymocarpus. Chirita fimbrisepala Hand.-Mazz. = Primulina fimbrisepala (Hand.Chirita hiepii Kiew = Primulina hiepii (Kiew) Mich. Moller & A. Weber Mazz.) Y.Z. Wang Chirita fimbrisepala Hand.-Mazz. var. mollis W.T. Wang = Primulina Chirita hochiensis C.C. Huang & X.X. Chen = Primulina hochiensis (C.C. Huang & X.X. Chen) Mich. Moller & A. Weber fimbrisepala var. mollis (W.T. Wang) Mich. Moller & A. Weber Chirita hookeri C.B. Clarke = Henckelia hookeri (C.B. Clarke) D.J. Chirita flava Wall, ex R. Br., nom. superfl. = Henckelia pumila (D. Don) Middleton & Mich. Moller A. Dietr. Chirita flava Steud., nom. nud. = Henckelia pumila (D. Don) A. Dietr. Chirita flavimaculata W.T. Wang = Primulina fiavimaculata (W.T. Wang) Chirita horsfieldii R. Br. = Liebigia horsfieldii (R. Br.) Mich. Moller & A. Weber Mich. Moller & A. Weber Chirita horsfieldii R. Br. var. scaberrima (R. Br.) C.B. Clarke = Liebigia Chirita floribunda W.T. Wang = Primulina floribunda (W.T. Wang) Mich.horsfieldii (R. Br.) Mich. Moller & A. Weber Moller & A. Weber Chirita humilis Miq., Fl. Ned. Ind. 2: 729. 1858 = Codonoboea humilis (Miq.) A. Weber, comb. nov. Chirita forbesii S. Moore = Liebigia speciosa (Blume) Endl. Chirita infundibuliformis W.T. Wang = Henckelia infundibuliformis (W.T. Chirita forbesii Ridl., nom. illeg. = Liebigia speciosa (Blume) Endl. Wang) D.J. Middleton & Mich. Moller Ch irita fordii (Hemsl.) D. Wood = Primulina fordii (Hemsl.) Y.Z. Wang Chirita insignis C.B. Clarke = Henckelia insignis (C.B. Clarke) D.J. Chirita fordii (Hemsl.) D. Wood var. dolichotricha (W.T. Wang) W.T. Middleton & Mich. Moller Wang = Primulina fordii var. dolichotricha (W.T. Wang) Mich. Moller & A. Weber Chirita Integra Barnett = Damrongia integra (Barnett) D.J. Middleton Chirita forrestii J. Anthony = Henckelia forrestii (J. Anthony) D.J. & A. Weber Middleton & Mich. Moller Chirita involucellata Craib = Microchirita involucrata (Craib) Y.Z. Wang Chirita involucrata Craib = Microchirita involucrata (Craib) Y.Z. Wang Chirita forrestii J. Anthony var. acutidentata W.T. Wang = Henckelia Chirita jiuwanshanica W.T. Wang = Primulina jiuwanshanica (W.T. forrestii (J. Anthony) D.J. Middleton & Mich. Moller Wang) Y.Z. Wang Chirita fruticola H.W. Li = Henckelia fruticola (H.W. Li) D.J. Middleton & Mich. Moller Chirita juliae Hance = Primulina juliae (Hance) Mich. Moller & A. Weber Chirita kerrii Craib = Microchirita rupestris (Ridl.) A. Weber & Rafidah Chirita fulva Barnett = Damrongia fulva (Barnett) D.J. Middleton & A. Weber Chirita kurzii C.B. Clarke = Briggsia kurzii (C.B. Clarke) W.E. Evans (see Wood, 1974; Wang & al., 1998) Chirita fusca C.B. Clarke = Microchirita caliginosa (C.B. Clarke) Y.Z. Chirita lacei (W.W. Sm.) B.L. Burtt = Henckelia lacei (W.W. Sm.) D.J. Wang Middleton & Mich. Moller Chirita gemella D. Wood = Primulina gemell a (D. Wood) Y.Z. Wang Chirita lachenensis C.B. Clarke = Henckelia lachenensis (C.B. Clarke) Chirita geoffrayi Pellegr. = Microchirita involucrata (Craib) Y.Z. Wang D.J. Middleton & Mich. Moller Chirita glabra Miq. = Liebigia glabra (Miq.) Mich. Moller & A. Weber Chirita lacunosa (Hook, f.) B.L. Burtt = Damrongia lacunosa (Hook, f.) IChirita glabra Miq. var. intermedia Ridl. = Liebigia polyneura (Miq.) Mich. Moller & A. Weber D.J. Middleton & A. Weber Chirita glabrescens W.T. Wang & D.Y. Chen = Primulina glabrescensChirita laifengensis W.T. Wang = Primulina obtusidentata (W.T. Wang) Mich. Moller & A. Weber (W.T. Wang & D.Y. Chen) Mich. Moller & A. Weber Chirita langshanica W.T. Wang = Primulina langshanica (W.T. Wang) Chiritopsis glandulosa D. Fang, L. Zeng & D.H. Qin = Primulina Y.Z. Wang glandulosa (D. Fang, L. Zeng & D.H. Qin) Y.Z. Wang Chirita latinervis W.T. Wang = Primulina latinervis (W.T. Wang) Mich. Chiritopsis glandulosa D. Fang, L. Zeng & D.H. Qin var. vangshuoensis Moller & A. Weber F. Wen, Yue Wang & Q.X. Zhang = Primulina glandulosa var. yangshuoensis (F. Wen, Yue Wang & Q.X. Zhang) Mich. Moller Chirita lavandulacea Stapf = Microchirita lavandulacea (Stapf) Y.Z. Wang & A. Weber Chirita laxiflora W.T. Wang = Primulina laxiflora (W.T. Wang) Y.Z. Wang Chirita leeii F. Wen, Yue Wang & Q.X. Zhang = Primulina leeii (F. Wen, Chirita glasgovii Ridl. = Microchirita rupestris (Ridl.) A. Weber & Rafidah Yue Wang & Q.X. Zhang) Mich. Moller & A. Weber Chirita grandibracteata J.M. Li & Mich. Moller = Primulina Chirita leiophvlla W.T. Wang = Primulina leiophylla (W.T. Wang) Y.Z. grandibracteata (J.M. Li & Mich. Moller) Mich. Moller & A. Weber Wang Chirita grandidentata W.T. Wang = Didymocarpus grandidentatus (W.T. Chirita leuserensis Hilliard = Liebigia leuserensis (Hilliard) Mich. Moller Wang) W.T. Wang (see Wang & al., 1998) & A. Weber Chirita grandiflora Wall. = Henckelia urticifolia (D. Don.) A. Dietr. Chirita liboensis W.T. Wang & D.Y. Chen = Primulina liboensis (W.T. Chirita griffithii Hook. f. & Thomson ex C.B. Clarke = Didymocarpus Wang & D.Y. Chen) Mich. Moller & A. Weber griffithii Wight = Loxostigma griffithii (Wight) C.B. Clarke (see Chirita lienxienensis W.T. Wang = Primulina lienxienensis (W.T. Wang) Wood, 1974; Wang & al., 1998) Mich. Moller & A. Weber Chirita guangxiensis S.Z. Huang = Pseudochirita guangxiensis (S.Z. Chirita liguliformis W.T. Wang = Primulina liguliformis (W.T. Wang) Huang) W.T. Wang (see Wang, 1983a; Wang & al., 1998) Mich. Moller & A. Weber Chirita gueilinensis W.T. Wang = Primulina gueilinensis (W.T. Wang) Chirita lilacina Lem. = Monopyle sp. (see Wood, 1 974) Y.Z. Wang Chirita limans (Miq.) B.L. Burtt = Liebigia limans (Miq.) Mich. Moller Chirita gueilinensis W.T. Wang var. brachvcarpa W.T. Wang = Primulina & A. Weber juliae (Hance) Mich. Moller & A. Weber Chirita linearifolia W.T. Wang = Primulina linearifolia (W.T. Wang) Y.Z. Chirita gueilinensis W.T. Wang var. dolichotricha = W.T. Wang Primulina Wang fordii var. dolichotricha (W.T. Wang) Mich. Moller & A. Weber Chiritopsis lingchuanensis Yan Liu & Y.G. Wei = Primulina Chirita guihaiensis Y.G. Wei, B. Pan & W.X. Tang = Primulina guihaiensis lingchuanensis (Yan Liu & Y.G. Wei) Mich. Moller & A. Weber (Y.G. Wei, B. Pan & W.X. Tang) Mich. Moller & A. Weber Chirita linglingensis W.T. Wang = Primulina linglingensis (W.T. Wang) Chirita halongensis Kiew & T.H. Nguyen = Primulina halongensis (Kiew Mich. Moller & A. Weber & T.H. Nguyen) Mich. Moller & A. Weber Chirita liujiangensis = Primulina liujiangensis D. Fang & D.H. Qin Chirita hamosa R. Br. = Microchirita hamosa (R. Br.) Y.Z. Wang Chiritopsis lobulata W.T. Wang = Primulina lobulata (W.T. Wang) Mich. Chirita hamosa var. finlavsonia " C.B. Clarke = Microchirita hamosa Moller & A. Weber (R. Br.) Y.Z. Wang " Chirita longgangensis W.T. Wang = Primulina longgangensis (W.T. Chirita hamosa R. Br. var. unifolia C.B. Clarke = Microchirita hamosa Wang) Y.Z. Wang (R.Br.) Y.Z. Wang Chirita longgangensis W.T. Wang var. hongvao S.Z. Huang = Primulina Chirita hedvotidea (Chun) W.T. Wang = Primulina hedyotidea (Chun) longgangensis var. hongyao (S.Z. Huang) Mich. Moller & A. Weber Y.Z. Wang Chirita longicalyx J.M. Li & Yin Z. Wang = Primulina longicalyx (J.M. Li Chirita heterostigma B.L. Burtt = Henckelia heterostigma (B.L. Burtt) D.J. Middleton & Mich. Moller & Y.Z. Wang) Mich. Moller & A. Weber 788 This content downloaded from 142.51.238.106 on Fri, 09 Jun 2017 20:39:35 UTC All use subject to http://about.jstor.org/terms TAXON 60 (3) • June 201 1 : 767-790 Weber & al. • Remodelling of Chirita Appendix 1. Continued. Chirita Hilliard = Liebigia neoforbesii (Hilliard) Mich. Moller Chirita longii Z. Yu Li = Primulina longii (Z. Yu Li)neoforbesii Y.Z. Wang & A. Weber Chirita longipedicellata B.L. Burtt = Henckelia longipedicellata (B.L. Burtt) D.J. Middleton & Mich. Moller Chirita oblongifolia (Roxb.) J. Sinclair = Henckelia oblongifolia (Roxb.) D.J. Middleton & Mich. Moller Chirita longipedunculata W.T. Wang = Lysionotus longipedunculatus (Roxb.) B.L. Burtt, comb, superfl. = Henckelia (W.T. Wang) W.T. Wang (see Wang, 1983b; Chirita Wangoblongifolia & al., 1998) Hemiboeopsis longisepala (H.W. Li) W.T. Wang = oblongifolia Henckelia (Roxb.) longisepala D.J. Middleton & Mich. Moller (H.W. Li) D.J. Middleton & Mich. Moller Chirita obtusa C.B. Clarke = Didymostigma obtusum (C.B. Clarke) W.T. Chirita longistyla W.T. Wang = Primulina fordii Wang (Hemsl.) Y.Z.1984, Wang (see Wang, Wang & al., 1998) Wentsaiboea luochengensis Yan Liu & W.B. Xu Chirita = Primulina obtusidentata W.T. Wang = Primulina obtusidentata (W.T. Wang) Mich. & Moller A. Weber luochengensis (Yan Liu & W.B. Xu) Mich. Moller A.& Weber Chirita lunglinensis W.T. Wang = Primulina lunglinensis (W.T. Wang) Chirita obtusidentata W.T. Wang var. mollipes W.T. Wang = Primulina Mich. Moller & A. Weber obtusidentata var. mollipes (W.T. Wang) Mich. Moller & A. Weber Chirita oculata Craib = Microchirita oculata (Craib) A. Weber & D.J. Chirita lunglinensis W.T. Wang var. amblyosepala W.T. Wang = Primulina lunglinensis var. amblyosepala (W.T. Wang) Mich. Moller & A. WeberMiddleton Chirita lungzhouensis W.T. Wang = Primulina lungzhouensis (W.T. Chirita ophiopogoides D. Fang & W.T. Wang = Primulina ophiopogoides (D. Fang & W.T. Wang) Y.Z. Wang Wang) Mich. Moller & A. Weber Chirita lutea Yan Liu & Y.G. Wei = Primulina lutea (Yan Liu & Y.G.Chirita Wei) orbicularis W.W. Sm. = Henckelia dielsii (Borza) D.J. Middleton Mich. Moller & A. Weber & Mich. Moller Chirita luzhaiensis Yan Liu, Y.S. Huang & W.B. Xu = Primulina luzhai-Chirita orthandra W.T. Wang = Primulina orthandra (W.T. Wang) Mich. Moller & A. Weber ensis (Yan Liu, Y.S. Huang & W.B. Xu) Mich. Moller & A. Weber Chirita macrodonta D. Fang & D.H. Qin = Primulina macrodonta Chirita parvifolia W.T. Wang = Primulina parvifolia (W.T. Wang) Y.Z. Wang (D. Fang & D.H. Qin) Mich. Moller & A. Weber Chirita macrophylla Wall. = Henckelia grandifolia A. Dietr. Chirita parvula Ridl. = Microchirita caliginosa (C.B. Clarke) Y.Z. Wang Chirita macrophylla Wall. var. tirapensis Panigrahi = Henckelia Chirita peduncularis B.L. Burtt = Henckelia peduncularis (B.L. Burtt) D.J. Middleton & Mich. Moller grandifolia A. Dietr. Chirita macrorhiza D. Fang & D.H. Qin = Primulina macrorhiza Chirita pellegriniana B.L. Burtt = Primulina swinglei (Merr.) Mich. Moller & A. Weber (D. Fang & D.H. Qin) Mich. Moller & A. Weber Chirita macrosiphon Hance = Raphiocarpus macrosiphon (Hance) B.L. Chirita pinnata W.T. Wang = Primulina pinnata (W.T. Wang) Y.Z. Wang Burtt (see Weber & Burtt, 1998) Chirita pinnatifida (Hand.-Mazz.) B.L. Burtt = Primulina pinnatifida Chirita maguanensis Z. Yu Li, H. Jiang & H. Xu = Primulina (Hand.-Mazz.) Y.Z. Wang maguanensis (Z. Yu Li, H. Jiang & H. Xu) Mich. Moller & A. Weber Chirita poilanei Pellegr. = Primulina poilanei (Pellegr.) Mich. Moller & A. Weber Chirita mangshanensis W.T. Wang = Primulina juliae (Hance) Mich. Moller & A. Weber Chirita polycarpa Steud. = Lysionotus serratus D. Don (see Wood, 1 974) Chirita mangshanensis W.T. Wang var. lasiandra W.T. Wang = Primulina Chirita polycephala (Chun) W.T. Wang = Primulina polycephala (Chun) Mich. Moller & A. Weber juliae (Hance) Mich. Moller & A. Weber Chirita marcanii Craib = Microchirita marcanii (Craib) A. Weber & D.J. Chirita polyneura Miq. = Liebigia polyneura (Miq.) Mich. Moller Middleton & A. Weber Chirita medica D. Fang = Primulina medica (D. Fang) Y.Z. Wang Chirita micromusa B.L. Burtt = Microchirita micromusa (B.L. Burtt) A. Weber & D.J. Middleton Chirita minutihamata D. Wood Mich. Moller & A. Weber Chirita minutimaculata D. Fang minutimaculata (D. Fang & Chirita minutiserrulata Hayata Middleton & Mich. Moller Chirita polyneura Miq. var. albiflora Miq. = Liebigia polyneura (Miq.) Mich. Moller & A. Weber Chirita polyneura Miq. var. amabilis C.B. Clarke = Henckelia dimidiata = Primulina minutihamata (D. Wood) (Wall, ex C.B. Clarke) D.J. Middleton & Mich. Moller Chirita polyneura Miq. var. thomsonii C.B. Clarke = Henckelia pumila (D. Don) A. Dietr. & W.T. Wang = Primulina W.T. Wang) Y.Z. Wang Chirita praeterita Hilliard = Liebigia praeterita (Hilliard) Mich. Moller & A. Weber = Henckelia anachoreta (Hance) D.J. Chirita primulacea C.B. Clarke = Henckelia primulacea (C.B. Clarke) Chirita mirabilis (Pamp.) Jafri, error for Christolea mirabilis (Pamp.) Jafri D.J. Middleton & Mich. Moller (Brassicaceae) (see Skog & Boggan, 2007) Chirita primuloides (Miq.) Ohwi = Opithandra primuloides (Miq.) B.L. Burtt (see Wood, 1974) (Debbarm. ex Biswas) D.J. Middleton & Mich. Moller Chirita pseudoeburnea D. Fang & W.T. Wang = Primulina Chirita modesta Kiew & T.H. Nguyen = Primulina modesta (Kiew & T.H. pseudoeburnea (D. Fang & W.T. Wang) Mich. Moller & A. Weber Nguyen) Mich. Moller & A. Weber Chirita pseudoheterotricha T.J. Zhou, B. Pan & W.B. Xu = Primulina Chirita mollifolia D. Fang, Y.G. Wei & J. Murata = Primulina weii Mich. pseudoheterotricha (T.J. Zhou, B. Pan & W.B. Xu) Mich. Moller Chirita mishmiensis Debbarm. ex Biswas = Henckelia mishmiensis Moller & A. Weber & A. Weber Chiritopsis mollifolia D. Fang & W.T. Wang = Primulina mollifolia (D. Chirita pteropoda W.T. Wang = Primulina pteropoda (W.T. Wang) Y.Z. Fang & W.T. Wang) Y.Z. Wang Wang Chirita mollis Miq. = Codonoboea sp. - There is a note by Olive Hilliard Chirita puerensis Y.Y. Qian = Henckelia puerensis (Y.Y. Qian) D.J. Middleton & Mich. Moller on the isotype in Leiden saying this is a Henckelia (sensu Weber & Burtt, 1 998). This must now be treated as a species of Codonoboea Chirita pumila D. Don = Henckelia pumila (D. Don) A. Dietr. as will be explained in a forthcoming paper. Chirita pungentisepala W.T. Wang = Primulina pungentisepala (W.T. Chirita mollissima Ridl. = Microchirita mollissima (Ridl.) A. Weber Wang) Mich. Moller & A. Weber & D.J. Middleton Chirita purpureolineata (Kerr ex Craib) D. Wood = Damrongia Chirita monantha W.T. Wang = Henckelia monantha (W.T. Wang) D.J. purpureolineata Kerr ex Craib Middleton & Mich. Moller Chirita pycnantha W.T. Wang = Henckelia pycnantha (W.T. Wang) D.J. Middleton & Mich. Moller Chirita monophylla C.B. Clarke = Henckelia monophylla (C.B. Clarke) D.J. Middleton & Mich. Moller Chirita quercifolia D. Wood = Primulina pinnatifida (Hand.-Mazz.) Y.Z. Chirita moonii Gardner = Henckelia moonii (Gardner) D.J. Middleton Wang & Mich. Moller Wentsaiboea renifolia D. Fang & D.H. Qin = Primulina renifolia (D. Fang Chirita nandanensis S.X. Huang, Y.G. Wei & W.H. Luo = Primulina & D.H. Qin) Y.Z. Wang nandanensis (S.X. Huang, Y.G. Wei & W.H. Luo) Mich. Moller Chiritopsis repanda W.T. Wang = Primulina repanda (W.T. Wang) Y.Z. & A. Weber Wang Chirita napoensis Z. Yu Li = Primulina napoensis (Z. Yu Li) Mich. Chiritopsis repanda var. guilinensis W.T. Wang = Primulina repanda var. Moller & A. Weber guilinensis (W.T. Wang) Mich. Moller & A. Weber 789 This content downloaded from 142.51.238.106 on Fri, 09 Jun 2017 20:39:35 UTC All use subject to http://about.jstor.org/terms Weber & al. • Remodelling of Chirita TAXON 60 (3) • June 201 1 : 767-790 Appendix 1. Continued. Chirita swinglei (Merr.) W.T. Wang = Primulina swinglei (Merr.) Mich. Chirita reptans B.L. Burtt & Panigrahi = Henckelia burttii D.J. Middleton Moller & A. Weber & Mich. Moller Primulina tabacum Hance Chirita ridleyana S. Moore = Liebigia speciosa (Blume) Endl. Chirita tamiana B.L. Burtt = Primulina tamiana (B.L. Burtt) Mich. Chirita ronganensis D. Fang & Y.G. Wei = Primulina ronganensis (D. Moller & A. Weber Fang & Y.G. Wei) Mich. Moller & A. Weber Chirita tenuifolia W.T. Wang = Primulina tenuifolia (W.T. Wang) Y.Z. Chirita roseoalba W.T. Wang = Primulina roseoalba (W.T. Wang) Mich. Moller & A. Weber Wang Chirita rotundata Barnett = Henckelia rotundata (Barnett) D.J. Middleton Chirita tenuipes Hilliard = Liebigia tenuipes (Hilliard) Mich. Moller & A. Weber & Mich. Moller Chirita tenuituba (W.T. Wang) W.T. Wang = Primulina tenuituba (W.T. Chirita rotundifolia (Hemsl.) D. Wood = Primulina rotundifolia (Hemsl.) Mich. Moller & A. Weber Wang) Y.Z. Wang Chirita ternifolia Buch.-Ham. ex D. Don = Lysionotus serratus D. Don Chirita rupestris Ridl. = Microchirita rupestris (Ridl.) A. Weber & Rafidah Chirita sahyadriensis Punekar & Lakshmin. = Microchirita sahyadriensis (see Wood, 1974) Chirita tibetica (Franch.) B.L. Burtt = Henckelia tibetica (Franch.) D.J. (Punekar & Lakshmin.) A. Weber & D.J. Middleton Middleton & Mich. Moller Chirita scaberrima R. Br. = Liebigia horsfieldii (R. Br.) Mich. Moller & A. Weber Chirita tobaensis Hilliard = Liebigia tobaensis (Hilliard) Mich. Moller & A. Weber Chirita scabra Royle ex C.B. Clarke, nom. illeg. pro syn. = Henckelia Chirita trailliana Forrest & W.W. Sm. = Henckelia speciosa (Kurz) D.J. bifolia (D. Don) A. Dietr. Middleton & Mich. Moller Chirita sclerophylla W.T. Wang = Primulina sclerophylla (W.T. Wang) Chirita tribracteata W.T. Wang = Primulina tribracteata (W.T. Wang) Y.Z. Wang Mich. Moller & A. Weber Chirita secundiflora (Chun) W.T. Wang = Primulina secundiflora (Chun) Mich. Moller & A. Weber, Chirita tribracteata W.T. Wang var. zhuana Z. Yu Li, Q. Xing & Yuan B. Li = Primulina tribracteata var. zhuana (Z. Yu Li, Q. Xing Chirita semicontorta Pellegr. = Primulina semicontorta (Pellegr.) Mich. Moller & A. Weber & Yuan B. Li) Mich. Moller & A. Weber Chirita sericea Ridl. = Microchirita sericea (Ridl.) A. Weber & Rafidah Chirita trisepala Barnett = Damrongia trisepala (Barnett) D.J. Middleton & A. Weber Chirita sericea H. Lev. & Vaniot, nom. illeg. = Oreocharis auricula Chirita tubulosa Craib = Microchirita tubulosa (Craib) A. Weber & D.J. (S. Moore) C.B. Clarke var. auricula (see Wang & al., 1998) Middleton Chirita sericea var. scortechinii Ridl. = Microchirita sericea (Ridl.) A. Weber & Rafidah Chirita umbricola W.W. Sm. = Henckelia lachenensis (C.B. Clarke) D.J. Middleton & Mich. Moller Chirita shennungjiaensis W.T. Wang = Primulina tenuituba (W.T. Wang) Chirita umbrophila C.Y. Wu ex H.W. Li: generic placement uncertain Y.Z. Wang (see Wang & al., 1 998: 311) Chirita shouchengensis Z. Yu Li = Primulina shouchengensis (Z. Yu Li) Chirita uniflora Ridl.: excluded from Chirita in the revision of Wood Y.Z. Wang (1974) and referred to "Didymocarpus" . From its locality and flower Chirita shuii Z. Yu Li = Henckelia shuii (Z. Yu Li) D.J. Middleton & Mich. Moller description it is most likely a Codonoboea, probably the species currently called Henckelia hirta (Ridl.) A. Weber. Chirita sichuanensis W.T. Wang = Primulina sichuanensis (W.T. Wang) Mich. Moller & A. Weber Chirita urticifolia Buch.-Ham. ex D. Don = Henckelia urticifolia (D. Chirita sinensis Lindl. = Primulina sinensis (Lindl.) Y.Z. Wang, nom. illeg. Don.) A. Dietr. Chirita varicolor D. Fang & D.H. Qin = Primulina varicolor (D. Fang = Primulina dryas (Dunn) Mich. Moller & A. Weber &D.H.Qin) Y.Z. Wang Chirita sinensis var. angustifolia Dunn = Primulina dryas (Dunn) Mich. Moller & A. Weber Chirita verecunda (Chun) W.T. Wang = Primulina verecunda (Chun) Mich. Moller & A. Weber Chirita sinensis var. bodinieri H. Lev. = Primulina dryas (Dunn) Mich. Moller & A. Weber Chirita vestita D. Wood = Primulina vestita (D. Wood) Mich. Moller & A. Weber Primulina sinensis Hook, f., nom. illeg. = Primulina tabacum Hance Chirita skogiana Z. Yu Li = Primulina skogiana (Z.Y. Li) Mich. Moller Chirita villosissima W.T. Wang = Primulina villosissima (W.T. Wang) Mich. Moller & A. Weber & A. Weber Chirita smitinandii B.L. Burtt = Henckelia smitinandii (B.L. Burtt) D.J.Chirita viola Ridl. = Microchirita viola (Ridl.) A. Weber & Rafidah Middleton & Mich. Moller Chirita violoides (Wall.) M.R. Almeida = Platystemma violoides Wall. Chirita vulgaris Morren, nom. nud. = Henckelia communis (Gardner) D.J. Chirita spadiciformis W.T. Wang = Primulina spadiciformis (W.T. Wang) Middleton & Mich. Moller Mich. Moller & A. Weber Chirita speciosa Kurz = Henckelia speciosa (Kurz) D.J. Middleton & Mich. Moller Chirita walkerae Gardner = Henckelia walkerae (Gardner) D.J. Middleton & Mich. Moller Chirita walkerae Gardner var. parviflora C.B. Clarke = Henckelia Chirita spectabilis Miq. = Liebigia speciosa (Blume) Endl. Chirita spectabilis Var. 6? pedicellata C.B. Clarke = Liebigia polyneura walkerae (Gardner) D.J. Middleton & Mich. Moller Chirita walkerae Gardner subsp. parviflora (C.B. Clarke) W.L. Theob. (Miq.) Mich. Moller & A. Weber Chirita speluncae (Hand.-Mazz.) D. Wood = Primulina speluncae (Hand.-& Grupe = Henckelia walkerae (Gardner) D.J. Middleton & Mich. Moller Mazz.) Mich. Moller & A. Weber Chirita sphagnicola H. Lev. & Vaniot = Henckelia pumila (D. Don) Chirita wangiana = Primulina wangiana (Z.Y. Li) Mich. Moller A. Dietr. & A. Weber Chirita wentsaii D. Fang & L. Zeng = Primulina wentsaii (D. Fang Chirita spinulosa D. Fang & W.T. Wang = Primulina spinulosa (D. Fang & L. Zeng) Y.Z. Wang & W.T. Wang) Y.Z. Wang Chirita xinningensis W.T. Wang = Primulina xinningensis (W.T. Wang) Chirita stolonifera C.E.C. Fisch. & Kaul = Henckelia lachenensis (C.B. Mich. Moller & A. Weber Clarke) D.J. Middleton & Mich. Moller Chirita subacaulis (Hand.-Mazz.) B.L. Burtt = Hemiboea subacaulisChiritopsis xiuningensis X.L. Liu & X.H. Guo = Primulina xiuningensis (X.L. Liu & X.H. Guo) Mich. Moller & A. Weber Hand.-Mazz. (see Wood, 1974, Wang & al., 1998) Chirita yungfuensis W.T. Wang = Primulina yungfuensis (W.T. Wang) Chirita subrhomboidea W.T. Wang = Primulina subrhomboidea (W.T. Mich. Moller & A. Weber Wang) Y.Z. Wang Chirita zeylanica Hook. = Henckelia communis (Gardner) D.J. Middleton Chiritopsis subulata W.T. Wang = Primulina subulata (W.T. Wang) Mich. Moller & A. Weber & Mich. Moller Chiritopsis subulata var. yangchunensis W.T. Wang = Primulina subulataChirita zeylanica Hook. var. angusta C.B. Clarke = Henckelia angusta (C.B. Clarke) D.J. Middleton & Mich. Moller var. yangchunensis (W.T. Wang) Mich. Moller & A. Weber Chirita zollingeri C.B. Clarke = Microchirita caerulea (R. Br.) Y.Z. Wang Chirita subulatisepala W.T. Wang = Primulina subulatisepala (W.T. Wang) Mich. Moller & A. Weber 790 This content downloaded from 142.51.238.106 on Fri, 09 Jun 2017 20:39:35 UTC All use subject to http://about.jstor.org/terms