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The polyphyletic genus Chirita is remodelled after an extensive molecular phylogenetic study of species assigned to it and to other associated genera. Most of Chirita sect. Chirita and the monotypic Hemiboeopsis are amalgamated with Henckelia sect. Henckelia, resulting in a very differently circumscribed genus Henckelia and the synonymisation of Chirita. The remain- ing species of Chirita sect. Chirita are accommodated in the revived genus Damrongia. Chirita sect. Liebigia is recognised as the genus Liebigia. Chirita sect. Microchirita is recognised as the genus Microchirita. Chirita sect. Gibbosaccus is, together with Chiritopsis and Wentsaiboea, included in the originally monotypic and now enormously expanded genus Primulina. The necessary combinations are made and a general list showing the present accommodation of the species previously described under Chirita, Chiritopsis, Hemiboeopsis, Primulina and Wentsaiboea is provided.
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767
Weber & al. •
Remodelling of Chirita
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60 (3) • June 2011: 767–790
INTRODUCTION
In its traditional sense, the genus Chirita Buch.-Ham. ex
D. Don, established formally by Don (1822) and revised by Can-
dolle (1845), Clarke (1883) and Wood (1974), is a large genus of
perennial or annual herbs or subshrubs ranging from Sri Lanka,
the Himalayas and southern China through continental South-
east Asia and into Malesia as far as Sumatra and Java. Wood
(1974) recognised 77 species in three sections but the genus has
grown rapidly in recent years, particularly through the addition
of many new species from China (see Wang & al., 1998; Li &
Möller, 2009). Wang (1985a,b), Wang & al. (1998) and Wang &
al. (2011) estimated that there were about 140 species and Li &
Möller (2009) 150 species, two-thirds of which belong to Chirita
sect. Gibbosaccus C.B. Clarke and most of which are endemic to
China. Weber (2004) gave a very large range of 80–140 species,
marked the genus as highly heterogeneous and noted there were
problems in the delimitation of the genus and the subdivisions
within it. The great variance in the estimate of the number of
species is also testament to the considerable problems of specific
delimitation in Chirita but this problem will need to be addressed
in subsequent revisions of the species rather than in this work.
The genus has mostly (Wood, 1974; Wang, 1985a,b; Wang & al.,
1998; Li & Wang, 2004) been divided into three sections: Chirita
sect. Chirita, Chirita sect. Gibbosaccus and Chirita sect. Micro-
chirita C.B . Clarke. Wang (1985a ,b) also recog nis ed subsec tions
and series in the genus and Hilliard (2004) resurrected Chirita
sect. Liebigia (Endl.) C.B. Clarke from synonymy of Chirita sect.
Chirita to create a fourth section.
In this paper we set out to investigate the relationships of
Chirita to the other didymocarpoid genera (see Weber, 2004;
Möller & al., 2009, 2011) and to establish a stable framework that
can be used as a str uct u re for invest igat ing specie s deli mitation.
There have been several molecular phylogenetic studies that have
included species of Chirita, beginning with Mayer & al. (2003),
and continuing with Li & Wang (2007), Möller & al. (2009),
Wang & al. (2011) and Möller & al. (2011). The focus of the pa-
per by Li & Wang (2007) was to study the relationship bet ween
Chirita sect. Gibbosaccus and Chiritopsis W.T. Wang. They did
not include representatives of other genera except as outgroups.
Of the three sections recognised by Wood (1974) and Wang &
al. (1998), Chirita sect. Chirita, Chirita sect. Microchirita and
Chirita sect. Gibbosaccus, Li & Wang (2007) found the first
two to be monophyletic and the last to be par ap hyletic due to the
inclusion of Chiritopsis. However, this result was more or less
inevitable given the lack of sampling of other didymocarpoid
Molecular systematics and remodelling of Chirita and associated
genera (Gesneriaceae)
Anton Weber,1 David J. Middleton,2 Alan Forrest, 2 Ruth Kiew,3 Chung Lu Lim, 3 A.R. Rafidah,3 Susanne
Sontag,1 Pramote Triboun,4 Yi-Gang Wei,5 Tze Leong Yao3 & Michael Möller2
1 University of Vienna, Faculty Centre of Biodiversity, Depart ment of Structural and Functional Botany, Rennweg 14,
1030 Vienna, Austria
2 Royal Botanic Garden Edinburgh, 20A 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 Variet y Protection Division, Department of Agriculture,
Chatuchak, Bangkok 10900, Thailand
5 Guangxi Institute of Botany, Guangxi Zhuang Autonomous Region and the Chinese Academy of Sciences, Guilin 541006, China
Author for correspondence: David J. Middleton, d.middleton@rbge.ac.uk
Abstract
The polyphyletic genus Chirita is re mo de ll ed af te r an exte nsive molecu la r ph yloge net ic study of sp ecies assig ne d to it
and to other associated genera. Most of Chirita se ct . Chirita and the monotypic Hemiboeopsis are am al ga mat ed wit h Henckelia
sect. Henckelia, resulting in a very differently circumscribed genus Henckelia an d the syn on ymisa ti on of Chirita. The re ma in-
ing species of Chirita sect . Chirita are accommodated in the revived genus Damrongia. Chirita sect. Liebigia is recogn ised as
the genus Liebigia. Chirita sect. Microchirita is recognised as the genus Microchirita. Chirita sect. Gibbosaccus is, together
with Chiritopsis and Wentsaiboea, included in the originally monotypic and now enormously expanded genus Primulina. The
necessary combinations are made and a general list showing the present accommodation of the species previously described
under Chirita, Chiritopsis, Hemiboeopsis, Primulina and Wentsaiboea is provided.
Keywords
Bayesian inference analysis; Chirita; Chiritopsis; Damrongia; Hemiboeopsis; Henckelia; ITS; Liebigia;
maximu m parsimony; Microchirita; molecular phylogeny; Primulina; taxonomy; trnL-F intron-spacer; Wentsaiboea
Supplementary Material
Diagnostics of the Bayesian inference analysis are available in Electronic Supplement 1
and Figures S1 and S2 and Table S1 are available in Electronic Supplement 2 to the online version of this article
(http://www.ingentaconnect.com/content/iapt/tax).
768
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Remodelling of Chirita
genera. In Möller & al. (2009, 2011), with more limited sam-
pling in Chirita but considerably more in other genera, only
Chirita sect. Microchirita remained monophyletic, albeit with
low sampling in all sections. Chirita sect. Chirita (in the concept
of Wood, 1974) was represented only by five species in Möller
& al. (2009) and these did not group together. One of these was
Chirita asperifolia (Blume) B.L. Burtt, the type of Chirita sect.
Liebigia (Endl.) C.B. Clarke which Hilliard (2004) resurrected.
The other species were Chirita lacunosa (Hook. f.) B.L. Burtt,
C. walkerae Gardner (as C. walkeri), C. pumila D. Don, and
C. urticifolia Buch.-Ham. ex D. Don, the type of Chirita. In
Möller & al. (2009) four species of Chirita sect. Gibbosaccus
were included but these were found to be par aphyletic due to the
inclusion of the monotypic Primulina Hanc e and a single sp ecies
of Chiritopsis. These results were confirmed in a larger analysis
of didymocarpoid Gesneriaceae (Möller & al., 2011).
Li & Wang (2007) did not include Primulina in their study.
The largest published study to date is by Wang & al. (2011), a
paper submitted for publication after we submitted ours to Taxo n
but which was reviewed and published in just a few weeks.
1
In their paper they included a larger sample of Chiritopsis and
Chirita sect. Gibbosaccus species, combined with sequences
from Möller & al. (2009) for the crucial Chinese species Prim-
ulina tabacum Hance and Petrocodon dealbatus Hance and for
some non-Chinese taxa. They confirmed the basic topology of
the previous findings by Möller & al. (2009) and proceeded to
raise Chirita sect. Microchirita and Chirita sect. Liebigia to ge-
nus rank and to reduce Chirita sect. Gibbosaccus, Chiritopsis
and Wentsaiboea D. Fang & D.H. Qi n to sy nonymy of Primulina.
They did not address the problem of Chirita lacunosa nor the re-
lationship between species of Chirita sec t. Chirita and Henckelia
from Sri Lanka which were highlighted by Möller & al. (2009).
Chirita is morphologically quite variable (Wood, 1974;
Weber, 2004). Clarke (1883, 1884) commented on how Chirita
“is with difficulty distinguished from Didymocarpus”. Wood
(1974: 126) remarked that the difference between the apically
fused anthers in Chirita sect. Microchirita, in contra st to face-
to-face fusion in the other sections, “together with the possi-
bility that the syndrome of bilobed stigma and geniculate fila-
ments has originated several times independently, suggests that
Chirita may be polyphyletic.” Clarke’s and Wood’s comments
may have been pertinent indeed and an investigation of Chirita
clearly needs to take into account any possible relationships the
species may have with other didymocarpoid genera. Clarke
(1883, 1884) wa s lar gely unawar e of the large nu mb er of spe cies
of Chirita sect. Gibbosaccus that were yet to be collected in
China and his comments mainly reflected his understanding of
the species in Chirita sect. Chirita. Chirita sect. Gibbosaccus
has since become by far the largest of the sections in Chirita
(Wang & al., 1998; Li & Wang, 2004).
From the comments in the older literature and from the
preliminary findings of Möller & al. (2009, 2011), the ques-
tions that need to be addressed for a better understanding of
Chirita are: (1) Will a better sampling of Chirita confirm that
1 An invita tion fr om us to combi ne th e two pa per s be fore publication
remained unanswered.
the genus is polyphyletic? (2) Are the three (Wood, 1974) or
four (Hilliard, 2004) sections of Chirita each monophyletic?
(3) What are the relationships of any possible new groupings
of Chirita species to other didymocarpoid genera, particularly
to Henckelia, Primulina and Chiritopsis ?
In this paper the responsibility for the various aspects of the
paper are as follows: D.J. Middleton and A. Weber initiated the
need to further investigate generic delimitation in Chirita and
have compiled the taxonomic sections (with contributions by
A.R. Rafidah in Microchirita); R. Kiew, C.L . Lim, D.J. Middle-
ton, M. Möller, A.R. Rafidah, P. Triboun, Y.-G. Wei and T.L. Yao
collected and identified most of the new material sequenced;
A. Forrest, C.L. Lim, M. Möller and T.L. Yao generated the se-
quence data; S. Sontag helped to compile the formal taxonomy
and procured the literature required; and M. Möller oversaw all
aspects of the molecular work and analysed the molecular data.
MATERIAL AND METHODS
Plant material. —
The samples included in the present
study are those used in a recent study (Möller & al., 2011), plus
41 samples from additional species of Chirita, Chiritopsis and
Henckelia. Some sequences came from GenBank, others from
re cent field collections (Table S1 in Elec t ro nic Suppl. 2). In total
232 samples have been inclu de d covering 66 out of the 80 gen-
era classified in the Old World didymocarpoid Gesneriaceae
(We ber & Skog , 20 07 onward s; Wei & al ., 2010), or 58 out of 61
of the advanced Asiatic and Malesian genera to which Chirita
belongs (Möller & al., 2011). Details of the sample coverage,
in group and outg roup select io n and rooti ng ar e give n in Möller
& al. (2011). Here we focus on the genus Chirita.
Sampling within the highly polyphyletic genus Chirita has
been expanded from 15 to 42 samples (representing 33 species
and three undescribed species), with nine species (plus one un-
described species) from section Chirita (ou t of 40 de scr ibed), 14
species from section Gibbosaccus (out of 117 described), nine
species (plus two undescribed species) from section Micro-
chirita (out of 19 described) and one species from section Liebi-
gia (out of 12 species described, all of which were previously
included in synonymy of a variable Chirita asperifolia).
The genus Chiritopsis was earlier shown to be nested in
Chirita sect. Gibbosaccus (Li & Wang, 2007; Möller & al.,
2009, 2011). To investigate closer the relationships of Chiri-
topsis with Chirita, five out of the nine described species have
also been included.
The polyphyletic Henckelia is represented by 15 species
(out of ca. 180). We have increased the sampling in Henckelia
because in Möller & al. (2009, 2011), with very low sampling,
some species of Henckelia were nested in Chirita sect. Chirita.
These 15 samples have been selected from a very much larger
sample of Henckelia species, the results of which will be pre-
sented in follow-up papers concentrating on that genus (results
of the larger Henckelia stud y confirm the re sults pre sente d with
the smaller 15 species sample included in this paper).
DNA extraction, PCR, and direct sequencing. —
Mo-
lecular methods and protocols basically followed Möller & al.
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(2009, 2011), acquiring sequences of the plastid trnL-F intron-
spacer (trnL-F) and the nuclear ribosomal internal transribed
spacer (ITS) region.
Phylogenetic analysis. —
For this study 27 trnL-F and ITS
sequences were newly generated and an additional 14 trnL-F
and ITS sequences were obtained from GenBank (both sources
given in bold in Appendix 2 and distinguished in the last two
columns). Both sequence regions were available for all samples
included (Appendix 2). The newly acquired and GenBank se-
quences were added to existing matrices (Möller & al., 2011).
The trnL-F alignment was unproblematic, only 66 characters
had to be excluded at the 5′ end due to the poor quality of se-
quences, particularly those retrieved from GenBank. For the
ITS matrix, initially aligned automatically and then adjusted by
eye (see Möller & al., 2009), 155 cha racters had to be excluded
due to alignment ambiguities.
The phylogenetic analysis basically followed Möller &
al. (2009, 2011). In brief, two analyses, maximum parsimony
(MP) implemented in PAUP* v.4.0b10 (Swofford, 2002), and
Bayesian inference (BI) in MrBayes v.3.1.2 (Huelsenbeck &
Ronquist, 2001, 2007) were carried out. The trnL-F and ITS
sequences were analysed together after their combinability was
checked by the incong r uence length differenc e (IL D; P = 0.07)
test implemented in PAUP*.
For the MP analysis characters were unordered and un-
weighted with alignment gaps treated as missing data. Start-
ing trees were found by parsimony ratchet (Nixon, 1999), in
PAUPRat (Sikes & Lewis, 2001) and PAUP*, and the saved
trees fu r the r op tim ise d in PAUP*, with bot h TBR and Multre es
on. Statistical branch support was obtained from 10,000 heu-
ristic bo ot strap re pl icates (Möl le r & al., 2009, 2011) in PAUP*.
For the BI analysis models and parameter prior settings
were obtained independently for the trnL-F, the ITS spacers
an d 5.8S seque nces using MrMod elte st v.2.3 (Nyla nd er, 20 04),
and were GTR + G, GTR + I + G and SYM + I + G, respectively, as
suggested by the Akaike Information Criterion (AIC; Akaike,
1974). Five million generations were run in two independent
analyses each with four Markov chain Monte Carlo (MCMC)
chains (Electronic Supplement 1A–D). One tree was sampled
every 250 generations (= 20,000 trees), and the first 100,000
generations (the first 400 trees) discarded as burn-in, deter-
mined after plotting the generations against the Log(n) likeli-
hoods (Electronic Supplement 1E–F). The remaining trees were
used to reconstruct a majority rule consensus tree in PAUP*,
and posterior probabilities (PP) obtained. The PP branch sup-
port values showed a high correlation between the two parallel
Bayesian runs (Electronic Supplement 1G).
RES U LTS
Matrix characteristics. —
After the exclusion of ambig-
uous characters, the combined matrix had a length of 1966
characters, 1132 for trnL-F and 834 for ITS. In the combined
analysis 906 of the remaining 1966 characters were constant,
309 variable but parsimony-uninformative, and 751 (38.2%)
parsimony-informative.
Phylogenetic analyses. —
The MP analysis on the com-
bined data resulted in 117,344 most parsimonious trees of 6711
steps length, a consistency index of 0.2845 and retention index
of 0.66 48. The majority-rule co nsensu s tre e was hig hly resolve d
and was similar in topology to the previous phylogeny of Möller
& al. (2011) where the cla des received hig h bra nch suppo rt (Fig.
S1 in Electronic Suppl. 2). The BI tree recovered was basically
congruent to the MP tree, in areas where the branches were well
supported (Fig. S2 in Electronic Suppl. 2). A detailed report
and discussion of the inter- and intra-generic relationships of
the advanced didymocarpoid Gesneriaceae is given in Möller
& al. (2011) and we here refrain from repeating these. Instead
we focus on the positions of the Chirita sa mple s and ass ociated
genera. The results of the MP and BI analyses are summarised
in a simplified tree (Fig. 1) and subtrees (Figs. 2–6), but given
in full as Figs. S1 and S2 (in Electronic Suppl. 2).
Chirita samples fell in six different clades. Several nodes
separating these clades received medium to high support values
(Fig. 1, grey squares), supporting the separate phylogenetic
positions of the six respective Chirita positions; i.e., they rep-
resent separate evolutionary lineages. The branches leading
to the six clades with Chirita samples (Fig. 1, grey circles) all
received high support values; these clades represent well de-
fined separate genetic entities. This can also be inferred from
the long branches leading to these clades in the BI tree (Fig.
S2 in Electronic Suppl. 2).
Chirita elata Ridl. was nested inside the Malesian Henck-
elia clade, as sister to Henckelia nana A. Weber (BS = 100%,
PP = 1.0) (Fig. 1; Figs. S1 and S2 in Electronic Suppl. 2). Chirita
lacunosa, C. purpureolineata (Kerr ex Craib) D. Wood and an
undescribed Chirita sample formed a clade (BS = 100%, PP =
1.0) sister to a highly supported clade of Paraboea (C.B. Clarke)
Ridl., Trisepalum C.B. Clarke and Phylloboea Benth. samples
(BS = 98%, PP = 1.0), but with no statistical support for this
sister relationship (Figs. 1 and 2).
The 11 Chirita sect. Microchirita samples (not including
Chirita elata which is di scuss ed above), re prese nti ng eig ht spe-
cies (plus two undescribed), were placed in a highly supported
clade (BS = 100%, PP = 1.0) (Fig. 3) forming the basal branch
among the straight-fruited advanced Asiatic and Malesian gen-
era (Fig. 1; Fig. S1 in Electron ic Suppl. 2), though this position
received no branch support, and was not present in the BI tree
(Fig. S2 in Electronic Suppl. 2). Within the Microchirita clade
the two samples of Chirita hamosa R. Br. were sister (BS =
100%, PP = 1.0).
Ten samples of seven species of Chirita sect. Chirita
formed 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
& B.L. Burtt and H. incana (Vahl) Spreng. (Fig. 4). The rela-
tionships between Hemiboeopsis and Chirita urticifolia and
C. walkerae received no branch support. The two Henckelia
samples were sister (BS = 100%, PP = 1.0), and this pair was
sister to five Chirita species (BS = 83%, PP = 1.0).
The grouping of Chirita asperifolia with Allocheilos
W.T. Wang, Gyrocheilos W.T. Wang and Didymocarpus Wall.
was highly supported (BS = 99%, PP = 1.0) (Figs. 1 and 5),
770
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Remodelling of Chirita
Agalmyla biflora
Basal Asian, European
and Litostigma
Streptocarpus
Chirita sect. Chirita 2
Henckelia+Chirita elata
Hemiboea/Lysionotus-dom.
Briggsiopsis delavayi
Raphiocarpus macrosiphon
Lagarosolen-dominated
Chirita sect. Gibbosaccus
Loxostigma/Petrocosmea-dom.
Didymocarpus and
Gyrocheilos-dominated
Didymostigma obtusum
Didymostigma trichanthera
Chirita sect. Liebigia
Metapetrocosmea peltata
Oreocharis-dominated
Agalmyla
Aeschynanthus
Cyrtandra/Ridleyandra-dom.
Chirita sect. Microchirita
Damrongia purpureo lineata 2
Didissandra frutescens
Boea hygrometrica
Chirita sect. Chirita 1
Paraboea dominated
Henckelia sect. Loxocarpus
remaining Boea clade
100
100
100
100
100
73
100
100
100 100
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100
100
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100 100
100; 100/1.0
100
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100
100
100
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100
100
100; 100/1.0
100
100 100
100
100
100
100
95
95
100/1.0
*/*
100/1.0
92/1.0
*/*
*/0.55
100/
1.0
86 /1.0
*/0.99 */0.87
*/0.99
100/1.0
100/1.0
88/
1.0
*/0.81
53 /1.0
68 /1.0
63 /1.0
*/*
*/* 96/0.99
*/*
*/0.74
99 /1.0
*/*
*/*
*/*
100/1.0
*/*
*/*
54 /0.91
82 /
1.0 */*
100/1.0
98 /1.0
*/*
*/*
*/*
*/*
100
100/1.0
100
99/1.0
Boea philippensis
Cathayanthe biflora
Raphiocarpus sp.
Chirita purpureolineata 1
Chirita purpureolineata 2
Chirita sp. nov. 2
Chirita lacunosa 1
Chirita lacunosa 2
Trisepalum birmanicum
Phylloboea glandulosa
Paraboea crassifolia
Paraboea umbellata
Paraboea acutifolia
Paraboea capitata
100
100
100
100
100
100
100
100
100 100
52 /0.96
60 /0.98
100/1.0
98 /1.0
100/1.0
82 /1.0
100/1.0
100/1.0
*/*
84 /1.0
Fig. .
Simplif ied max imum pa rsimony majority-rule consensu s tr ee ba sed on combined trnL-F and ITS sequences. Mai n cla des are repr ese nte d
by names of genera dominating these clades. Chirita clades are highlighted in grey. Numbers above branches are majority-rule frequencies,
numbers below branches are bootstrap values (bold italics) and Bayesian posterior probabilities (bold). Asterisks indicate branches with <50%
bootstrap or <0.5 posterior probability values. Numbers in grey circles indicate branches suppor ting the monophyly of individual Chirita clades.
Numbers in grey squares indicate branches supporting the independent origins and relationships of the different Chirita clades.
Fig. .
Cladogram of the majority-rule consensus tree
from the full analysis on combined trnL-F and ITS
sequences of 232 didymocarpoid Gesneriaceae samples,
depicting the Chirita sect. Chirita clade 1 (→ Damron-
gia). For further explanation see Fig. 1.
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60 (3) • June 2011: 767–790
Chirita lavandulacea
Chirita involucrata
Chirita rupestris
Chirita mollissima
Chirita caliginosa
Chirita sp. nov. 1
Chirita tubulosa
Chirita hamosa 1
Chirita hamosa 2
Chirita viola
Chirita sp. nov. 3
100
100
100
100
100
100
100
100
100 100
88 /1.0
100/1.0
92 /0.99
95 /1.0
100/1.0
*/*
*/*
99 /1.0
93 /1.0
100/1.0
Chirita walkerae
Hemi
boeopsis longisepala
Chirita urticifolia 1
Chirita urticifolia 2
Chirita urticifolia 3
Henckelia floccosa
Henckelia incana
Chirita pumila 1
Chirita pumila 2
Chirita dielsii
Chirita anachoreta
Chirita macrophylla
Chirita bifolia
100
100
100 100
100
100
100
100
100
100 100 100
92 /1.0
*/*
*/*
100/1.0
63/0.77
83/1.0
100/1.0
72 /0.85
100/1.0
85/1.0
63/0.93
79/1.0
Chirita asperifolia
Allocheilos guangxiensis
Didymocarpus cortusifolius
Gyrocheilos chorisepalus var.synsepalus
Gyrocheilos retrotrichus
Gyrocheilos lasiocalyx
Gyrocheilos retrotrichus var. oligolobus
Didymocarpus citrinus
Didymocarpus antirrhinoides
Didymocarpus cordatus
Didymocarpus purpureobracteatus
Didymocarpus stenanthos
Didymocarpus podocarpus
Didymocarpus villosus
100
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100
100
100
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100
*/*
100/1.0
52/0.99
100/1.0
74/0.81
58/1.0
100/1.0
100/1.0
72/0.88
100/1.0
100/1.0
68/0.79
100
99/1.0
Chirita gemella
Chirita heterotricha
Chirita pteropoda
Chiritopsis bipinnatifida
Chiritopsis repanda var. guilinensis
Chiritopsis glandulosa
Chiritopsis gland. var
. yangshuoensis
Chirita sinensis
Chirita pinnata
Chirita pinnatifida
Chirita spadiciformis
Chiritopsis cordifolia
Chirita minutimaculata
Chirita ophiopogoides
Chirita wentsaii
Chirita spinulosa
Wentsaiboea renifolia
Wentsaiboea luochengensis
Chiritopsis mollifolia
Primulina tabacum
Chirita mollifolia
Chirita longgangensis
Chirita linearifolia
100
100
100
100
100
100
100
100
100
100
60
100
100
100
100
100
100
100
100
100
100
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100/1.0
*/*
100/1.0
95/1.0
100/1.0
*/0.52
86/1.0
55/0.87
87/1.0
95/1.0
*/0.98
84/0.93
*/*
99/1.0
100/1.0
*/*
*/*
76/*
85/1.0
*/*
80/1.0
100/1.0
Fig. .
Cladogram of the majority-rule consensus tree
from the full analysis on combined trnL-F and ITS
sequences of 232 didymocar poid Gesneriaceae samples,
depicting the Chirita sect. Microchirita clade (→ Micro-
chirita). For further explanation see Fig. 1.
Fig. .
Cladogram of the majority-rule consensus
tree from the full analysis on combined trnL-F and
ITS sequences of 232 didymocarpoid Gesneriaceae
samples, depicting the Chirita sect. Chirita clade 2
(→ Henckelia). For fur ther explanation see Fig. 1.
Fig. .
Cladogram of the majority-rule
consensus tree from the full analysis
on combined trnL-F and ITS sequenc-
es of 232 didymocarpoid Gesneria-
ceae samples, depicting the Chirita
sect. Liebigia clade (→ Liebigia). For
fur ther explanation see Fig. 1.
Fig. .
Cladogram of the majority-rule
consensus tree from the full analysis on
combined trnL-F and ITS sequences of
232 didymocarpoid Gesneriaceae samples,
depicting the Chirita sect. Gibbosaccus
clade (→ Primulina). For fu rther explana-
tion see Fig. 1.
772
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60 (3) • June 2011: 767–790Weber & al. •
Remodelling of Chirita
though the position among these genera is unclear. In the MP
tree (Fig. S1 in Elect ron ic Sup pl. 2) Chirita asperifolia is siste r
to all three genera, though the branch supporting this relation-
ship received no bootstrap support. In the BI tree (Fig. S2 in
Electronic Suppl. 2) it was in a sister position to the Didymo-
carpus samples (BS=100%, PP=0.99).
The clade including the samples of Chirita se ct. Gibbosac-
cus formed a strongly supported sister relationship to a clade
dominated by Lagarosolen W.T. Wang samples (B S = 100% , PP
= 1.0). The Chirita sect. Gibbosaccus clade itsel f rec eived high
br anch supp ort val ue s (BS = 100%, PP = 1.0). As well as the 14
species of Chritia sect. Gibbosaccus it included six species of
Chiritopsis, two species of Wentsaiboea and the monoty pic ge-
nus Primulina (Fig. 6). The topology within the clade differed
between the MP and BI analysis mainly due to the changing
position of Primulina tabacum Hance and Wentsaiboea renifo-
lia D. Fang & D.H . Qin (Figs. S1 and S2 in Elec tronic Suppl. 2).
The Chiritopsis samples appeared in three places, in strongly
supported relationships with Chirita samples or Wentsaiboea
luochengensis Yan Liu & W.B. Xu (Fig. 6).
DISCUSSION
This work is an extension of the recent most comprehen-
sive study on the “didymocarpoid Gesneriaceae” (Möller &
al., 2011), with an additional 27 Chirita, four Chiritopsis and
10 Henckelia sa mples added. The present st udy focuses on the
highly polyphyletic genus Chirita which appears in six differ-
ent locations in the resulting phylogenies. Despite the differ-
ences in topology between the MP and BI analysis, which is
mainly the result of a poorly supported backbone structure, the
separate positions of the Chirita samples were hig hly sup por ted
and are not likely artefacts. Although the intergeneric and clade
relationships cannot be elucidated from these results, the high
support values for the individual clades containing Chirita spe-
cies means that we can with some confidence propose redefined
genera for these different clades, in some cases agreeing with
the very recent changes proposed by Wang & al. (2011) and in
some cases disagreeing.
Chirita sect. Chirita 1 clade. —
The clade labelled Chirita
sect. Chirita 1 in Fig. 1 contains Chirita purpureolineata, C. la-
cunosa and an undescribed species. Chirita purpureolineata
was originally described as Damrongia purpureolineata Kerr
ex Craib and is the type species of the genus Damrongia. This
genus name is available for this clade. This clade was not found
by Wang & al. (2011), nor did they discuss the isolated position
of Chirita lacunosa in Möller & al. (2009).
Chirita sect. Chirita 2 clade. —
The clade labelled Chirita
sect. Chirita 2 in Fig. 1 contains the type species of the genera
Chirita (C. urticifolia), Henckelia (H. incana) and Hemiboeopsis
(H. longisepala (H.W. Li) W.T. Wang, the only species). The other
species of Henckelia included, H. floccosa, is also a member of
Henckelia sect. Henckelia which is a group of 14 rather similar
species in South India and Sri Lanka. The other taxa in this clade
are all species of Chirita sect. Chirita. This clade can be recog-
nised as the genus Henckelia with Chirita and Hemiboeopsis in
synonymy. Given that both Henckelia and Chirita were large
genera this conclusion has far-reaching implications, not only
because Chirita now disappears altogether, but also due to the
radical change in the delimitation of Henckelia. Henckelia now
includes only the species of Henckelia from South India and
Sri Lanka plus most of the species of Chirita sect. Chirita and
excludes all species formerly placed in Henckelia from T haila nd
and Malesia. Unfortunately the Thai and Malesian species of
Henckelia made up the very large majority of species. Manu-
scripts on the fate of the se spe cies, along with fu r ther mole cu la r
phylogenetic work, are in preparation. Wang & al. (2011) did not
include any species of Henckelia in their study and concluded
that this clade corresponded to Chirita sect. Chirita as defined
by Wood (1974) except for Liebigia (see below).
Chirita sect. Microchirita clade. —
Chirita sect. Micro-
chirita is a morphologically well defined group that, surpris-
ingly, had never been recognised at generic rank before being
raised by Wang & al. (2011). We agree that generic rank is
appropriate for Microchirita. Although the sampling within
the 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-
some numbers in the section (C. involucrata Craib, C. caligi-
nosa C.B. Clarke and C. rupestris Ridl. belong to the group
with x = 9, whi le C. lavandulacea Stapf and C. hamosa belong
to the group having x = 17—see Kiehn & al., 1998) as well as
the geographical distribution (all diploids with x = 9 occur in
Peninsular Malaysia and Southern Thailand, while the species
with x = 17, possibly dys-tetraploids, derived from 2n = 18,
have a more northerly distribution in Central and Northern
Thailand, Vietnam and Southern China (Wood’s, 1974 records
of C. hamosa for Southern Thailand and Northern Peninsular
Malaysia are erroneous due to confusion with C. viola Ridl.).
The species (including the undescribed species) which have
not had their chromosomes counted also divide into the same
northern and southern clades. The group of species highlighted
by Wood (1974) as having the unusual feature of the presence
of hairs with swollen tips on the upper surface of the corolla
throat, C. caliginosa, C. mollissima Rid l. and C. viola, do come
out as a group sister to C. rupestris. Chirita sp. nov. 3, which
is very close to C. sericea, is also in this group.
Chirita elata. —
The generic position of Chirita elata has
be en problematic since it wa s firs t descr ibe d (see Raf idah & al.,
2011). Our results show it nested firmly amongst the Malesian
species of Henckelia (in the sense of Weber & Burtt, 1998),
which is in agreement with the diandrous flowers.
Chirita sect. Liebigia clade. —
Chirita sect. Liebigia was
relatively recently resurrected as a distinct section within
Chirita by Hilliard (2004), having been included in Chirita
sect. Chirita by Wood (1974). Wood (1974) recognise d only one
widespread and variable species, Chirita asperifolia, but Hill-
iard (2004) recognised 12 species, 10 of them based on material
also available to Woods (who freely admitted that “the account
of Chirita asperifolia is probably the least satisfactory in this
revision”). We have sampled only material identified by Hill-
iard as Chirita asperifolia, the type species of the section, and
it is quite distinct from the other parts of Chirita sect. Chirita
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and from other didymocarpoid genera. Liebigia, at genus rank,
is already available for this group.
Chirita sect. Gibbosaccus clade. —
Chirita sect. Gibbosac-
cus is by far the largest se ction wit hin Chirita. Our result s sh ow
all species in a well-supported clade but with the species of
Chiritopsis, Wentsaiboea renifolia, W. luochengensis and Prim-
ulina nested within. In this group the genus name Primulina,
previously monotypic but very recently expanded to include
these groups by Wang & al. (2011), has priority.
TAXONOMIC IMPLICATIONS
Each of the six clades identified above has already been
highlighted in previous taxonomic work as morphologically
distinct. The molecular work presented here lends weight to the
recognition of each of the groups as distinct taxa (or in a differ-
ent genus in the case of Chirita elata). However, these groups
are not always easily distinguished from each other and not
easily distinguished from other didymocarpoid Gesneriaceae,
a not uncommon problem in the family.
The single character which was used to unite the species
and sections of Chirita into one genus and distinguish it from
other genera was the lack of development of the upper lobe of
the stigma and the usually bifid development of the lower lobe
(henceforth referred to as the chiritoid stigma). It was already
recognised that this character, although useful, had its prob-
lems and led to confusion over the correct placement of some
species. Below we discuss the morphological basis of each of
the genera we now recognise but acknowledge that further
research is necessary in the didymocarpoid Gesneriaceae to
better understand the morphological as well as molecular evo-
lution in the group.
Below we list all of the taxa we now recognise within each
genus, including the many necessary new combinations. We
list heterotypic synonyms when the synonym was published
in Chirita, but we must emphasise that we have not critically
checked the status of these synonyms. Instead we have gener-
ally adopted heterotypic synonyms for names in Chirita as they
have appeared in recent revisions, especially by Wood (1974)
and Wang & al. (1998). Further work may show that some
names presently found in synonymy must be re-established
and transferred to one of the present genera. In Appendix 1
we give a complete list of all names ever published in Chirita
and the current name for each of them.
It should be borne in mind that we have moved all cur-
rently recognised taxa out of Chirita even when we cannot be
sure we have made the correct decision. Chirita as a genus is
a synonym of Henckelia and no currently recognised species
can bear this generic name. To leave species in Chirita due to
some uncertainties would result in competing and confusing
classification frameworks for this group. Instead we have made
new combinations for all taxa based on our best knowledge of
where each taxon now belongs. We freely admit that in some
cases we may be mistaken and have highlighted these taxa in
the Appendices. These taxa, however, were already previously
poorly understood and we have merely ensured that they are
accounted for in the new classification framework and hope
their status can be better resolved in the future.
One new combination is made in the genus Codonoboea
for Chirita humilis Miq. Chirita humilis was previously also
included in Henckelia, the Malesian members of which must
now be excluded from the genus. Manuscripts focusing on this
group are in preparation but for the sake of completeness this
new combination is proposed in Appendix 1.
Henckelia Spreng., Anleit. Kenntn. Gew., ed. 2, 2(1): 402. 1817
Henckelia sect. Henckelia Weber & Bu rtt in Beitr. Biol.
Pflanzen 70: 334. 1998 (‘1997’) Didymocarpus sect.
Ortho boea Benth. in Benthem & Hooker, Gen. Pl. 2(2):
1022. 1876 – Type: Henckelia incana (Vahl) Spreng.
= Chirita Buch.-Ham . ex D. Don, Edi nbu rgh Phi lo s. J. 7: 83. 1822
Chirita sect. Euchirita C.B. Clarke in Candolle & Can-
dolle, Monogr. Phan. 5(1): 111. 1883 ≡ Didymocarpus sect.
Euchirita (C.B. Cla rke) Chun in Su nyatse nia 6: 294. 1946
Roettlera sect. Euchirita (C.B. Clarke) Fritsch in Engler &
Prantl, Nat. Pflanzenfam. IV/3b: 148. 1895 – Type (lectotype
Burtt, 1954): Chirita urticifolia Buch.-Ham. ex D. Don (
Henckelia urticifolia (Buch.-Ham. ex D. Don.) A. Dietr.).
= Calosacme Wall., Numer. List: 800–806. 1829. All species
names established in Calosacme by Wallich (1829) are
nomina nuda.
= Babactes DC. ex Meisn., Pl. Vasc. Gen. 1: tab. diag. 302,
Comm. 211. 1840 – Type: Babactes oblongifolia ( Roxb.)
DC. (≡ Henckelia oblongifolia (Roxb.) D.J. Middleton &
Mich. Möller).
= Gonatostemon Regel in Gartenflora 15: 353. 1866 – Type:
Gonatostemon boucheanum Regel (≡ Henckelia urticifolia
(Buch.-Ham. ex D. Don.) A. Dietr.).
= Ceratoscyphus Chun in Sunyatsenia 6: 276. 1946 – Type:
Ceratoscyphus caerulea Chun (≡ Henckelia ceratoscyphus
(B.L. Burtt) D.J. Middleton & Mich. Möller).
= Hemiboeopsis W.T. Wang in Acta Bot. Yunnan. 6: 397. 1984
– Type: Hemiboeopsis longisepala (H.W. Li) W.T. Wang (≡
Henckelia longisepala (H.W. Li) D.J. Middleton & Mich.
Möller).
Perennial or annual herbs, sometimes woody at base,
caulescent or acaulescent, rarely creeping. Leaves opposite,
alternate or whorled, sometimes clustered at the stem apex, or
reduced to just one or two leaves, petiolate, lamina variable,
sometimes pinnately lobed. Inflorescences axillary (or some-
times appearing terminal by reduction of vegetative shoot),
flowers 1–15; bracts paired or in whorls, free or connate at
base, orbicular to linear, narrowly ovate or narrowly triangular,
sometimes early caducous. Calyx of 5 free lobes or fused into a
tube for part of length, when fused often more deeply divided
on dor sal side, seg ments tr iangular or nar rowly tria ng u la r. Co-
rolla infundibuliform, more rarely tubular, tube often slightly
pouched, sometimes rather constricted in throat; bilabiate with
2-lobed upper lip and 3-lobed lower lip; colouration variable,
often with yellow markings in throat. Stamens 2; filaments
geniculate or straight; anthers fused face to face, glabrous or
pubescent. Disc a simple annular ring or 5-lobed, often very
small. Ovary shortly stipitate or not; stigma chiritoid, often
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weakly so and rarely almost truncate, lower lobe bifid or not.
Fruit a capsule splitting into 2 valves or opening only along the
dorsal side, stipitate or not, plagiocarpic or not, calyx persistent
or not in fruit. Seeds minute, ellipsoid.
Distribution: Sri Lanka, Southern and Northeastern India,
Nepal, Bhutan, Southern China, Northern Viet nam, Northern
Laos, Norther n Thailand. — As far as is known Henckelia
species are found on soils and rocks but not on limestone. Ap-
proximately 56 species.
Clarke (1883, 1884) noted the difficulty of distinguishing
Didymocarpus and Chirita but it is likely th at he wa s, in part icu-
lar, referr ing to those species which were later (Weber & Burtt,
1998) removed from Didymocarpus to a resurr ecte d Henckelia.
Henckelia was distinguished by having a fruit which dehisces
only along the dorsal side, whilst Didymocarpus has fruits which
dehisce into two valves. It was noted by Weber & Burtt (1998)
that the South Indian and Sri Lankan species were distinct
enough to form a section, Henckelia sect. Henckelia, the type
se ction of the genus. Its morphology is described by these authors
in some detail under “South Indian Didymocarpus” (Weber &
Burtt, 1998: 316–317). Study of specimens and evaluation of the
older literature and illustrations have revealed that stigma shape
is variable in this group, ranging from truncate-capitate to defi-
nitely chiritoid with a short or undeveloped upper stigma lobe
and a large lower lobe. In one individual of Henckelia incana it
has even been observed that the lower lobe is bifid, and the same
is suggested in the description of the stigma of Henckelia repens
by Clarke (1883, as Didymocarpus repens). Therefore, there is no
sharp boundary between Henckelia sect. Henckelia and Chirita
with regard to the crucial stigma character. A chiritoid stigma,
however, has never been observed in the Malesian species placed
in Henckelia by Weber & Bur t t (1998). In addition the fr u it cha r-
acter used to characterise Henckelia, dehiscence only along the
dorsal side, has also been observed in Chirita smitinandii B.L.
Burtt and C. lachenensis C.B. Clarke and may occur in other
Chirita species in fresher material (drying of specimens often
makes this character difficult to observe, particularly in those
species with very long and thin fruits such as Chirita pumila). All
but one species of Henckelia sect. Henckelia s.str. are acaules-
cent. Most species of Chirita sect. Chirita (without Damrongia)
are caulescent but acaulescent species of Chirita sect. Chirita
are known such as C. briggsioides W.T. Wang , C. dielsii (Borza)
B.L. Bur tt, C. lachenensis and C. primulacea C.B. Clarke. It
should also be noted that both caulescent and acaulescent spe-
cies are known from other Gesneriaceae genera (e.g., Paraboea).
Recorded chromosome numbers are n = 4, 9, 10, 14, 16, 17, 27
and ±45 (see “Webcyte”, Möller & al., 2002 onwards). The newly
defined genus Henckelia is variable but no more so than other
large didymocarpoid genera. It occurs in South India and Sri
Lanka, the Himalayas and continental Southeast Asia, excluding
Peninsular Thailand and Malaysia. It does not get into Malesia.
Some of the species we include in Henckelia are known
to us only from herbarium specimens and a few only from the
literature. Habitat preference is something not well documented
on specimens or in the literature with most species recorded as
occurring on rocks. We have the impression from those species
that we do know from the field that most, if not all, species in
the newly circumscribed Henckelia do not occur on limestone
but rather on sandstone, granite and other non-limestone rocks.
This is in marked contrast to Damrongia, Microchirita and
Primulina which are almost exclusively found on limestone.
Although the name Hemiboeopsis suggests a close affin ity
to Hemiboea, it relates only to the similarity in habit. Wang
(1984) himself stressed a possible relationship with Chirita in
the diagnosis (“fortasse affinis Chiritae D. Don”). Th e st a mens,
the stigma shape (though with short upper lobe), as well as the
long, cylindrical and orthocarpic capsules are not similar to
Hemiboea but rather confirm Wang’s suggestion of a relat ion-
ship to Chirita. The molecular data show that it is part of the
Chirita sect. Chirita clade.
Wang & al. (2011) included several species of Chirita sect.
Chirita in their analysis but as they did not include any species
of Henckelia they incorrectly maintained the genus Chirita to ac-
commodate these species. They also did not include Hemiboeop-
sis in their analysis so did not reduce this genus to synonymy.
There are a number of species which we include in Henck-
elia but which have not be en included in the molecul ar an alyses
and for which their inclusion could be debated. These include
Henckelia forrestii (J. Anthony) D.J. Mid dleto n & Mi ch . Möller,
H. smitinandii (B.L. Burtt) D.J. Middleton & Mich.ller.
and H. tibetica (Franch.) D.J. Middleton & Mich. Möller. Only
a more focused study on these species can resolve this issue
but until contrary evidence is found they should be included in
Henckelia along with most other species in Chirita se ct. Chirita.
In its new sense, Henckelia di ffers fu nda mentally from the
(purely morphological) concept of Weber & Burtt (1998): it is
an amalgamation of the type sections of Henckelia and Chirita
(excluding the species placed in Damrongia), but excludes all
other infrageneric taxa referred to Henckelia by these authors
(se cts . Loxocarpus, Didymanthus, Heteroboea and Glossad-
enia). The fate of the species in the se sect io ns wi ll be dis cusse d
in forthcoming papers.
Henckelia adenocalyx (Chatterjee) D.J. Middleton & Mich.
Möller, comb. nov. Chirita adenocalyx Chatterjee in
Kew Bull. 1948: 63. 1948. — Distr.: India (Arunachal
Pradesh), Northern Burma, China (Yunnan).
Henckelia anachoreta (Hance) D.J. Middl eton & Mich. Möller,
comb. nov. Chirita anachoreta Hance in Ann. Sci. Nat.,
Bot., sér. 5, 5: 231. 1866 Didymocarpus anachoretus
(Hance) H. Lév. in Compt. Rend. Assoc. Franç. 34: 427.
1906 (‘1905’) = Chirita minutiserrulata Hayata, Icon. Pl.
Formosan. 5: 133. 1915. — Distr.: India (Sikkim), Burma,
Northern Thailand, China (Guangdong, Guangxi, Hunan,
Yunnan, Xizang), Taiwan, Laos, Northern Vietnam.
Henckelia angusta (C.B. Clarke) D.J. Middlet on & Mich. Möl ler,
comb. nov. Chirita zeylanica var. angusta C.B. Clarke
in Candolle & Candolle, Monogr. Phan. 5(1): 113. 1883 ≡
Chirita angusta (C.B. Clarke) W.L . Theob. & Gr upe in Ce y-
lon J. Sci., Biol. Sci. 10(1): 70. 1972. — Distr.: Sri Lanka.
Henckelia auriculata (J.M. Li & S.X. Zhu) D.J. Middleton
& Mich. Möller, comb. nov. Chirita auriculata J.M. Li
& S.X. Zhu in Ann. Bot. Fenn. 47: 222. 2010. — Distr.:
China (Yunnan).
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Henckelia bifolia (D. Don) A. Dietr., Sp. Pl., ed. 6, 1: 574.
1831 Chirita bifolia D. Don, Prodr. Fl. Nepal.: 90. 1825
= Chirita amplectens Steud., Nomencl. Bot., ed. 2, 1: 351.
1840, nom. nud. = Chirita scabra Royle ex C.B. Clarke in
Candolle & Candolle, Monogr. Phan. 5(1): 113. 1883, nom.
illeg. pro syn. — Distr.: India (Himachal Pradesh, Uttar
Pradesh), Nepal, Bhutan.
Henckelia briggsioides (W.T. Wang) D.J. Middleton & Mich.
Möller, comb. nov. Chirita briggsioides W.T. Wang in
Bull. Bot. Res., Harbin 2(2): 138. 1982. — Distr.: China
(Hubei).
Henckelia burttii D.J. Middleton & Mich. Möller, nom. nov.
Chirita reptans B.L. Burtt & Panigrahi in Notes Roy.
Bot. Gard. Edinburgh 26: 265. 1965, non Henckelia reptans
(Jack) Spreng. (1827). — Distr.: Northern Burma.
Henckelia calva (C.B. Cl arke) D.J. Mi ddleton & Mic h. Möl le r,
comb. nov. Chirita calva C.B. Clarke in Candolle &
Candolle, Monogr. Phan. 5(1): 116. 1883. — Distr.: India
(Assam, Sikkim), Northern Burma.
Henckelia ceratoscyphus (B.L . Bur t t) D.J. Mid dleto n & Mich.
Möller, comb. nov. Chirita ceratoscyphus B.L. Bur tt in
Notes Roy. Bot. Gard. Edinburgh 21: 212. 1954 ≡ Cerato-
scyphus caeruleus Chun in Sunyatsenia 6: 278. 1946, non
Chirita caerulea R. Br. (1840), non Henckelia caerulea
(Ridl.) A. Weber (1998) = Chirita corniculata Pellegr. in
Bull. Soc. Bot. France 73: 420. 1926, non Henckelia cor-
niculata (Jack) Spren g. (1827). — Di str.: Ch ina (Guangxi),
Northern Vietnam.
Henckelia communis (Ga rd ner) D.J. Middleton & Mich.
Möller, comb. nov.Chirita communis Gardner in Cal-
cutta J. Nat. Hist. 4: 481. 1846 = Chirita zeylanica Hook.
in Bot. Mag. 71: t. 4182. 1845, non Henckelia zeylanica
(R. Br.) A. Weber & B.L. Burtt (1998) =
Chirita vulgaris
Morren in
Belgique Hort. 3: 238, t. 36. 1853, nom. nud.
— Distr.: Sri Lanka.
Henckelia dibangensis (B.L. Burtt, S.K. Srivastava & Mehro-
tra) D.J. Middleton & Mich. Möller, comb. nov. Chirita
dibangensis B.L. Burtt, S.K. Srivastava & Mehrotra in
Notes Roy. Bot. Gar d. Ed inbur gh 45(3): 469. 1989 (‘1988’).
— Distr.: India (Arunachal Pradesh).
Henckelia dielsii (Borza) D.J. Middleton & Mich. Möller,
comb. nov. Chirita dielsii (Borza) B.L. Burtt in Notes
Roy. Bot. Gard. Edinburgh 23: 97. 1960 = Chirita orbicu-
laris W.W. Sm. in Notes Roy. Bot. Gard. Edinburgh 9: 94.
1916. — Distr.: China (Yunnan, Sichuan).
Henckelia dimidiata (Wall. ex C.B. Clarke) D.J. Middleton
& Mich. Möller, comb. nov. Chirita dimidiata Wall.
ex C.B. Clarke in Candolle & Candolle, Monogr. Phan.
5(1): 115. 1883 = Chirita polyneura Miq. var. amabilis
C.B. Clarke, Commelyn. Cyrtandr. Bengal.: t. 76. 1874 =
Chirita dimidiata Steud ., Nomen cl. Bot., ed . 2, 1: 351. 1840 ,
nom. nud. — Distr.: India (Assam, Sikkim).
Henckelia fasciculif lora (W.T. Wang), D.J. Mid dleto n & Mich.
Möller, comb. nov.Chirita fasciculif lora W.T. Wang in
Bull. Bot. Res., Harbin 2(2): 139. 1982. — Distr.: China
(Yunn a n).
Henckelia fischeri (Gamble) A. Weber & B.L. Burtt in Beitr.
Biol. Pf lanzen 90: 344. 1998 (‘1997’) Didymocarpus
fischeri Gamble in Bull. Misc. Inform. Kew 1923: 117.
1923. — Distr.: South India.
Henckelia floccosa (Thwaites) A. Weber & B.L. Burtt in Beitr.
Biol. Pf lanzen 90: 344. 1998 (‘1997’) Didymocarpus
floccosus Thwaites, Enum. Pl. Zeyl.: 207. 1860. — Distr.:
Sri Lanka.
Henckelia forrestii (J. Anthony) D.J. Middleton & Mich.
Möller, co mb . nov.Chirita forrestii J. Ant hony in Not es
Roy. Bot. Gard. Edinburgh 18: 192. 1934 Didymocarpus
forrestii (J. Anthony) Hand.-Mazz., Symb. Sin. 7: 882.
1936 = Chirita forrestii var. acutidentata W.T. Wang in
Acta Phytotax. Sin. 13(3): 103. 1975. Distr.: China
(NW Yunnan). — Chirita forrestii was referr ed to Chirita
sect. Chirita by Wood (1974), but to Chirita sect. Gib-
bosaccus by Wang & al. (1998) and Li & Wang (2004)
which would result in it now being transferred to Prim-
ulina. We conclude that it is a Henckelia but further st udy
is clearly necessary to resolve the issue of its correct
placement.
Henckelia fruticola (H.W. Li) D.J. Middlet on & Mich. Möl le r,
comb. nov.Chirita fruticola H.W. Li in Bull. Bot. Res.,
Harbin 3(2): 37. 1983. — Distr.: China (Yunnan)
Henckelia gambleana (C.E.C. Fisch.) A. Weber & B.L. Burtt
in Beitr. Biol. Pflanzen 90: 345. 1998 (‘1997’) ≡ Didymo-
carpus gambleanus C.E.C. Fisch. in Bull. Misc. Inform.
Kew 1938: 36. 1938. — Distr.: South India.
Henckelia grandifolia A. Dietr., Sp. Pl., ed. 6, 1: 576. 1831 ≡
Chirita macrophylla Wall., Pl. Asiat. Rar. 1: 56, t. 72. 1830),
non Henckelia macrophylla (D. Don) Spreng. = Chirita
macrophylla Wall. var. tirapensis Panigrahi in Bull. Bot.
Soc. Bengal. 21: 32. 1967. — Distr.: Nepal, India (Assam,
Sikkim, West Bengal), Bhutan, Northern Burma, China
(Yunnan), Northern Thailand.
Henckelia heterostigma (B.L. Burtt) D.J. Middleton & Mich.
Möller, comb. nov. Chirita heterostigma B.L. Burtt in
Notes Roy. Bot. Gard. Edinburgh 26: 262. 1965. — Distr.:
Burma.
Henckelia hookeri (C.B. Clarke) D.J. Middleton & Mich.
Möller, comb. nov. Chirita hookeri C.B. Clarke,
Commelyn. Cyrtandr. Bengal.: t. 70. 1874. — Distr.: India
(Assam).
Henckelia humboldtiana (Gardner) A. Weber & B.L. Burtt in
Beitr. Biol. Pflanzen 90: 346. 1998 (‘1997’) ≡ Didymocar-
pus humboldtianus Gardner in Calcutta J. Nat. Hist. 6: 477.
1846. — Distr.: Sri Lanka.
Henckelia incana (Vahl) Spreng., Syst. Veg. 1: 38. 1824
Rott lera incana Vahl, Enum. Pl. 1: 88. 1804. — Distr.:
South India.
Henckelia infundibuliformis (W.T. Wang) D.J. Middleton
& Mich. Möller, comb. nov.Chirita infundibuliformis
W.T. Wang in Acta Phytotax. Sin. 17(1): 111. 1979.
Distr.: China (Xizang).
Henckelia innominata (B.L. Burtt) A. Weber & B.L. Burtt
in Beitr. Biol. Pflanzen 90: 347. 1998 (‘1997’) ≡ Didymo-
carpus innominatus B.L. Burtt in Notes Roy. Bot. Gard.
Edinburgh 21: 201. 1954. — Distr.: South India.
776
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60 (3) • June 2011: 767–790Weber & al. •
Remodelling of Chirita
Henckelia insignis (C.B. Clar ke) D.J. Midd le ton & Mich. Möller,
comb. nov.Chirita insignis C.B. Clarke in Candolle &
Candolle, Monogr. Phan. 5(1): 121. 1883. — Distr.: Burma.
Henckelia lacei (W.W. Sm.) D.J. Middleton & Mich. Möller,
comb. nov. Didymocarpus lacei W.W. Sm. in Rec.
Bot. Surv. India 6: 41. 1913 Chirita lacei (W.W. Sm .)
B.L. Burtt in Notes Roy. Bot. Gard. Edinburgh 21: 187.
1954. — Distr.: Burma. — Wood (1974: 161–162) placed
the species tentatively in Chirita sect. Chirita, but ear-
marked it as “another anomalous” species, whose position
“remains uncertain”. We make the transfer to Henckelia
but note that further study is required.
Henckelia lachenensis (C.B. Clarke) D.J. Middleton & Mich.
Möller, comb. nov.Chirita lachenensis C.B. Clarke in
Candolle & Candolle, Monogr. Phan. 5(1): 118. 1883 ≡
Chirita clarkei Hook. f., Fl. Brit. India 4: 359. 1884, nom.
illeg. = Chirita umbricola W.W. Sm. in Notes Roy. Bot.
Ga rd. Edinbu rgh 10: 172. 1918 = Chirita stolonifera C.E.C.
Fisch. & Kaul in Kew Bull. Misc. Inform. 1940: 196.
1940. — Distr.: Bhutan, India (Assam, Sikkim), Norther n
Burma, China (Yunnan).
Henckelia longipedicellata (B.L. Burtt) D.J. Middleton &
Mich. Möller, comb. nov. Chirita longipedicellata
B.L. Burtt in Notes Roy. Bot. Gard. Edinburgh 26: 264.
1965. — Distr.: Burma.
Henckelia longisepala (H.W. Li) D.J. Middleton & Mich.
Möller, comb . nov.Lysionotus longisepalus H.W. Li,
Bu ll. Bot. Res., Harbin 3(2): 1. 1983Hemiboeopsis longi-
sepala (H.W. Li) W.T. Wang in Acta Bot. Yunna n. 6(4): 397.
1984. — Distr.: China (Yunnan), Laos.
Henckelia lyrata (Wig ht) A. Weber & B.L. Burtt in Beitr. Biol.
Pflanzen 90: 349. 1998 (‘1997’) ≡ Didymocarpus lyratus
Wight, Icon. Pl. Ind. Orient. 42: 10, t. 1350. 1848. — Dist r.:
South India.
Henckelia macrostachya (E. Barnes) A. Weber & B.L. Burtt
in Beitr. Biol. Pflanzen 90: 349. 1998 (‘1997’) ≡ Didymo-
carpus macrostachyus E. Barnes in Bull. Misc. Inform.
Kew 1938: 37. 1938. — Distr.: South India.
Henckelia meeboldii (W.W. Smith & Ramaswami) A. Weber
& B.L. Burtt in Beitr. Biol. Pflanzen 90: 349. 1998 (‘1997’)
Didymocarpus meeboldii W.W. Sm it h & Ramaswami in
Rec. Bot. Surv. India 6: 43. 1913. — Distr.: South India.
Henckelia mishmiensis (Debba r m. ex Bisw as) D.J. Midd leton
& Mich. Möller, comb. nov.Chirita mishmiensis Deb-
barm. ex Biswas in Assam Forest Rec., Bot. 1: 21. 1934.
— Distr.: India (Assam).
Henckelia missionis (Wall. ex R. Br.) A. Weber & B.L. Burtt
in Beitr. Biol. Pflanzen 90: 350. 1998 (‘1997’) ≡ Didymo-
carpus missionis Wall. ex R. Br., Cyrtandreae: 119. 1839.
— Distr.: South India.
Henckelia monantha (W.T. Wang) D.J. Middleton & Mich.
Möller, comb. nov.Chirita monantha W.T. Wang in Bull.
Bot. Res., Ha rbin 2(4): 53. 1982. — Distr.: Ch ina (Hu nan).
Henckelia monophylla (C.B. Clarke) D.J. Middleton & Mich.
Möller, comb. nov.Chirita monophylla C.B. Clarke in
J.D. Hooker, Fl. Brit. India 4: 360. 1884. — Distr.: India
(Assam).
Henckelia moonii (Gardner) D.J. Middleton & Mich. Möller,
comb. nov. Chirita moonii Gard ner, Calcut ta J. Nat. Hist .
6: 479. 1846. — Distr.: Sri Lanka.
Henckelia oblongifolia (Roxb.) D.J. Midd le ton & Mich. Möller,
comb. nov.Incarvillea oblongifolia Roxb., Fl. Ind., ed.
2, 3: 113. 1832 ≡ Chirita oblongifolia (Roxb.) J. Sinclair in
Bull. Bot. Soc. Bengal 9: 102. 1957 ≡ Chirita oblongifo-
lia (Roxb.) B.L. Burtt in Notes Roy. Bot. Gard. Edinburgh
22: 307. 1958, comb. superfl. = Chirita acuminata Wall. ex
R. Br., Cyrtandreae: 117. 1839, nom. nud. = Chirita acumi-
nata Steud., Nomencl. Bot., ed. 2, 1: 351. 1840, nom. nud.
— Distr.: India (Assam), Northern Burma, China (Yunnan).
Henckelia ovalifolia (Wight) A. Weber & B.L. Bur tt in Beitr.
Biol. Pf lanzen 90: 351. 1998 (‘1997’) Didymocarpus
ovalifolius Wight, Icon. Pl. Ind. Orient. 4(2): 10, t. 1351.
1848. — Distr.: South India.
Henckelia peduncularis (B.L. Burtt) D.J. Middleton & Mich.
Möller, comb. nov. Chirita peduncularis B.L. Burtt in
Notes Roy. Bot. Gard. Edinburgh 26: 265. 1965. — Distr.:
Northern Burma.
Henckelia primulacea (C.B. Clarke) D.J. Middleton & Mich.
Möller, comb. nov. Chirita primulacea C.B. Clarke,
Commelyn. Cyrtandr. Bengal.: t. 82. 1874. — Distr.: India
(Sikkim), Nepal.
Henckelia puerensis (Y.Y. Qian) D.J. Middleton & Mich.
Möller, comb. nov.Chirita puerensis Y.Y. Qian in Acta
Phyt ota x. Sin. 34(2): 216. 1996. — Dist r.: China (Yunnan).
Henckelia pumila (D. Don) A. Dietr., Sp. Pl., ed. 6, 1: 574.
1831 ≡ Chirita pumila D. Don, Prodr. Fl. Nepal.: 90. 1825
= Chirita edgeworthii A. DC., Pro dr. 9: 269. 1845 = Chirita
polyneura Miq. var. thomsonii C.B. Clarke, Commelyn.
Cyrtandr. Bengal.: t. 75. 1874 = Chirita sphagnicola
H. Lév. & Vaniot in Bull. Soc. Bot. France 53: 550. 1906
= Chirita diaphana Royle, Il l. Bot . Him al. Mts.: 294. 1836,
nom. nud . = Chirita flava Wall . ex R. Br., Cyr t and rea e: 117.
1839, nom. superf l. = Chirita flava Steud., Nomencl. Bot.,
ed. 2, 1: 351. 1840, nom. nud. Distr.: Northern India
(Himachal Pradesh to Assam), Nepal, Bhutan, Northern
Burma, Thailand, Southern China (Yunnan, Guizhou),
Northern Vietnam.
Henckelia pycnantha (W.T. Wang) D.J. Middleton & Mich.
Möller, comb. nov.Chirita pycnantha W.T. Wang in
Novon 7(4): 425. 1998 (‘1997’). — Distr.: China (Yunnan).
Henckelia repens (Bed d.) A. Webe r & B.L. Bu r tt in Beit r. Biol.
Pflanzen 90: 354. 1998 (‘1997’) ≡ Didymocarpus repens
Bedd., Icon. Pl. Ind. Orient. 1: 24, t. 120. 1874. — Distr.:
South India.
Henckelia rotundata (Barnett) D.J. Middleton & Mich. Möller,
comb. nov.Chirita rotundata Barnett in Kew Bull. 15:
254. 1961. — Distr.: Northern Thailand.
Henckelia shuii (Z. Yu Li) D.J. Middleton & Mich. Möller,
comb. nov.Chirita shuii Z. Yu Li in Novon 7(4): 427.
1998 (‘1997’). — Distr.: China (Yunnan).
Henckelia smitinandii (B.L. Burtt) D.J. Middleton & Mich.
Möller, comb. nov. Chirita smitinandii B.L. Burtt in Thai
Forest Bull., Bot. 29: 89. 2001. — Distr.: Eastern Thailand
(Khao Yai N.P.). — Burtt (2001) did not give the sectional
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60 (3) • June 2011: 767–790
placement of the species but suggested it was related to
Chirita poilanei Pellegr., a species we place in Primulina.
However, in its distribution, thin leaves, dorsally dehis-
cent capsule and habitat preference (not on limestone) it
would appear to be more similar to species now placed in
Henckelia. Further study is necessary.
Henckelia speciosa (Kurz) D.J. Middleton & Mich. Möller,
comb. nov. Chirita speciosa Kurz in J. Bot . 11: 195. 1873
= Chirita brevipes C.B. Clarke in Candolle & Candolle,
Monogr. Phan. 5(1): 120. 1883 = Chirita trailliana Forrest
& W.W. Sm. in Notes Roy. Bot. Gard. Edinburgh 9: 95.
1916. — Distr.: Southern China (Yunnan).
Henckelia tibetica (Franch.) D.J. Middleton & Mich. Möller,
comb. nov. Roettlera tibetica Franch. in Bull. Mus.
Hist. Nat. (Paris) 5: 251. 1899 ≡ Didymocarpus tibeticus
(Franch.) Hand.-Mazz., Symb. Sin. 7: 881. 1936 ≡ Chirita
tibetica (Fr anch.) B.L. Bu r tt in Not es Roy. Bot . Gard. Edin -
burgh 23: 99. 1960. — Distr.: China (Yunnan, Sichuan).
Burtt (1960: 99) discussed the transfer of Hemiboea
subacaulis Hand.-Mazz. and Roettlera tibetica Franch. to
Chirita and he regarded the two species as closely allied.
Wang & al. (1998) did not accept the transfer of Hemiboea
subacaulis to Chirita, and thus the two species are cur-
rently placed in different genera. The placement of the
species thus is most problematic and requires further study.
Given that to us it has closer affinities with Chirita sect.
Chirita than to Chirita sect. Gibbosaccus we make the
transfer to Henckelia pending these studies.
Henckelia urticifolia (D. Don.) A. Dietr., Sp. Pl., ed. 6, 1: 574.
1831 ≡ Chirita urticifolia Buch.-Ham. ex D. Don, Prodr.
Fl. Nepal.: 90. 1825 = Chirita grandiflora Wall., Pl. Asiat.
Rar. 1: 43. 1830. — Distr.: India (Assam, Sikkim), Nepal,
Bhutan, Northern Burma, Southern China (Yunnan).
Henckelia walkerae (Ga rd ner) D.J. Midd leton & Mich. Möller,
comb. nov. Chirita walkerae Gardner in Calcutta J. Nat.
Hi st. 6: 480. 1846 (‘walkeri’) = Chirita walkerae va r. parvi-
flora C.B. Clarke in Candolle & Candolle, Monogr. Phan.
5(1): 112. 1883 ≡ Chirita walkerae Gardner subsp. parvi-
flora (C.B. Clarke) W.L. The ob. & Grupe in Ceylo n J. Sci.,
Biol. Sci. 10(1): 70. 1972. — Distr.: Sri Lanka. — The spe-
cific epithet has previously almost always been spelt ‘walk-
eri’ but Gardner makes very clear that his plant is named
after a Mrs. Walker so must be corrected to ‘walkerae’
under Art. 60.11 of the ICBN (McNeill & al., 2006).
Henckelia wightii (C.B. Clarke) A. Weber & B.L. Burtt in
Beitr. Biol. Pflanzen 90: 359. 1998 (1997’) Didymocarpus
rottlerianus va r. wightii C.B. Clarke in Candolle & Can-
dolle, Monogr. Phan. 5(1): 101. 1883. — Dist r.: South Indi a.
Henckelia zeylanica (R. Br.) A. Weber & B.L. Burtt in Beitr.
Biol. Pflanzen 90: 359. 1998 (‘1997’) Didymocarpus zey-
lanicus R. Br., Cyrta ndr eae: 119. 1839. — Dist r.: Sri La n ka.
Damrongia Kerr ex Craib in Bull. Misc. Inform. Kew 1918: 364.
1918 – Type: Damrongia purpureolineata Kerr ex Craib.
Perennial herbs, acaulescent. Leaves whorled, petiolate,
lamina ovate or elliptic. Inflorescences axillary, scapose,
flowers 1–10; bracts paired or ternate, free or fused along one
margin, linear to ovate. Calyx of 5 free lobes or fused into a tube
for part of length, segments triangular or narrowly triangular,
sometimes reduced to 3 lobes by fusion of upper 3 lobes. Corolla
infundibuliform; bilabiate with 2-lobed upper lip and 3-lobed
lower lip; white to deep purple, sometimes with yellow markings
in throat. Stamens 2; filaments straight or curved; anthers fused
face to face, glabrous. Stigma chiritoid, lower lobe bifid or not.
Fruit a capsule splitting into 2 valves. Seeds minute, ellipsoid.
Distribution: Thailand and Peninsular Malaysia. — Found
on limestone rocks, usually in shade. Approximately 6 species.
Damrongia was coined by Kerr but published by Craib
(1918) for the accommodation of a single species, D. pur-
pureolineata and was later transferred to Chirita by Wood
(1972). Wood did not provide any argument for his decision to
reduce the genus to a synonym of Chirita. A second species,
D. kerrii, was incorrectly attributed to the genus by Pellegrin
(1930) and is now recognised in Petrocosmea. The genus name
honours Damrong Raja Nubhap (1862–1943) of Thailand who
“has done so much for the advancement of education in his
country” (Craib, 1918: 365).
Damrongia purpureolineata belongs to a small group of
similar species restricted to Thailand and Peninsular Malaysia.
The most-widespread species is perhaps Damrongia lacunosa,
originally described as Didymocarpus lacunosus. It was tra ns-
ferred to Chirita, along with Didymocarpus cyaneus, by Burtt
(1965). Wood (1974) combined the two species under Chirita
lacunosa and suggested it was closely related to Chirita fulva,
C. integra and C. trisepala, distinguishable “from the other
species in the section [sect. Chirita] by an aca ulescent habit and
a scap ose inflore scenc e”. Th ree spe cies of the gro up have be en
included in the molecular analysis under their names in Chirita:
Chirita purpureolineata (the type species of Damrongia),
Chirita lacunosa and an as yet undescribed species, Chirita
sp. nov. 2. The clade, with all sp ecies pos sessi ng stra ig ht fr u its,
is nevertheless nested amongst the twisted-fruited genera. At
present, only the chromosome number of Damrongia lacunosa
is known with n = 9 (Kiehn & al., 1998).
Wang & al. (2011) included several Chirita samples from
Möller & al. (2009) in their analysis but, curiously, did not
include Chirita lacunosa despite it s isolat ed position in Möller
& al. (2009). They, therefore, incorrectly included Damrongia
in synonymy of their maintained genus Chirita.
Damrongia cyanea
(R idl.)
D.J. Middlet on & A. Webe r, comb.
nov.
Didymocarpus cyaneus Ridl. in J. Bot. 38: 68.
1900
Chirita cyanea (Ridl.) B.L. Burtt in
Notes Roy.
Bot. Gard. Edinburgh 26: 267. 1965.
— Distr.
: Southern
Thailand (Surat Thani).
Damrongia fulva (Barnett) D.J. Middleton & A. Weber, comb.
nov.Chirita fulva Barnett in
Nat. Hist. Bull. Siam Soc.
20: 15. 1961.
Distr.
: Southern Thailand (Nakhon Si
Thammarat, Surat Thani).
Damrongia integra (Barnett) D.J. Middleton & A. Weber, comb.
nov.Chirita integra Barnett in
Nat. Hist. Bull. Siam Soc.
20: 16. 1961.
— Distr.
: Southern Thailand (Krabi).
Damrongia lacunosa (Hook. f.) D.J. Middleton & A. Weber,
comb. nov. Didymocarpus lacunosus Hook. f. in Bot.
778
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Remodelling of Chirita
Mag. 1892: t. 7236. 1892 Chirita lacunosa ( Ho ok . f.)
B.L. Burtt in
Notes Roy. Bot. Gard. Edinburgh 26: 267.
1965.
Distr.
: Peninsular Malaysia (Pahang, Perak),
Southern Thailand (Krabi, Satun).
Damrongia purpureolineata Kerr ex Craib in Bull. Misc. In-
form. Kew 1918: 364. 1918 ≡ Chirita purpureolineata ( Ke r r
ex Craib) D. Wood in
Notes Roy. Bot. Gard. Edinburgh
31(3): 371. 1972.
— Distr.
: Northern Thailand (Lamphun).
Damrongia trisepala (Barnett) D.J. Middleton & A. Weber,
comb. nov.Chirita trisepala Barnett in
Nat. Hist. Bull.
Siam Soc. 20: 18. 1961; et in Kew Bull 15: 254. 1961.
Distr.
: Central Thailand (Chanthaburi, Nakhon Nayok).
Microchirita (C.B. Clarke) Y.Z. Wang in J. Syst. Evol. 49: 59.
2011 Chirita sect. Microchirita C.B. Clarke in Candolle
& Candolle
, Monogr. Phan. 5: 127. 1883
Roettlera sect.
Microchirita (C.B. Clarke) Fritsch in Engler & Prantl,
Nat. Pflanzenfam. IV/3b: 148. 1895 ≡ Didymocarpus se ct.
Microchirita (C.B. Clarke) Chun in Sunyatsenia 6: 290.
1946 – Type (lectotype Burtt, 1954): Microchirita hamosa
(R. Br.) Y.Z. Wang.
Annual or longer lived (flowering and fruiting for many
years in a ± aseasonal climate, dying down to a rootstock in
the dry season of more seasonal climates), caulescent, stems
often fleshy or juicy. Leaves in distant pairs, except lowermost
leaf solitary (macrocotyledon). Inflorescences usually two or
several in a leaf axil, often displaced onto the petiole, often
consisting of a short-stalked serial f lower pair only, but this
repeated several times (serial arrangement, “inflorescence
crest”). Calyx lobes free to base, narrowly triangular to nar-
rowly ovate, appressed to fruit. Corolla usually infundibuli-
form, more rarely tubular or funnel-shaped; limb bilabiate,
upper lip 2-lobed, lower lip 3-lobed, lobes rounded; colour
white, bluish, blue, purple, yellow, orange or reddish. Stamens
2, anthers usually joined by an apical ligature (projections on
the connectives). Ovary not stipitate, stigma chiritoid. Fruit
an elongate, straight or curved capsule, usually splitting into
2 valves. Seeds minute, ellipsoid; (always?) with conspicuous
knobs on the testa cells.
Distribution: Ranging from the Western Ghats of India, the
foothills of the Himalayas, through continental Southeast Asia
to Borneo, Sumatra and Java. Rock plants, on limestone,
growing in forest or on exposed rocks in the open or in light
shade. Approximately 18 species.
Microchirita is perhaps the most readily recognisable of
the genera to arise from Chirita. The highest concentration
of species is in Thailand. Chromosome numbers of n = 9, 17,
18 have been recorded (see ‘Webcyte’, Möller & al., 2002 on-
wards). All species currently known are found only on karst
li mesto ne rocks, being su n-t oler ant to so me deg ree . Wang & al.
(2011), using sequence data from Li & Wang (2007) and Möller
& al. (2009), raised this group to genus level.
Microchirita aratriformis (D. Wood) A. Weber & D.J. Midd le -
ton, comb. nov. Chirita aratriformis D. Wood in Notes
Roy. Bot. Gard. Edinburgh 31(3): 367. 1972. — Distr.:
Northern Vietnam.
Microchirita barbata (Sprague) A. Weber & D.J. Middleton,
comb. nov.Chirita barbata Sprague in Bot. Mag. 134:
t. 8200. 1908. — Distr.: Only known from cultivated mate-
rial but probably from Indochina.
Microchirita bimaculata (D. Wood) A. Weber & D.J. Mid dle-
ton, comb. nov. Chirita bimaculata D. Wood in Notes
Roy. Bot. Gard. Edinburgh 31(3): 368. 1972. — Distr.:
Thailand (Chiang Mai).
Microchirita caerulea (R. Br.) Y.Z. Wang in J. Syst. Evol. 49:
60. 2011 Chirita caerulea R. Br., Cyr t and rea e: 117. 1839
= Chirita zollingeri C.B. Clarke in Candolle & Candolle,
Monogr. Phan. 5(1): 125. 1883. — Distr.: Indonesia (Bali,
Java, Sumba).
Microchirita caliginosa (C.B. Clarke) Y.Z. Wang in J. Syst.
Evol. 49: 60. 2011 Chirita caliginosa C.B. Clar ke in Can -
dolle & Candolle, Monogr. Phan. 5(1) 122. 1883 = Chirita
fusca C.B. Clarke in Candolle & Candolle, Monogr. Phan.
5(1) 129. 1883 = Chirita parvula Ridl. in J. Fed. Malay
States Mus. 10: 149. 1920. — Distr.: Peninsular Malaysia
(Perak, Pahang, Selangor), Sarawak.
Microchirita elphinstonia (Craib) A. Web er & D.J. Midd leton ,
co mb . nov.Chirita elphinstonia Craib in Bull. Misc.
Inform. Kew 1932: 149. 1932. — Distr.: Thailand (Prachin
Bur i).
Microchirita hamosa (R. Br.) Y.Z. Wang in J. Syst. Evol. 49:
60. 2011 Chirita hamosa R. Br., Cyr tand reae: 117. 1839 =
Chirita hamosa va r. finlaysonia C.B. Clarke in Candolle &
Candolle, Monogr. Phan. 5(1) 128. 1883 = Chirita hamosa
R. Br. var. unifolia C.B. Clarke in Candolle & Candolle,
Monogr. Phan. 5(1) 129. 1883 = Chirita cristata (Dalzell)
B.L. Burtt in Notes Roy. Bot. Gard. Edinburgh 23: 96. 1960.
— Distr.: Northern and Northeastern India, China (Guangxi,
Yunnan), Burma, Thailand, Laos, Northern Vietnam.
Microchirita involucrata (Craib) Y.Z. Wang in J. Syst. Evol.
49: 60. 2011 (involucrate”) ≡ Chirita involucrata Craib in
Gard. Chron., ser. 3, 83: 140. 1928 = Chirita involucellata
Crai b in Ga rd. Chron., se r. 3, 83: 140. 1928 = Chirita capitis
Craib in Bull. Misc. Inform. Kew 1930: 173. 1930 = Chirita
geoffrayi Pellegr. in Lecomte & al., Fl. Indo-Chine 4: 529.
1930. — Distr.: Central and southern Thailand, Cambodia,
Peninsular Malaysia (Kedah, Kelantan, Penang, Perak,
Pahang).
Microchirita lavandulacea (Stapf ) Y.Z. Wang in J. Syst. Evol.
49: 60. 2011 Chirita lavandulacea Stapf in Bot. Mag.
150: t. 9047. 1925. — Distr.: China, Northern Vietnam.
Microchirita marcanii (Craib) A. Weber & D.J. Middleton,
comb. nov. Chirita marcanii Craib in Bull. Misc. In fo rm.
Kew 1926: 171. 1926. — Distr.: Thailand (Saraburi).
Microchirita micromusa (B.L. Bur t t) A. Weber & D.J. Mid dle-
ton, comb. nov. Chirita micromusa B.L. Burtt in J. Roy.
Hort. Soc. 85: 28. 1960. — Distr.: Thailand (Nakhon Nayok).
Microchirita mollissima (Ridl.) A. Weber & D.J. Middleton,
comb. nov. Chirita mollissima Ridl. in J. Li nn. Soc., Bot.
32: 517. 1896. — Distr.: Thailand (Phangnga, Surat Thani).
Microchirita oculata (Craib) A. Weber & D.J. Middleton,
comb. nov. Chirita oculata Craib in Bull. Misc. Infor m.
Kew 1930: 174. 1930. — Distr.: Thailand (Prachin Buri).
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60 (3) • June 2011: 767–790
Microchirita rupestris (Ridl .) A. Webe r & Raf ida h, com b. nov.
Chirita rupestris Rid l. in J. Stra its Bra nch Roy. Asia t. Soc.
44: 59. 1905 = Chirita glasgovii Ridl. in J. Straits Branch
Roy. Asiat. Soc. 44: 60. 1905. = Chirita kerrii Craib in Bul l.
Misc. Inform. Kew 1914: 129. 1914. — Distr.: Southern
Thailand, Peninsular Malaysia (Perak, Perlis, Kedah).
Microchirita sahyadriensis (Punekar & Lakshmin.) A. Weber
& D.J. Middleton, comb. nov. Chirita sahyadriensis
Punekar & Lakshmin. in Fol. Malaysiana 10: 18. 2009. —
Distr.: India (Western Ghats).
Microchirita sericea (Ridl.) A. Weber & Rafidah, comb. nov.
Chirita sericea Ridl. in J. Straits Branch Roy. Asiat.
Soc. 44: 59. 1905 = Chirita sericea var. scortechinii Ridl.
in J. Straits Branch Roy. Asiat. Soc. 43: 59. 1905. — Distr.:
Peninsular Malaysia (Kedah, Perak, Kelantan).
Microchirita tubulosa (Craib) A. Weber & D.J. Middleton,
comb. nov. Chirita tubulosa Cra ib in Bul l. Misc. Infor m.
Kew 1922: 173. 1922. — Distr.: Southern Thailand.
Microchirita viola (Ridl.) A. Weber & Rafidah, comb. nov.
Chirita viola Ridl., J. Linn. Soc., Bot. 32: 516. 1896. —
Distr.: S Thailand, Peninsular Malaysia (Kedah: Langkawi
Isla nds).
Liebigia Endl.,
Gen. Pl.: 1407. Feb
Mar. 1841 ≡
Tromsdorf-
fia Blume, Bijdr. Fl. Ned. Ind.: 762. 1826, non Bernhardi
(1800), nec Martius (1826)
Morstdorffia Steud., No-
mencl. Bot., ed. 2, 2: 162. Mar. 1841
Chirita sect. Liebi-
gia (Endl.) C.B. Clarke in Candolle & Ca nd ol le
, Monogr.
Phan. 5(1): 122. 1883; Hilliard in Edinburgh J. Bot. 60(3):
364. 2004 (‘2003’)
Type: Liegibia speciosa ( Blume)
Endl.
=
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.ller &
A. Weber).
Coarse perennial herbs or subshr ubs, 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. Inf lorescences axillary, sometimes
arising from small brachyblasts, bracts paired, either linear-
lanceolate and free from one another or broadly lanceolate to
suborbicula r and marg ins con tig uo us and pa r tly con nat e. Calyx
funnel-shaped or tubular, subequally 5-lobed or spathaceous
(split op en on one side). Co rolla broad -tu bed , ve nt r ic ose, wh it e
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
be ard; anthers cohering fa ce to face at tips. Ovary shor tly stipi-
tate, stigma chiritoid, lower lobe deeply bilobed (fish-tailed).
Fruit an elongate capsule splitting into 2 valves; seeds minute,
ellipsoid.
Distribution: Sumatra, Java and Bali. — Forest plants, also
in dist u rbed forest , open plac es and fore st marg ins , river bank s
etc.; probably growing in acid soil (limestone not recorded,
but ecological information generally scanty). Twelve species
in Hilliard (2004).
Liebigia is morphologically and biogeographically well
defined. The species represent large, coarse perennial herbs,
with elongate stems (to 2 m long) and scabrid leaves in distant
pairs. For a detailed description of the morphology see Hilliard
(2004) and the generic description above. Chromosome num-
bers have been reported as n = 14 (Lee, 1962, as C. blumei)
and 16 (Ratter & Prentice, 1967).
Liebigia adenonema (Hilliard) Mich. Möller & A. Weber,
comb. nov. Chirita adenonema Hilliard in
Edinburgh
J. Bot. 60: 381. 2004 (‘2003’).
— Distr.
: Indonesia: Su-
matra (North Sumatra).
Liebigia dissimilis (Hilliard) Mich. Möller & A. Weber, comb.
nov. Chirita dissimilis Hilliard in
Edinburgh J. Bot. 60:
372. 2004 (‘2003’).
— Distr.
: Indonesia: Sumatra (West
Suma tra).
Liebigia glabra (Miq.) Mich. Möller & A. Weber, comb. nov.
Chirita glabra Miq.,
Fl. Ind. Bat. 2: 729. 1858.
— Dis tr.
:
Indonesia: Sumatra (West Sumatra, Jambi).
Liebigia horsfieldii (R. Br.) Mich. Möller
& A. Weber
,
co mb . nov.
Chirita horsfieldii R. Br., Cyrtandreae:
117. 1839 = Chirita scaberrima R. Br.,
Cyrtandre a e: 117.
1839
Chirita horsfieldii R. Br. var. scaberrima (R. Br.)
C.B. Clarke in Candolle & Candolle, Monogr. Phan. 5(1):
124. 1883 =
Liebigia bracteosa Zoll. & Moritzi, Natuu r-
Geneesk. Arch. Ned.-Indië 2: 574. 1845
Chirita brac-
teosa (Zoll. & Moritzi) Miq., Fl. Ned. Ind. 2: 728. 1858.
— Distr.
: Indonesia: Java (West Java), Bali.
Liebigia leuserensis (Hilliard) Mich. Möller & A. Weber,
comb. nov. Chirita leuserensis Hilliard in
Edinburgh
J. Bot. 60: 383. 2004 (‘2003’).
— Distr.
: Indonesia: Su-
matra (North Sumatra).
Liebigia limans (Miq.) Mich. Möller & 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. Möller & A. Weber,
comb. nov.
Chirita neoforbesii Hilliard in
Edinburgh
J. Bo t. 60: 368. 20 04 (‘2003’)
Chirita blumei C.B. Cla rke
var. cordifolia S. Moore in
J. Bo t. 63 (Su ppl.): 74. 1925.
Distr.
: Indonesia: Sumatra (South Sumatra).
Liebigia polyneura (Miq.) Mich. Möller
& 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. ß? pedicellata
C.B. Clarke in Candolle & Candolle
, Monogr. Phan. 5(1):
125. 1883 =
Chirita alba Rid l. in
J. Fed. Malay States Mus.
8(4): 68. 1917 = ?Chirita glabra Miq. var. intermedia
Ridl. in J. Fed. Malay St at es Mus. 8(4): 68. 1917.
— Distr.
:
Indonesia: Sumatra (West Sumatra).
Liebigia praeterita (Hilliard) Mich. Möller & A. Weber, comb.
nov. Chirita praeterita Hilliard in
Edinburgh J. Bot. 60:
370. 2004 (‘2003’).
— Distr.
: Java.
Liebigia speciosa (Blume) Endl., Gen. Pl., Suppl. I: 1407. 1841
Tromsdorffia speciosa Blume, Bijdr. Fl. Ned. Ind.: 763.
780
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1826
Chirita blumei C.B. Clarke in Candolle & Can-
dolle,
Monogr. Phan. 5(1): 122. 1883 ≡
Chirita asperifolia
(Blume) B.L. Burtt in
Notes Roy. Bot. Gard. Edinburgh
24: 41. 1962
= Chirita spectabilis Miq.,
Fl. Ned. Ind. 3:
564. 1861
= Chirita forbesii S. Moore in
J. Bot. 37: 175.
189 9
= Chirita forbesii Ridl. in
J. St raits Branch Roy.
Asiat. Soc. 44: 60. 1905,
nom. illeg.
= Chirita ridleyana
S. Moore,
J. Bot. 63 (Suppl.): 74. 1925.
— Dist r.
: Indone-
sia: Sumatra (Benkulu, South Sumatra), Java, Krakatau
island.
Liebigia tenuipes (Hilliard) Mich. Möller & A. Weber, comb.
nov. Chirita tenuipes Hilliard in
Edinburgh J. Bot. 60:
384. 2004 (‘2003’).
— Distr.
: Indonesia: Sumatra (North
Suma tra).
Liebigia tobaensis (Hilliard) Mich. Möller & A. Weber, comb.
nov. Chirita tobaensis Hilliard in
Edinburgh J. Bot. 60:
379. 200 4 (‘2003’).
— Dis tr.
: Indonesia : Sumatra (North
Suma tra).
Primulina Hance in J. Bot. 21: 169. 1883 – Type: Primulina
tabacum Hance.
= Chirita sec t. Gibbosaccus C.B. Cla rke in Ca ndol le & Can-
dolle, Monogr. Phan. 5(1): 130. 1883 – Type: Chirita sin-
ensis Lindl. (≡
Primulina dryas ( Dun n) Mich. Möller &
A. Weber
).
= Deltocheilos W.T. Wang in Bull. Bot. Res., Harbin, 1(3): 39.
1981 – Type: Deltocheilos tenuitubum W.T. Wang (
Prim-
ulina tenuituba (W.T. Wang) Y.Z. Wang).
= Chiritopsis W.T. Wang in Bull. Bot. Res., Harbin, 1: 21.
1981 – Type: Chiritopsis repanda W.T. Wang (
Primulina
repanda (W.T. Wang) Y.Z. Wang).
= Wentsaiboea D. Fang & D.H. Qin in
Acta Phytotax. Sin.
42(6): 534. 2004 – Type:
Wentsaiboea renifolia D. Fang
& D.H. Qin (
Primulina renifolia (D. Fang & D.H. Qin)
Y.Z. Wang).
Perennial rhizomatous herbs, acaulescent, rarely with a
short woody stem, rhizome usually stout. Leaves in a basal
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).
Inf lorescences axillary, scapose, flowers in pair-flowered
cymes, often umbel-like; bracts usually present, small to con-
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 infun-
dibuliform, 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 usu-
ally 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. Approxi-
mately 131 species (but see comments below).
Primulina, now comprising Chiritopsis, Wentsaiboea (ex-
cept W. tiandengensis Yan Liu & B. Pan, a species found to be
more closely related to Lagarosolen [Möller & al., 2011] and
which will be dealt with in a forthcoming paper) and the large
number of species described in Chirita sect. Gibbosaccus, is a
genus of perennial, acaulescent rosette plants, with no excep-
tions known so far. The inflorescences are axillary and scapose.
Phyllotaxis is primarily opposite (easily seen in the type species
Primulina tabacum and P. dryas = Chirita sinensis, the type of
Chirita sect. Gibbosaccussee photos of many other species in
Li & Wang, 2004), or in whorls of three, but this may become
blurred in many-leaved rosettes or phyllotaxy may change from
opposite to alternate in the course of development. The f low-
ers are usually infundibuliform or tubular, with a slightly to
distinctly bilabiate limb. Only Primulina tabacum has salver-
form f lowers with an almost actinomorphic limb, apparently
reflecting a different pollination syndrome (see below). The
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
(in the present sense), especially in western and southern China,
but the species of the two genera seem to be ecologically well
separated, those of Primulina growing in limestone areas, those
of Henckelia not. There is no overlap with Damrongia and
Liebigia. Only one species of Microchirita, the widespread
M. hamosa, is found within the range of Primulina.
Available information suggests that Primulina is essen-
tially a calciphilous genus with typical habitats being the karst
limestone towers in southern China. Guangxi province is par-
ticularly rich in species with about half of the species described
occurring there, either as local endemics or also occurring in
other provinces such as Guizhou, Yunnan, Hunan, etc.
It may be that some of the species transferred to Primulina
here may prove misplaced in future studies. Targeted studies
are necessary to identify problematic species and eventually
to find a better place for them. It is also our impression that
too many species have been described with numerous pairs or
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,
will have been sunk into synonymy.
The relationship of Chiritopsis to Chirita (sec t. Gib-
bosaccus) 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, Möller & al., 2009 and
Wang & al., 2011) and Primulina, respectively.
Though Primulina tabacum, previously the sole repre-
sentative of Primulina, looks quite different from the species
781
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60 (3) • June 2011: 767–790
of Chirita sect. Gibbosaccus, especially in the narrow-tubed
flowers with an almost actinomorphic patent limb (a salverform
flower), the relationship is uncontroversial. The species has
a typical chiritoid stigma and the rosette habit with opposite
leaves also suggests a close relationship. In the treatments of
Wang & al. (1990, 1998) and Li & Wang (2004) Primulina is
directly associated with Chirita. The functional interpretation
of the different flower form seems rather easy. Primulina taba-
cum has appare ntly swit ch ed fr om bee- polli nat io n to butt erfly-
pollination. In their form and vivid colouration the flowers
exhibit the typical syndrome of psychophilous f lowers. Field
studies are urgently needed to confirm or reject this hypothesis.
The recently described genus Wentsaiboea was said to be
similar to Dolicholoma and Chiritopsis by Fang & Qin (2004).
The differences to the latter, now in Primulina, in the pal mat ely
nerved leaves, the campanulate corolla tube and the “obliquely
hippocrepiform” stigma would appear, from the molecular re-
sults, not to be very significant. Its position within Primulina
thus does not pose a great problem.
Despite the large number of species described, the chro-
mosome number of only one species, Chirita sinensis (which
is the type species of Chirita sect. Gibbosaccus and which is
now Primulina dryas: n = 18) has so far been publi sh ed. Fur t her
counts, con f irming th at number, have bee n made by F. Chri st ie
at the Royal Botanic Garden Edinburgh (unpub.).
Primulina annamensis (Pellegr.) Mich. Möller & A. Weber,
comb. nov. Chirita annamensis Pellegr. in Lecomte &
al., Fl. Indo-Chine 4: 530. 1930. — Distr.: Central Vietnam.
Primulina atroglandulosa (W.T. Wang) Mich. Möll er & A. We-
ber, comb. nov. Chirita atroglandulosa W.T. Wang in
Novon 7(4): 423. 1998 (‘1997’). — Distr.: China (Guangxi).
Primulina atropurpurea (W.T. Wang) Mich. Möller & A. We-
ber, comb. nov. Chirita atropurpurea W.T. Wang in Bull.
Bot. Res., Har bi n 1(4): 60. 1981. — Dist r.: China (Guangxi).
Primulina baishouensis (Y.G. Wei, H.Q. Wen & S.H. Zhong)
Y.Z. Wang in J. Syst. Evol. 49: 60. 2011 Chirita baisho-
uensis Y.G. Wei, H.Q. Wen & S.H. Zhong in Acta Phytotax.
Sin. 38(3): 299. 2000. — Distr.: China (Guangxi).
Primulina balansae (Drake) Mich. Möller & A. Weber, comb.
nov. Chirita balansae Dr ake in Bull. Annuel Soc. Philom.
Paris, sér. 8, 2: 138. 1890. — Distr.: Northern Vietnam.
Primulina bicolor (W.T. Wang) Mich. Möller & A. Weber,
comb. nov. Chirita bicolor W.T. Wang in Bull. Bot. Res .,
Harbin 5(3): 59. 1985. — Distr.: China (Guangdong).
Primulina bipinnatifida (W.T. Wang) Y.Z. Wang in J. Syst.
Evol. 49: 60. 2011 ≡ Chiritopsis bipinnatifida W.T. Wang
in Bull. Bot. Res., Harbin 1(3): 26. 1981. — Distr.: China
(Guangxi).
Primulina bogneriana (B. L. Burtt) Mich. Möller & A. Web er,
comb. nov. Chirita bogneriana B.L. Bu r tt in Gard . Bull.
Singapore 54: 239. 2002. — Distr.: Vietnam.
Primulina brachystigma ( W.T. Wan g) M i c h . l l e r & A . Webe r,
comb. nov. Chirita brachystigma W.T. Wa ng in Bull. Bot.
Res., Harbin 4(1): 22. 1984. — Distr.: China (Guangxi).
Primulina brachytricha (W.T. Wang & D.Y. Chen) Y.Z. Wang
in J. Syst. Evol. 49: 60. 2011. Chirita brachytricha
W.T. Wang & D.Y. Chen in Bull. Bot. Res., Harbin 5(3):
54. 1985. — Distr.: China (Guizhou).
Primulina brachytricha va r. magnibracteata (W.T. Wang &
D.Y. Chen) Mich. Möller & A. Weber, comb. nov. Chirita
brachytricha var. magnibracteata W.T. Wang & D.Y. Chen
in Bull. Bot. Res., Harbin 5(3): 55. 1985. — Distr.: China
(Guizhou).
Primulina brassicoides (W.T. Wang) Mich. Möller & A. We-
ber, comb. nov. Chirita brassicoides W.T. Wan g in Bull.
Bot. Res ., Harbin 1(4): 63. 1981. — Di str.: Ch ina (Gu ang xi).
Primulina brunnea (W.T. Wang) Mich. Möller & A. Weber,
comb. nov. Chirita brunnea W.T. Wang in Bull. Bot.
Res., Harbin 3(4): 48. 1983. — Distr.: Vietnam.
Primulina carnosifolia (C.Y. Wu ex H.W. Li) Y.Z. Wang in
J. Sy st . Evol. 49: 60. 2011 ≡ Chirita carnosifolia C.Y. Wu ex
H.W. Li in Bull. Bot. Res., Harbin 3(2): 36. 1983. — Distr.:
China (Yunnan).
Primulina colaniae (Pellegr.) Mich. Möller & A. Weber, comb.
nov. Chirita colaniae Pellegr. in Bull. Soc. Bot. France
73: 418. 1926. — Distr.: Northern Vietnam.
Primulina confertif lora (W.T. Wang) Mich. Möller & A. We-
ber, comb. nov. Chiritopsis confertiflora W.T. Wang
in Bull. Bot. Res., Harbin 1(3): 25. 1981. — Distr.: China
(Guangdong).
Primulina cordata Mich. Möller & A. Weber, nom. nov.
Chirita cordifolia W.T. Wang in Bull. Bot. Res., Harbin
1(4): 51. 1981. — Distr.: China (Guangxi).
Primulina cordifolia (D. Fang & W.T. Wanga) Y.Z. Wang in
J. Syst. Evol. 49: 61. 2011 ≡ Chiritopsis cordifolia D. Fang
& W.T. Wang in Bull. Bot. Re s., Ha rbin 2(4): 54. 1982, non
Chirita cordifolia W.T. Wang. — Dist r.: Chi na (Guangxi).
Primulina crassituba (W.T. Wang) Mich. Möller & A. Weber,
comb. nov. Chirita crassituba W.T. Wang in Bull. Bot.
Res., Harbin 9(4): 24. 1989. — Distr.: China (Hunan).
Primulina cruciformis (Chun) Mich. Möller & A. Weber,
comb. nov. Didymocarpus cruciformis Chu n in Sunyat-
se nia 6: 296. 1946 ≡ Chirita cruciformis (Chu n) W.T. Wang
in Bull. Bot. Res., Harbin 1(4): 45. 1981. — Distr.: China
(Hunan).
Primulina cycnostyla (B.L. Burtt) Mich. Möller & A. Weber,
comb. nov. Chirita cycnostyla B.L. Burtt in Notes Roy.
Bot. Gard. Ed inburg h 23: 96. 1960. — Di str.: Ce nt ral Viet-
nam.
Primulina cyrtocarpa (D. Fang & L. Zeng) Mich. Möller &
A. Weber, comb. nov. Chirita cyrtocarpa D. Fang &
L. Zeng in Acta Phytotax. Sin. 31(5): 468. 1993. — Distr.:
China (Guangxi).
Primulina demissa (Hance) Mich. Möller & A. Weber, comb.
nov. Didymocarpus demissus Hance in J. Bot. 21: 166.
1883 ≡ Chirita demissa (Ha nc e) W.T. Wang in Novon 7(4):
424. 1998 (‘1997’). — Distr.: China (Guangdong). — Fur-
ther studies are required to confirm its generic placement.
Primulina depressa (Hook. f.) Mich. Möller & A. Weber, comb.
nov. Chirita depressa Hook. f. in Bot. Mag. t. 7213. 1892.
— Distr.: China (Guangdong).
Primulina drakei (B.L. Burtt) Mich. ller & A. Weber,
co mb . nov.Chirita bracteosa Drake in Bull. Annuel
782
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60 (3) • June 2011: 767–790Weber & al. •
Remodelling of Chirita
Soc. Philom. Paris, sér. 8, 2: 39. 1890, nom. illeg., non
Chirita bracteosa (Zoll. & Moritzi) Miq. (1858) ≡ Chirita
drakei B.L. Burt t in Notes Roy. Bot. Gard. Edinburgh 23:
98. 1960. — Distr.: Northern Vietnam.
Primulina dryas (D unn) Mich. Möller & A. Weber, comb.
nov. Chirita dryas Dunn in Bull. Misc. Inform. Kew,
Addit. Ser. 10: 195. 1912 = Chirita sinensis Lindl. in Ed-
ward s’s Bot. Reg. 30: t. 59. 1844. Didymocarpus sinensis
(Lindl.) H. Lév. in Compt. Rend. Assoc. Franç. Avancem.
Sci. 34: 427. 1906 ≡ Primulina sinensis (Li nd l.) Y.Z. Wang
in J. Syst. Evol. 49: 62. 2011, nom. illeg., non Primulina sin-
ensis Hook. f. in Bot. Mag. 116: pl. 7117. 1890, nom. illeg. =
Chirita sinensis var. angustifolia Dunn in J. Bot. (London)
45: 403. 1907 = Chirita sinensis var. bodinieri H. Lév. in
Fed des Rep ert . 9: 328. 1911. — Distr.: Ch ina (Guangd ong).
Primulina eberhardtii (Pellegr.) Mich. Möller & A. Weber,
comb. nov. Chirita eberhardtii Pellegr. in Bull. Soc. Bot.
France 73: 418. 1926. — Distr.: Central Vietnam.
Primulina eburnea (Hance) Y.Z. Wang in J. Syst. Evol. 49: 61.
2011 ≡ Chirita eburnea Hance in J. Bot. 21: 168. 1883 =
Chirita fauriei Franch. in Bull. Mens. Soc. Linn. Paris 1:
45. 1885. — Distr.: China (Guangdong, Guangxi, Guizhou,
Hubei, Hunan, Sichuan).
Primulina fangii (W.T. Wang) Mich. Möller & A. Weber,
comb. nov. Chirita fangii W.T. Wang in Bull. Bot. Res.,
Harbin 2(4): 51. 1982. — Distr.: China (Sichuan).
Primulina fimbrisepala (Hand.-Mazz.) Y.Z. Wang in J. Syst.
Evol. 49: 61. 2011 Chirita fimbrisepala Hand.-Mazz.
in Anz. Akad. Wiss. Wien, Math.-Naturwiss. Kl. 62: 65.
1925. — Distr.: China (Fujian, Guangdong, Guangxi,
Guizhou, Hunan, Jiangxi).
Primulina fimbrisepala var. mollis (W.T. Wang) Mich. Möller
& A. Weber, comb. nov. Chirita fimbrisepala Hand.-
Mazz. var. mollis W.T. Wang in Bull. Bot