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Abstract

Based on a phylogenetic analysis of Asian Gesneriaceae with the most comprehensive coverage at the genus level to date, the new genus Chayamaritia is established and described in subfamily Didymocarpoideae, tribe Trichosporeae, subtribe Didymocarpinae. It contains two species, of which one, Chayamaritia smitinandii (B.L.Burtt) D.J.Middleton, was formerly placed in the genera Chirita and Henckelia. The other, Chayamaritia banksiae D.J.Middleton, is newly described. The exclusion of Chayamaritia smitinandii from Henckelia further clarifies the taxonomic and biogeographic limits of Henckelia following its considerable recircumscription during the recent remodelling and synonymisation of Chirita.
1 23
Plant Systematics and Evolution
ISSN 0378-2697
Volume 301
Number 7
Plant Syst Evol (2015) 301:1947-1966
DOI 10.1007/s00606-015-1213-2
Chayamaritia (Gesneriaceae:
Didymocarpoideae), a new genus from
Southeast Asia
David J.Middleton, Kanae Nishii,
Carmen Puglisi, Laura L.Forrest &
Michael Möller
1 23
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ORIGINAL ARTICLE
Chayamaritia (Gesneriaceae: Didymocarpoideae), a new genus
from Southeast Asia
David J. Middleton
1
Kanae Nishii
2
Carmen Puglisi
2
Laura L. Forrest
2
Michael Mo
¨ller
2
Received: 8 July 2014 / Accepted: 16 March 2015 / Published online: 30 April 2015
Springer-Verlag Wien 2015
Abstract Based on a phylogenetic analysis of Asian
Gesneriaceae with the most comprehensive coverage at the
genus level to date, the new genus Chayamaritia is estab-
lished and described in subfamily Didymocarpoideae, tribe
Trichosporeae, subtribe Didymocarpinae. It contains two
species, of which one, Chayamaritia smitinandii
(B.L.Burtt) D.J.Middleton, was formerly placed in the
genera Chirita and Henckelia. The other, Chayamaritia
banksiae D.J.Middleton, is newly described. The exclusion
of Chayamaritia smitinandii from Henckelia further clari-
fies the taxonomic and biogeographic limits of Henckelia
following its considerable recircumscription during the
recent remodelling and synonymisation of Chirita.
Keywords Biogeography Chayamaritia
Gesneriaceae New genus Molecular phylogeny
Southeast Asia
Introduction
The genus Chirita Buch.-Ham. ex D.Don was remodelled
and split into five genera by Weber et al. (2011a). Chirita
included a heterogeneous assemblage of species that was
united by the possession of the so-called ‘chiritoid stigma’.
The chiritoid stigma is where the upper lobe of the stigma is
not developed and the lower lobe is enlarged and often bifid
(see Wu and Raven 2000, p 326). Despite a very diverse
range in other characters, the possession of a chiritoid stigma
was considered diagnostic for Chirita until Mo
¨ller et al.
(2009,2011) demonstrated that the included species did not
form a monophyletic group. The species were divided be-
tween redefined genera by Weber et al. (2011a) who also
noted that there was a number of species for which the correct
generic placement in the new system was problematic. This
was largely due to these species not having been included in
the molecular phylogenetic analyses and because the avail-
able herbarium material was rather sparse and/or poor to
properly study the morphology. One such highlighted spe-
cies was Chirita smitinandii B.L.Burtt. This species was
moved into Henckelia Spreng., even though its relationships
were not well understood, largely because no species could
be left behind in Chirita when this genus was synonymised
under Henckelia. Since Weber et al. (2011a) we have had the
opportunity to collect new material of Henckelia smitinandii
(B.L.Burtt) D.J.Middleton & Mich.Mo
¨ller and study it much
more closely, morphologically and with DNA sequence data.
This paper presents our conclusions on this species and a new
species clearly related to it.
Handling editor: Yunpeng Zhao.
Electronic supplementary material The online version of this
article (doi:10.1007/s00606-015-1213-2) contains supplementary
material, which is available to authorized users.
&David J. Middleton
David_MIDDLETON@nparks.gov.sg
Michael Mo
¨ller
m.moeller@rbge.ac.uk
1
Singapore Botanic Gardens, National Parks Board, 1 Cluny
Road, 259569 Singapore, Singapore
2
Royal Botanic Garden Edinburgh, 20A Inverleith Row,
EH3 5LR Edinburgh, Scotland, UK
123
Plant Syst Evol (2015) 301:1947–1966
DOI 10.1007/s00606-015-1213-2
Author's personal copy
Materials and methods
Plant material
Herbarium material has been studied from the following
herbaria: BKF, CMU, E, K, L, P, QBG [abbreviations from
Thiers (continuously updated)].
For the phylogenetic analyses, the extensive Old World
Gesneriaceae matrix of Middleton et al. (2014) with 225
samples of tribe Trichosporeae (Weber et al. 2013), cov-
ering 52 out of 67 genera recognized in the tribe at present
(Mo
¨ller et al. 2011,2014; Weber et al. 2013; Middleton
et al. 2014), was extended by two Tribounia D.J.Middleton
samples, four Didissandra C.B.Clarke samples, two sam-
ples of Henckelia smitinandii and one sample of an unde-
scribed but similar species (Table 1). Not included were
the African genera sensu Weber et al. (2013), and the Asian
Deinostigma W.T.Wang, Championia Gardner (for both of
which no material was available) and Beccarinda Kuntze.
Beccarinda and Championia likely belong to the basal
lineages of tribe Trichosporeae, in subtribe Leptoboeinae
(Weber et al. 2013). The trees were rooted on Tetraphyllum
Griff. ex C.B.Clarke (Mo
¨ller et al. 2009).
This wide sampling across tribe Trichosporeae was
deemed prudent to correctly place the three samples under
investigation within the tribe, since the morphological
characteristics of the Henckelia smitinandii group do not
suggest a placement near or in any existing Asian genus.
Phylogenetic analysis
The sequences of the nuclear ribosomal internal transcribed
spacers (ITS) and plastid trnL-F intron-spacer (trnL-F) for
the Didissandra and Henckelia smitinandii group samples
were acquired as described in Weber et al. (2011a). The
newly acquired sequences have been submitted to Gen-
Bank (Table 1) and sequence matrices and tree files to
TreeBase (http://purl.org/phylo/treebase/phylows/study/TB2:
S17032).
Combinability tests of the two datasets, trnL-F and ITS,
for phylogenetic analyses, and calculation of branch sup-
port were carried out as in Weber et al. (2011a), with the
difference in sampling frequency in the Bayesian inference
(BI) analysis (every 1000th generation) and the burn-in
(250,000 generations, 5 %).
Results
The combined ITS and trnL-F matrix was 2209 characters
long (1000 for ITS, 1209 for trnL-F). Of these, 66 had to be
removed from the beginning of the plastid matrix due to
sequencing artefacts, and 145 characters had to be excluded
from the ITS matrix because of alignment ambiguities in
hypervariable regions. Of the remaining 1998 characters,
939 were constant, 292 autapomorphic and 767 (38.4 %)
parsimony informative. The two matrices were analysed
together because the partition homogeneity test (PHT,
P=0.39) indicated no incongruence in phylogenetic sig-
nal between the two data sets.
The parsimony analysis resulted in 35,844 parsimonious
trees (length =6712 steps, consistency index =0.2869,
retention index =0.6763). The characteristics of the
Bayesian inference analysis indicated a good convergence
of the two independent runs (Online Resources 1–5). There
were differences in topology between the parsimony tree
and Bayesian inference tree (Online Resources 6–8).
However, these concern areas which received little or no
branch support.
The three Henckelia smitinandii group samples formed a
strongly supported clade (PP =1; BS =100 %) (Fig. 1)
that is quite distant from the rest of Henckelia and not
nested within any other genera. We, therefore, recognise
them here as the new genus Chayamaritia, and the samples
are labelled as such in Fig. 1. The genus is placed in
subtribe Didymocarpinae, as well supported sister to
Petrocosmea Oliv. (PP =1; BS =82 %). The clades of
these two genera have long stems, indicating that the
genera represent distinct units (Online Resources 6, 8). The
two genera together are sister to Allostigma W.T.Wang
(PP =1; BS =69 %) and these together sister (PP =96;
BS B50 %) to a clade comprised of Pseudochirita
W.T.Wang and Loxostigma C.B.Clarke (PP =1;
BS =73 %). The latter clade relationship, however, was
not retrieved in the MP analysis (Online Resource 7).
However, the sister relationship of the clade that includes
Chayamaritia (and Petrocosmea) to a clade including the
six genera Glabrella Mich.Mo
¨ller & W.H.Chen, Brig-
gsiopsis K.Y.Pan, Anna Pellegr., Raphiocarpus Chun,
Lysionotus D.Don and Hemiboea C.B.Clarke was identical
between the two phylogenetic analyses, including almost
all relationships amongst the six genera. All of these, ex-
cept for the two Glabrella species, are caulescent.
Discussion
We have assembled the largest dataset at the genus level to
date for the family Gesneriaceae to investigate the position
of a morphologically distinct lineage that was suspected to
represent a new evolutionary unit. Indeed, our results
showed that the samples formed a separate lineage and we,
therefore, establish a new genus: Chayamaritia. The phy-
logenetic position of Chayamaritia in the molecular phy-
logeny is interesting, since most members of this clade
combine a caulescent habit with an opposite decussate leaf
1948 D. J. Middleton et al.
123
Author's personal copy
arrangement, except Chayamaritia and Petrocosmea.
Chayamaritia, with its thick, short and prostrate stem and
alternate leaf arrangement, could be seen as a transition
from a caulescent habit to the rosette forms of Petrocos-
mea. Several species of Petrocosmea sect. Deinanthera
W.T.Wang have long-petioled and pubescent leaves
Fig. 1 Bayesian inference tree with genera collapsed (based on tree
Online Resource 8), except Chayamaritia, based on combined
analysis of ITS and trnL-F data. Numbers along branches are
posterior probabilities (italics) and parsimony bootstrap values (bold).
Asterisk indicates values \50 %
Chayamaritia, a new genus from Southeast Asia 1949
123
Author's personal copy
Table 1 List of the 234 Gesneriaceae samples of tribe Trichosporeae included in the phylogenetic analysis, including voucher number and deposition, origin information and respective
GenBank accession numbers
Taxon Voucher number Deposited
in
Origin trnL-F ITS or
ITS1/ITS2
Aeschynanthus bracteatus Wall. ex A.DC. Y.Z. Wang 991113 PE China, Yunnan, Xichou county FJ501501
Aeschynanthus bracteatus Wall. ex A.DC. R. Cherry 123 [Cult. RBGE
19970165]
E Vietnam; Lao Cai AF349203/AF349284
Aeschynanthus lancilimbus W.T.Wang Y.Z. Wang S-10868 PE China, unknown locality FJ501499 HQ632992
Aeschynanthus micranthus C.B.Clarke M. Mo¨ller and Y.D. Qi MMO
01-79
E, PE, WU China, Yunnan, Hekou county FJ501500
Aeschynanthus micranthus C.B.Clarke A. Reid and J. Fernie 004
[Cult. RBGE 19951561]
E China, Yunnan, Xishuangbanna Dai
Aut. Pref.
– AF349218/AF349299
Aeschynanthus rhododendron Ridl. P. Woods 600 [Cult. RBGE
19680624]
E Peninsular Malaysia, Genting
Highlands
HQ632895 FJ501333
Aeschynanthus roseoflorus Mendum G. Argent 87/14 E Indonesia, Seram HQ632896 HQ632993
Agalmyla biflora (Elmer) Hilliard &
B.L.Burtt
RBGE-PNH1998 25435
[Cult. RBGE 19980287]
E Philippines, Palawan, Near summit
of Cleopatra Needle
FJ501541 –
Agalmyla biflora (Elmer) Hilliard &
B.L.Burtt
RBGE-PNH1998-25517
[Cult. RBGE 19980292]
E Philippines, Palawan, near Thumb
Peak
– FJ501361
Agalmyla bilirana Hilliard & B.L.Burtt RBGE-PNHE 1999 12 E Philippines, Leyte Island HQ632891 HQ632988
Agalmyla clarkei (Elmer) B.L.Burtt RBGE-PNH1999(P99) 13
[Cult. RBGE 19991911]
E Philippines, Leyte Island, Mt. Lobi FJ501540
Agalmyla clarkei (Elmer) B.L.Burtt RBGE-PNH1997 IS26 [Cult.
RBGE 19972530A]
E Philippines, Luzon, Barangay
Penicuason
– FJ501360
Agalmyla glabra (Merr.) Hilliard &
B.L.Burtt
RBGE-PNHE 1999 28 E Philippines, Camiguin Island HQ632892 HQ632989
Agalmyla paucipilosa Hilliard & B.L.Burtt P. Smith and L. Galloway 261 E Indonesia, Sulawesi, Mt Rantemario HQ632893 HQ632990
Agalmyla sojoliana Hilliard & B.L.Burtt P. Smith and L. Galloway 321 E Indonesia, Sulawesi, Mt Sojol HQ632894 HQ632991
Allocheilos guangxiensis H.Q.Wen,
Y.G.Wei & S.H.Zhong
Y.G. Wei 06-02 IBK China, Guangxi, Yongfu county HQ632897 HQ632994
Allostigma guangxiense W.T.Wang M. Mo¨ller and Y.G. Wei
MMO 05-755
E, IBK China, Guangxi, Longzhou county HQ632880 HQ632977
Anna mollifolia (W.T.Wang) W.T.Wang &
K.Y.Pan
M. Mo¨ller and Y.D. Qi MMO
01-146
E, PE, WU China, Guangxi, Napo county FJ501543 AF055050/AF055051
Anna ophiorrhizoides (Hemsl.) B.L.Burtt &
R.A.Davidson
M. Mo¨ller and Y.G. Wei
MMO 08-1280
E, IBK China, Sichuan, Emei Shan HQ632937 HQ633034
Anna submontana Pellegr. M. Mo¨ller and Y.D. Qi MMO
01-85
E, PE, WU China, Yunnan, Maguan county FJ501542 FJ501362
Billolivia longipetiolata D.J.Middleton &
Luu
L.H. Truong and P.H. Nhan
BD624
E Vietnam, Lam Dong Province,
Bidoup–Nui Ba National Park
Middleton et al.
(2014)
Middleton et al.
(2014) (ITS1 only)
Billolivia minutiflora D.J.Middleton &
H.Atkins
L.N. Sam LY498 E Vietnam, Lam Dong Province, Da
Hoai District
Middleton et al.
(2014)
Middleton et al.
(2014) (ITS1 only)
Billolivia vietnamensis D.J.Middleton & Luu L.H. Truong and N.Q. Dat
BGM1601
E Vietnam, Binh Phuoc, Bu Gia Map
National Park
Middleton et al.
(2014)
(only trnL-F spacer)
Middleton et al.
(2014) (ITS1 only)
1950 D. J. Middleton et al.
123
Author's personal copy
Table 1 continued
Taxon Voucher number Deposited
in
Origin trnL-F ITS or
ITS1/ITS2
Billolivia violacea D.J.Middleton &
H.Atkins
D.J. Middleton 4210 E Vietnam, Lam Dong, Duc Trong
District
Middleton et al.
(2014)
Middleton et al.
(2014)
Boea hygrometrica (Bunge) R.Br. Z.J. Gu 01-6184 KUN China, unknown locality FJ501476 FJ501319
Boea philippensis C.B.Clarke S. Scott 02-142 E Indonesia, Sulawesi, Gunung Ali HQ632862 HQ632953
Boeica ferruginea Drake M. Mo¨ller and Y.D. Qi MMO
01-182B ex Z.C. Qin
200012
E, PE, WU China, SE Yunnan FJ501440 Wei et al. (2010)
Boeica multinervia K.Y.Pan Y.Z. Wang 015 PE China, Yunnan, Yingjiang HQ632861 HQ632951
Briggsiopsis delavayi (Franch.) K.Y.Pan W. Fang 1 IBK China, Chongqing, Nanchuan county HQ632879 HQ632976
Cathayanthe biflora Chun M. Mo¨ller and Y.G. Wei
MMO 08-1327
E, IBK China, Hainan, Tongshi county HQ632899 HQ632996
Codonoboea albomarginata (Hemsl.) Kiew A. Weber 840805-1/12 WU Peninsular Malaysia, Perak,
Maxwell’s Hill
AJ492297 HQ632961
Chayamaritia banksiae D.J.Middleton D.J. Middleton 5220 and M.
Newman et al. LAO1428
E Laos, Khammouan, Nakai Nam
Theun
KP325433 KP325426
Chayamaritia smitinandii (B.L.Burtt)
D.J.Middleton & Mich.Mo
¨ller
D.J. Middleton et al. 5632 E Thailand, Nakhon Nayok, Khao Yai
NP
KP325431 KP325424
Chayamaritia smitinandii (B.L.Burtt)
D.J.Middleton & Mich.Mo
¨ller
D.J. Middleton et al. 5652 E Thailand, Nakhon Nayok, Khao Yai
NP
KP325432 KP325425
Codonoboea codonion (Kiew) C.L.Lim C.L. Lim FRI 65040 KEP Malaysia, Terengganu, Jerangau F.R. JF912538 JF912565
Codonoboea corrugata (Mendum)
D.J.Middleton
RBGE-PNHE 1998s.n. E Philippines, Palawan FJ501484 HQ632962
Codonoboea elata (Ridl.) Rafidah A.R. Rafidah FRI 64321 KEP Malaysia, Perak, Maxwell’s Hill JF912523 JF912550
Codonoboea floribunda (M.R.Hend.)
C.L.Lim
C.L. Lim FRI 64971 KEP Malaysia, Terengganu, Sg. Nipah
F.R.
JF912539 JF912566
Codonoboea leucocodon (Ridl.) Ridl. C.L. Lim FRI 64821 KEP Malaysia, Pahang, Gunung Tahan JF912540 JF912567
Codonoboea malayana (Hook.f.) Kiew R. Kiew and D.J. Middleton
FRI 57513
KEP Malaysia, Pahang, Fraser’s Hill JF912541 JF912568
Codonoboea pumila (Ridl.) C.L.Lim T.L. Yao FRI 55963 KEP Malaysia, Pahang, Fraser’s Hill JF912543 JF912570
Codonoboea racemosa (Jack) A.Weber P.S. Smith SMTSU 110/110 E Indonesia, Sumatra, Aceh JF912544 JF912571
Codonoboea venusta (Ridl.) Kiew R. Kiew RK 5430 KEP Malaysia, Fraser’s Hill JF912545 JF912572
Conandron ramondioides Siebold and Zucc. Takeda Herbal Garden Kyoto
[Cult. RBGE 19691267]
E Japan, unknown locality FJ501515 FJ501340
Cyrtandra cumingii C.B.Clarke G. Kokubugata 11134 TNS Japan, Ruykyus, Iriomote Island HQ632905 HQ633002
Cyrtandra cupulata Ridl. A. Weber 840806-2/4 WU Pensinsular Malaysia, Perak,
Maxwell’s Hill
FJ501532 AY818826/AY818861
Cyrtandra glabra Banks ex C.F.Gaertn. Q.C.B. Cronk and D. Percy
T91
E French Polynesia: Society Is.: Tahiti:
Mt. Tearoa Col
AY423136 FJ501353
Cyrtandra kusaimontana Hosok. NTBG 960873 PTBG Federated States of Micronesia,
Caroline Islands
HQ632907 –
Cyrtandra kusaimontana Hosok. Flynn 5995 PTBG Federated States of Micronesia,
Caroline Islands
– EU919945
Chayamaritia, a new genus from Southeast Asia 1951
123
Author's personal copy
Table 1 continued
Taxon Voucher number Deposited
in
Origin trnL-F ITS or
ITS1/ITS2
Cyrtandra longifolia (Wawra) Hillebr. ex
C.B.Clarke
M. Kiehn 920825-2/1 [Cult.
HBV]
WU USA, Hawaii, Kauai FJ501531 EU919939
Cyrtandra pendula Blume A. Weber and S. Anthonysamy
860730-1/2 [Cult. HBV]
WU Peninsular Malaysia, Negeri
Sembilan, Kuala Pilah distr., Jeram
Toi
FJ501530 FJ501354
Cyrtandra pulchella O.Rich ex A.Gray Lorence 8525 PTBG Samoa Islands HQ632906 EU919941
Damrongia fulva (Barnett) D.J.Middleton &
A.Weber
P. Triboun s.n. BK Thailand, Khampaeng Phet JF912536 JF912563
Damrongia lacunosa (Hook.f.)
D.J.Middleton & A.Weber 1
K. Imin et al. FRI 63238 KEP Peninsular Malaysia, Perak,
Temengor F.R., Pulau Batu Putih
JF912530 JF912557
Damrongia lacunosa (Hook.f.)
D.J.Middleton & A.Weber 2
A. Weber 870510-1/8 WU Peninsular Malaysia, Pahang, Lipis
distr., Gua Rusa
FJ501458 FJ501308
Damrongia purpureolineata Kerr ex Craib 1 P. Triboun s.n. BK Thailand, Lamphun, Li JF912534 JF912561
Damrongia purpureolineata Kerr ex Craib 2 D.J. Middleton et al. 4812 E Thailand, Lamphun, Li JF912535 JF912562
Didissandra elongata ssp. minor (Ridl.)
A.Weber & B.L.Burtt
C. Puglisi, M. Hughes, D.
Girmansyah Suryadi 186
E Indonesia, Sumatra, Bengkulu,
Gunung Kemumu
KP325427 KP325420
Didissandra frutescens (Jack) C.B.Clarke A. Weber 840805-1/2 WU Peninsular Malaysia, Perak,
Maxwell’s Hill
FJ501522 –
Didissandra frutescens (Jack) C.B.Clarke A.R. Rafidah FRI 64355 KEP Peninsular Malaysia, Perak, Kuala
Kangsar
– HQ632952
Didissandra sp. C. Puglisi, M. Hughes, D.
Girmansyah Roki 69
E Indonesia, West Sumatra, road to
Melampah
KP325428 KP325421
Didissandra sp.12 C. Puglisi, M. Hughes, D.
Girmansyah Roki 130
E Indonesia, West Sumatra, Solok
Ambah, Sijunjung
KP325429 KP325422
Didissandra sp.16 C.Puglisi, M. Hughes,
D.Girmansyah Roki 101
E Indonesia, West Sumatra, Bukit
Gagoan
KP325430 KP325423
Didymocarpus antirrhinoides A.Weber K. Jong 9009 [Cult. RBGE
19650167]
E Peninsular Malaysia, Perak, Bujong
Melaka, Ipoh.
FJ501513 DQ912671
Didymocarpus citrinus Ridl. P. Davis 69437 [Cult. RBGE
19830510]
E Peninsular Malaysia, Perlis, Kedah
Peak
AJ492293 DQ912669
Didymocarpus cordatus Wall. ex A.DC. A. Weber 860816-2/1 WU Peninsular Malaysia, Perak,
Maxwell’s Hill
AJ492294 DQ912673
Didymocarpus podocarpus C.B.Clarke H. Noltie, Pradhan, Sherub
and Wangdi 193
E Bhutan, Deothang District FJ501514 DQ912688
Didymocarpus purpureobracteatus
W.W.Sm.
Y.Z. Wang 991106 PE China, Yunnan, Pingbian county FJ501510
Didymocarpus purpureobracteatus
W.W.Sm.
M. Mo¨ller and Y.D. Qi MMO
01-70
E, PE, WU China: Yunnan, Pingbian county DQ912676
Didymocarpus stenanthos C.B.Clarke M. Mo¨ller and Y.D. Qi MMO
01-156
E, PE, WU China, Yunnan, Binchuan county FJ501512 DQ912687
Didymocarpus villosus D.Don B. Adhikari SB 9 E Nepal, Sundarijal HQ632904 HQ633001
1952 D. J. Middleton et al.
123
Author's personal copy
Table 1 continued
Taxon Voucher number Deposited
in
Origin trnL-F ITS or
ITS1/ITS2
Didymostigma obtusum (C.B.Clarke)
W.T.Wang
M. Mo¨ller and Y.G. Wei
MMO 08-1310
E, IBK China, Guangdong, Fengkai county HQ632875 HQ632971
Didymostigma trichanthera C.X.Ye &
X.G.Shi
M. Mo¨ller and Y.G. Wei
MMO 08-1335
E, IBK China, Guangdong, Longmen county HQ632876 HQ632972
Emarhendia bettiana (M.R.Hend.) Kiew,
A.Weber & B.L.Burtt
R. Kiew FRI 55716 KEP Peninsular Malaysia HQ632864 HQ632955
Glabrella longipes (Hemsl. ex Oliv.)
Mich.Mo
¨ller & W.H.Chen
M. Mo¨ller and Y.D. Qi MMO
01-122
E, PE, WU China, Yunnan, Xichou county FJ501545 AF055052/AF055053
Glabrella mihieri (Franch.) Mich.Mo
¨ller &
W.H.Chen
Y.Z. Wang 11315B PE China, Chongqing, Nanchuan county FJ501544 FJ501363
Gyrocheilos chorisepalus W.T.Wang var.
synsepalus W.T.Wang
Y.G. Wei 07-708 IBK China, Guangdong, Xinyi county HQ632900 HQ632997
Gyrocheilos lasiocalyx W.T.Wang M. Mo¨ller and Y.G. Wei
MMO 06-881
E China, Guangxi, Guiping county HQ632901 HQ632998
Gyrocheilos retrotrichus W.T.Wang M. Mo¨ller and Y.G. Wei
MMO 07-1136
E, IBK China, Guangxi, Wuming county HQ632902 HQ632999
Gyrocheilos retrotrichus W.T.Wang var.
oligolobus W.T.Wang
Y.G. Wei 06-208 IBK China, Guangxi, Rongshui county,
Sirong town
HQ632903 HQ633000
Haberlea rhodopensis Friv. Voucher from Cult. RBGE
19754106
E (Greece) AJ492296 Mo
¨ller and Cronk (2001)
Hemiboea bicornuta (Hayata) Ohwi Voucher from Cult. RBGE
19951207
E unknown origin FJ501534 FJ501356
Hemiboea cavaleriei H.Le
´v. Z.J. Gu G3 KUN China, unknown locality FJ501533 FJ501355
Hemiboea fangii Chun ex Z.Yu Li M. Mo¨ ller and Y.G. Wei
MMO 08-1284
E, IBK China, Sichuan, Emei Shan HQ632882 HQ632979
Hemiboea follicularis C.B.Clarke Y.G. Wei G03 IBK China, Guangxi, Huanjiang county HQ632885 HQ632982
Hemiboea gracilis Franch. Y.Z. Wang 11317 PE China, Chongqing, Nanchuan county FJ501536 Wei et al. (2010)
Hemiboea longgangensis Z.Yu Li Y.G. Wei 07-550 IBK China, Guangxi Longzhou county HQ632889 HQ632986
Hemiboea longzhouensis W.T.Wang M. Mo¨ller and Y.G. Wei
MMO 07-1127
E, IBK China, Guangxi, Longan county HQ632888 HQ632985
Hemiboea magnibracteata Y.G.Wei &
H.Q.Wen
M. Mo¨ller and Y.G. Wei
MMO 08-1347
E, IBK China, Guangxi, introduced from
Huanjiang county to Guilin city
HQ632887 HQ632984
Hemiboea omeiensis W.T.Wang M. Mo¨ller and Y.G. Wei
MMO 08-1271
E, IBK China, Sichuan, Emei Shan HQ632886 HQ632983
Hemiboea ovalifolia (W.T.Wang) A.Weber
& Mich.Mo
¨ller
B.M. Nong 06-1IBK China, Guangxi, Napo county,
Nonghua
HQ632883 HQ632980
Hemiboea purpureotincta (W.T.Wang)
A.Weber & Mich.Mo
¨ller
M. Mo¨ller and Y.G. Wei
MMO 06-813
E, IBK China, Guangxi, Tian Ling county HQ632884 HQ632981
Hemiboea rubribracteata Z.Yu Li & Yan
Liu
M. Mo¨ller and Y.G. Wei
MMO 07-1093
E, IBK China, Guangxi, introduced from
Jingxi to Guilin city
HQ632890 HQ632987
Hemiboea subcapitata C.B.Clarke Y.Z. Wang 11306 PE China, Chongqing, Chengkou county FJ501535 FJ501357
Henckelia anachoreta (Hance)
D.J.Middleton & Mich.Mo
¨ller
D.J. Middleton et al. 4480 E Thailand, Chiang Mai, Doi Suthep HQ632870 HQ632966
Chayamaritia, a new genus from Southeast Asia 1953
123
Author's personal copy
Table 1 continued
Taxon Voucher number Deposited
in
Origin trnL-F ITS or
ITS1/ITS2
Henckelia bifolia (D.Don) A.Dietr. B. Adhikari L2B6 E Nepal, Chyalding, near Sybrubesi JF912522 JF912549
Henckelia dielsii (Borza) D.J.Middleton &
Mich.Mo
¨ller
M. Mo¨ller et al. MMO 08-
1211
E, KUN China, Yunnan, Jingdong county HQ632871 HQ632967
Henckelia floccosa (Thwaites) A.Weber &
B.L.Burtt
C.G. Jang s.n. [G 157] WU Sri Lanka FJ501486 HQ632964
Henckelia grandifolia A.Dietr. M. Mo¨ller et al. MMO 08-
1222
E, KUN China, Yunnan, Jingdong county JF912527 JF912554
Henckelia incana (Vahl) Spreng. S. Vogel SVG s.n. E India, Nilghiri mts HQ632869 HQ632965
Henckelia longisepala (H.W.Li)
D.J.Middleton & Mich.Mo
¨ller
Y.M. Shui 73170 KUN China, Yunnan, Jinping county HQ632890 HQ632963
Henckelia pumila (D.Don) A.Dietr. 1 D.J. Middleton et al. 4505 E Thailand, Chiang Mai, Doi Inthanon JF912529 JF912556
Henckelia pumila (D.Don) A.Dietr. 2 Gaoligong Shan Expedition
1996 7938 [Cult. RBGE
19962271]
E China, Yunnan, Nujiang Lisu Aut.
Pref., Fugong county
FJ501491 FJ501327
Henckelia urticifolia (D.Don) A.Dietr. 1 J.M. Li 05851 PE China, Yunnan DQ872821 DQ872835
Henckelia urticifolia (D.Don) A.Dietr. 2 NPSW 110 E Bhutan, Tashigang distr. JF912532 JF912559
Henckelia urticifolia (D.Don) A.Dietr. 3 EMAK 109 H (Edinburgh-
Makalu Expedition 1991)
E Nepal, Sankhuwasabha distr., Arun
valley
FJ501492 FJ501328
Henckelia walkerae (Gardner)
D.J.Middleton & Mich.Mo
¨ller
L. Skog 7736 (US 590934)
[Cult. Smithsonian 94-250]
US Sri Lanka; leg. in US 11.03.1996 FJ501490 FJ501326
Hexatheca fulva C.B.Clarke J. Sang and C. Geri S99358 E Sarawak, Bau, Fairy Cave HQ632873 HQ632969
Jancaea heldreichii Boiss. E.G. Cairns s.n. [Cult. RBGE
19771605]
photo E Greece, Mt Olympus FJ501439 Mo
¨ller et al. (1999)
Kaisupeea herbacea (C.B.Clarke) B.L.Burtt K. Larsen 44272 [Cult.
RBGE 19972918]
E Thailand, Chachoengsao, Khao Tak
Groep
FJ501459 FJ501309
Leptoboea multiflora (C.B.Clarke) Gamble
subsp. grandifolia B.L.Burtt
K. Larsen et al. 32065 E Thailand, Chanthaburi, Khaso Phra
Bat
FJ501442 Wei et al. (2010)
Liebigia barbata (Jack) D.J.Middleton P. Woods 1071 (C6570) E Indonesia, Java, forest above
Cibodas Garden
FJ501538 JF501359
Litostigma coriaceifolium Y.G.Wei, F.Wen
& M.Mo
¨ller
Y.G. Wei MMO 07-1162B E, IBK China, Guizhou, Xingyi county Wei et al. (2010) Wei et al. (2010)
Litostigma crystallinum Y.M.Shui &
W.H.Chen
Y.M. Shui 43865 KUN China, Yunnan, Malipo county Wei et al. (2010) Wei et al. (2010)
Loxocarpus argenteus B.L.Burtt T.L. Yao FRI 57975 KEP Malaysia, Sarawak, Bako National
Park
JF912537 JF912564
Loxocarpus violoides (C.B.Clarke) T.L.Yao T.L. Yao FRI 65458 KEP Malaysia, Sayap, Kinabalu Park JF912546 JF912573
Loxostigma fimbrisepalum K.Y.Pan Y.Z. Wang 991005 PE China, Yunnan, Jinping county FJ501507 Wei et al. (2010)
Loxostigma glabrifolium D.Fang & K.Y.Pan Y.G. Wei 709 IBK China, Guangxi, Napo county HQ632910 HQ633006
Loxostigma griffithii (Wight) C.B.Clarke Kew/Edinburgh
Kanchenjunga Expedition
(1989) 940 [Cult. RBGE
19892473A]
E Nepal, Yamphudin FJ501508 FJ501338
1954 D. J. Middleton et al.
123
Author's personal copy
Table 1 continued
Taxon Voucher number Deposited
in
Origin trnL-F ITS or
ITS1/ITS2
Loxostigma sp. Gaoligong Shan Expedition
1996 7668
E China, Yunnan AY423137 HQ633005
Lysionotus chingii Chun ex W.T.Wang Y.Z. Wang S-10669 PE China, unknown locality FJ501498 FJ501332
Lysionotus forrestii W.W.Sm. Gaoligong Shan Expedition
1996 7925 [Cult. RBGE
19962269A]
E China, Yunnan, Nujiang Lisu Aut.
Pref.,
FJ501495 AF349152/AF349233
Lysionotus pauciflorus Maxim. M. Mo¨ller and Y.D. Qi MMO
01-101
E, PE, WU China, Yunnan, Xichou county,
Cheng Jia Po
FJ501497 FJ501331
Lysionotus petelotii Pellegr. M. Mo¨ller and Y.D. Qi MMO
01-100/4
E, PE China, Yunnan, road to Xichou FJ501496 HQ632974
Metapetrocosmea peltata (Merr. & Chun)
W.T.Wang
Y.G. Wei 07-702 IBK China, Hainan,Wuzhi Shan HQ632872 HQ632968
Microchirita caliginosa (C.B.Clarke) Yin Z.
Wang
ex HB Mu
¨nchen-
Nymphenburg; M. Kiehn
and M. Pfosser 2000-1
[Cult. HBV GS-96-02]
WU Peninsular Malaysia FJ501488 FJ501325
Microchirita hamosa (R.Br.) Yin Z. Wang 1 M. Mo¨ller and Y.G. Wei
MMO 05-753
E, IBK China, Guangxi, Longzhou county JF912524 JF912551
Microchirita hamosa (R.Br.) Yin Z. Wang 2 J.M. Li LJM1181 PE China, unknown locality DQ872822 DQ872822
Microchirita involucrata (Craib) Yin Z.
Wang 1
A.R. Rafidah FRI 64447 KEP Malaysia, Kelantan, Gunung Reng JF912525 JF912552
Microchirita involucrata (Craib) Yin Z.
Wang 2
K. Imin et al. FRI 63180 KEP Peninsular Malaysia, Kedah, Baling JF912526 JF912553
Microchirita lavandulacea (Stapf) Yin Z.
Wang
Voucher from Cult. RBGE
20000897
E China, unknown locality FJ501487 FJ501324
Microchirita mollissima (Ridl.) A.Weber &
D.J.Middleton
D.J. Middleton et al. 4361 E Thailand, Surat Thani, Khlong
Phanom
JF912528 JF912555
Microchirita sericea (Ridl.) A.Weber &
Rafidah
A.R. Rafidah FRI 64328 KEP Malaysia, Kelantan, Gunung Reng JF912548 JF912521
Microchirita sp. nov. 1 D.J. Middleton et al. 4849 E Thailand, Tak, Mae Sot JF912520 JF912547
Microchirita tubulosa (Craib) A.Weber &
D.J.Middleton
D.J. Middleton et al. 4809 E Thailand, Nakhon Sawan, Wat Thep
Satha Phon
JF912531 JF912558
Microchirita viola (Ridl.) A.Weber &
Rafidah
A.R. Rafidah FRI 64388 KEP Malaysia, Kedah, P. Langkawi JF912533 JF912560
Orchadocarpa lilacina Ridl. R. Kiew RK 5410 KEP Peninsular Malaysia, Pahang,
Fraser’s Hill
HQ632863 HQ632954
Oreocharis acaulis (Merr.) Mich.Mo
¨ller &
A.Weber
M. Mo¨ller and Y.G. Wei
MMO 08-1328
E, IBK China, Guangdong, Zhaoqin county HQ632916 HQ633012
Oreocharis argyreia Chun ex K.Y.Pan M. Mo¨ ller and Y.G. Wei
MMO 07-1131
E, IBK China, Guangxi, Wuming county HQ632919 HQ633015
Oreocharis aurea Dunn M. Mo¨ller and L.M. Gao
MMO 06-980
E, IBK China, Yunnan, Jinping county HQ632920 HQ633016
Chayamaritia, a new genus from Southeast Asia 1955
123
Author's personal copy
Table 1 continued
Taxon Voucher number Deposited
in
Origin trnL-F ITS or
ITS1/ITS2
Oreocharis auricula (S.Moore) C.B.Clarke M. Mo¨ ller and L.M. Gao
MMO 03-304
E, KUN China; Guizhou, Jiangkou county FJ5011481 FJ501323
Oreocharis begoniifolia (H.W.Li)
Mich.Mo
¨ller & A.Weber
M. Mo¨ller et al. MMO 08-
1221
E, KUN China, Yunnan, Jing Dong county HQ632929 HQ633025
Oreocharis concava (Craib) Mich.Mo
¨ller &
A.Weber
M. Mo¨ller and Y.D. Qi MMO
01-153
E, PE, WU China, Yunnan, Binchuan county FJ501505 FJ501336
Oreocharis convexa (Craib) Mich.Mo
¨ller &
A.Weber
M. Mo¨ller and Y.D. Qi MMO
01-176
E, PE, WU China, Yunnan, Dali county FJ501506 FJ501337
Oreocharis cotinifolia (W.T.Wang)
Mich.Mo
¨ller & A.Weber
Q.M. Chuan 01 IBK China, Guangxi, Dayaoshan, Jinxiu
conuty
HQ632914 HQ633010
Oreocharis craibii Mich.Mo
¨ller & A.Weber M. Mo¨ller and L.M. Gao
MMO 07-1072
E, KUN China, Sichuan, Pan Zhi Hua county HQ632921 HQ633017
Oreocharis dasyantha Chun var. ferruginosa
K.Y.Pan
Y.G. Wei 07-700 IBK China, Hainan, Delong HQ632918 HQ633014
Oreocharis esquirolii H.Le
´v. D.W. Zhang 723 IBK China, Guizhou, An Long county,
Longtoushan
HQ632915 HQ633011
Oreocharis jiangxiensis (W.T.Wang)
Mich.Mo
¨ller & A.Weber
M. Mo¨ller et al. MMO 09-
1451
E China, Fujian, Jiangle county HQ632914 HQ633029
Oreocharis lancifolia (Franch.) Mich.Mo
¨ller
& A.Weber
M. Mo¨ller and P. Zhou MMO
09-1624
E China, Sichuan, Mianning county HQ632924 HQ633020
Oreocharis longifolia (Craib) Mich.Mo
¨ller
& A.Weber
M. Mo¨ller et al. MMO 08-
1239
E, KUN China, Yunnan, Jingdong county HQ632934 HQ633030
Oreocharis lungshengensis (W.T.Wang)
Mich.Mo
¨ller & A.Weber
M. Mo¨ller and Y.G. Wei
MMO 06-916
E, IBK China, Guangxi, Longsheng county HQ632917 HQ633013
Oreocharis magnidens Chun ex K.Y.Pan M. Mo¨ ller and Y.G. Wei
MMO 06-896
E, IBK China, Guangxi, Jinxiu county HQ632930 HQ633026
Oreocharis mileensis (W.T.Wang)
Mich.Mo
¨ller & A.Weber
Y.M. Shui 65214 KUN China, Yunnan, Shilin county HQ632928 HQ633024
Oreocharis muscicola (Diels) Mich.Mo
¨ller
& A.Weber
Kew (1995-2229) K unknown origin FJ501548 FJ501366
Oreocharis pankaiyuae Mich.Mo
¨ller &
A.Weber
Voucher from Cult. RBGE
20060865
E China, unknown locality HQ632925 HQ633021
Oreocharis primuliflora (Batalin)
Mich.Mo
¨ller & A.Weber
M. Mo¨ller and P. Zhou MMO
09-1605
E China, Sichuan, Danba county HQ632923 HQ633019
Oreocharis primuloides (Miq.) Benth. &
Hook.f. ex C.B.Clarke
T. Tsuzuki s.n. [Cult. RBGE
19842178A]
E Japan, unknown locality FJ501546 FJ501364
Oreocharis ronganensis (K.Y.Pan)
Mich.Mo
¨ller & A.Weber
M. Mo¨ller and Y.G. Wei
MMO 06-776
E, IBK China, Guangxi, Rong An county HQ632927 HQ633023
Oreocharis rosthornii (Diels) Mich.Mo
¨ller
& A.Weber
Sino-American Bryological
Expedition, no. 398 (US
229325)
US China, Guizhou, Jiangkou Xian FJ501547 FJ501365
Oreocharis sinensis (Oliv.) Mich.Mo
¨ller &
A.Weber
M. Mo¨ller and Y.G. Wei
MMO 08-1329
E, IBK China, Guangdong, Bolou county HQ632912 HQ633008
1956 D. J. Middleton et al.
123
Author's personal copy
Table 1 continued
Taxon Voucher number Deposited
in
Origin trnL-F ITS or
ITS1/ITS2
Oreocharis sinohenryi (Chun) Mich.Mo
¨ller
& A.Weber
M. Mo¨ller and Y.G. Wei
MMO 07-1150
E, IBK China, Guangxi, Fangcheng county HQ632913 HQ633009
Oreocharis stewardii (Chun) Mich.Mo
¨ller &
A.Weber
M. Mo¨ller and Y.G. Wei
MMO 06-917
E, IBK China, Guangxi,Shanjiang county HQ632926 HQ633022
Oreocharis urceolata (K.Y.Pan)
Mich.Mo
¨ller & A.Weber
M. Mo¨ller and P. Zhou MMO
09-1633
E China, Sichuan, Liangshan Yizu
county
HQ632922 HQ633018
Oreocharis xiangguiensis W.T.Wang &
K.Y.Pan 1
M. Mo¨ller and Y.G. Wei
MMO 05-741
E, IBK China, Guangxi, Lin Gui county HQ632932 HQ633028
Oreocharis xiangguiensis W.T.Wang &
K.Y.Pan 2
M. Mo¨ller and Y.G. Wei
MMO 06-915
E, IBK China, Guangxi, Longsheng county HQ632931 HQ633027
Ornithoboea arachnoidea (Diels) Craib Voucher from Cult. RBGE
19972903
E Thailand, Chiang Mai, Doi Chiang
Dao
FJ501461 FJ501312
Ornithoboea wildeana Craib Y.Z. Wang 00401 PE China, Yunnan, Xichou county FJ501462 FJ501313
Paraboea acutifolia (Ridl.) B.L.Burtt A. Weber 86805-2/1 WU Peninsular Malaysia, Kedah, Pulau
Langkawi, Bukit Terbak
FJ501464 FJ501314
Paraboea birmanica (Craib) C.Puglisi M. Mo¨ller and Y.G. Wei
MMO 06-862
E, IBK China, Guangxi, Jingxi county HQ632866 HQ632958
Paraboea capitata Ridl. A. Weber 870522-5/2 [Cult.
HBV]
WU Peninsular Malaysia, Perak, Kinta
district
AJ492298 FJ501315
Paraboea crassifolia (Hemsl.) B.L.Burtt M. Mo¨ller and Y.D. Qi MMO
01-83
E, PE, WU China, Yunnan, Maguan county FJ501472 FJ501318
Paraboea glandulosa (B.L.Burtt) C.Puglisi D.J. Middleton and P.Triboun
5202
BK, E Thailand, Kanchanaburi, Thong Pha
Phum, Ti Pugae
HQ632867 HQ632959
Paraboea umbellata (Drake) B.L.Burtt M. Mo¨ller and Y.D. Qi MMO
01-147
E, PE, WU China, Guangxi, Napo county FJ501470 FJ501317
Petrocodon ainsliifolius W.H.Chen &
Y.M.Shui
Y.M. Shui et al. 44071 KUN China, Yunnan, Maguan county HQ632941 HQ633038
Petrocodon coccinea (C.Y.Wu ex H.W.Li)
Yin Z.Wang
M. Mo¨ller and Y.D. Qi MMO
01-141
E, PE, WU China, Guangxi, Napo county FJ501516 FJ501365
Petrocodon coriaceifolius (Y.G.Wei)
Y.G.Wei & Mich.Mo
¨ller
M. Mo¨ller and Y.G. Wei
MMO 06-913
E, IBK China, Guangxi, Yangshuo county HQ632943 HQ633040
Petrocodon dealbatus Hance Q.J. Xie J-042 (US 422841) US China, Guangdong, Lianxian county FJ501537 FJ501358
Petrocodon fangianus (Y.G.Wei) J.M.Li &
Yin Z.Wang
Y.G. Wei MMO 07-1168 IBK China, Guangxi, Napo county Wei et al. (2010) Wei et al. (2010)
Petrocodon ferrugineus Y.G.Wei M. Mo¨ller and Y.G. Wei
MMO 06-784
E, IBK China, Guangxi, Xincheng county HQ632946 HQ633043
Petrocodon hancei (Hemsl.) A.Weber &
Mich.Mo
¨ller
M. Mo¨ller and Y.G. Wei
MMO 08-1342
E, IBK China, Guangxi, He Zhou city HQ632944 HQ633041
Petrocodon hechiensis (Y.G.Wei, Yan Liu
& F.Wen) Y.G.Wei & Mich.Mo
¨ller
M. Mo¨ller and Y.G. Wei
MMO 07-1077
E, IBK China, Guangxi, Hechi city HQ632942 HQ633039
Petrocodon integrifolius (D.Fang & L.Zeng)
A.Weber & Mich.Mo
¨ller
M. Mo¨ller and Y.G. Wei
MMO 06-865
E, IBK China, Guangxi, Longzhou county HQ632940 HQ633037
Chayamaritia, a new genus from Southeast Asia 1957
123
Author's personal copy
Table 1 continued
Taxon Voucher number Deposited
in
Origin trnL-F ITS or
ITS1/ITS2
Petrocodon jasminiflorus (D.Fang &
W.T.Wang) A.Weber & Mich.Mo
¨ller
M. Mo¨ller and Y.G. Wei
MMO 06-851
E, IBK China, Guangxi, Napo county Wei et al. (2010) Wei et al. (2010)
Petrocodon lui (Yan Liu & W.B.Xu)
A.Weber & Mich.Mo
¨ller
Y.G. Wei 8012 IBK China, Guangxi, Jingxi county,
Wuping town, Xunma village
HQ632938 HQ633035
Petrocodon scopulorum (Chun) Yin Z.Wang W. Fang 2010-02 IBK China, Guizhou, Xiuwen county,
Maochong village
HQ632947 HQ633044
Petrocodon tiandengensis (Yan Liu & B.
Pan) A.Weber & Mich.Mo
¨ller
Y.G. Wei MMO 07-1164 E, IBK China, Guangxi, Tiandeng county, HQ632945 HQ633042
Petrocodon viridescens W.H.Chen,
Mich.Mo
¨ller & Y.M.Shui
Y.M. Shui et al. 82661 E China, Yunnan, Maguan county HQ632939 HQ633036
Petrocosmea kerrii Craib Voucher from Cult. RBGE
19715592
E unknown origin FJ501502 FJ501334
Petrocosmea minor Hemsl. Sino-Amer. Bot.
Expedition, no. 1574 (US
56119)
US China, Yunnan, Lunan Xian FJ501504 Wei et al. (2010)
Petrocosmea nervosa Craib Smithsonian Institute 78-057
[Cult. RBGE 19933232]
E, US China, N Yunnan AJ492299 FJ501335
Petrocosmea sericea C.Y.Wu ex H.W.Li Z.J. Gu 99-1104 KUN China, unknown locality FJ501503 Wei et al. (2010)
Platystemma violoides Wall. Projektteam 197-241 WU Nepal, SE Kathmandu, Pulchoki FJ501443 Wei et al. (2010)
Primulina weii Mich.Mo
¨ller & A.Weber J.M. Li Ljm-04-42 PE China, Guangxi DQ872811 DQ872832
Primulina bipinnatifida (W.T.Wang) Yin
Z.Wang
PE China, Guangxi DQ872806 DQ872842
Primulina cordifolia (D.Fang & W.T.Wang)
Yin Z.Wang
J.M. Li 05561 PE China, Guangxi DQ872803 DQ872845
Primulina dryas (Dunn) Mich.Mo
¨ller &
A.Weber
T.C. Godfrey 369 [Cult.
RBGE 19791050]
E China, Hong Kong FJ501524 FJ501348
Primulina gemella (D.Wood) Yin Z.Wang L. Averyanov 1987 [Cult.
RBGE 19941913]
E Vietnam, Hong Quang Special
Region, Cat Hai
FJ501523 FJ501345
Primulina glandulosa (D.Fang et al.) Yin
Z.Wang var. yangshuoensis (F.Wen,
Q.X.Zhang & Yue Wang) Mich.Mo
¨ller &
A.Weber
M. Mo¨ller and Y.G. Wei
MMO 06-912
E, IBK China, Guangxi, Yang Shuo county HQ632948 HQ633045
Primulina glandulosa (D.Fang et al.) Yin
Z.Wang
J.M. Li 054291 PE China, Guangxi DQ872804 DQ872841
Primulina heterotricha (Merr.) Yin Z.Wang Y.Z. Wang 067311 PE China, Guangxi DQ872816 DQ872826
Primulina linearifolia (W.T.Wang) Yin
Z.Wang
J.M. Li 11121 PE China, Guangxi DQ872810 DQ872834
Primulina longgangensis (W.T.Wang) Yin
Z.Wang
A. Takhtajan and N. Aruzytov
1975 [Cult. RBGE
19941915]
E Vietnam, unknown locality AJ492290 FJ501347
Primulina luochengensis (Yan Liu &
W.B.Xu) Mich.Mo
¨ller & A.Weber
Y.G. Wei MMO 07-1163 IBK China, Guangxi, Luocheng county,
Xiaochangan town
HQ632949 HQ633046
1958 D. J. Middleton et al.
123
Author's personal copy
Table 1 continued
Taxon Voucher number Deposited
in
Origin trnL-F ITS or
ITS1/ITS2
Primulina minutimaculata (D.Fang et
W.T.Wang) Yin Z.Wang
J.M. Li 067134 PE China, Guangxi DQ872815 DQ872828
Primulina mollifolia (D.Fang et W.T.Wang)
Yin Z.Wang
J.M. Li 054281 PE China, Guangxi DQ872802 DQ872847
Primulina ophiopogoides (D.Fang et
W.T.Wang) Yin Z.Wang
Y.Z. Wang 067134 PE China, Guangxi DQ872814 DQ872829
Primulina pinnata (W.T.Wang) Yin Z.Wang Expedition Beijing 896526
(US 294374)
US China, Guangxi, Rongshui Xian FJ501526 FJ501349
Primulina pinnatifida (Hand.-Mazz.) Yin
Z.Wang
Q.J. Xie J-037 (US 422838) US China, Guangdong, Lianxian county FJ501527 FJ501350
Primulina pteropoda (W.T.Wang) Yin
Z.Wang
Y.Z. Wang 067312 PE China, Guangxi DQ872817 DQ872827
Primulina renifolia (D.Fang & D.H.Qin) Yin
Z.Wang
M. Mo¨ller and Y.G. Wei
MMO 06-791
E, IBK China, Guangxi, Duan county Wei et al. (2010) Wei et al. (2010)
Primulina repanda (W.T.Wang) Yin
Z.Wang var. guilinensis (W.T.Wang)
Mich.Mo
¨ller & A.Weber
ex Smithsonian Institute 94-
083 [Cult. RBGE
19951206]
E China, Guangxi AJ492292 FJ501351
Primulina spadiciformis (W.T.Wang)
Mich.Mo
¨ller & A.Weber
ex Smithsonian Institute 94-
087 [Cult. RBGE
19951205]
E China, unknown locality AJ492291 FJ501346
Primulina spinulosa (D.Fang et W.T.Wang)
Yin Z.Wang
Y.Z. Wang 067133 PE China, Guangxi DQ872813 DQ872830
Primulina tabacum Hance Q.J. Xie and C.X. Ye s.n.
[Cult. RBGE 19951540]
E China, Guangdong, Lian River AJ492300 FJ501352
Primulina wentsaii (D.Fang et L.Zeng) Yin
Z.Wang
J.M. Li 11630 PE China, Guangxi DQ872812 DQ872831
Pseudochirita guangxiensis (S.Z.Huang)
W.T.Wang
M. Mo¨ller and Y.G. Wei
MMO 06-798
E, IBK China, Guangxi, Mashan county HQ632908 HQ633003
Pseudochirita guangxiensis (S.Z.Huang)
W.T.Wang var. glauca Y.G.Wei & Yan
Liu
M. Mo¨ller and Y.G. Wei
MMO 05-751
E, IBK China, Guangxi, Jingxi county HQ632909 HQ633004
Ramonda myconi (L.) Rchb. Voucher from Cult. RBGE
19711477
E Spain, Pyrenees AJ492301 Mo
¨ller et al. (1999)
Ramonda nathaliae Panc
ˇic
´& Petrovic
ˇVoucher from Cult. RBGE
19784020
E Macedonia, unknown locality AJ501438 Mo
¨ller et al. (1999)
Raphiocarpus sinicus Chun M. Mo¨ller and Y.G. Wei
MMO 07-1141
E, IBK China, Guangxi, Shangsi county HQ632877 HQ632973
Rhabdothamnopsis sinensis Hemsl. Voucher from Cult. Kew
1988 4866
K China, unknown locality AJ492302
Rhabdothamnopsis sinensis Hemsl. M. Mo¨ller and L.M. Gao
MMO 08-1059
E, KUN China, Sichuan, Mianning county HQ632960
Rhynchotechum discolor (Maxim.) B.L.Burtt RBGE-PNH Expedition
1997/SM8 [Cult. RBGE
19972562]
E Philippines, Luzon, Isabela FJ501436 Wei et al. (2010)
Chayamaritia, a new genus from Southeast Asia 1959
123
Author's personal copy
Table 1 continued
Taxon Voucher number Deposited
in
Origin trnL-F ITS or
ITS1/ITS2
Rhynchotechum parviflorum Blume M. Mendum, G. Argent and
Hendrian 00148
E Central Sulawesi, Mt. Sojol FJ501437 Wei et al. (2010)
Ridleyandra petiolata (Ridl.) A.Weber M.A. Mohd.Hairul FRI 60092 KEP Peninsular Malaysia, G. Inas HQ632935 HQ633032
Ridleyandra porphyrantha (A.Weber &
Kiew) A.Weber
A. Weber 870420-2/4 WU Malaysia, Pahang, side ridge of
Gunung Bunga Buah
FJ501520 HQ633031
Ridleyandra quercifolia (Ridl.) A.Weber T.L. Yao FRI 65405 KEP Peninsular Malaysia, Perak, Maxwell
Hill
HQ632936 HQ633033
Senyumia minutiflora (Ridl.) Kiew, A.Weber
& B.L.Burtt
A.R. Rafidah, R. Kiew and
M.A. Mohd.Hairul FRI
55722
KEP Peninsular Malaysia, Pahang,
Gunung Senyum
HQ632865 HQ632957
Spelaeanthus chinii Kiew, A.Weber &
B.L.Burtt
A. Weber 860709-2/2 WU Peninsular Malaysia, Pahang,
Jerantut distr., Taman Negara
FJ501457 FJ501307
Streptocarpus andohahelensis Humbert M. Mo¨ller and G.
Rafamatanantsoa MM 9717
E, TAN Madagascar, Tule
´ar, Ranomafana FJ501449 AF316903
Streptocarpus beampingaratrensis Humbert M. Mo¨ller and G.
Rafamatanantsoa MM 9715
E, TAN Madagascar, Tule
´ar, Ranomafana FJ501448 AF316905
Streptocarpus dunnii Hook.f. I. LaCroix s.n. [Cult. RBGE
19941745]
E Swaziland, Mbabane FJ501456 AF316951
Streptocarpus hilsenbergii R.Br. B.L. Burtt s.n. [Cult. RBGE
19631505]
E Madagascar, Mandrake valley FJ501450 AF316907
Streptocarpus holstii Engl. Cornell University (Bail.
Hort) [Cult. RBGE
19592272]
E Tanzania, unknown locality AJ492304 AF316917
Streptocarpus ibityensis Humbert E. Fischer 250/93 [Cult.
RBGE 19932867]
E Madagascar, Antananarivo FJ501455 AF316926
Streptocarpus papangae Humbert M. Mo¨ller and G.
Rafamatanantsoa MM 9718
E, TAN Madagascar, Tule
´ar, Ranomafana FJ501444 AF316929
Streptocarpus rexii (Hook.) Lindl. K. Jong s.n. [Cult. RBGE
19870333]
E South Africa, NE Cape,
Grahamstown
AJ492305 AF316979
Streptocarpus saxorum Engl. A. Chautems and M. Perret
01-023
G cult. CJBG FJ501447
Streptocarpus saxorum Engl. I.C. Mather 4 [Cult. RBGE
19721499]
E Tanzania, Tanga region AF316914
Tetraphyllum roseum Stapf H.K. Kurzweil 798 WU Thailand, Krabi Province FJ501434 HQ632950
Tribounia grandiflora D.J.Middleton D.J. Middleton and P.
Triboun 5205
E Thailand, Kanchanaburi, Muang,
Wat Tham Khao Pun
JX839281 JX839280
Tribounia venosa (Barnett) D.J.Middleton D.J. Middleton and P.
Triboun 4589
E Thailand, Lampang, Mae Prik, Tham
Nam Pha Phangam
JX839282 JX839283
1960 D. J. Middleton et al.
123
Author's personal copy
somewhat resembling those of Chayamaritia. However, the
flowers of the two genera are very different, clearly
forming a long corolla tube in Chayamaritia and often flat-
faced or with only a short tube in Petrocosmea. In fact, the
corolla of the former is rather similar in shape and size to
those found among most other genera in the clade, such as
Pseudochirita and Allostigma. In addition, the imbricate
sepals of Chayamaritia are rather characteristic and ap-
parently uncommon in Asian Gesneriaceae. Although we
have not done an extensive survey, we are aware of im-
bricate sepals only in some species of Petrocosmea (e.g. P.
bicolor D.J.Middleton & Triboun) and Paraboea (e.g. P.
sinensis (Oliv.) B.L.Burtt).
An alternate leaf arrangement occurs in genera across
the family Gesneriaceae and, amongst those from the Old
World, can be found in tribe Trichosporeae, subtribe
Leptoboeinae, in most species of Boeica C.B.Clarke and
some species of Rhynchotechum Blume, and in subtribe
Didymocarpinae in the recently described Billolivia
D.J.Middleton (Middleton et al. 2014) and in Cathayanthe
W.Y.Chun from Hainan. None of these have been shown to
be closely related to Chayamaritia (Fig. 1).
Henckelia is likely to remain a confusing concept for
some time to come due to its very radical realignment
following its treatment by Weber et al. (2011a). It was a
large genus both before and after being redefined by Weber
et al. (2011a), but only a minority of the species is common
to both circumscriptions. The other result of the new genus
concept for Henckelia is that it shifted from being a genus
with two centres of diversity in South India/Sri Lanka and
Malesia to being a genus of India, Sri Lanka and the Hi-
malayas with extensions into northern Thailand, northern
Laos and northern Vietnam. In the old concept, most of its
species were found in the aseasonal wet tropics of Malesia
south of 9N, and in the new concept all species bar one in
Southeast Asia are only found north of 18N. Indeed, if it
were not for the extremely widespread Henckelia ana-
choreta (Hance) D.J.Middleton & Mich.Mo
¨ller, which is
found from around 30N in China to around 13Nin
Thailand, there would be no overlap with Chayamaritia
which occupies a zone between the old and most of the new
concepts of Henckelia at 14–18N. There is a similar
biogeographic split with its sister genus Petrocosmea, all
species of which are only found north of 18N except for
the poorly known Petrocosmea condorensis Pellegr., which
occurs on Con Son Island off the south coast of Vietnam.
This species is almost certainly misplaced in Petrocosmea
but requires further study.
Chayamaritia is the fifth new genus of Gesneriaceae
described from Southeast Asia since 2012, the others being
Somrania D.J.Middleton (Middleton and Triboun 2012)
and Tribounia (Middleton and Mo
¨ller 2012), both from
Thailand, Billolivia (Middleton et al. 2014), from Vietnam,
and Glabrella from South China (Mo
¨ller et al. 2014).
Somrania and Billolivia were described from species that
were also previously unknown. The situation with Chaya-
maritia is more akin to Tribounia and Glabrella in that the
type species of each of them were previously described in
other genera. In the cases of Chayamaritia and Tribounia,
the discovery of a new but clearly related species to these
original two species was the spur to investigate their rela-
tionships that then resulted in new genera. All of these new
genera, each supported by molecular phylogenetic studies
(Mo
¨ller et al. 2009,2011; Middleton and Mo
¨ller 2012;
Middleton et al. 2014; Puglisi 2014), present a contrast to
the other recent trend in Gesneriaceae systematics where
large numbers of genera have been synonymised (Mo
¨ller
et al. 2011; Weber et al. 2011b,2011c) or radically re-
aligned (Weber et al. 2011a; Middleton et al. 2013).
Although there are unlikely to be more large realignments
of genera, there are still a number of genera where the
limits are in need of clarification, including other species
assemblages formerly placed in Chirita (see Weber et al.
2011a).
Conservation assessments using IUCN (2012) guideli-
nes are calculated for one species. For the other species,
Chayamaritia banksiae, it is concluded that there are in-
sufficient data to give an assessment.
Taxonomic treatment
Chayamaritia D.J.Middleton & Mich.Mo
¨ller, gen. nov.
TYPE: Chayamaritia smitinandii (B.L.Burtt)
D.J.Middleton.
Description: Herb with thick, short, somewhat fleshy
stem firmly attached to rock. Leaves simple, congested,
alternately arranged around thick stem, petiolate, pubescent
above and beneath, venation pinnate, eucamptodromous,
tertiary venation alternate percurrent. Inflorescences axil-
lary, scapose, 2–12-flowered, pubescent throughout, flow-
ers somewhat pendent. Calyx of five lobes divided to base,
strongly imbricate, upper lobe generally larger, margins
minutely to coarsely dentate. Corolla with a long broad
tube and 2-lipped limb with slightly spreading lobes; tube
dorsally with 2 furrows with a ridge inbetween; upper lip
2-lobed, lobes rounded; lower lip 3-lobed, lobes rounded.
Fertile stamens 2, the anterior ones; filaments inserted
around or slightly below middle of corolla tube, bent
around the middle, widest in middle; anther with 2 thecae,
thecae parallel, glabrous, adnate tip to tip, touching face to
face; lateral staminodes 2, large; medial staminode small.
Disc a small 5-lobed or 5-crenate ring. Pistil with short
stipe, ovary and style, held in a wide groove in the dorsal
side of the corolla tube; ovary cylindrical, bicarpellate,
unilocular with two parietal placentae, ovules numerous;
stigma chiritoid, with a spathulate lower lobe, this bifid
Chayamaritia, a new genus from Southeast Asia 1961
123
Author's personal copy
with blunt lobes. Fruit a fusiform capsule, shortly stipitate,
not twisted, dehiscence loculicidal along upper side. Seeds
small, numerous, unappendaged.
Diagnosis: Similar to both Henckelia and Primulina in
the tubular corollas, two fertile stamens and chiritoid
stigma but differing from both in the combination of
thickened rhizomatous prostrate stem, the alternately ar-
ranged leaves and the imbricate sepals.
Distribution: Central and eastern Thailand, Laos.
Etymology: The genus is named in honour of Dr
Kongkanda Chayamarit of the Forest Herbarium in Bang-
kok. She has been a tireless advocate of research on the
Thai flora and has been instrumental in the increase in
publication rate of the Flora of Thailand series in recent
years. She has done this through finding sources of funding
for field work and family accounts and through ensuring
that authors of accounts do not forget their promises!
Currently only known from two species, one of them
newly described here.
Key to the species in Chayamaritia
1. Calyx lobes 7–15 mm wide; corolla tube whitish
ventrally, lobe margins dentate; leaves 1.5–1.9 times as
long as wide, base rounded to subcordateC. banksiae.
1. Calyx lobes 0.9–4.2 mm wide; corolla tube purple
ventrally, lobe margins more or less entire; leaves 1.8–5.6
times as long as wide, base cuneate to attenuate or rarely
roundedC. smitinandii.
Chayamaritia banksiae D.J.Middleton, sp. nov.
TYPE: Laos, Khammouan, Nam On catchment, Phou Ak
escarpment, Nakai Nam Theun NBCA, 936 m a. s. l., 22
May 2006, collected only as a living collection Newman
et al. LAO1428, grown on at the Royal Botanic Garden
Edinburgh as accession number 20060845*A and vou-
chered for the herbarium and as the holotype as Middleton
5220 (Holotype E) (Figs. 2,3a–c, 4).
Description: Basal parts, petioles and leaf blades above
and below densely covered with short white appressed
hairs, these somewhat reddish when young. Leaves con-
gested; petioles 6–16 cm long; blade ovate, 8.8–18 9
5.7–10.7 cm, 1.5–1.9 times as long as wide, base rounded
to subcordate, sometimes slightly asymmetrical, apex
shortly acuminate, margin minutely dentate, midrib im-
pressed above, strongly prominent beneath, 8–9 veins on
each side of midrib, these impressed above, very prominent
beneath, tertiary venation obscure above, prominent be-
neath. Inflorescence 4–6-flowered, all axes and bracts pale
green with red appressed hairs; peduncle to 15.5 cm long;
bracts ovate, apex acuminate, margin dentate, c.
291.9 cm; pedicels 1.2–1.5 mm. Calyx lobes ovate,
15–21 97–15 mm, apex acuminate, pale green with red
and green appressed hairs, inside with white appressed
hairs in upper half, margin coarsely dentate but obscured
by ciliate hairs. Corolla 4.8–6.2 cm long, tube white ven-
trally and purple dorsally outside, white to pale purple with
2 parallel yellow lines ventrally inside, lobes purple outside
and inside, paler at base; tube 3.5–4.5 cm long; upper lip
with each lobe orbicular, apex rounded, margins minutely
dentate and ciliate, 10–14 912.5–16 mm; lower lip with
each lobe orbicular, apex rounded, margins minutely den-
tate and ciliate, lateral lobes 12.5–13 911–16 mm, middle
lobe 11–12 912–14 mm; outside of corolla short glandu-
lar pubescent throughout except at very base, glandular heads
of hairs very small, inside glabrous except for short glandular
hairs at sinus of upper lobes and sessile glands on ventral
throat. Fertile stamens inserted at 15–21 mm from corolla
base; filaments white, 11.5–15 mm long, geniculate and
widening at 6–8 mm, minutely and sparsely glandular pu-
berulent; anthers cream-coloured, 3 92.2–3 mm; lateral
staminodes 5.5–11 mm long, middle staminode 0.5–1.2 mm
long. Disc 5-lobed, c. 1.5 mm high, with sparse minute
glandular hairs. Ovary and style green, densely covered with
long eglandular hairs; ovary shortly stipitate, 20–28 mm long;
style 8 mm long; stigma only of lower lobe, c. 5 mm long,
this bifid with blunt lobes. Fruit unknown.
Diagnosis: Most similar to Chayamaritia smitinandii but
differing in the outside of the corolla being white ventrally
(purple in C. smitinandii), the leaf blade index mostly being
lower (1.5–1.9 vs. 1.8–5.6 times as long as wide in C. smiti-
nandii), the blade base being rounded to subcordate (cuneate to
attenuate or only rarely rounded in C. smitinandii), the calyx
lobes being broader and more dentate (7–15 mm vs.
0.9–4.2 mm wid e in C. smitinandii) andthe corolla lobes being
minutely dentate along margin (entire in C. smitinandii).
Distribution: Endemic to Laos (Khammouan Province).
Habitat: Growing on sides of boulders in evergreen
forest.
Etymology: Named after the botanical illustrator Claire
Banks.
Proposed IUCN conservation assessment: Data Deficient.
Chayamaritia banksiae is only known from one collection,
and its complete distribution is unknown. The type specimen
was collected from a cultivated plant with known provenance
in Nakai Nam Theun National Biodiversity Conservation
Area in Khammouan Province in Laos. Although the collec-
tion locality is not particularly far from the populations of
Chayamaritia smitinandii in northeastern Thailand, the two
species are easily distinguished, most obviously in leaf and
sepal shapes as noted in the key and diagnosis.
Chayamaritia smitinandii (B.L.Burtt) D.J.Middleton,
comb. nov.—TYPE: Thailand, Nakhon Ratchasima, Khao
Yai National Park, 6 Oct 1962, T. Smitinand 7491 (Holo-
type BKF) (Figs. 3c, d, 4)
:Chirita smitinandii B.L.Burtt in Thai Forest Bull.,
Bot. 29: 89. 2001
1962 D. J. Middleton et al.
123
Author's personal copy
:Henckelia smitinandii (B.L.Burtt) D.J.Middleton &
Mich.Mo
¨ller, Taxon 60: 776. 2011.
Description: Leaves congested; petioles (1.1–)4–13 cm
long, densely pubescent; blades (3.2–)13–21 9(0.8–)
2.5–10.5 cm, 1.8–5.6 times as long as wide, base attenuate
to cuneate or rarely rounded, apex acuminate, margin
minutely dentate, secondary veins 6–9, pale pubescent all
over above and beneath, more densely so beneath. Inflo-
rescences 2–12-flowered, all axes densely pubescent; pe-
duncles 11.5–26 cm long; bracts narrowly elliptic to
lanceolate, somewhat falcate, 7.7–34 91.8–8 mm, apex
acuminate, densely pubescent; pedicels 6.5–11 mm long.
Fig. 2 Chayamaritia banksiae D.J.Middleton. aHabit; bflower,
lateral view; ccalyx opened out showing imbricate lobes; dcorolla
dissection; eclose-up of corolla lobe margin; fpistil; gclose-up of
stigma showing enlarged and slightly bifid lower lobe. Drawn from
Middleton 5220 (e) by Claire Banks. Scale bars a=5 cm; bd,f,
g=2 cm; e=2mm
Chayamaritia, a new genus from Southeast Asia 1963
123
Author's personal copy
Calyx slightly zygomorphic with upper lobe longer than
others, lobes lanceolate or narrowly elliptic,
5.3–15 90.9–4.2 mm, apex acute or acuminate, margin
slightly toothed or appearing as large sessile glands on
margin, densely pubescent. Corolla 3.8–6 cm long, outside
deep purple throughout, paler proximally, inside with 2
ventral yellow lines in line with sinuses on lower lip and
down into tube; tube 3.5–4.5 cm long; upper lip with each
Fig. 3 Chayamaritia banksiae D.J.Middleton. aWhole plant in
cultivation at the Royal Botanic Garden Edinburgh; bClose-up of
flower showing strongly imbricate sepals; cClose-up of corolla throat
and stigma. Chayamaritia smitinandii (B.L.Burtt) D.J.Middleton.
dFruiting population of Middleton et al. 5632. eFlowering plant of
Middleton et al. 5652. Scale bars in c,d=1cm
1964 D. J. Middleton et al.
123
Author's personal copy
lobe squarish, apex rounded, margins ±entire,
8–11.5 99–11.5 mm; lower lip with each lobe orbicular to
obovate, apices rounded, margins ±entire, lateral lobes
5.5–8 910 mm, middle lobe 7–10 97–10 mm; outside
of corolla densely covered in gland-tipped purple hairs,
inside glabrous except for gland-tipped hairs along inside
of each furrow dorsally. Stamens inserted at 16–26 mm
from corolla base; filaments 9.5–13 mm long, white,
widest in middle, glabrous; anthers 2.1–2.7 92.1–3.1 mm;
lateral staminodes 4–5 mm long, medial staminode
0.7 mm long. Disc 0.9–1.4 mm, 5-crenate at margin. Pistil
40–47 mm long; ovary shortly stipitate, 26.5–30 mm long,
green, densely eglandular pubescent and with few subses-
sile glands; style 13.5–17 mm long, white, densely eglan-
dular pubescent intermixed with fewer and shorter gland-
tipped hairs; stigma with upper lip absent and an enlarged
spathulate entire lower lip, 2–3 mm long, weakly bifid at
apex. Fruit 3–5 cm long, 2.2–2.7 mm wide.
Distribution: Thailand (Nakhon Ratchasima, Nakhon
Nayok, Buengkan and Nakhon Phanom Provinces).
Habitat: In evergreen and submontane forest in deep
shade at 150–1200 m altitude.
Etymology: Named after the Thai botanist Tem Smiti-
nand (1920–1995) who was also a maternal uncle of
Kongkanda Chayamarit after whom the genus is named.
Proposed IUCN conservation assessment: Vulnerable
VU B1 ab(iii). This species has an Extent of Occurrence of
less than 20,000 km
2
, and the northeastern populations are
widely separated from the central populations by the pre-
dominant agricultural landscape of northeastern Thailand.
In addition, the known populations can be subject to
disturbance by the high visitor numbers found in the areas
where this species is found in Khao Yai National Park.
Additional specimens studied: Thailand: Nakhon
Nayok: Muang Nakhon Nayok, Hin Tang subdistrict, Pah
Dah Baek Falls, 12 Sep 2002, J.F. Maxwell 02-353 (BKF,
CMU, L); ibid, 19 Oct 2003, P. Palee 643 (CMU); Muang
Nakhon Nayok, Khao Yai National Park, Near Pha Ta-
Baeg Waterfall, 14 Oct 1984, G. Murata et al. T-52442
(BKF); Muang Nakhon Nayok, Khao Yai National Park,
Pah Da Bek trail to river, 20 Aug 2012, D.J. Middleton
et al. 5632 (E); Muang Nakhon Nayok, Khao Yai National
Park, Pha Diao Dai Nature Trail, 23 Aug 2012, D.J. Mid-
dleton et al. 5652 (E). Nakhon Ratchasima: Khao Yai
National Park, Khao Khiew, Pha Tom Chai, 14 Sep 1999,
S. Watthana 695 (QBG); Khao Yai National Park, Khao
Laem, 19 Oct 1969, C.F.v. Beusekom and C. Charoenphol
1758 (BKF, E, K, L, P). Nakhon Phanom: Phu Langka
National Park, Tadkham Falls, 30 Oct 1998, T. Wong-
prasert s.n. (BKF). Buengkan: Seka district, Phu Wua
Wildlife Sanctuary, Bang Bart stream, 10 Oct 2013, S.
Suddee et al. 4595 (BKF).
Until fairly recently, this species was only known from
600–1200 m altitude in Khao Yai National Park in Nakhon
Ratchasima and Nakhon Nayok Provinces. Recent collec-
tions from some distance to the northeast in Buengkan and
Nakhon Phanom Provinces were collected at much lower
altitudes.
Acknowledgements We thank the directors and the curators of the
herbaria that have loaned material or hosted visits of the first author;
the curatorial staff of the Royal Botanic Garden Edinburgh for help in
managing the specimens; the staff of the Forest Herbarium Bangkok
for assistance with field work; Dr Pramote Triboun for advice and
help in the field; Dr Martin Pullan for assistance with the database;
Claire Banks for the illustration; RBGE for the use of their molecular
laboratories and financial contributions for lab consumables, and the
Sibbald Trust for support to KN. The Royal Botanic Garden Edin-
burgh is supported by the Rural and Environment Science and Ana-
lytical Services division (RESAS) in the Scottish Government.
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directory of public herbaria and associated staff. New York
Botanical Garden’s Virtual Herbarium. Available at: http://
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... The genus Chayamaritia D.J. Middleton & Mich.Möller (Gesneriaceae) was originally established and described, based on molecular and morphological data (Middleton et al. 2015). The genus Chayamaritia comprises two species and is hitherto known only in Laos and Thailand. ...
... The genus Chayamaritia comprises two species and is hitherto known only in Laos and Thailand. A thickened rhizomatous prostrate stem, along with alternately arranged leaves and the imbricate sepals characterise the genus (Middleton et al. 2015). The type species, Chayamaritia smitinandii (B.L.Burtt) D.J.Middleton, was initially placed in the genus Chirita Buch.-Ham. ...
... (Burtt, 2001), then transferred to the genus Henckelia Spreng. (Weber et al. 2011) and eventually separated as a new genus in 2015, together with the newly-described species, Chayamaritia banksiae D.J.Middleton (Middleton et al. 2015). Chayamaritia banksiae and C. smitinandii are endemic to Laos and Thailand, respectively (Middleton et al. 2015). ...
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Chayamaritia vietnamensis , a new species from Son La Province, northern Vietnam, is described and illustrated. The phylogenetic study revealed that the new species is most closely related to C. banksiae and C. smitinandii . The morphological comparison suggests it as the third new species of Chayamaritia and distinguished from C. banksiae and C. smitinandii by a combination of morphological characters of leaf blades, bracts, calyx and corolla, especially its peltate leaf blades. This species is provisionally assessed as endangered (EN B1ab(iii), B2ab(iii)) using IUCN Categories and Criteria. Information on ecology, phenology and an identification key for the known Chayamaritia species are also provided.
... Data for the phylogenetic analyses were downloaded from GenBank (including one additional sample of Primulina minutihamata from China which we are treating here as Chirita cicatricosa). These basically represented a reduced matrix of Old World Gesneriaceae as presented in Middleton et al. (2015). Altogether 132 samples (129 species) were included, covering all 32 genera currently recognised in the subtribe Didymocarpinae of tribe Trichosporeae (Möller et al., 2009 Weber et al., 2013; Middleton et al., 2014a Middleton et al., , 2015). ...
... These basically represented a reduced matrix of Old World Gesneriaceae as presented in Middleton et al. (2015). Altogether 132 samples (129 species) were included, covering all 32 genera currently recognised in the subtribe Didymocarpinae of tribe Trichosporeae (Möller et al., 2009 Weber et al., 2013; Middleton et al., 2014a Middleton et al., , 2015). The sampling included 20 samples of 19 species of Primulina, including the type, P. tabacum (Table 1). ...
... Editing and assemblage of sequencing results were performed using the programs Sequencher 4.5 (Gene Codes Corp, Ann Arbor, USA). The newly acquired sequences were added to the reduced matrix of Middleton et al. (2015) and the matrices realigned manually. They were subsequently submitted to GenBank (Table 1). ...
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Based on molecular, morphological and cytological studies the previously monotypic genus Deinostigma W.T.Wang & Z.Y.Li has been expanded to include several species previously ascribed to Primulina Hance. Deinostigma now comprises seven species, including one previously placed in synonymy. The new combinations Deinostigma cicatricosa are made. Deinostigma eberhardtii is lectotypified. The genus is defined by a combination of an alternate leaf arrangement, hooked hairs on many plant parts, flowers with the pedicel inserted at an angle and off-centre on the receptacle, and, where known, a somatic chromosome number (2n) of < 36. This new circumscription of the genus expands its distribution from Vietnam into South China.
... The generic delimitations in Asian Gesneriaceae are often ambiguous due to significant overlap in characters between genera (Burtt 1977), which has led to many revisions including synonymization of small and monotypic genera (Möller et al. 2011a, b;Puglisi et al. 2011;Weber et al. 2011a, b;Möller et al. 2016) and establishment or resurrection of new or previously synonymized genera in recent years (Wei et al. 2010;Weber et al. 2011c;Middleton and Triboun 2012;Middleton and Möller 2012;Middleton et al. 2014Middleton et al. , 2015Middleton et al. , 2018. The combined phylogenetic-morphological approach was therefore performed as it takes both molecular and morphological evidence into account (Wei et al. 2010). ...
... The phylogenetic relationship was largely congruent with previous studies (Möller et al. 2009(Möller et al. , 2016Middleton et al. 2015Middleton et al. , 2018. Michaelmoelleria vietnamensis and its morphologically similar genus Deinostigma is recovered within a polytomy but the phylogenetic relationship of them is distant. ...
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Michaelmoelleria , a new genus from southern Vietnam is described with a single species, M. vietnamensis . The new genus is morphologically most similar to Deinostigma and Tribounia but it differs from the latter two by having four fertile stamens. Nuclear ribosomal internal transcribed spacer (ITS) region and plastid trnL-F intron spacer ( trnL-F ) DNA sequence data from the new genus and eighty-seven species representing 42 genera within tribe Didymocarpeae are used to resolve its generic placement. The molecular evidence reveals that it is most closely related to Cathayanthe rather than Deinostigma and Tribounia . The chromosome number is counted as 2n = 36 that further clarified its distinction comparing to the related genera within tribe Didymocarpeae. A global conservation assessment is also performed and classifies Michaelmoelleria vietnamensis as Critically Endangered (CR).
... In A Checklist of Vascular Plants of Lao PDR, 11 genera and 28 species of Gesneriaceae are listed (Newman et al., 2007). Subsequently, Chayamaritia D.J. Middleton & Mich.Möller was confirmed to be a new genus distributed in Lao PDR and Thailand, it appear placed in a subfamily Didymocarpoideae with genus Didymocarpus, which Chayamaritia banksiae D.J.Middleton endemic to Lao PDR (Middleton et al., 2015). ...
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Didymocarpus albiflorus, a new species from central Lao PDR, is described and illustrated with photographs. The new species is similar to D. middletonii and D. brevipedunculatus, but can be distinguished by a combination of characters (see diagnosis and note). A detailed description, illustration, photographs, distribution, ecology and provisional conservation assessment and key to the species of Didymocarpus in the flora of Lao PDR are provided.
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Based on an updated taxonomy of Gesneriaceae, the biogeography and evolution of the Asian Gesneriaceae are outlined and discussed. Most of the Asian Gesneriaceae belongs to Didymocarpoideae, except Titanotrichum was recently moved into Gesnerioideae. Most basal taxa of the Asian Gesneriaceae are found in the Indian subcontinent and Indo-China Peninsula, suggesting Didymocarpoideae might originate in these regions. Four species diversification centers were recognized, i.e. Sino-Vietnam regions, Malay Peninsula, North Borneo and Northwest Yunnan (Hengduan Mountains). The first three regions are dominated by limestone landscapes, while the Northwest Yunnan is well-known for its numerous deep gorges and high mountains. The places with at least 25% species are neoendemics (newly evolved and narrowly endemic) which were determined as evolutionary hotspots, including Hengduan Mountains, boundary areas of Yunnan-Guizhou-Guangxi in Southwest China, North Borneo, Pahang and Terengganu in Malay Peninsula, and mountainous areas in North Thailand, North Sulawesi Island. Finally, the underlying mechanisms for biogeographical patterns and species diversification of the Asian Gesneriaceae are discussed.
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Rachunia, a new genus of Gesneriaceae from Thailand, is described with a single species, Rachunia cymbiformis. Its relationship to the rest of subtribe Didymocarpinae is investigated through a phylogenetic study based on Bayesian Inference and Parsimony analyses of nuclear ITS and plastid trnL‐trnF (intron‐spacer) sequences. Morphologically, Rachunia differs from the related genera Codonoboea in the large boat‐shaped bracts and orthocarpic vs plagiocarpic fruit; from Microchirita in the bracts, wiry vs fleshy stem, the campanulate vs tubular corolla and the clavate vs chiritoid stigma, and from Henckelia in the clavate vs chiritoid stigma, large boat‐shaped bracts in the inflorescence, free and imbricate sepals, short and campanulate corolla, clavate stigma, and relatively robust orthocarpic fruit.
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The new species Ornithoboea grandiflora D.J. Middleton and new variety Ornithoboea maxwellii var. minutiflora D.J. Middleton are described.
Article
The genus Henckelia Sprengel (1817: 402) was re-established by Weber & Burtt (1997), with approximately 180 species, to include most of the Malesian, Sri Lankan and south Indian species of Didymocarpus Wallich (1819: 378). Weber et al. (2011) and Middleton et al. (2013) based on molecular data redefined the genus and included only 56 species. Subsequently three more species were added to the genus, H. pradeepiana Nampy, Manudev & Weber in Manudev et al. (2012: 119), H. sivagiriensis (Rajakumar et al. 2009: 481) Kumar (2014: 149) and H. bracteata Janeesha & Nampy (2015: 53) while Middleton et al. (2015) transferred H. smitinandii (Burtt 2001: 89) Middleton & Möller in Weber et al. (2011: 776) to the genus Chayamaritia Middleton & Möller (2015: 1961). At present the genus includes 58 species distributed in Sri Lanka, southern and northeastern India, Nepal, Bhutan, southern China, northern Vietnam, northern Laos and northern Thailand.
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Based on molecular studies, the small Chinese genus Petrocodon (two species and one variety) has been recently enlarged to include the monotypic genera Calcareoboea, Paralagarosolen and Tengia. It is shown here that the (6-7) species of Lagarosolen, the monotypic Dolicholoma, a few species of Didymocarpus, and a number of new species that have recently been published (but not formally described) under Petrocodon and Lagarosolen should be included in this genus. This raises the size of the genus from five to around 20 species. With respect to the floral diversity (corolla form, size, and coloration; with the exception of Tengia, the androecium is always diandrous) and inferred pollination syndromes (different forms of melittophily, ornithophily, psycho- and/or sphingophily), Petrocodon represents one of the most varied genera of Old World Gesneriaceae, comparable to some New World genera.
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A new formal classification of Gesneriaceae is proposed. It is the first detailed and overall classification of the family that is essentially based on molecular phylogenetic studies. Three subfamilies are recognized: Sanangoideae (monospecific with Sanango racemosum), Gesnerioideae and Didymocarpoideae. As to recent molecular data, Sanango/Sanangoideae (New World) is sister to Gesnerioideae + Didymocarpoideae. Its inclusion in the Gesneriaceae amends the traditional concept of the family and makes the family distinctly older. Subfam. Gesnerioideae (New World, if not stated otherwise with the tribes) is subdivided into five tribes: Titanotricheae (monospecific, East Asia), Napeantheae (monogeneric), Beslerieae (with two subtribes: Besleriinae and Anetanthinae), Coronanthereae (with three subtribes: Coronantherinae, Mitrariinae and Negriinae; southern hemisphere), and Gesnerieae [with five subtribes: Gesneriinae, Gloxiniinae, Columneinae (5the traditional Episcieae), Sphaerorrhizinae (5the traditional Sphaerorhizeae, monogeneric), and Ligeriinae (5the traditional Sinningieae)]. In the Didymocarpoideae (almost exclusively Old World, especially E and SE Asia/Malesia) two tribes are recognized: Epithemateae [with four small, but morphologically and genetically very distinctive subtribes: Loxotidinae (monogeneric with Rhynchoglossum), Monophyllaeinae, Loxoniinae and Epithematinae (monogeneric)] and Trichosporeae (the earliest name at tribal rank for the ‘‘Didymocarpoid Gesneriaceae’’). The last is subdivided into ten subtribes: Jerdoniinae (monospecific), Corallodiscinae (monogeneric), Tetraphyllinae (monogeneric), Leptoboeinae, Ramondinae (Europe), Litostigminae (monogeneric), Streptocarpinae (Africa and Madagascar), Didissandrinae, Loxocarpinae and Didymocarpinae. Didymocarpinae is the largest subtribe (ca. 30 genera and .1600 species) and still requires intensive study. It includes the most speciose genera such as Cyrtandra, Aeschynanthus, Agalmyla, Didymocarpus, Henckelia, Codonoboea, Oreocharis and Primulina and the types of the traditional tribes Didymocarpeae, Trichosporeae and Cyrtandreae.
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Based on molecular data and a morphological evaluation, evidence is provided that the species of eleven, mostly small-sized and monotypic genera of Chinese Gesneriaceae (Ancylostemon, Bournea, Briggsia s.str., Dayaoshania, Deinocheilos, Isometrum, Opithandra, Oreocharis, Paraisometrum, Thamnocharis, Tremacron) form a highly-supported group in which the species interrelationships run across traditional generic boundaries. The data confirm previous doubts on the naturalness of some of these genera and, after a detailed discussion of the particular genera, the conclusion is reached that the whole group is best regarded as a single genus, Oreocharis, which is thus expanded to comprise over 80 species. A list of the species is given and the necessary transfers are made. The new delimitation provides a framework for studying the species relationships and working out an infrageneric classification. Oreocharis provides an excellent example of a major monophyletic group that has experienced a rapid radiation early in its evolution and shows manifold convergences in floral characters (corolla form and coloration, fertility of stamens, anther shape and dehiscence mode), apparently reflecting different pollination strategies, but has little variation in vegetative habit and fruit structure.
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The new genus Tribounia is described with two species, Tribounia venosa (Barnett) D.J. Middleton, a new combintion, and Tribounia grandiflora D.J. Middleton, a new species. A key to the species and conservation assessments are provided.
Article
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Following recent molecular phylogenetic studies in Old World Gesneriaceae the nomenclatural implications for names in Henckelia are examined. New combinations are made in Codonoboea and Loxocarpus to account for species now excluded from Henckelia. A list is presented in which the current position of all species hitherto assigned to Henckelia is given, including the new combination Henckelia rotundata (Barnett) D.J.Middleton & Mich.Möller. A new combination in Oreocharis is made.
Thesis
Since it was first published, many additional species have been ascribed to the genus Boea Comm. ex Lam. As the genus grew in size, it also grew in morphological diversity until it was recircumscribed and became the source of a number of new segregate genera. Today, the Boea group (i.e. Boea, the segregate genera and other close relatives) comprises over 200 species in some 15 genera, found from China to Australia and throughout Malesia from Sumatra to the Solomon Islands. Previous molecular studies suggested a much more complex structure to the clade than previously thought. Here the most up to date phylogeny, covering all the genera known to belong to the Boea group, is presented. Parsimony and Bayesian Inference were the chosen approaches to the phylogenetic analysis of nine matrices generated using DNA data from 277 accessions. The markers used were the nuclear ITS and the chloroplast regions trnL-trnF (intron and spacer) and ndhF-rpl32-trnLUAG. The results show important discrepancies between the current taxonomy of the group and the clades delineated by the phylogeny. In an attempt to establish a natural classification of Boea and its allies, taxonomic and nomenclatural work was carried out on most of the genera found to be non-monophyletic. Boea Comm. ex Lam. is divided into two genera. The recircumscribed Boea is restricted to the group of taxa found in New Guinea, Solomon Islands and Australia. The genus is fully revised and contains 11 species, including the newly described Boea morobensis C.Puglisi. The Southeast Asian group of species formerly attributed to Boea, centred in Thailand, is given the resurrected name Dorcoceras Bunge. To the four species traditionally known to belong to this group, three new ones are added. These are Dorcoceras brunneum C.Puglisi, D. glabrum C.Puglisi and D. petiolatum C.Puglisi. Damrongia is deeply transformed with the synonymisation of D. cyanantha Triboun in D. trisepala (Barnett) D.J.Middleton & A.Weber, and the inclusion of Boea clarkeana Hemsl. and the three Asian species of Streptocarpus Lindl., S. orientalis Craib, S. burmanicus Craib and S. sumatranus B.L.Burtt. As a result of the phylogenetic study, a new genus, Middletonia C.Puglisi, is segregated from Paraboea. Middletonia consists of five species, including the newly described Middletonia glebosa C.Puglisi, and has its centre of distribution in Thailand. Finally, Paraboea (C.B.Clarke) Ridl. is recircumscribed with the inclusion of the genera Trisepalum C.B.Clarke and Phylloboea Benth. In order to limit the number of new combinations needed and maintain clarity, the name Paraboea was conserved against both Phylloboea and Trisepalum. In addition to the 15 new combinations in Paraboea, a new species from the Philippines, P. zamboangana C.Puglisi, is described.
Article
Based on molecular data and morphology, Metabriggsia is reduced to synonymy with Hemiboea and its two species transferred to that genus.
Article
Based on molecular and morphological data, the new genus Billolivia with five new species, B. longipetiolata, B. minutiflora, B. poilanei, B. vietnamensis and B. violacea, is described. IUCN conservation assessments are provided for the species.