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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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Accepted by M. Weigend: 15 Apr. 2011; published: 16 May 2011 1
PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Copyright © 2011 Magnolia Press
Phytotaxa 23: 136 (2011)
www.mapress.com/phytotaxa/Article
A new delineation for Oreocharis incorporating an additional ten genera of
Chinese Gesneriaceae
MICHAEL MÖLLER1,5, DAVID MIDDLETON1, KANAE NISHII1,2, YI-GANG WEI3, SUSANNE
SONTAG4 & ANTON WEBER4,5
1 Royal Botanic Garden Edinburgh, 20A Inverleith Row, Edinburgh EH3 5LR, Scotland, UK. m.moeller@rbge.org.uk
2 Institute of Ecology and Evolutionary Biology, Department of Life Science, National Taiwan University, Taipei 10617, Taiwan.
3 Guangxi Institute of Botany, Guangxi Zhuang Autonomous Region and the Chinese Academy of Sciences, Guilin 541006, China
4 University of Vienna, Faculty Centre of Biodiversity, Department of Structural and Functional Botany,
Rennweg 14, 1030 Vienna (Wien), Austria.
5 Authors for correspondence (anton.weber@univie.ac.at) (m.moeller@rbge.ac.uk)
Abstract
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.
Key words: Ancylostemon, Bournea, Briggsia, Dayaoshania, Deinocheilos, Isometrum, Lamiales, molecular
systematics, Opithandra, Paraisometrum, pollination syndromes, Thamnocharis, Tremacron
Introduction
Recent molecular phylogenetic work on the Old World Gesneriaceae has revealed a high level of polyphyly
on the one side, and species relationships running across traditional generic boundaries on the other (Möller et
al. 2009, 2011). Examples include the several different lineages of Chirita Buch.-Ham. ex Don (1825: 89)—
the one including the type now being included in Henckelia Sprengel (1817: 402)—and the expanded
definitions of Hemiboea Clarke (1888: t. 1798) and Petrocodon Hance (1883: 167; Wang et al. 2011, Weber et
al. 2011a, b, c).
In the present paper we focus on a case where molecular data suggest an extreme restructuring of existing
genera and their union into one: Oreocharis. In this paper, we will define this genus anew to include species
from 11 previously accepted genera (or 14 if the previously synonymised genera Dasydesmus, Perantha and
Schistolobos are taken into account). A strongly supported clade comprising samples of Oreocharis mixed
with species from several other genera was already found by Möller et al. (2011). In that work the clade
included 29 samples of 28 species. Here, we increase the sampling in this Oreocharis-dominated clade to 43
samples, representing 39 species. The clade, and the new generic definition based upon it, is characterised by
MÖLLER ET AL.2 Phytotaxa 23 © 2011 Magnolia Press
rather uniform vegetative habit and fruit characters, but high diversity in floral traits, particularly in corolla
shape and colour, and in stamen fertility.
We use molecular data of two gene regions, the chloroplast trnL-F intron-spacer and the nuclear
ribosomal ITS regions, for the reconstruction of phylogenetic relationships. These have proven useful in many
previous analyses of Gesneriaceae at genus and species level (Möller et al. 1999, 2009, 2011, Zimmer et al.
2002, Smith et al. 2004, Roalson et al. 2005, Li & Wang 2007, Wei et al. 2010a, Weber et al. 2011a, b, c).
Prior to the analysis presented here, we included the newly added samples in the large matrices of Weber
et al. (2011a) to confirm that these fall in the Oreocharis-dominated clade. This top-down approach is critical,
particularly for an analysis of Old World didymocarpoid Gesneriaceae, where it is sometimes difficult to
predict which species or even genera to include in a particular analysis, since generic delineation, particularly
for those found in China, is poorly supported by morphology. Such a case was previously found in the clade
dominated by Lagarosolen Wang (1984: 11), which included species from seven genera and gave rise to a new
definition of Petrocodon (Weber et al. 2011b).
Material and Methods
Plant material
Silica gel dried leaf material for newly acquired sequences was obtained during recent field collections (Table
1). The bulk of the data came from previous work (Möller et al. 2009, 2011, Weber et al. 2011a). Data for
eight samples were newly acquired and seven downloaded from GenBank (Table 1). In total, 55 samples were
included in the analyses, representing 51 species (including one new species), one variety and three duplicate
species samples (Dayaoshania serrulata sp. ined., Oreocharis auricula, Paraisometrum mileense).
TABLE 1: List of the 55 didymocarpoid Gesneriaceae samples included in the phylogenetic analysis, including voucher number and
deposition, origin information and respective GenBank accession numbers. Names in bold denote generic type species.
Taxon Voucher number Deposited
in Origin trnL-F ITS or
ITS1 / ITS2
Aeschynanthus lancilimbus
W.T.Wang (1975: 64) Y.Z.Wang S-10868 PE China, unknown locality FJ501499 HQ632992
Aeschynanthus rhododendron Ridley
(1896: 500) P.Woods 600 [Cult.
RBGE 19680624] E Peninsular Malaysia,
Genting Highlands HQ632895 FJ501333
Aeschynanthus mi cranthus C.B.Clarke
(1883: v. 27) M.Möller MMO 01-79 E, WU China, Yunnan, Hekou
county FJ501500 -
Aeschynanthus mi cranthus C.B.Clarke
(1883: v. 27) A.Reid & J. Fernie 004
[Cult. RBGE 19951561] E China, Yunnan,
Xishuangbanna Da i Aut.
Pref.
- AF349218 /
AF349299
Agalmyla clarkei (Elmer) B.L.Burtt
(1968: 224) RBGE-PNH1999(P99)
13 [Cult. RBGE
19991911]
E Philippines, Leyte
Island, Mt. Lobi FJ501540 -
Agalmyla clarkei (Elmer) B.L.Burtt
(1968: 224) RBGE-PNH1997 IS26
[Cult. RBGE
19972530A]
E Philippines, Luzon,
Barangay Penicuason - FJ501360
Agalmyla paucipilosa Hilliard &
B.L.Burtt (2002: 80) Smith & Galloway 261 E Indonesia, Sulawesi, Mt
Rantemario HQ632893 HQ632990
Agalmyla sojoliana Hilliard &
B.L.Burtt (2002: 78) Smith & Galloway 321 E Indonesia, Sulawesi, Mt
Sojol HQ632894 HQ632991
...... continued on the next page
Phytotaxa 23 © 2011 Magnolia Press 3
A NEW DELINEATION FOR OREOCHARIS
TABLE 1 (continued)
Taxon Voucher number Deposited
in Origin trnL-F ITS or
ITS1 / ITS2
Ancylostemon aureus (Franch.)
B.L.Burtt (1958b: 305). M.Möller MMO 01-153 E, WU China, Yunnan,
Binchuan county FJ501505 FJ501336
Ancylostemon convexus Craib (1920a:
235) M.Möller MMO 01-176 E, WU China, Yunnan, Dali
county FJ501506 FJ501337
Ancylostemon humilis W.T.Wang
(1975: 100) Liang R.H.SC-YB PE China, Sichuan GU350665 GU350633
Ancylostemon mairei (H.Lév.) Craib
(1920b: 267) Liang R.H.YN-Qj PE China, Yunnan GU350689 GU350658
Ancylostemon rhombifolius K.Y.Pan
(1988a: 435) Liang, R.H. LRH-07-01 PE China, Sichuan GU350664 GU350632
Ancylostemon ronganensis K.Y.Pan
(1988a: 436) M.Möller MMO 06-776 E China, Guangxi, Rongan
county HQ632927 HQ633023
Bournea leiophylla (W.T.Wang)
W.T.Wang & K.Y.Pan in Wang et al.
(1990: 135)
Zhou X.R. ZXR-05-01 PE China, Fujian GU350676 GU350644
Bournea sinensis Oliver in Hooker
(1893: t. 2254) M.Möller MMO 08-
1329 E China, Guangdong,
Bolou county HQ632912 HQ633008
Briggsia longifolia Craib (1920a:
238) M.Möller MMO 08-
1239 E China, Yunnan,
Jingdong county HQ632934 HQ633030
Briggsia muscicola Craib (1920b:
264). Kew (1995-2229) K unknown origin FJ501548 FJ501366
Briggsia rosthornii (Diels) B.L.Burtt
(1958b: 306) Sino-American
Bryological Expedition,
no. 398 (US 229325)
US China, Guizhou,
Jiangkou county FJ501547 FJ501365
Briggsia stewardii Chun (1946: 303) M.Möller MMO 06-917 E China, Guangxi,
Shanjiang county HQ632926 HQ633022
Cyrtandra cumingi i C.B.Clarke (1883:
v. 263) G.Kokubugata 11134 TNS Japan, Ruykyus,
Iriomote Island HQ632905 HQ633002
Cyrtandra pendula Blume (1826: 768) A.Weber &
Anthonysamy 860730-1/
2 [Cult. HBV]
WU Peninsular Malaysia,
Negeri Sembilan, Kuala
Pilah distr., Jeram Toi
FJ501530 FJ501354
Cyrtandra pulchella W.P.Rich ex
A.Gray (1862: 38) Lorence 8525 PTBG Samoa Islands HQ632906 EU919941
Dayaoshania cotinifolia W.T.Wang
(1983a: 320) Q.M.Chuan 01 IBK China, Guangxi, Jinxiu
conuty HQ632914 HQ633010
Dayaoshania serrulata Yan Liu &
W.B.Xu sp.ined. 1 F.Wen 0901 IBK China, Guangxi,
Wuzhou City JF687580 JF697568
Dayaoshania serrulata Yan Liu &
W.B.Xu sp.ined. 2 F.Wen 2010-04 IBK China, Guangxi,
Wuzhou City JF697581 JF697569
Deinocheilos jiangxiense W.T.Wang
(1986: 4) M.Möller MMO 09-
1451 E China, Fujian, Jiangle
county HQ632933 HQ633029
Didymocarpus citrinus Ridley (1896:
508) P.Davis 69437 [Cult.
RBGE 19830510] E Peninsular Malaysia,
Perlis, Kedah Peak AJ492293 DQ912669
Didymocarpus purpureobracteatus
W.W.Smith (1912: 153) Y.Z.Wang 991106 PE China, Yunnan,
Pingbian county FJ501510 -
...... continued on the next page
MÖLLER ET AL.4 Phytotaxa 23 © 2011 Magnolia Press
TABLE 1 (continued)
Taxon Voucher number Deposited
in Origin trnL-F ITS or
ITS1 / ITS2
Didymocarpus purpureobracteatus
W.W. Smith (1912: 153) M.Möller MMO 01-70 E, WU China: Yunnan,
Pingbian county - DQ912676
Didymocarpus stenanthos C.B.Clarke
(1888: t. 1799) M.Möller MMO 01-156 E, WU China, Yunnan,
Binchuan county FJ501512 DQ912687
Isometrum farreri Craib (1920a:
250). Zhou Ping ZP 2010-020 E China, Shaanxi, Mian
county JF697585 JF697573
Isometrum lancifolium (Franch.)
K.Y.Pan (1986: 30) M.Möller MMO 09-
1624 E China, Sichuan,
Mianning county HQ632924 HQ633020
Isometrum lungshengense
(W.T.Wang) W.T.Wang & K.Y.Pan in
Wang et al. (1990: 187)
M.Möller MMO 06-916 E China, Guangxi,
Longsheng county HQ632917 HQ633013
Isometrum primuliflorum (Batalin)
B.L.Burtt (1960: 93) M.Möller MMO 09-
1605 E China, Sichuan, Danba
county HQ632923 HQ633019
Opithandra acaulis (Merr.) B.L.Burtt
(1958a: 303) M.Möller MMO 08-
1328 E China, Guangdong,
Zhaoqin county HQ632916 HQ633012
Opithandra burttii W.T.Wang in
Weitzman et al. (1998: 430) F.Wen 2010-05 IBK China, Guangdong,
Wengyuan county JF697582 JF697570
Opithandra dalzielii (W.W.Sm.)
B.L.Burtt (1958a: 303) F.Wen 2010-06 IBK China, introduced from
Guangdong, Shantou
county
JF697583 JF697571
Opithandra dinghushanensis
W.T.Wang (1987: 10) Lin Q.B. LQB06-01 PE China, Guangdong,
Zhaoqing GU350675 GU350643
Opithandra primuloides (Miq.)
B.L.Burtt (1956: 162) T.Tsuzuki [Cult. RBGE
19842178A] E Japan, unknown locality FJ501546 FJ501364
Opithandra sinohenryi (Chun)
B.L.Burtt (1958a: 303) M.Möller MMO 07-
1150 E China, Guangxi,
Fangcheng county HQ632913 HQ633009
Oreocharis argyrei a Chun ex K.Y.Pan
(1987: 283) M.Möller MMO 07-
1131 E China, Guangxi,
Wuming county HQ632919 HQ633015
Oreocharis argyreia var. angustifolia
K.Y.Pan (1987: 285) Liang R.H. GX-SS-01 PE China, Guangxi GU350671 GU350639
Oreocharis aurea Dunn (1908b: 19) M.Möller MMO 06-980 E China, Yunnan, Jinping
county HQ632920 HQ633016
Oreocharis auricula 1 (S.Moore)
C.B.Clarke (1883: 64) M.Möller MMO 03-304 E China; Guizhou,
Jiangkou county FJ5011481 FJ501323
Oreocharis auricula 2 (S.Moore)
C.B.Clarke (1883: 64) Liang R.H. GX-LG-02 PE China; Guangxi GU350672 GU350640
Oreocharis benthamii C.B.Clarke
(1883: 63) M.Möller MMO 08-
1317 E China, Guangdong,
Zhao Qin JF697584 JF697572
Oreocharis dasyantha var.
ferruginosa K.Y.Pan (1987: 283) Y.G.Wei 07-700 E China, Hainan, Delong HQ632918 HQ633014
Oreocharis henryana Oliver in
Hooker (1890: t. 1944) M.Möller MMO 10-
1691 E China, Sichuan, Lushan
county JF697586 JF697574
Oreocharis magnidens Chun ex
K.Y.Pan (1987: 276) M.Möller MMO 06-896 E China, Guangxi, Jinxiu
county HQ632930 HQ633026
...... continued on the next page
Phytotaxa 23 © 2011 Magnolia Press 5
A NEW DELINEATION FOR OREOCHARIS
In previous analyses the Oreocharis-dominated clade was shown to be highly supported and had the
longest branch length leading to the clade (Möller et al. 2011, Weber et al. 2011a). To ensure the newly
acquired sequences fell in the Oreocharis-dominated clade, we used a top-down approach and added these to
the comprehensive didymocarpoid Gesneriaceae matrix of Weber et al. (2011a; in total then comprising 259
samples), and found that all 15 newly added samples fell inside the Oreocharis-dominated clade (data not
shown). This gave us confidence in the validity of our analysis here, which is focussed on the Oreocharis-
dominated clade. Based on the results from the extended analysis and previously published data (Möller et al.
2009, 2011, Weber et al. 2011a), we selected three species each of the genera Agalmyla Blume (1826: 766),
Aeschynanthus Jack (1823: 42), Cyrtandra Forster & Forster (1776: t. 3) and Didymocarpus Wallich (1819:
378) as outgroup samples, and the trees were rooted on Didymocarpus.
The ingroup comprised 43 samples. These belonged to 11 genera. For 10 genera all samples included fell
within the ingroup; Ancylostemon (6 included/12 described), Bournea (2/2), Dayaoshania (2/2), Deinocheilos
(1/2), Isometrum (4/14), Opithandra (6/10), Oreocharis (8/28), Paraisometrum (1/1), Thamnocharis (1/1),
and Tremacron (4/7). Only four of the eight included species of Briggsia in previous work (out of 22
described) fell in the Oreocharis-dominated clade (B. longifolia, B. muscicola, B. rosthornii, B. stewardii)
(Möller et al. 2011).
DNA extraction, PCR, and sequencing
The molecular methods followed previously published protocols (Möller et al. 2009, 2011, Weber et al.
2011a). For all samples sequences of both the trnL-F intron-spacer (trnL-F) and the ITS region were available
for analysis. Newly acquired sequences were deposited in GenBank.
Phylogenetic analysis
Our top-down-approach taken here has several advantages: (a) the inclusion of all species and genera that
belong to the Oreocharis-clade. In Möller et al. (2009) samples of Ancylostemon, Briggsia, Opithandra and
Oreocharis were first implicated in this clade. In Wang et al. (2010) only species of Bournea, Isometrum and
Thamnocharis were additionally included without explicit justification. Only comprehensive analyses (e.g.
TABLE 1 (continued)
Taxon Voucher number Deposited
in Origin trnL-F ITS or
ITS1 / ITS2
Oreocharis xian gguinensis W.T.Wang
& K.Y.Pan in Pan (1987: 285) M.Möller MMO 05-741 E China, Guangxi, Lingui
county HQ632932 HQ633028
Paraisometrum mileense 1
W.T.Wang in Weitzman et al.
(1998: 434)
Y.M.Shui 65214 KUN China, Yunnan, Shilin
county HQ632928 HQ633024
Paraisometrum mileense 2
W.T.Wang in Weitzman et al.
(1998: 434)
Y.G.Wei 0901 IBK China, Guangxi, Longlin JF697587 JF697575
Thamnocharis esquirolii (H.Lév.)
W.T.Wang (1981: 485) D.W.Zhang 723 IBK China, Guizhou, Anlong
county HQ632915 HQ633011
Tremacron aurantiacum K.Y.Pan
(1988a: 431) Voucher from Cult.
RBGE 20060865 E China, unknown locality HQ632925 HQ633021
Tremacron begoniifolium H.W.Li
(1983: 12) M.Möller MMO 08-
1221 E China, Yunnan,
Jingdong county HQ632929 HQ633025
Tremacron forrestii Craib (1918: 217) M.Möller MMO 07-
1072 E China, Sichuan, Pan Zhi
Hua city HQ632921 HQ633017
Tremacron urceolatum K.Y.Pan
(1988a: 429) M.Möller MMO 09-
1633 E China, Sichuan,
Liangshan Yizu county HQ632922 HQ633018
MÖLLER ET AL.6 Phytotaxa 23 © 2011 Magnolia Press
Möller et al. 2011, Weber et al. 2011a, data not shown) including all but one genus [i.e. Deinostigma Wang &
Li (1992: 356)] of the advanced didymocarpoid Gesneriaceae, showed the full extent of the genera belonging
to the Oreocharis clade in that species of an additional four genera (i.e. Dayaoshania, Deinocheilos,
Paraisometrum and Tremacron) also need to be considered; (b) the inclusion of only the species relevant for
the Oreocharis clade (i.e. only a few but not all samples of Briggsia); (c) the inclusion of all molecular
characters (i.e. hypervariable regions that had to be excluded in the larger analyses due to alignment
ambiguities, particularly in the ITS spacer region); and (d) due to the above more accurate phylogenetic
reconstruction results. The narrower taxonomic range of the reduced Oreocharis-focussed analyses here
allowed a re-alignment of the matrices and a more accurate determination of primary sequence homologies,
reducing the homoplasy resulting in fewer parsimony trees retrieved.
Maximum parsimony (MP) and Bayesian inference (BI) analyses were performed as described previously
(Möller et al. 2009, 2011, Weber et al. 2011a), using PAUP* v4.0b10 (Swofford 2002), and MrBayes v3.1.2
(Huelsenbeck & Ronquist 2001, 2007). Only combined trnL-F and ITS sequences were analysed. Their
combinability was tested using the incongruence length difference (ILD; P=0.52) test implemented as the
partition homogeneity test (PHT) in PAUP*. Alignment gaps with consistent boundaries were coded as
additional characters according to the simple method of Simmons & Ochoterena (2000), 9 in trnL-F, 12 in
ITS. MP branch support came from bootstrap analyses as described previously (Möller et al. 2009, 2011) with
10000 replicates, TBR on and MulTrees off.
Best-fitting models for MrBayes were obtained separately for trnL-F, the ITS spacers and the 5.8S gene
through AIC in MrModeltest 2.3 (Nylander 2004), and were GTR+G, GTR+G and SYM+I, respectively. The
gaps were treated as standard characters. The analysis was run for 2.5 million generations for the 55 sample
matrix, with a burn-in of 4% determined by plotting likelihoods against generations (Appendix 1). Posterior
probabilities (PP) were taken from MrBayes 50% majority rule consensus trees using the ‘sumt’ command.
The PP branch support values showed a high correlation between the two parallel Bayesian runs (Appendix
1).
Results
Matrix characteristics
The 55 sample matrix was 1632 characters long (trnL-F: 870 characters, ITS: 762) which included 364
(22.3%) parsimony informative characters.
Phylogenetic analyses
The MP analysis on combined data resulted in 70 most parsimonious trees of 1245 steps (CI=0.6321,
RI=0.7243). The resulting majority rule consensus tree was highly resolved (Fig. 1). The phylogram
illustrates the distribution of genetic diversity and the primarily short backbone branches in the Oreocharis
clade (Fig. 2). The BI tree was less resolved but fully congruent (virtually identical) where the MP branches
received bootstrap support (Fig. 3). MP branches with bootstrap support <50% collapsed in the BI tree.
The monophyly of each of the outgroup genera was highly supported (BS=98-100%; PP=1.00), although
the intergeneric relationships between Aeschynanthus, Agalmyla and Cyrtandra received no or low support
values and the relationships differed between the MP and BI analysis. The Oreocharis clade received
maximum branch support (BS=100%; PP=1.00). Within the clade the backbone structure received no MP
branch support or collapsed to a polytomy in the BI analysis.
Opithandra acaulis and O. dinghushanensis were sister (BS=100%; PP=1.00), and were resolved as sister
to the rest of the samples only in the MP analysis, but with no BS support. In the BI analysis this clade fell on
a backbone polytomy with several other lineages or individual samples, including the type species Oreocharis
benthamii. In the MP analysis the Oreocharis type formed a grade between the Opithandra clade and two
larger clades, Clade I and II. However, these received no support.
Phytotaxa 23 © 2011 Magnolia Press 7
A NEW DELINEATION FOR OREOCHARIS
FIGURE 1: Majority rule consensus tree of 70 most parsimonious trees, based on combined trnL-F and ITS data plus gap characters
(tree length 1245 steps, CI = 0.6321, RI = 0.7243). Numbers along branches, above are majority rule frequencies, below (bold, italics)
bootstrap values, * indicates branches with <50% bootstrap values. Number of fertile stamens (stam.: a, anterior pair; p, posterior
pair), corolla shape, colour and anther coherence (anther) indicated in box.
In the MP Clade I, Briggsia stewardii and Ancylostemon ronganensis were strongly supported as a sister
pair (BS=100%; PP=1.00), as were the three samples of Dayaoshania (BS=95%; PP=1.00). The two
Dayaoshania serrulata sp. ined. samples were closest (BS=100%; PP=1.00) in this genus. The two varieties
of Oreocharis argyreia included were sister (BS=100%; PP=1.00). A group of samples forming a highly
supported clade Ib (BS=97%; PP=1.00), included Bournea sinensis, Deinocheilos jiangxiense, two
Opithandra and three Oreocharis species. Within this clade the two Opithandra, O. burttii and O. dalzielii,
formed a sister pair with high branch support (BS=99%; PP=1.00). The two Oreocharis auricula samples plus
O. xiangguinensis formed a highly supported clade (BS=99%; PP=1.00), with Bournea sinensis as sister
(BS=99; PP=1.00).
Within the MP Clade II, few relationships received branch support. Clade IIc, with 19 samples from 6
genera (Ancylostemon, Briggsia, Isometrum, Oreocharis, Paraisometrum, Tremacron) received some branch
support, high in the BI analysis (BS=58%; PP=1.00). Within this clade, the BI tree showed higher resolution
with medium branch support. Oreocharis aurea and Briggsia longifolia fell on a polytomy sister to the 17
remaining samples (PP=0.79). On the next grade with medium support (PP=0.80) fell a well supported clade
(BS=70%; PP=1.00) with six samples. Within this BI clade, Oreocharis henryana and Ancylostemon mairei
MÖLLER ET AL.8 Phytotaxa 23 © 2011 Magnolia Press
were sister (BS=71%; PP=0.95), as were two Tremacron species, T. forrestii and T. urceolatum (BS=68%;
PP=0.92). The remaining 11 samples formed a clade, only weakly supported in the BI analysis (PP=0.80).
Among these samples, only few relationships received high support; the two Paraisometrum mileense
samples (BS=100%; PP=1.00), Isometrum farreri and I. primuliflorum (BS=100%; PP=1.00) and four of the
six Ancylostemon species here included (BS=100%; PP=1.00). Within this Ancylostemon clade, A. aureus and
A. convexus were sister (BS=84%; PP=1.00), as were A. humilis and A. rhombifolius (BS=56%; PP=0.69).
The Bayesian posterior probabilities were often higher than the bootstrap support values (see also Möller et al.
2009).
FIGURE 2: One of 70 most parsimonious trees depicted as a phylogram to illustrate the length of the backbone branches. Shaded area
indicates period of rapid radiation.
Phytotaxa 23 © 2011 Magnolia Press 9
A NEW DELINEATION FOR OREOCHARIS
FIGURE 3: Bayesian inference tree with average branch lengths, based on combined trnL-F and ITS data plus gap characters.
Numbers above branches indicate posterior probabilities. Shaded area indicates period of rapid radiation.
Discussion
In the present molecular phylogenetic study species of no fewer than 11 extant Chinese gesneriad genera
(even 14, if genera in synonymy, Dasydesmus, Perantha and Schistolobos, are taken into account, see formal
MÖLLER ET AL.10 Phytotaxa 23 © 2011 Magnolia Press
treatment) form a monophyletic clade, the Oreocharis clade. This clade has maximum statistical branch
support. Within the clade, the well-supported species relationships do not conform to traditional generic
delimitations. The only exception is Dayaoshania, whose two species appear in a sister relationship.
However, the results of the phylogenetic study do not necessarily mean that the whole alliance should be
combined into a single genus. At first sight it may appear undesirable to do so, since it would result in a large
and, with respect to flower morphology, very variable genus of over 80 species. Therefore, our molecular
results require a detailed justification for such a measure. We justify our conclusions through: (1) molecular
analyses, (2) examination of weaknesses in the traditional generic delimitations, (3) an evaluation of
morphological characters hitherto used for generic definitions, and (4) a discussion of relationships between
particular species. We conclude that the overall results clearly indicate that the reduction of all these genera
under Oreocharis, the oldest generic name in the alliance, is indeed the most sensible solution.
(1) Molecular phylogenetic analyses
The present molecular analysis demonstrates that all species (investigated) of the genera Oreocharis (28
species / 8 included in the analysis), Ancylostemon (12/6), Opithandra (10/6), Isometrum (14/4), Tremacron
(7/4), Dayaoshania (2/2), Deinocheilos (2/1), Bournea (2/2), Thamnocharis (1/1) and Paraisometrum (1/1)
fall into the Oreocharis clade. In addition, four species of Briggsia, all with yellow flowers, were retrieved in
this clade, but other species of that genus (usually with different flower colours) were found in five other
places in the analyses of Möller et al. (2009, 2011). This suggests that the present delineation of Briggsia (c.
22 spp.) is highly artificial and needs to be resolved.
The present analysis includes sequence data for nearly half the species described in these genera (39 of
80). Given that we included most of the morphological diversity of these genera, the addition of further taxa,
though desirable, is not likely to alter our conclusions.
All non-monotypic genera with multiple samples here included were found to be non-monophyletic
(except the two Dayaoshania species), with mixed-generic clades often receiving high branch support (Figs.
1, 3). This is a further strong indication that the addition of more samples will not likely increase the level of
monophyly among the 11 genera in the Oreocharis clade. We are thus confident that our systematic
conclusions and taxonomical consequences are justified.
(2) Weaknesses of traditional delimitations
The weaknesses of delimiting the core genera of the present alliance (Oreocharis, Opithandra, Briggsia,
Ancylostemon, Isometrum, Tremacron) are well known. Before addressing these in some detail, it may be
useful to briefly survey their taxonomic history. This started with the original wide definition of
Didymocarpus, which included species with di- and tetrandous flowers, coherent or free anthers and a huge
geographical range in Asia (later extended to Africa, Madagascar and Australia) (see Weber & Burtt 1998b).
Bentham (1876) suggested that Didymocarpus oreocharis and two other species be moved to his new genus
Oreocharis. The formal transfer was not made by Bentham himself, but by Clarke (1883) who accepted
Oreocharis as a genus with four fertile and non-coherent anthers and established the new name Oreocharis
benthamii for Didymocarpus oreocharis. Clarke (1883) also separated another alliance from Didymocarpus,
also with four stamens, but these cohering in pairs, or all together. This was named Didissandra Clarke (1883:
65). The seven species included were very heterogeneous and Clarke (1883) established no fewer than four
sections for their accommodation. Ridley (1905), working with the Malesian species of Didissandra, was the
first to suggest the likely artificial nature of the genus, but he did not take any action to rectify it. Craib (1918,
1920a, b), working with the species in the northern part of the distribution range (N Thailand, China), split
Didissandra up into smaller entities, including Briggsia, Ancylostemon, Isometrum and Tremacron. Genera
established later (Opithandra by Burtt 1956, Thamnocharis by Wang 1981) have their roots either in
Oreocharis (Bournea was described by Oliver in Hooker 1893 as an independent genus, but also placed in the
vicinity of Oreocharis), or in Didissandra (Paraisometrum by Wang in Weitzmann et al. 1998). The essential
difference of the two lineages is the non-coherence vs. coherence of the anthers. The only genus described
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A NEW DELINEATION FOR OREOCHARIS
independently from these two lines was Dayaoshania, which was thought to be allied to Petrocosmea Oliver
in Hooker (1887: t. 1716) (Wang 1983a). However, the molecular data unequivocally places the genus in the
Oreocharis clade.
The difficulties in separating the genera of the alliance under consideration were clearly seen by Wang et
al. (1998). Statements such as “Using only vegetative or fruit characters, it [Oreocharis rhytidophylla] cannot
be placed definitely in Oreocharis, since other genera (e.g. Tremacron, Isometrum and Ancylostemon) are
virtually indistinguishable in these characters” or “Oreocharis is related to Ancylostemon, Tremacron and
Isometrum. As noted above it can be difficult to distinguish these genera, and some species, notably O.
delavayi [Franch.], are intermediate between them. Oreocharis delavayi has exserted stamens and adaxial
corolla lobes much smaller than abaxial ones, characters which relate it to Tremacron” (Wang et al. 1998:
252) give testimony to the difficulties of placing particular species correctly.
The difficulty of separating Briggsia from Oreocharis was addressed by Burtt (1954a: 188) as follows:
“There are some fourteen species in this genus [Briggsia], but they are of varied habit and appearance, and are
associated entirely on their floral characters: large, ventricose corollas, arched filaments and anthers cohering
in pairs tip to tip. Even this last feature is not quite constant, for we now have species in cultivation which
have all the characteristics of Briggsia except that on dehiscence the anthers separate and the filaments
straighten. There is, then, at full flowering, very little to separate these from Oreocharis”.
In several genera sections have been established (Opithandra: 5 sections for some 10 species, Oreocharis:
4, Isometrum: 3), sometimes based on characters (such as anther coherence, see below) otherwise used for
generic delimitation. Some sections are monotypic [Opithandra sect. Microstigma Wang (1987: 9),
Opithandra sect. Stenosiphon Wang (1987: 9), Opithandra sect. Schistolobos Wang (1992: 291), Isometrum
sect. Chorianthera W.T.Wang & K.Y.Pan in Pan (1988a: 432)], and Opithandra sect. Schistolobos was
previously described as a genus. Thus, the question arises, what is the difference between the monotypic
sections and the monotypic genera? The answer is simple: there are a number of rather isolated species that
cannot be placed satisfactorily and only make sense in a widely defined genus.
(3) Character evaluation
As stated above, the genera of the Oreocharis clade scarcely differ in vegetative habit and in fruit
characteristics (loculicidal, bivalved capsules, usually long and cylindrical, occasionally ovoidal). All are
rosette plants (with spirally arranged leaves, Fig. 4) with scapose, axillary inflorescences. The distinctive
features are in the flower, particularly in corolla shape and colour, and in stamen morphology (number and
placement of fertile stamens, anther coherence, anther shape and dehiscence). Traditional taxonomy is heavily
based on floral characters and this has remained so in Old World Gesneriaceae until now. Through the work of
Wiehler (1983), and manifold molecular studies, the situation is otherwise in Neotropical Gesneriaceae (e.g.,
Kvist & Skog 1996, Skog & Kvist 2000, Perret et al. 2001, 2003, Roalson et al. 2002, 2003). In the New
World floral similarity lost its paramount importance for defining genera as it became increasingly clear that
floral characters are evolutionarily plastic (and, therefore, taxonomically unreliable) due to their link to the
functional constraints of pollination. The floral characters of the Oreocharis alliance should be reconsidered
in the same light.
Clade IIc includes all taxa with yellow flowers. This suggests that this trait has evolved once with perhaps
a few reversals to purple. However, the flower and anther morphologies are highly diverse (see below).
Preliminary investigations on seed testa cell ornamentations revealed that the clade includes seeds with striate,
reticulate and verruculose patterns, while all taxa in clade I which have been investigated have verruculose
testa cells (data not shown).
The reason for this lack of character correlation possibly lies in the early evolution of the Oreocharis
clade. The short backbone branches (Figs. 2, 3) support a hypothesis of rapid radiation early on in the
diversification of the lineage. This is not conducive for a detectable phylogenetic hierarchy to become
established. Rather, this clade has been shaped by a mixture of independently evolving characteristics due to
selection pressures from different pollinators, often several times independently.
MÖLLER ET AL.12 Phytotaxa 23 © 2011 Magnolia Press
FIGURE 4: Examples of habit of various taxa belonging to the Oreocharis clade: A) Ancylostemon ronganensis; B) Briggsia
stewardii; C) Bournea leiophylla; D) Oreocharis magnidens; E) Dayaoshania cotinifolia; F) dehisced fruit of Deinocheilos
jiangxiensis.
Flower symmetry and corolla features
Corolla symmetry, shape and coloration are the most important elements to define pollination syndromes.
Among the samples included in the Oreocharis clade there are narrow- and wide-tubed, laterally flattened,
urceolate and radially symmetrical corolla forms, predominantly purple or yellow (Figs. 5, 6). These features
suggest different pollination syndromes. It is difficult to discuss the evolution of these syndromes across the
phylogenetic tree, as the internal branches are short and receive only low branch support. There is some
probability that the yellow flower colour has evolved from a purple colour, since a subclade (clade IIc) in
derived position includes all yellow-flowered taxa (plus a few purple ones), from five genera (Ancylostemon,
Briggsia, Oreocharis, Paraisometrum, Tremacron). However, these include a considerable variety of flower
shapes (narrow and wide tube, urceolate, laterally flattened), suggesting that the adaptation to different
pollinators has played a strong role in the diversification of the taxa.
A very similar situation is encountered in the African genus Streptocarpus Lindley (1828: pl. 1173) The
Streptocarpus phylogeny shows similar short internal and backbone branches (Möller & Cronk 2001) and the
floral diversity suggests six different pollination syndromes, having sometimes clearly evolved in parallel
(Harrison et al. 1999, Hughes et al. 2006). Nobody has suggested splitting Streptocarpus into several genera.
In order to provide a consistent classification for Gesneriad genera, we suggest accepting a wider range of
floral morphological variation within genera and defining Oreocharis to include the florally divergent
lineages.
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A NEW DELINEATION FOR OREOCHARIS
FIGURE 5: Examples of floral diversity arranged by phylogenetic relationships of various taxa belonging to the Oreocharis Clade I:
A) Opithandra dinghushanensis; B) Oreocharis benthamii; C) Thamnocharis esquirolii; D) Ancylostemon ronganensis; E) Briggsia
stewardii; F) Dayaoshania cotinifolia; G) Isometrum lungshengense; H) Oreocharis argyreia; J&K) Deinocheilos jiangxiensis; L)
Opithandra burttii; M) Oreocharis magnidens; N) Oreocharis auricula.
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FIGURE 6: Examples of floral diversity arranged by phylogenetic relationships of various taxa belonging to the Oreocharis Clade II:
A) Opithandra sinohenryi; B) Oreocharis dasyantha var. ferruginosa; C) Bournea sinensis; D) Oreocharis aurea; E) Briggsia
longifolia; F) Tremacron aurantiacum; G) Oreocharis henryana; H&J) Tremacron begoniifolium; K&L) Tremacron urceolatum; M)
Briggsia rosthornii; N) Paraisometrum mileense; O) Isometrum primuliflorum.
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A NEW DELINEATION FOR OREOCHARIS
Number of fertile stamens
The taxa of the Oreocharis clade include species with two (Dayaoshania, Deinocheilos, Opithandra), five
(Bournea, Thamnocharis) and four fertile stamens (rest). The diandrous taxa differ in the position of the fertile
stamen pair in the flower. In Dayaoshania and Deinocheilos they are in the anterior position, while those in
Opithandra are in the posterior position. Posterior stamen fertility is a character that has evolved also outside
the Oreocharis clade in the South American Sarmienta Ruiz & Pavon (1794: 4) and the Asian/African
Epithema Blume (1826: 737).
Even taking into account that some relationships in the Oreocharis clade are not well supported, there is
evidence that the stamen position is a variable character, since a highly supported clade (clade Ib, Fig. 1)
includes taxa with four different androecial conditions, with five (Bournea), four (Oreocharis), two anterior
(Deinocheilos) respectively two posterior (Opithandra) stamens. This indicates that differences in the
androecium are not good characters at the generic level. Precedents for currently recognized monophyletic
genera showing variation in stamen characteristics are available and include Rhynchoglossum Blume (1826:
741) and Agalmyla (Burtt 1962, Hilliard & Burtt 2002). In the African Acanthonema strigosum Hooker (1862:
t. 5339) this characteristic is variable even within the species; different flowers on individual plants can have
two or four fertile stamens.
The possibly parallel stamen reduction in (and thus possible polyphyly of Opithandra has been explicitly
addressed by Burtt (1958a: 301), when adding some more species to his originally monotypic Opithandra):
“In leaf characters and in the form of the stigma, however, there is variation and it cannot be gainsaid that
Opithandra is a somewhat artificial genus with the single character of fertile posticous stamens the one
feature of paramount importance” and “It may well be suggested that these species have all been derived
independently from different tetrandrous ancestors, and consequently that the genus Opithandra as here
constituted is quite artificial”. The molecular data prove conclusively that Opithandra is indeed polyphyletic,
the species being scattered over clade I.
Free or fused anthers
The coherence of anthers is an important aspect of the functioning of the flower. They facilitate a targeted
deposition of pollen on the pollinator (e.g. head of birds in Aeschynanthus), or guide the amount of pollen
deposited (e.g. virtually all pollen is deposited on a single visit by long-tongued flies in Streptocarpus, MM
pers. observ.). Free anthers are often observed for flowers with constricted corollas (Deinocheilos, Tremacron)
or radially or almost radially symmetrical flowers (Bournea, Thamnocharis, some Oreocharis and
Isometrum). These characteristics have been used as generic (Bournea, Thamnocharis, Tremacron) or
sectional descriptors (Isometrum, Oreocharis, Opithandra), but as our phylogeny indicates, appear to be
highly plastic, and unsuited as characters for generic delimitation in the Oreocharis clade.
(4) Relationships between selected species within the Oreocharis clade
The artificial nature of the current generic system can be illustrated by statistically well supported examples of
intergeneric sister relationships. For example, Ancylostemon ronganensis and Briggsia stewardii have
identical trnL-F sequences and differ only by 4 nucleotide changes in ITS but share 2 ITS insertions unique in
the Oreocharis clade. They are genetically certainly closely related and morphologically difficult to
distinguish (Fig. 5D, E) especially in vegetative habit (Fig. 4A, B). Their fruits, unknown when they were first
described, are now known and do not differ significantly. The differences that are recorded are that B.
stewardii has 2 bracts (in A. ronganensis “apparently absent”), corolla blue to pale purple (v. pink), and
filaments sparsely glandular pubescent (v. glabrous), which can hardly be considered generic characters. It is
difficult to see why they were described in different genera, perhaps incomplete descriptions or specimens
masked the similarities.
Among the taxa in the yellow flowered clade is the monotypic Paraisometrum with an interesting history.
It was rediscovered after 100 years of absence from herbarium records (Shui & Jie 2007). W.T. Wang noticed
that a herbarium specimen collected in 1898 from the southeast of Yunnan was distinctive with regard to the
MÖLLER ET AL.16 Phytotaxa 23 © 2011 Magnolia Press
corolla lobes and stamen arrangements from other genera and with colleagues established a new genus
(Weitzman et al. 1998). The corolla has four upper lobes and one lower lobe and the anthers are “coherent in
pairs by adaxial surfaces, thecae parallel” (Wang et al. 1998). However, photographs of corollas of
Ancylostemon saxatilis in Li & Wang (2004) clearly show 4 upper lobes and one lower lobe [as do the flowers
of Calcareoboea C.Y.Wu in Li (1982: 241), a member of the Petrocodon clade, Weber et al. 2011b, fig. 3B].
In Ancylostemon, the anthers are described as “coherent in pairs, thecae divergent”. In our opinion, this does
not constitute a generic difference and we interpret this as support for our combination of these genera.
Taxonomic consequences
The molecular data, the well-known weaknesses in generic delimitations, and the unreliability of floral
characters for defining genera, have lead us to the conclusion that it is best to include the whole Oreocharis
clade into Oreocharis, which is the oldest name in the alliance. This circumscription of Oreocharis includes
all species of Ancylostemon, Bournea, Dayaoshania, Deinocheilos, Isometrum, Opithandra, Oreocharis,
Paraisometrum, Thamnocharis and Tremacron, but only four of Briggsia (including the type species). The
other Briggsia species cannot be placed generically at present and must await further studies. In the “formal
treatment” a list is given which includes the species of Oreocharis currently recognised and those here
transferred from the other genera to be included in Oreocharis. All are given in alphabetical order.
Outlook
It would be desirable to fully resolve, with statistical support, the relationships among the taxa of the
Oreocharis clade. However, simply adding more genes may not be the answer, as an example in the
Saxifragales suggests, unless a huge number of characters are added, in this case an increase from 9000 to
50000bp (Jian et al. 2008). How deep the problem might be lies in the density, or shortness of the branches
involved (or the period of time) of radiation. The shorter the branches the less likely it is that signals of a
possible phylogenetic hierarchy can be retrieved. In a very short burst of radiation no hierarchy may have
existed in the first place and any retrieved may be an artefact of factors such as long branch attraction (Siddall
& Whiting 1999). The backbone branches in the Oreocharis clade are very short (Figs. 2, 3), and it is not
certain whether a stable, robust phylogenetic hierarchy can ever be established.
Whether one can get to the stage of defining sections remains to be seen. Judging from a superimposition
of the sectional system of the largest genus, Oreocharis, the prospects are not promising, as only the members
of the type section Oreocharis and section Orthoanthera Pan (1987: 281) included in our analysis fall into a
“section”-clade. Members of Oreocharis section Stomactin (C.B.Clarke) Fritsch (1893–1894: 145) are
scattered in both main clades. Since the sections are based on corolla form (Pan 1987), and corolla form is
here found to be most unsuited for classification purposes, this is not a surprising result. A classification based
on phylogenetic hypotheses and character mapping (e.g. Palee et al. 2006) would be the most promising
approach.
Formal treatment
Oreocharis Bentham (1876: 995, 1021).
Lectotype: O. benthamii C.B.Clarke (selected by Burtt 1954b).
Heterotypic synonyms:
Bournea Oliver in Hooker (1893: tab. 2254). Type: B. sinensis Oliv.
Perantha Craib (1918: 212). Lectotype: P. forrestii Craib (selected by Burtt 1954b).
Tremacron Craib (1918: 217). Type: T. forrestii Craib
Ancylostemon Craib (1920a: 233). Lectotype: A. concavus Craib [= A. aureus (Franch.) B.L.Burtt] (selected
by Burtt 1954b).
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A NEW DELINEATION FOR OREOCHARIS
Briggsia Craib (1920a: 236), pro parte incl. type. Lectotype: B. longifolia Craib (selected by Burtt 1954b).
Isometrum Craib (1920a: 250). Lectotype: I. farreri Craib (selected by Burtt 1954b).
Dasydesmus Craib (1920a: 253). Type: D. bodinieri Craib
Opithandra Burtt (1956: 162). Type: O. primuloides (Miq.) B.L.Burtt
Thamnocharis Wang (1981: 485). Type: T. esquirolii (H.Lév.) W.T.Wang
Dayaoshania Wang (1983a: 319). Type: D. cotinifolia W.T.Wang
Schistolobos Wang (1983b: 16). Type: S. pumilus W.T.Wang
Deinocheilos Wang (1986: 1). Type: D. sichuanense W.T.Wang
Paraisometrum W.T.Wang in Weitzman et al. (1998: 431). Type: P. mileense W.T.Wang
The following list includes all species and infraspecific taxa hitherto described in Oreocharis and those
transferred to that genus on the basis of the present analysis. Arrangement is in alphabetical order. Notes are
given where appropriate.
Oreocharis acaulis (Merr.) Mich.Möller & A.Weber, comb. nov.
Basionym:—Chirita acaulis Merrill (1934: 47).
Homotypic synonym:—Opithandra acaulis (Merr.) B.L.Burtt (1958a: 303).
Distribution:—China (Guangdong).
Notes:—Type and only species of Opithandra sect. Microstigma.
Oreocharis amabilis Dunn (1908a: 362).
Distribution:—China (Yunnan).
Notes:—Referred to Oreocharis sect. Stomactin by Pan (1987), Wang et al. (1990) and Li & Wang (2004).
Oreocharis argyreia Chun ex Pan (1987: 283).
Distribution:—China (Guangxi, Guangdong).
Notes:—Referred to Oreocharis sect. Oreocharis by Pan (1987), Wang et al. (1990) and Li & Wang (2004).
Oreocharis argyreia var. angustifolia Pan (1987: 285).
Distribution:—China (Guangxi).
Oreocharis aurantiaca Franchet (1888: 716).
Basionym:—Perantha aurantiaca (Franch.) Pellegrin (1926: 873).
Heterotypic synonym:—Perantha forrestii Craib (1918: 213).
Distribution:—China (Yunnan).
Notes:—Referred to Oreocharis sect. Stomactin by Pan (1987), Wang et al. (1990) and Li & Wang (2004).
Oreocharis aurea Dunn (1908b: 19).
Distribution:—China (Yunnan), Vietnam.
Notes:—Referred to Oreocharis sect. Stomactin by Pan (1987), Wang et al. (1990), Li & Wang (2004) and
Wei et al. (2010b).
Oreocharis aurea var. cordato-ovata (C.Y.Wu ex H.W.Li) K.Y.Pan et al. in Weitzman et al. (1998: 430).
Basionym:—Oreocharis cordato-ovata C.Y.Wu ex Li (1983: 7).
Distribution:—China (Yunnan).
Oreocharis auricula (S.Moore) C.B.Clarke (1883: 64).
Basionym:—Didymocarpus auriculus Moore (1875: 229).
Homotypic synonym:—Perantha auricula (S.Moore) Pellegrin (1926: 873).
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Heterotypic synonyms:—Chirita sericea H.Lév. & Vaniot in Léveillé (1906: 428), non Ridley,
Didymocarpus sericeus Léveillé (1906: 427), non Ridley, Oreocharis sericea Léveillé (1911a: 329).
Heterotypic synonyms:—Oreocharis esquirolii Léveillé (1911b: 447), non Léveillé (1911a: 329), Oreocharis
leveilleana Fedde (1911: 64).
Distribution:—China (Anhui, Fujian, Guangdong, Guangxi, Guizhou, Hubei, Hunan, Jiangxi, Sichuan).
Notes:—Referred to Oreocharis sect. Stomactin by Pan (1987), Wang et al. (1990) and Li & Wang (2004).
Léveillé (1911b) erroneously gave two different species the same name, Oreocharis esquirolii. The mistake
was realised and corrected soon afterwards (Fedde 1911).
Oreocharis auricula var. denticulata Pan (1987: 276).
Distribution:—China (Fujian).
Oreocharis begoniifolia (H.W.Li) Mich.Möller & A.Weber, comb. nov.
Basionym:—Tremacron begoniifolium Li (1983: 12).
Distribution:—China (Yunnan).
Oreocharis benthamii C.B.Clarke (1883: 63, as: “benthami”).
Basionym:—Didymocarpus oreocharis Hance (1866: 230).
Distribution:—China (Guangdong, Guangxi, Hong Kong, Hunan, Jiangxi).
Notes:—Type species of Oreocharis. Referred to Oreocharis sect. Oreocharis by Pan (1987), Wang et al.
(1990) and Li & Wang (2004).
Oreocharis benthamii var. reticulata Dunn (1908a: 362).
Distribution:—China (Guangdong, Guangxi).
Oreocharis bodinieri Léveillé (1915b: 40).
Basionym:—Dasydesmus bodinieri (H.Lév.) Craib (1920a: 254).
Distribution:—China (Sichuan, Yunnan).
Notes:—Referred to Oreocharis sect. Oreocharis by Pan (1987), Wang et al. (1990) and Li & Wang (2004).
Type species of Dasydesmus.
Oreocharis bullata (W.T.Wang & K.Y.Pan) Mich.Möller & A.Weber, comb. nov.
Basionym:—Ancylostemon bullatus W.T.Wang & K.Y.Pan in Wang (1992: 289).
Distribution:—China (Yunnan).
Oreocharis burttii (W.T.Wang) Mich.Möller & A.Weber, comb. nov.
Basionym:—Opithandra burttii W.T.Wang in Weitzman et al. (1998: 430).
Distribution:—China (Jiangxi).
Notes:—Referred to Opithandra sect. Opithandra by Wang (1987; as “Opithandra sp. Burtt”) and Li & Wang
(2004).
Oreocharis cavalieri Léveillé (1909: 258).
Distribution:—China (Guizhou).
Notes:—Referred to Oreocharis sect. Orthoanthera by Pan (1987), Wang et al. (1990) and Li & Wang
(2004).
Oreocharis chienii (Chun) Mich.Möller & A.Weber, comb. nov.
Basionym:—Briggsia chienii Chun (1946: 300).
Distribution:—China (Anhui, Jiangxi, Zhejiang).
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Oreocharis cinerea (W.T.Wang) Mich.Möller & A.Weber, comb. nov.
Basionym:—Opithandra cinerea W.T.Wang (1982: 39).
Distribution:—China (Guizhou).
Notes:—Referred to Opithandra sect. Briggsioides by Wang (1987), Wang et al. (1990) and Li & Wang
(2004).
Oreocharis cinnamomea Anthony (1934: 200).
Distribution:—China (Yunnan).
Notes:—Referred to Oreocharis sect. Stomactin by Pan (1987), Wang et al. (1990) and Li & Wang (2004).
Oreocharis concava (Craib) Mich.Möller & A.Weber, comb. nov.
Basionym:—Ancylostemon concavus Craib (1920a: 234, as: “concavum”).
Heterotypic synonyms:—Roettlera aurea Franchet (1899b: 250), Didymocarpus aureus (Franch.) Diels
(1912b: 355), Ancylostemon aureus (Franch.) Burtt (1958b: 305).
Distribution:—China (Yunnan).
Notes:—Type species of Ancylostemon.
Oreocharis concava var. angustifolia (K.Y.Pan) Mich.Möller & A.Weber., comb. nov.
Basionym:—Ancylostemon aureus (Franch.) B.L.Burtt var. angustifolius Pan (1988a: 434).
Distribution:—China (Yunnan).
Oreocharis convexa (Craib) Mich.Möller & A.Weber, comb. nov.
Basionym:—Ancylostemon convexus Craib (1920a: 235).
Distribution:—China (Yunnan).
Oreocharis cordato-ovata C.Y.Wu ex Li (1983: 7) Oreocharis aurea var. cordato-ovata, see there.
Oreocharis cordatula (Craib) Pellegrin (1926: 873).
Basionym:—Perantha cordatula Craib (1918: 214).
Distribution:—China (Yunnan, Sichuan).
Notes:—Referred to Oreocharis sect. Stomactin by Pan (1987), Wang et al. (1990) and Li & Wang (2004).
Oreocharis cotinifolia (W.T.Wang) Mich.Möller & A.Weber, comb. nov.
Basionym:—Dayaoshania cotinifolia Wang (1983a: 320).
Distribution:—China (Guangxi).
Notes:—Type and hitherto only species of Dayaoshania. There is an additional species tentatively referred to
Dayaoshania, but not yet formally described: D. serrulata, sp. ined. (mentioned in Wei et al. 2010b: 152,
153). Two samples of this species have been included in the present analysis and they appear associated with
D. cotinifolia. It is suggested here that it should be formally published under Oreocharis.
Oreocharis craibii Mich.Möller & A.Weber, nom. nov.
Basionym:—Tremacron forrestii Craib (1918: 217).
Distribution:—China (Sichuan, Yunnan).
Notes:—Type species of Tremacron.
Oreocharis crenata (K.Y.Pan) Mich.Möller & A.Weber, comb. nov.
Basionym:—Isometrum crenatum Pan (1986: 27).
Distribution:—China (Hubei).
Notes:—Referred to Isometrum sect. Isometrum by Pan (1986), Wang et al. (1990) and Li & Wang (2004).
MÖLLER ET AL.20 Phytotaxa 23 © 2011 Magnolia Press
Oreocharis dalzielii (W.W.Sm.) Mich.Möller & A.Weber, comb. nov.
Basionym:—Chirita dalzielii Smith (1918: 171).
Homotypic synonym:—Opithandra dalzielii (W.W.Smith) B.L.Burtt (1958a: 303).
Distribution:—China (Fujian, Guangdong).
Notes:—Referred to Opithandra sect. Briggsioides by Wang (1987), Wang et al. (1990) and Li & Wang
(2004).
Oreocharis dasyantha Chun (1946: 287).
Distribution:—China (Hainan).
Notes:—Referred to Oreocharis sect. Orthoanthera by Pan (1987), Wang et al. (1990) and Li & Wang
(2004).
Oreocharis dasyantha var. ferruginosa Pan (1987: 283).
Distribution:—As for species.
Oreocharis delavayi Franchet (1888: 715).
Heterotypic synonym:—Oreocharis elliptica Anthony (1934: 200).
Heterotypic synonym:—Oreocharis elliptica var. parvifolia W.T.Wang & K.Y.Pan ex Pan (1987: 282).
Distribution:—China (Sichuan, Xizang, Yunnan).
Notes:—Wang et al. (1998) state that O. delavay is intermediate between Oreocharis, Ancylostemon,
Tremacron and Isometrum, approaching the latter particularly in the exserted stamens and adaxial corolla
lobes which are much smaller than the abaxial ones.
Oreocharis dentata A.L.Weitzman & L.E.Skog in Weitzman et al. (1998: 431).
Distribution:—China (Sichuan).
Notes:—The species is not explicitly referred to a section by the authors (Weitzman et al. 1998), but is said to
be most similar to O. aurantiaca and O. minor. Both belong to Oreocharis sect. Stomactin. Wang et al. (1990)
and Li & Wang (2004), therefore, placed it in that section.
Oreocharis dinghushanensis (W.T.Wang) Mich.Möller & A.Weber, comb. nov.
Basionym:—Opithandra dinghushanensis Wang (1987: 10).
Distribution:—China (Guangdong).
Notes:—Referred to Opithandra sect. Briggsioides by Wang (1987), Wang et al. (1990) and Li & Wang
(2004).
Oreocharis esquirolii Léveillé (1911a: 329).
Homotypic synonym:—Thamnocharis esquirolii (H.Lév.) Wang (1981: 485).
Distribution:—China (Guizhou).
Notes:—Flowers actinomorphic, 4- or 5 merous.
Oreocharis eximia (Chun ex K.Y.Pan) Mich.Möller & A.Weber, comb. nov.
Basionym:—Isometrum eximium Chun ex K.Y.Pan (1988a: 432).
Distribution:—China (Sichuan).
Notes:—Type and only species of sect. Isometrum sect. Chorianthera (Pan 1988a, Wang et al. 1990, Li &
Wang 2004).
Oreocharis fargesii (Franch.) Mich.Möller & A.Weber, comb. nov.
Basionym:—Didissandra fargesii Franchet (1899a: 123).
Homotypic synonym:—Isometrum fargesii (Franch.) Burtt (1960: 92).
Distribution:—China (Sichuan).
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A NEW DELINEATION FOR OREOCHARIS
Notes:—Referred to Isometrum sect. Pachysiphon Pan (1986: 26) by Pan (1986), Wang et al. (1990) and Li &
Wang (2004).
Oreocharis farreri (Craib) Mich.Möller & A.Weber, comb. nov.
Basionym:—Isometrum farreri Craib (1920a: 250).
Distribution:—China (Gansu, Shaanxi, Sichuan).
Notes:—Type species of Isometrum.
Oreocharis filipes Hance (1883: 166) Paraboea filipes (Hance) B.L.Burtt, see Burtt (1984).
Oreocharis flabellata (C.Y.Wu ex H.W.Li) Mich.Möller & A.Weber, comb. nov.
Basionym:—Ancylostemon flabellatus C.Y.Wu ex Li (1983: 16).
Distribution:—China (Yunnan).
Oreocharis flavida Merrill (1922: 354).
Distribution:—China (Hainan).
Notes:—Referred to Oreocharis sect. Platyanthera (Pan 1987: 289) by Pan (1987), Wang et al. (1990), Li &
Wang (2004) and Wei et al. (2010b).
Oreocharis fokienensis Franchet (1899c: 212) = Oreocharis maximowiczii C.B.Clarke, see there.
Oreocharis forrestii (Diels) Skan (1917: t. 8719).
Basionym:—Roettlera forrestii Diels (1912a: 224).
Distribution:—China (Sichuan, Yunnan).
Notes:—Referred to Oreocharis sect. Orthoanthera by Pan (1987), Wang et al. (1990) and Li & Wang
(2004).
Oreocharis gamosepala (K.Y.Pan) Mich.Möller & A.Weber, comb. nov.
Basionym:—Ancylostemon gamosepalus Pan (1988a: 434).
Distribution:—China (Sichuan).
Oreocharis georgei Anthony (1934: 202).
Distribution:—China (Yunnan, Sichuan).
Notes:—Referred to Oreocharis sect. Stomactin by Pan (1987), Wang et al. (1990) and Li & Wang (2004).
Oreocharis giraldii (Diels) Mich.Möller & A.Weber, comb. nov.
Basionym:—Didissandra giraldii Diels (1905: 98).
Homotypic synonym:—Isometrum giraldii (Diels) Burtt (1960: 92).
Distribution:—China (Shaanxi).
Notes:—Referred to Isometrum sect. Isometrum by Pan (1986), Wang et al. (1990) and Li & Wang (2004).
Oreocharis glandulosa (Batalin) Mich.Möller & A.Weber, comb. nov.
Basionym:—Didissandra glandulosa Batalin (1892: 175).
Homotypic synonym:—Isometrum glandulosum (Batalin) Craib (1920b: 267).
Distribution:—China (Gansu, Sichuan).
Notes:—Referred to Isometrum sect. Pachysiphon by Pan (1986), Wang et al. (1990) and Li & Wang (2004).
Oreocharis hekouensis (Y.M.Shui & W.H.Chen) Mich.Möller & A.Weber, comb. nov.
Basionym:—Ancylostemon hekouensis Y.M.Shui & W.H.Chen in Chen & Shui (2006: 448).
Distribution:—China (Yunnan).
MÖLLER ET AL.22 Phytotaxa 23 © 2011 Magnolia Press
Oreocharis henryana Oliver in Hooker (1890: tab. 1944).
Heterotypic synonym:—Oreocharis squamigera Léveillé (1915a: 24).
Distribution:—China (Gansu, Sichuan, Yunnan).
Notes:—Referred to Oreocharis sect. Orthoanthera by Pan (1987), Wang et al. (1990) and Li & Wang
(2004).
Oreocharis heterandra D.Fang & D.H.Qin in Fang et al. (1994: 563).
Distribution:—China (Guangxi).
Notes:—The species was not explicitly referred to a section by the authors, but was said to be allied to O.
aurea, which belongs to sect. Stomactin. Wang et al. (1990) and Li & Wang (2004) and Wei et al. (2010b),
therefore, placed it in that section.
Oreocharis hirsuta Barnett (1961: 9).
Distribution:—Thailand.
Notes:—Referred to Oreocharis sect. Oreocharis by Pan (1987), Wang et al. (1990) and Li & Wang (2004).
Oreocharis humilis (W.T.Wang) Mich.Möller & A.Weber, comb. nov.
Basionym:—Ancylostemon humilis Wang (1975: 100).
Heterotypic synonyms:Didissandra saxatilis var. microcalyx Hemsley (1890: 227), Ancylostemon saxatilis
(Hemsl.) Craib var. microcalyx (Hemsl.) Craib (1920b: 266).
Distribution:—China (Hubei, Sichuan).
Oreocharis jiangxiensis (W.T.Wang) Mich.Möller & A.Weber, comb. nov.
Basionym:—Deinocheilos jiangxiense Wang (1986: 4).
Oreocharis lancifolia (Franch.) Mich.Möller & A.Weber, comb. nov.
Basionym:—Didissandra lancifolia Franchet (1887–1888: 63).
Homotypic synonyms:—Isometrum lancifolium (Franch.) Pan (1986: 30). Ancylostemon lancifolius (Franch.)
Burtt (1958b: 305).
Heterotypic synonym:—Ancylostemon purpureus Burtt & Davidson (1954: 216).
Distribution:—China (Sichuan).
Notes:—Referred to Isometrum sect. Isometrum by Pan (1986), Wang et al. (1990) and Li & Wang (2004).
Oreocharis lancifolia var. mucronata (K.Y.Pan) Mich.Möller & A.Weber, comb. nov.
Basionym:—Isometrum lancifolium (Franch.) K.Y.Pan var. mucronatum Pan (1986: 30).
Distribution:—China (Sichuan).
Oreocharis leiophylla Wang (1975: 99).
Homotypic synonym:—Bournea leiophylla (W.T.Wang) W.T.Wang & K.Y.Pan in Wang et al. (1990: 135).
Distribution:—China (Fujian).
Notes:—Flowers actinomorphic, 5-merous.
Oreocharis leveilleana Fedde (1911: 64) = Oreocharis auricula (S.Moore) C.B.Clarke, see there.
Oreocharis leucantha (Diels) Mich.Möller & A.Weber, comb. nov.
Basionym:—Didissandra leucantha Diels in Pax (1922: 487).
Homotypic synonym:—Isometrum leucanthum (Diels) Burtt (1960: 92).
Distribution:—China (Sichuan).
Notes:—Referred to Isometrum sect. Isometrum by Pan (1986), Wang et al. (1990) and Li & Wang (2004).
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Oreocharis longifolia (Craib) Mich.Möller & A.Weber, comb. nov.
Briggsia longifolia Craib (1920a: 238).
Distribution:—China (Gansu, Sichuan, Yunnan), Myanmar.
Notes:—Type species of Briggsia. The type variety is found only in Yunnan and Myanmar.
Oreocharis longifolia var. multiflora (S.Y.Chen ex K.Y.Pan) Mich.Möller & A.Weber, comb. nov.
Basionym:—Briggsia longifolia var. multiflora S.Y.Chen ex Pan (1988b: 453).
Distribution:—China (Gansu, Sichuan).
Oreocharis lungshengensis (W.T.Wang) Mich.Möller & A.Weber, comb. nov.
Basionym:—Opithandra lungshengensis Wang (1975: 102).
Homotypic synonym:—Isometrum lungshengense (W.T.Wang) W.T.Wang & K.Y.Pan in Wang et al. (1990:
187).
Distribution:—China (Guangxi).
Notes:—Referred to Isometrum sect. Isometrum by Pan (1986), Wang et al. (1990) and Li & Wang (2004).
Oreocharis magnidens Chun ex Pan (1987: 276).
Distribution:—China (Guangxi).
Notes:—Referred to Oreocharis sect. Stomactin by Pan (1987), Wang et al. (1990) and Li & Wang (2004).
Oreocharis mairei Léveillé (1912: 301).
Basionym:—Tremacron mairei (H.Lév.) Craib (1918: 218).
Distribution:—China (Yunnan).
Notes:—This is not identical with Ancylostemon mairei, which is based on Didymocarpus mairei, described
in the same publication as Oreocharis mairei. Didymocarpus/Ancylostemon mairei is given the new name
Oreocharis wangwentsaii (see there).
Oreocharis maximowiczii C.B.Clarke (1883: 63).
Heterotypic synonym:—Oreocharis fokienensis Franchet (1899c: 212).
Distribution:—China (Fujian, Jiangxi).
Notes:—Referred to Oreocharis sect. Oreocharis by Pan (1987), Wang et al. (1990) and Li & Wang (2004).
Oreocharis micranthaveillé (1915a: 24) = Didymocarpus stenanthos C.B.Clarke var. stenanthos, see
Wang et al. (1990, 1998).
Oreocharis mileensis (W.T.Wang) Mich.Möller & A.Weber, comb. nov.
Basionym:—Paraisometrum mileense W.T.Wang in Weitzman et al. (1998: 434).
Oreocharis minor (Craib) Pellegrin (1926: 873).
Basionym:—Perantha minor Craib (1918: 213).
Distribution:—China (Sichuan, Yunnan).
Notes:—Referred to Oreocharis sect. Stomactin by Pan (1987), Wang et al. (1990) and Li & Wang (2004).
Oreocharis muscicola (Craib) Mich.Möller & A.Weber, comb. nov.
Basionym:—Briggsia muscicola Craib (1920b: 264).
Distribution:—China (Xizang, Yunnan), Bhutan, northeast India, Myanmar
Oreocharis nanchuanica (K.Y.Pan & Z.Y.Liu) Mich.Möller & A.Weber, comb. nov.
Basionym:—Isometrum nanchuanicum K.Y.Pan & Liu (1995: 100).
Distribution:—China (Sichuan).
MÖLLER ET AL.24 Phytotaxa 23 © 2011 Magnolia Press
Notes:—Referred to Isometrum sect. Pachysiphon by Pan (1986), Wang et al. (1990) and Li & Wang (2004).
Oreocharis nemoralis Chun (1946: 288).
Distribution:—China (Hunan, Guangdong).
Notes:—Referred to Oreocharis sect. Stomactin by Pan (1987), Wang et al. (1990) and Li & Wang (2004).
The type variety is found only in Hunan.
Oreocharis nemoralis var. lanata Zheng & Xia (2002: 34).
Distribution:—China (Guangdong).
Oreocharis notha Clarke (1883: 64). = Napeanthus primulifolius (Raddi) Sandw., see Leeuwenberg (1958).
Oreocharis notochlaena (H.Lév. & Vaniot) Léveillé (1911a: 330).
Basionym:—Didissandra notochlaena H.Lév. & Vaniot in Léveillé (1906: 425).
Homotypic synonyms:—Didymocarpus notochlaenus Léveillé (1906: 428; “notochlaena”), nom. inval., see
Vitek et al. (1998, 2000), Ancylostemon notochlaenus (H.Lév. & Vaniot) Craib (1920b: 266).
Distribution:—China (Guizhou).
Oreocharis obliqua C.Y.Wu ex Li (1983: 6).
Distribution:—China (Yunnan).
Notes:—Referred to Oreocharis sect. Stomactin by Pan (1987), Wang et al. (1990) and Li & Wang (2004).
Oreocharis obliquifolia (K.Y.Pan) Mich.Möller & A.Weber, comb. nov.
Basionym:—Tremacron obliquifolium Pan (1988a: 429).
Distribution:—China (Sichuan).
Oreocharis obovata Barnett (1961: 10). = Ridleyandra flammea (Ridl.) A.Weber, see Weber & Burtt
(1998a).
Oreocharis obtusidentata (W.T.Wang) Mich.Möller & A.Weber, comb. nov.
Basionym:—Opithandra obtusidentata Wang (1982: 38).
Distribution:—China (Hunan).
Notes:—Referred to Opithandra sect. Briggsioides by Wang (1987), Wang et al. (1990) and Li & Wang
(2004).
Oreocharis pankaiyuae Mich.Möller & A.Weber, nom. nov.
Basionym:—Tremacron aurantiacum Pan (1988a: 431).
Distribution:—China (Sichuan, Guizhou).
Notes:—The new name honours the Chinese botanist Pan Kai-Yu. Her many papers on Chinese Gesneriaceae
included revisions of Oreocharis (Pan 1987), Isometrum (Pan 1986) and Tremacron (Pan 1988a). The type
variety is found only in Sichuan.
Oreocharis pankaiyuae var. weiningense (S.Z.He & Q.W.Sun) Mich.Möller & A.Weber, comb. nov.
Basionym:—Tremacron aurantiacum var. weiningense S.Z.He & Q.W.Sun in He et al. (2010: 199).
Distribution:—China (Guizhou).
Oreocharis primuloides (Miq.) Benth. & Hook.f. ex Clarke (1883: 63).
Basionym:—Boea primuloides Miquel (1867: 190).
Homotypic synonyms:—Didymocarpus primuloides (Miq.) Maximowicz (1874: 536), Chirita primuloides
(Miq.) Ohwi (1936: 662), Opithandra primuloides (Miq.) B.L.Burtt (1956: 162).
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Heterotypic synonym:—Opithandra primuloides f. albiflora (Makino) Yamazaki (1993: 379).
Distribution:—Japan (Honshu, Shikoku, Kyushu).
Notes:—Type species of Opithandra and sect. Opithandra.
Oreocharis pumila (W.T.Wang) Mich.Möller & A.Weber, comb. nov.
Basionym:—Schistolobos pumilus Wang (1983b: 16).
Homotypic synonym:—Opithandra pumila (W.T.Wang) Wang (1992: 293).
Distribution:—China (Guangxi).
Notes:—Type and only species of Opithandra sect. Schistolobos (Wang 1992).
Oreocharis pinnatilobata (K.Y.Pan) Mich.Möller & A.Weber, comb. nov.
Basionym:—Isometrum pinnatilobatum Pan (1986: 34).
Distribution:—China (Hubei, Sichuan).
Notes:—Referred to Isometrum sect. Isometrum by Pan (1986), Wang et al. (1990) and Li & Wang (2004).
Oreocharis primuliflora (Batalin) Mich.Möller & A.Weber, comb. nov.
Basionym:—Didissandra primuliflora Batalin (1895: 176).
Homotypic synonym:—Isometrum primuliflorum (Batalin) Burtt (1960: 93).
Distribution:—China (Sichuan).
Notes:—Referred to Isometrum sect. Isometrum by Pan (1986), Wang et al. (1990) and Li & Wang (2004).
Oreocharis rhombifolia (K.Y.Pan) Mich.Möller & A.Weber, comb. nov.
Basionym:—Ancylostemon rhombifolius Pan (1988a: 435).
Distribution:—China (Sichuan).
Oreocharis rhytidophylla C.Y.Wu ex Li (1983: 9).
Distribution:—China (Yunnan).
Notes:—The status and sectional placement of this species is uncertain. It is known only from fruiting
specimens and, therefore, was omitted from the treatment in the Flora of China (Wang et al. 1998: 251).
Wang et al. (1998: 260) state that it is similar to Oreocharis benthamii var. reticulata.
Oreocharis ronganensis (K.Y.Pan) Mich.Möller & A.Weber, comb. nov.
Basionym:—Ancylostemon ronganensis Pan (1988a: 436).
Distribution:—China (Guangxi).
Oreocharis rosthornii (Diels) Mich.Möller & A.Weber, comb. nov.
Basionym:—Didissandra rosthornii Diels (1900: 574).
Homotypic synonym:—Briggsia rosthornii (Diels) Burtt (1958b: 306).
Heterotypic synonym:—Briggsia hians Chun (1946: 302).
Distribution:—China (Guizhou, Hubei, Sichuan, Yunnan).
Notes:—The type variety is found in Guizhou, Hubei, Sichuan.
Oreocharis rosthornii var. crenulata (Hand.-Mazz.) Mich.Möller & A.Weber, comb. nov.
Basionym:—Briggsia crenulata Handel-Mazzetti (1934: 20).
Homotypic synonym:—Briggsia rosthornii var. crenulata (Hand.-Mazz.) K.Y.Pan in Wang et al. (1990:
223).
Distribution:—China (Guizhou).
Oreocharis rosthornii var. wenshanensis (K.Y.Pan) Mich.Möller & A.Weber, comb. nov.
Basionym:—Briggsia rosthornii var. wenshanensis Pan (1988b: 456).
MÖLLER ET AL.26 Phytotaxa 23 © 2011 Magnolia Press
Distribution:—China (Yunnan).
Oreocharis rosthornii var. xingrenensis (K.Y.Pan) Mich.Möller & A.Weber, comb. nov.
Basionym:—Briggsia rosthornii var. xingrenensis Pan (1988b: 456).
Distribution:—China (Guizhou).
Oreocharis rotundifolia Pan (1987: 280).
Distribution:—China (Yunnan).
Notes:—Referred to Oreocharis sect. Stomactin by Pan (1987), Wang et al. (1990) and Li & Wang (2004).
Oreocharis rubra (Hand.-Mazz.) Mich.Möller & A.Weber, comb. nov.
Basionym:—Tremacron rubrum Handel-Mazzetti (1936: 877).
Distribution: China (Yunnan).
Oreocharis saxatilis (Hemsl.) Mich.Möller & A.Weber, comb. nov.
Basionym:—Didissandra saxatilis Hemsley (1890: 227).
Homotypic synonyms:—Didymocarpus saxatilis (Hemsl.) Léveillé (1906: 427), Ancylostemon saxatilis
(Hemsl.) Craib (1920b: 266).
Distribution:—China (Gansu, Hubei, Sichuan).
Oreocharis seguinii Léveillé (1906: 427, as: “seguini”), nom. nud. = Paraboea rufescens (Franch.) B.L.Burtt,
see Xu et al. (2008).
Oreocharis sericea Léveillé. (1911a: 329) = Oreocharis auricula (S.Moore) C.B.Clarke, see there.
Oreocharis sichuanensis (W.T.Wang) Mich.Möller & A.Weber, comb. nov.
Basionym:—Deinocheilos sichuanense Wang (1986: 2).
Distribution:—China (Sichuan, Jiangxi).
Oreocharis sichuanica (K.Y.Pan) Mich.Möller & A.Weber, comb. nov.
Basionym:—Isometrum sichuanicum Pan (1986: 33).
Distribution:—China (Sichuan).
Notes:—Referred to Isometrum sect. Isometrum by Pan (1986), Wang et al. (1990) and Li & Wang (2004).
Oreocharis sinensis (Oliv.) Mich.Möller & A.Weber, comb. nov.
Basionym:—Bournea sinensis Oliver in Hooker (1893: tab. 2254).
Distribution:—China (Guangdong).
Notes:—Type species of Bournea; distinctive in the actinomorphic, 4-merous flowers.
Oreocharis sinohenryi (Chun) Mich.Möller & A.Weber, comb. nov.
Basionym:—Didymocarpus sinohenryi Chun (1946: 290).
Homotypic synonym:—Opithandra sinohenryi (Chun) Burtt (1958a: 303).
Distribution:—China (Guangxi).
Notes:—Referred to Opithandra sect. Opithandra by Wang (1987), Wang et al. (1990) and Li & Wang
(2004).
Oreocharis squamigera Léveillé (1915a: 24) = Oreocharis henryana Oliv., see there.
Oreocharis stenosiphon Mich.Möller & A.Weber, nom. nov.
Basionym:—Rottlera fargesii Franchet (1899b: 251).
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A NEW DELINEATION FOR OREOCHARIS
Homotypic synonym:—Opithandra fargesii (Franch.) Burtt (1958a: 303).
Distribution:—China (Sichuan).
Notes:—Type and only species of Opithandra sect. Stenosiphon (Wang 1987, Wang et al. 1990, Li & Wang
2004).
Oreocharis stewardii (Chun) Mich.Möller & A.Weber, comb. nov.
Basionym:—Briggsia stewardii Chun (1946: 303).
Distribution:—China (Guangxi).
Oreocharis tonkinensis Kraenzlin (1928: 216) Didymocarpus tonkinensis (Kraenzl.) Handel-Mazzetti
(1936: 883) Boeica tonkinensis (Kraenzl.) Burtt (1977: 373) = Boeica porosa Clarke (1883: 136), see Wang
et al. (1990, 1998).
Oreocharis trichantha (B.L.Burtt & R.Davidson) Mich.Möller & A.Weber, comb. nov.
Basionym:—Ancylostemon trichanthus Burtt & Davidson (1954: 218).
Distribution:—China (Yunnan).
Oreocharis tubicella Franchet (1899b: 249).
Distribution:—China (Sichuan, Yunnan).
Notes:—Referred to Oreocharis sect. Oreocharis by Pan (1987), Wang et al. (1990) and Li & Wang (2004).
Oreocharis tubiflora Pan (1987: 287).
Distribution:—China (Fujian).
Notes:—Referred to Oreocharis sect. Oreocharis by Pan (1987), Wang et al. (1990) and Li & Wang (2004).
Oreocharis urceolata (K.Y.Pan) Mich.Möller & A.Weber, comb. nov.
Basionym:—Tremacron urceolatum Pan (1988a: 429).
Distribution:—China (Sichuan).
Oreocharis villosa (K.Y.Pan) Mich.Möller & A.Weber, comb. nov.
Basionym:—Isometrum villosum K.Y.Pan (1986: 31).
Distribution:—China (Sichuan).
Notes:—Referred to Isometrum sect. Isometrum by Pan (1986), Wang et al. (1990) and Li & Wang (2004).
Oreocharis vulpina (B.L.Burtt & R.Davidson) Mich.Möller & A.Weber, comb. nov.
Basionym:—Ancylostemon vulpinus Burtt & Davidson (1954: 217).
Distribution:—China (Yunnan).
Oreocharis wangwentsaii Mich.Möller & A.Weber, nom. nov.
Basionym:—Didymocarpus mairei Léveillé (1912: 301).
Homotypic synonym:—Ancylostemon mairei (H.Lév.) Craib (1920b: 267).
Distribution:—China (Yunnan, Sichuan).
Notes:—The new name is chosen in honour of the eminent Chinese gesneriologist Prof. Wang Wen-Tsai,
Beijing. See also notes under Oreocharis mairei. The type variety is found only in Yunnan.
Oreocharis wangwentsaii var. emeiensis (K.Y.Pan) Mich.Möller & A.Weber, comb. nov.
Basionym:—Ancylostemon mairei (H.Lév.) Craib var. emeiensis Pan (1988a: 434).
Distribution:—China (Sichuan).
MÖLLER ET AL.28 Phytotaxa 23 © 2011 Magnolia Press
Oreocharis wanshanensis (S.Z.He) Mich.Möller & A.Weber, comb. nov.
Basionym:—Isometrum wanshanense He (2006: 454).
Distribution:—China (Guizhou).
Notes:—The species is said to be closely related to Isometrum villosum by He (2006) and thus can be referred
to Isometrum sect. Isometrum.
Oreocharis wentsaii (Z.Yu Li) Mich.Möller & A.Weber, comb. nov.
Basionym:—Opithandra wentsaii Li (2003: 73).
Distribution:—China (Guizhou).
Notes:—Referred to Opithandra sect. Briggsioides by Li (2003), who says that “it is most closely related to
O. cinerea W.T.Wang”.
Oreocharis xiangguiensis W.T.Wang & K.Y.Pan in Pan (1987: 285).
Distribution:—China (Hunan, Guangxi).
Notes:—Referred to Oreocharis sect. Oreocharis by Pan (1987), Wang et al. (1990) and Li & Wang (2004).
Acknowledgements
The work was carried out in cooperation between the University of Vienna (supported by the Austrian Science
Fonds, FWF-Proj. No. P-13107-Bio) and the Royal Botanic Garden Edinburgh (RBGE). RBGE is supported
by the Scottish Government Rural and Environment Research and Analysis Directorate (RERAD). Fieldwork
of MM was supported by the RBGE Expedition Fund. YGW benefitted from support through the Science &
Technology Innovation Program of the Guangxi Academy of Sciences Fund. We are grateful to Dr Wen Fang
for the provision of DNA samples of Dayaoshania and Opithandra.
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Phytotaxa 23 © 2011 Magnolia Press 33
A NEW DELINEATION FOR OREOCHARIS
Appendix 1: Diagnostics of the Bayesian inference analysis of the Oreocharis dataset of combine
d
trnL-F and ITS sequence data plus alignment gap matrix.
Number of taxa = 55
Number of characters = 1632
Number of generations = 2500000
Average standard deviation of split frequencies: 0.010898
Analysis completed in 44263 seconds
Analysis used 44262.85 seconds of CPU time
Likelihood of best state for "cold" chain of run 1 was -9387.23
Likelihood of best state for "cold" chain of run 2 was -9396.93
Acceptance rates for the moves in the "cold" chain of run 1:
With prob. Chain accepted changes to
54.02 % param. 1 (revmat) with Dirichlet proposal
20.50 % param. 2 (revmat) with Dirichlet proposal
57.63 % param. 3 (revmat) with Dirichlet proposal
19.96 % param. 4 (state frequencies) with Dirichlet proposal
15.31 % param. 6 (state frequencies) with Dirichlet proposal
19.86 % param. 9 (gamma shape) with multiplier
45.24 % param. 10 (gamma shape) with multiplier
69.87 % param. 11 (prop. invar. sites) with sliding window
14.45 % param. 12 (topology and branch lengths) with extending TBR
23.78 % param. 12 (topology and branch lengths) with LOCAL
Acceptance rates for the moves in the "cold" chain of run 2:
With prob. Chain accepted changes to
50.90 % param. 1 (revmat) with Dirichlet proposal
20.95 % param. 2 (revmat) with Dirichlet proposal
58.89 % param. 3 (revmat) with Dirichlet proposal
19.81 % param. 4 (state frequencies) with Dirichlet proposal
15.09 % param. 6 (state frequencies) with Dirichlet proposal
20.48 % param. 9 (gamma shape) with multiplier
45.24 % param. 10 (gamma shape) with multiplier
69.81 % param. 11 (prop. invar. sites) with sliding window
14.29 % param. 12 (topology and branch lengths) with extending TBR
23.78 % param. 12 (topology and branch lengths) with LOCAL
Chain swap information for run 1:
1 2 3 4
----------------------------------
1 | 0.19 0.01 0.00
2 | 415635 0.23 0.02
3 | 416899 417311 0.30
4 | 416255 416695 417205
Chain swap information for run 2:
1 2 3 4
----------------------------------
1 | 0.16 0.01 0.00
2 | 416978 0.24 0.03
3 | 417053 415321 0.31
4 | 417026 416960 416662
Upper diagonal: Proportion of successful state exchanges between chains
Lower diagonal: Number of attempted state exchanges between chains
MÖLLER ET AL.34 Phytotaxa 23 © 2011 Magnolia Press
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burn-in 4%
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Phytotaxa 23 © 2011 Magnolia Press 35
A NEW DELINEATION FOR OREOCHARIS
Oreocharis combined data. Posterior probabilities run 1 versus run 2.
Oreocharis combined data: Symmetric tree differences within and between run 1 vs run 2
.
MÖLLER ET AL.36 Phytotaxa 23 © 2011 Magnolia Press
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... Recent molecular and morphology-based analyses demonstrated that the traditionally-defined Oreocharis was phylogenetically intertwined with nine previously defined small genera and acaulescent, rosette forming members of Briggsia Craib. The re-circumscribed Oreocharis is a strongly supported monophyletic group and placed in the subfamily Didymocarpoideae (Möller et al. 2011;Middleton et al. 2013). Since its re-definition in 2011, Oreocharis s.l. ...
... Most of the approximately 130 species are found in southern and south-western China, with a few species also in northern Vietnam, India, Bhutan, Myanmar, Thailand and Japan (e.g. Pham 2000;Vu 2005Vu , 2018Wei et al. 2010;Möller et al. 2011Möller et al. , 2016Möller et al. , 2018Do et al. 2017;Chen et al. 2017Chen et al. , 2018Yang et al. 2019;Cai et al. 2019;Jin et al. 2021). ...
... 30 species and four varieties and was mainly distributed in Bhutan, China, India, Myanmar and Vietnam (Wang et al. 1990(Wang et al. , 1998Vu 2018). Nineteen species and four varieties of acaulescent, rosette forming Briggsia (Craib 1920;Pan 1988) were moved to Oreocharis s.l. in a later revision (Möller et al. 2011. Of which, O. phuongii is most similar to O. longifolia (Craib) Mich.Möller & A.Weber in having peduncles up to 22 cm long, bracts 2, zygomorphic, yellow flowers with tubular corolla, stamen 4 with two pairs of coherent anthers and capsules up to 6 cm long, but it clearly differs from the latter in the shape of leaf blade, leaf base, leaf apex, leaf margin, number of flowers, shape and size of calyx lobes, inner surface of corolla tube and inner surface of three lower corolla lobes. ...
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... Oreocharis Bentham (1876Bentham ( : 1021 was recently re-delimited to include ten small or monotypic genera and the species number reach to over 160 species, based on extensive molecular phylogenetic studies (Möller et al. 2011, Weber et al. 2013, while Bournea Oliver (in Hooker 1893: 2254) was re-instated as an independent genus (Chen et al. 2020). Oreocharis belongs to tribe Trichosporeae Nees (1825: 143), subfamily Didymocarpoideae Arnott (1832: 121) of Gesneriaceae, and it is phylogenetic closed to Aeschynanthus Jack (1823: 42), Cyrtandra Forster & Forster (1776: t. 3) and Agalmyla Blume (1826: 766). ...
... To identify the systematic position of the putative new species, we followed Möller et al. (2011) and Ling et al. (2020a) and used 57 other Oreocharis species with available DNA sequences in the study. Finally, a total of 62 species were included in the phylogenetic analysis (Table 2). ...
... Our Molecular phylogeny showed that Oreocharis could be separated into six main clades, Clade I was mainly distributed in the mountains of Southwest China, Clades II-VI were mainly distributed in South to Southeast China (Fig. 3), which is roughly congruent with Ling et al. (2020a), Kong et al. (2021) and Lv et al. (in press some phylogenetic relationships in our result are different from previously works, which may cause by incomplete sampling of Oreocharis and the unequal aligned matrix information sites among studies. This phenomenon indicates that the Oreocharis underwent an early and rapid evolution radiation (Möller et al. 2011, Kong et al. 2021. ...
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Oreocharis hainanensis, a new species of Gesneriaceae is described and illustrated from low-altitudual karst areas in Hainan Island, South China. The new species is easily distinguished from its closely-related O. jasminina by campanulate floral tube, zygomorphic corolla and exserted stamens. It also shows different habitats from the four currently-recognized Oreocharis taxa on the island. Molecular phylogeny analysis based on plastid trnL-trnF and nuclear ITS1/2 sequences supported the delimitation of the new species, which forms a monophyletic clade with all the other Oreocharis taxa from Hainan Island. The roles of habitat and floral isolation in the evolution of the new species and its affinities are discussed. The species was assessed as Vulnerable (VU C1 and D2) according to IUCN Red List Categories and Criteria.
... & Juss. (Möller et al. 2011;Weber et al. 2013). Oreocharis was previously a genus of ca. ...
... Oreocharis was previously a genus of ca. 28 species distributed mainly in southern China (Li 1996;Wang et al. 1998a), based on molecular data and a morphological evaluation, Möller et al. (2011) demonstrate that the traditionally defined Oreocharis was phylogenetically intertwined with ten small and sometimes monotypic Chinese genera: Ancylostemon W.G. Craib (1919), Bournea Oliv. (1893), Dayaoshania W.T. Wang (1983), Deinocheilos W.T. Wang (1986), Isometrum W.G. Craib (1919), Opithandra B.L. Burtt (1956), Paraisometrum W.T. Wang (1997), Thamnocharis W.T. Wang (1981), Tremacron W.G. Craib (1918) and Briggsia W.G. Craib (1919). ...
... According to molecular phylogenetic studies, made the additional palynological observation of Bournea to support the taxonomic reassessment of the genus Bournea in Gesneriaceae, and eventually Bournea were isolated (Chen et al. 2020). However, according to the molecular and morphological data, several new taxa of Oreocharis were described recently, the Bournea was restored, and Oreocharis comprises about 150 species at present , most of them are endemic to China, mainly distributed in south and southwest China, as well as in northern Vietnam, Myanmar, northeast India, Bhutan, Japan and Thailand (Möller et al. 2011(Möller et al. , 2016Jin et al. 2021). ...
Preprint
Oreocharis repenticaulis , a new species of Gesneriaceae from Guangxi, China, is described and illustrated. The new species is most similar to O.argyreia Chun ex K.Y. Pan, but it differs from the latter by its leaf shape, umbrella inflorescence, corolla color, filament, phenology and some obvious characteristics of rhizomes. Detailed morphological description, illustrations and morphological comparison with similar species are provided.
... & Juss. (Möller et al. 2011, Weber et al. 2013. The species of this redefined genus are all rosette herbs with spirally-arranged leaves, scapose inflorescences and loculicidally dehiscent capsules but they show great variation in floral characters (i.e. ...
... The species of this redefined genus are all rosette herbs with spirally-arranged leaves, scapose inflorescences and loculicidally dehiscent capsules but they show great variation in floral characters (i.e. corolla form and coloration, number of stamens, anther shape and dehiscence mode) (Möller et al. 2011, Yang et al. 2020. At present, the genus comprise about 150 species (Wen et al. 2021), most of which occur in southern and southwestern China and northern Vietnam, with a few extending to Myanmar, India, Bhutan, Japan and Thailand (Möller et al. 2011). ...
... corolla form and coloration, number of stamens, anther shape and dehiscence mode) (Möller et al. 2011, Yang et al. 2020. At present, the genus comprise about 150 species (Wen et al. 2021), most of which occur in southern and southwestern China and northern Vietnam, with a few extending to Myanmar, India, Bhutan, Japan and Thailand (Möller et al. 2011). Especially southwestern China is the center of species diversity of Oreocharis s.l., and numerous new species have been found in this area in recent years (Chen et al. 2015, 2016, Tan et al. 2015, Cai et al. 2017, 2019. ...
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Based on morphological observation and literature consultation, a new species of Oreocharis (Gesneriaceae), O. reticuliflora Li H. Yang & X.Z. Shi, is described and illustrated. This new species resembles O. auricula, but differs by its conspicuous and reticular secondary veins, corolla with a network of violet stripes on each lip lobe, glan- dular-pubescent ovary, shorter capsule and being densely brown woolly on the abaxial leaf surface, outside of bracts and calyx lobes. A detailed morphological description and photographic illustration of the new species are presented.
... After Oreocharis Bentham was redefined by Möller et al. (2011), it has become a large and morphologically diverse genus in Gesneriaceae Rich. & Juss. ...
... & Juss. (Möller et al. 2011Chen et al. 2014). There are 136 species and 15 varieties of Oreocharis in China, mainly in southern and south-western China (Wen et al. 2021;Yang and Shi 2021;GRC. ...
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A new species of Oreocharis , O. guangwushanensis from the Sichuan Province of south-western China, is described and illustrated here. This new species has a pink corolla that is different from other species of Oreocharis in southwest China and, although it is morphologically similar to O. ronganensis and O. reticuliflora , it has significant differences in the colour and shape of the corolla, the apex of the corolla limb, shape and indumentum of the filaments and a shorter pistil. A detailed description, colour photographs, distribution and habitat, as well as the IUCN conservation status, are also provided.
... funnel-form in the former and tubular in the latter (Wei et al. 2010, Averyanov et al. 2020. While overall the flower shape among Primulina species is relatively conserved compared to other genera such as Oreocharis Bentham (1876) and Petrocodon Hance (1883) (Möller et al. 2011, Weber et al. 2011, with some exception in floral tube curvature, e.g. Primulina curvituba B.Pan, L.H. Yang & M.Kang (Yang et al. 2017) and P. flexusa F. Wen, Tao Peng & B. Pan (Peng et al. 2020), there are some distinct features that are not present in all species: 1) two ridges on the floor of the corolla tube, often associated with 2) two yellow stripes, perhaps acting as nectar guides for pollinators and 3) conspicuously coloured longitudinal ridges or sometimes 'flaps', bilobed patches on the inner dorsal surface of the corolla tube where the upper two lobes fuse forming a hump or 'boss' on the dorsal side of the tube (similar to Hemiboea C.B.Clarke (1888: pl. ...
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We transfer Chirita crassifolia to Primulina as P. crassifolia and provide detailed amended descriptions for this species and for P. quanbaensis, both species found in the karst habitats in the Bat Dai Son mountains in Ha Giang province, northern Vietnam.
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Actinostephanus , a new genus from southern China, is described and colorfully illustrated with a single species, A. enpingensis . This new genus is morphologically most similar to Boeica and Leptoboea , nevertheless, it can be easily distinguished from the latter two by the following characteristics, such as leaves in whorls of three, all closely clustered at the top; corolla bowl-shaped, 5-lobed, actinomorphic; capsule hard, oblong-ovoid, short, 3–4 mm long, densely appressed villous, wrapped by persistent densely pubescent calyx lobes, style persistent. The new genus and related genera were sequenced using the next-generation sequencing technique. The whole plastid genome of the new genus is 154, 315 - 154, 344 bp in length. We reconstructed phylogenetic trees using the dataset of 80 encoded protein genes of the whole plastid genome from 47 accessions based on ML and BI analyses. The result revealed that the new genus was recovering in a polytomy including Boeica , Rhynchotechum , and Leptoboea with strong support, congruent to the morphological evidence. A global conservation assessment was also performed and classifies A. enpingensis as Least Concern (LC). In addition, after a review of recently described species of Gesneriaceae, we propose that plant enthusiasts, especially Gesneriad fans, have been playing an increasingly important role in the process of new taxa-discoveries.
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A new species of Gesneriaceae, Ancylostemon hekouensis Y.M. Shui & W.H. Chen, is described and illustrated from Yunnan, China. The new species is similar to A. notochlaenus (H. Léveillé & Vaniot) Craib in the leaves and flower shape, but differs by its longer calyx lobes and longer corolla lobes.
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