全 文 : Guihaia Jan. 2016ꎬ 36(1):44-60
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DOI: 10.11931 / guihaia.gxzw201512015
MÖLLER Michaelꎬ韦毅刚ꎬ温放ꎬ等. 得与失: 苦苣苔科新的属级界定与分类系统———中国该科植物之变迁 [J]. 广西植物ꎬ2016ꎬ36(1):44-60
MÖLLER MꎬWEI YGꎬWEN Fꎬet al. You win some you lose some: updated generic delineations and classification of Gesneriaceae ̄implications for the family
in China [J]. Guihaiaꎬ2016ꎬ36(1):44-60
You win some you lose some: updated generic
delineations and classification of Gesneriaceae ̄
implications for the family in China
MÖLLER Michael1∗ꎬ WEI Yi ̄Gang2ꎬ WEN Fang2ꎬ CLARK John L.3ꎬ WEBER Anton4
( 1. Royal Botanic Garden Edinburghꎬ Edinburgh EH3 5LRꎬ UKꎻ 2. Guangxi Key Laboratory of Plant Conservation and Restoration Ecology in Karst
Terrainꎬ Guangxi Institute of Botanyꎬ Guilin 541006ꎬ Chinaꎻ 3. Department of Biological Sciencesꎬ The University of Alabamaꎬ Tuscaloosaꎬ
AL 35487ꎬ USAꎻ 4. Department of Botany and Biodiversity Researchꎬ University of Viennaꎬ Rennweg 14ꎬ A ̄1030 Viennaꎬ Austria )
Abstract: Over the last two decades molecular phylogenetic research on Gesneriaceae has greatly advanced our under ̄
standing of species relationships and generic delimitations. It has allowed the proposal of a new classification of the family
that is thought to reflect the natural relationships of the taxa better than traditional morphological classifications. Dramatic
taxonomic changes were implemented affecting the classification of Gesneriaceae in China. Many traditional genera have
been splitꎬ merged or newly defined. Additionallyꎬ new genera have been established based on recently collected materi ̄
alꎬ illustrating on the one hand gaps in fieldwork in Chinaꎬ and on the other hand the biological richness of the Gesneri ̄
aceae in China. Hereꎬ we summarize and present an overview of our work and the taxonomic consequences.
Key words: Chinaꎬ classificationꎬ Gesneriaceaeꎬ molecular phylogeniesꎬ systematics
CLC number: Q949.4 Document code: A Article ID: 1000 ̄3142(2016)01 ̄0044 ̄17
得与失: 苦苣苔科新的属级界定与分类系统
———中国该科植物之变迁
MÖLLER Michael1∗ꎬ 韦毅刚2ꎬ 温 放2ꎬ CLARK John L.3ꎬ WEBER Anton4
(1. 英国皇家爱丁堡植物园ꎬ 爱丁堡ꎬ 英国ꎻ 2. 广西喀斯特植物保育与恢复生态学重点实验室ꎬ 广西壮族自治区中 国 科 学 院 广西植物研究所ꎬ
广西 桂林 541006ꎬ 中国ꎻ 3. 生命科学部ꎬ 阿拉巴马大学ꎬ 塔斯卡卢萨ꎬ 阿拉巴马州ꎬ 美国ꎻ
4. 植物学与生物多样性研究部ꎬ 维也纳大学ꎬ 维也纳ꎬ 奥地利 )
摘 要: 过去 20年对苦苣苔科植物的分子系统学研究已经极大地拓展了对这个科的种间关系与属一级之界定
的理解ꎮ 该文提供了一个苦苣苔科植物新分类系统ꎬ与传统的经典形态学分类系统比较ꎬ这一新系统被认为能
更好地反映科下分类单元彼此之间的自然关系ꎮ 众多传统意义上的属被分割、合并或者重新定义ꎬ这些巨大的
分类变动正在影响着中国的苦苣苔科植物系统分类ꎮ 此外ꎬ基于最近采集的材料ꎬ一些新属得以建立ꎬ一方面说
Received date: 2015 ̄12 ̄21 Accepted date: 2016 ̄01 ̄07
Foundation item: the Chinese Academy of Sciences Visiting Professorship Scheme (2011T1S20)ꎻ the National Natural Science Foundation of China
(31000258ꎬ 31470306ꎬ 31260038ꎬ 31460159)ꎻ the Key Laboratory of Plant Diversity and Biogeography of East Asiaꎬ Kunming Institute of Botany (KIB)
& the Chinese Academy of Sciences (2014CB954100) and Guangxi Natural Science Foundation (2015GXNSFBB139004 & 2015GXNSFBA139105)ꎻ
Guangxi Forest Science & Technology Foundation (Gui Lin Ke Zi〔2014〕No.27).
Biography: MÖLLER Michael (1960 ̄ )ꎬ Maleꎬ Germanꎬ Ph.D. & Professorꎬ focuses on the Old World Gesneriaceaeꎬ research interests on Phylogenetics & Sys ̄
tematicsꎬ Finescale Biogeographyꎬ Character Evolution & Evolutionary Developmentꎬ Genome & Chromosome Evolutionꎬ etc. (E ̄mail) m.moeller@ rbge.ac.uk.
∗Corresponding author
明在中国还有相当多的野外工作需要做ꎻ而另一方面则再次证明了中国的苦苣苔科植物多样性是如此之丰富ꎮ
在此ꎬ针对目前已经完成的工作和分类结论ꎬ对中国苦苣苔科植物的新分类系统进行了总结与概述ꎬ并深入地讨
论了部分传统分类学上存在困惑的疑难属ꎮ
关键词: 中国ꎬ 分类ꎬ 苦苣苔科ꎬ 分子系统学ꎬ 系统分类
Gesneriaceae are a medium ̄sized family in the or ̄
der Lamialesꎬ with main distribution in the tropics and
subtropics of both the Old and the New World. Particu ̄
larly rich in Gesneriaceae is China. In the original
(Chinese) version of the “Flora of China”ꎬ Wang et al
(1990) listed 413 species in 56 generaꎬ in the later
English version (Wang et alꎬ 1998) the number in ̄
creased to 442 speciesꎬ while the number of genera re ̄
mained the same: 354 species and 25 genera (many of
them monotypic)ꎬ respectivelyꎬ were considered to be
endemic. Without a doubtꎬ China is a significant centre
of diversity of the familyꎬ with the majority of taxa found
in China’ s South and Southwestꎬ in the provinces of
Yunnanꎬ Guizhouꎬ Guangdong and Guangxi Autono ̄
mous Region (Wang et alꎬ 1992). The species density
decreases northwards and the furthest locality North are
the Yenshan Mts at 41°N.
In the last two decadesꎬ the implementation of mo ̄
lecular methods has resulted in worldwide dramatic
changes in the definition and infrafamilial classification
of angiosperm families and in the delineation of genera
attributed to them. This is particularly true for the Chi ̄
nese Gesneriaceaeꎬ in which an unproportionally high
number of taxonomic changes has been made. In the fol ̄
lowingꎬ the partly dramatic changes of traditional gener ̄
ic concepts of Chinese Gesneriaceae are surveyedꎬ and
the position of the Chinese Gesneriaceae in recent clas ̄
sifications are shown and discussed.
1 Classifications of Gesneriaceae and
amendments at generic level
Traditionallyꎬ the family Gesneriaceae was subdivi ̄
ded into two subfamiliesꎬ Gesnerioideae and Cyrtan ̄
droideae (e. g.ꎬ Benthamꎬ 1876ꎻFritschꎬ 1893-1894ꎻ
Burttꎬ 1963). The last formal classification based on
morphology ( and partly cytogenetic data) was that of
Burtt and Wiehler (1995)ꎬ who recognized three sub ̄
families: (1) Gesnerioideae (56 generaꎻ predominantly
Neotropical)ꎻ (2) Coronantheroideae (9 generaꎬ south ̄
ern hemisphere: temperate South America to Austral ̄
ia)ꎻ (3) Cyrtandroideae ( = now Didymocarpoideaeꎻ
82 generaꎬ chiefly paleotropicalꎬ including three genera
in Europe). The distribution of the c. 3 400 species is
roughly equal between the Neotropics and Palaeotropics.
Traditional classifications were based on a few key
morphological features such as seed anatomyꎬ post ̄ger ̄
mination characteristics of the seedlingsꎬ nectary struc ̄
ture and ovary position. Thusꎬ the Neotropical Gesneri ̄
aceae were characterized by the presence of seed endo ̄
spermꎬ isocotyly ( both cotyledons showing equal and
limited growth after germination)ꎬ a nectary often con ̄
sisting of separate glandsꎬ and often partly or fully infe ̄
rior ovary positionꎬ while the Palaeotropical Gesneriace ̄
ae were characterized by the lack of endospermꎬ by con ̄
tinued growth of one cotyledon after germination (aniso ̄
cotylyꎻ Fritschꎬ 1904ꎻ Burttꎬ1963ꎬ1970ꎻ Jongꎬ 1970ꎻ
Nishii et alꎬ 2004ꎻ Mantegazza et alꎬ 2007)ꎬ a basical ̄
ly ring ̄shaped nectary and superior ovary position. Sub ̄
fam. Coronantheroideae was characterized by isocotylous
seedlingsꎬ a nectary ‘adnate’ to the ovary baseꎬ and
superior ovary position.
Burtt & Wiehler (1995) subdivided the three sub ̄
families into the following tribes (number of genera in
brackets): [1] Gesnerioideae: Gloxinieae (23)ꎬ Epi ̄
scieae (21)ꎬ Beslerieae (8)ꎬ Napeantheae (1) and
Gesnerieae (2)ꎻ [2] Coronantheroideae: only Coronan ̄
thereae (9)ꎬ[3] Cyrtandroideae: Klugieae (7)ꎬ Didy ̄
mocarpeae ( 64 )ꎬ Trichosporeae ( 6 )ꎬ Cyrtandreae
(3)ꎬ Titanotricheae (1).
The first classification taking to some extent molec ̄
ular data into accountꎬ was that of Weber (2004). How ̄
everꎬ particulary for the Palaeotropical Gesneriaceae the
molecular data situation was insufficient and no formal
classification was presented. Weber (2004)ꎬ thereforeꎬ
proposed a provisional classificationꎬ recognising four
informal groups: ( 1 ) Coronantheroid Gesneriaceaeꎻ
541期 MÖLLER Michael et al. 得与失: 苦苣苔科新的属级界定与分类系统———中国该科植物之变迁
( 2 ) Gesnerioid Gesneriaceaeꎻ ( 3 ) Epithematoid
Gesneriaceaeꎻ (4) Didymocarpoid Gesneriaceae. The
latter and the largest group was subdivided into morpho ̄
geographical assemblages: ‘ Basal Asiatic genera ’ꎬ
‘European genera’ꎬ ‘African and Madagascan genera’
and ‘Advanced Asiatic and Malesian genera’.
Especially by the work of the first author (MMO)
and associatesꎬ in the last 10 years a comprehensive da ̄
ta basis was established for the palaeotropical Gesneri ̄
aceae that enabled a better understanding of the phylo ̄
genetic diversification of that large and enormously di ̄
versified group. In parallelꎬ also considerable progress
was made with regard to Neotropical Gesneriaceaeꎬ in ̄
cluding the work of one of the authors (JLC). In 2013ꎬ
a new formal classification was published that was en ̄
tirely based on molecular data (Weber et alꎬ 2013).
By the inclusion of Sanangoꎬ which was repeatedly
shown to be sister to Gesneriaceae in previous molecular
studiesꎬ three subfamilies were recognised: Sanan ̄
goideae ( only Sanango)ꎬ Gesnerioideae and Didymo ̄
carpoideae. Wiehler’ s subfam. Coronantheroideae was
included as a tribe (Coronanthereae) in Gesnerioideae.
Apart from the traditional Gesnerioideae (subdivided in ̄
to the tribes Beslerieaeꎬ Napenatheae and Gesnerieae)
a fifthꎬ and rather aberrantꎬ tribe was included in
Gesnerioideae: the monospecific Titanotricheae from E
Asia. This is the only tribe that does not occur in the
Neotropics or the southern hemisphereꎬ but with a single
species (Titanotrichum oldhamii) in East Asia (E Chi ̄
naꎬ Taiwan region of Chinaꎬ S Japan). The former Cyr ̄
tandoideaeꎬ now referred to as Didymocarpoideaeꎬ were
subdivided into two tribes: Epithemateae and Trichos ̄
poreae. The former is apparently an ancient assemblage
of few and rather isolated genera that fall into four sub ̄
tribesꎬ while the latter includes a large number of gene ̄
ra and is obviously still in a stage of active evolution. In
this tribe provisionally 10 subtribes have been recog ̄
nisedꎬ the by far largest being subtribe Didymocarpinae.
Apart from a new infrafamilial classification of the
familyꎬ another important aspect of the molecular work
is the improvement in the understanding of generic de ̄
limitations. In factꎬ a wealth of taxonomic changes re ̄
sulted from the molecular studiesꎬ both with respect to
Neotropical and Palaeotropical taxa. On the one handꎬ
many genera have been reduced to synonymy ( e. g.
Möller et alꎬ 2011bꎻ Weber et alꎬ 2011bꎬcꎻ Nishii et
alꎬ 2015ꎻ Puglisi et alꎬ in press)ꎬ on the other handꎬ
some genera have been revived ( e. g. Centrosolenia:
Mora & Clarkꎬ 2016ꎻ Crantzia: Clarkꎬ 2005ꎻ Clark et
alꎬ 2006ꎻ Damrongia: Weber et alꎬ 2011aꎻ Glossolo ̄
ma: Clark et alꎬ 2006ꎻ Clarkꎬ 2009ꎻ Liebigia: Weber
et alꎬ 2011aꎻ Mandirola: Roalson et alꎬ 2005aꎬbꎻ Bog ̄
gan 2006ꎻ Seemannia: Roalson et alꎬ 2005aꎬbꎻ Boggan
2006ꎻ Trichodrymonia: Mora & Clark 2016) or newly
described (e. g. Billoliviaꎬ Middleton et alꎬ 2014ꎻ Cha ̄
utemsia: Araujo et alꎬ 2010ꎻ Christopheria: Smith &
Clark 2013ꎻ Chayamaritia: Middleton et alꎬ 2015ꎻ Gla ̄
brella: Möller et alꎬ 2014ꎻ Gloxinella: Roalson et alꎬ
2005aꎬbꎬ Bogganꎬ 2006ꎻ Gloxiniopsis: Roalson et alꎬ
2005aꎬbꎻ Bogganꎬ 2006ꎻ Lesia: Smith & Clarkꎬ 2013ꎻ
Litostigma: Wei et alꎬ 2010ꎻ Pachycaulos: Smith &
Clarkꎬ 2013ꎻ Pagothyra: Smith & Clarkꎬ 2013ꎻ Somra ̄
nia: Middleton & Triboun ꎬ2012ꎻ Sphaerorrhiza: Roal ̄
son et alꎬ 2005aꎬbꎻ Tribounia: Middleton & Möllerꎬ
2012). In many cases the species content changed dra ̄
matically (e. g.ꎬ Primulina: increment from 1 to >150
speciesꎻ Alloplectus: reduction from c. 140 to 5 species:
Clark et alꎬ 2006).
2 Position of Chinese Generiaceae in
the classification of Weber et al (2013)
In Table 1 the classification of Weber et al (2013)
is givenꎬ with indication of the position of genera pres ̄
ently recognised for China. Distribution is either endem ̄
ic or includes adjacent countries ( some genera with
main distribution further South and reaching China with
a few or a single species only).
A most striking discrepancy with former classifica ̄
tions is the position of Titanotrichum. This monotypic
genus was established (based on Rehmannia oldhamii
Hemsl.) by Solereder in 1909 and placed in Scrophular ̄
iaceae. It is characterised by a terminal racemose inflo ̄
rescence and the presence of masses of seedling ̄like
propagules in the upper part of the inflorescence (Wang
et alꎬ 2004b). Burtt ( 1963ꎬ 1977) earmarked Tit ̄
anotrichum as a ‘genus anomalum’ within Gesneriace ̄
64 广 西 植 物 36卷
Table 1 Classification of Gesneriaceae according to Weber et al (2013) and taxa represented in China
Rank No. of genera / species no. / genera represented in China
Gesneriaceae Rich. & Juss. in DC.
1. Subfam. Sanangoideae A. Weberꎬ J. L. Clark & Mich. Möller 1 / 1
2. Subfam. Gesnerioideae Burnett
2.1. Tribe Titanotricheae Yamaz. ex W. T. Wang 1/ 1 Titanotrichum Soler.
2.2. Tribe Napeantheae Wiehler 1/ 20+
2.3. Tribe Beslerieae Bartl.
2.3.1. Subtribe Besleriinae G. Don 4/ 239+
2.3.2. Subtribe Anetanthinae A. Weber & J. L. Clark 5/ 12+
2.4. Tribe Coronanthereae Fritsch
2.4.1. Subtribe Coronantherinae Fritsch 2/ 14-21
2.4.2. Subtribe Mitrariinae Hanst. 4 / 4
2.4.3. Subtribe Negriinae V. L. Wooꎬ J. F. Smith & Garn.-Jones 3/ 3
2.5.Tribe Gesnerieae Dumort. (1829)
2.5.1. Subtribe Gesneriinae Link 4/ 100
2.5.2. Subtribe Gloxiniinae G. Don 21/ 200+
2.5.3. Subtribe Columneinae Hanst. 26/ 525+
2.5.4. Subtribe Sphaerorrhizinae A. Weber & J. L. Clark 1/ 2
2.5.5. Subtribe Ligeriinae Hanst. 3 / 91
3. Subfam. Didymocarpoideae Arn.
3.1.Tribe Epithemateae C. B. Clarke
3.1.1. Subtribe Loxotidinae G. Don 1/ ≈15 Rhynchoglossum Blume
3.1.2. Subtribe Monophyllaeinae A. Weber & Mich. Möller 2/ 38+ Whytockia W. W. Sm.
3.1.3. Subtribe Loxoniinae A. DC. 2(3) / 9+ ? Gyrogyne W. T. Wangꎬ Stauranthera Benth.
3.1.4. Subtribe Epithematinae DC. ex Meisn. 1/ 20+ Epithema Blume
3.2. Tribe Trichosporeae Nees
3.2.01. Subtribe Jerdoniinae A. Weber & Mich. Möller 1/ 1 1
3.2.02. Subtribe Corallodiscinae A. Weber & Mich. Möller 1/ 3-5 Corallodiscus Batalin
3.2.03. Subtribe Tetraphyllinae A. Weber & Mich. Möller 1/ 3
3.2.04. Subtribe Leptoboeinae C. B. Clarke 6/ 43 Beccarinda Kuntzeꎬ Boeica C.B.Clarkeꎬ Leptoboea Benth.ꎬ
Platystemma Wall.ꎬ Rhynchotechum Blume
3.2.05. Subtribe Ramondinae DC. ex Meisn. 3(2) / 5
3.2.06. Subtribe Litostigminae A. Weber & Mich. Möller 1/ 2 Litostigma G. Weiꎬ F. Wen & Mich. Möller
3.2.07. Subtribe Streptocarpinae Ivanina 1/ 177
3.2.08. Subtribe Didissandrinae A. Weber & Mich. Möller 2/ 10
3.2.09. Subtribe Loxocarpinae A. DC. 14/ 202+ Damrongia Kerrꎬ Dorcoceras BungeꎬMiddletonia C. Puglisiꎬ
Ornithoboea Parish ex C. B. Clarkeꎬ Paraboea ( C. B.
Clarke) Ridl.ꎬ Rhabdothamnopsis Hemsl.
3.2.10. Subtribe Didymocarpinae G. Don 32 1660- Aeschynanthus Jackꎬ Allocheilos W. T. Wangꎬ Allostigma
1830 W. T. Wangꎬ Anna Pellegr.ꎬBriggsiopsis K. Y. Panꎬ Cathay ̄
anthe Chunꎬ Conandron Sieb. & Zucc.ꎬ Cyrtandra J. R.
Forst. & G. Forst.ꎬ Didymocarpus Wall.ꎬ Didymostigma W.
T. Wangꎬ Glabrella Mich. Möller & W. H. Chenꎬ Gyrocheilos
W. T. Wangꎬ Hemiboea C. B. Clarkeꎬ Henckelia Spreng.ꎬ
Loxostigma C. B. Clarkeꎬ Lysionotus D. Donꎬ Metapetro ̄
cosmea W. T. Wangꎬ Microchirita (C. B. Clarke) Yin Z.
Wangꎬ Oreocharis Benth.ꎬ Petrocodon Hanceꎬ Petrocosmea
Oliv.ꎬ Primulina Hanceꎬ Pseudochirita W. T. Wangꎬ
Raphiocarpus Chun
Bold face: Taxon represented in Chinaꎻ Underlined: Endemic to Chinaꎻ Spaced print: New genus for Chinaꎻ ?: Genus maybe extinct.
ae. Based on the aberrant morphologyꎬ Wang et al
(1990) established a separate tribe in subfam. Cyrtan ̄
droideae (now Didymocarpoideae) for its accomodationꎬ
while Weber (2004) proposed to exclude it from Gesne ̄
riaceae. The molecular study of Wang et al (2004a)
confirmed its position within Gesneriaceaeꎬ but yielded
the surprising result that it is more closely allied to the
New World than to the Old World Gesneriaceae. Weber
et al (2013)ꎬ thereforeꎬ included Titanotrichum in sub ̄
fam. Gesnerioideae as a monospecific tribe. Apart from
the geographical distribution (SE mainland Chinaꎬ Tai ̄
wan region of Chinaꎬ S Japan) which is difficult to ex ̄
plain for a member of subfam. Gesnerioideaeꎬ this taxo ̄
nomic placement is in line with the isocotylous seedlings
(the interpretation of Wang et alꎬ 2002 as anisocotylous
is erroneous). Whether the distribution of Titanotrichum
is to be regarded as a northern outlier of the southern
hemisphere Gesneriaceae or an early immigrant from the
Neotropics remains unexplained. In the isocotylous seed ̄
lings it is similar to both groupsꎬ but other charactersꎬ
including the curious propagulesꎬ indicate an isolated
position.
741期 MÖLLER Michael et al. 得与失: 苦苣苔科新的属级界定与分类系统———中国该科植物之变迁
Table 2 List of present and past genera of Chinese Gesneriaceae.
Genus
(alphabet.)
(Chinese name)
Infrafamilial
position:
Subfamily
Tribe Subtribe
Distribution Speciesnumber
Species no.
in China
(endemic)
Tax. status Reference Remarks
Aeschynanthus Jack
芒毛苣苔属
Didymocarpoideae
Trichosporeae
Didymocarpinae
From S Chinaꎬ N & S
India throughout
Malesia to New
Guinea and the
Solomon Islands
≈185 ≈34
(≈13)
Emended by inclusion
of Micraeschynanthus
Middletonꎬ
2007
The inclusion of
the monotypic
genus Micraeschy ̄
nanthus (Malay
peninsula) has
little bearing on
the definition of
Aeschynanthus
Allocheilos
W.T.Wang
异唇苣苔属
Didymocarpoideae
Trichosporeae
Didymocarpinae
S China (Guizhouꎬ E
Yunnan)
2 2
(2)
No change
Allostigma W.T.
Wang
异片苣苔属
Didymocarpoideae
Trichosporeae
Didymocarpinae
S China 1 1
(1)
No change See text under
Oreocharis
Ancylostemon
Craib
直瓣苣苔属
- - Sunk into Oreocharis Möller et alꎬ
2011bꎻ Chen
et alꎬ 2014b
See text under
Oreocharis
Anna Pellegr.
大苞苣苔属
Didymocarpoideae
Trichosporeae
Didymocarpinae
Chinaꎬ N Vietnam 4 4
(3)
No change
Beccarinda Kuntze
横蒴苣苔属
Didymocarpoideae
Trichosporeae
Leptoboeinae
NE Indiaꎬ Burmaꎬ S
Chinaꎬ Vietnamꎬ Su ̄
matra
≈8 5
(4)
No change
Boea Comm. ex
Lam.
旋蒴苣苔属
10 - Redefinedꎻ Chinese
spp. now in Dorcoceras
and Damrongia
Puglisi et alꎬ
in press
See text under
Boea
Boeica C.B.
Clarke
短筒苣苔属
Didymocarpoideae
Trichosporeae
Leptoboeinae
Bhutanꎬ S Chinaꎬ N
& NE Indiaꎬ Myan ̄
marꎬ N Vietnamꎬ NW
Malaya
≈12 7
(3)
No change
Bournea Oliv.
四数苣苔属
- - Sunk into Oreocharis Möller et alꎬ
2011b
See text under
Oreocharis
Briggsia Craib
粗筒苣苔属
- - Partly sunk into Oreo ̄
charisꎬ partly transferred
to Loxostigmaꎬ and two
spp. forming the new
genus Glabrella
Möller et alꎬ
2011bꎻ Chen
et alꎬ 2014bꎻ
Möller et alꎬ
2014
See text under
Briggsia and
Oreocharis
Briggsiopsis
K.Y.Pan
筒花苣苔属
Didymocarpoideae
Trichosporeae
Didymocarpinae
S China (C & S Si ̄
chuanꎬ NE Yunnanꎬ
Guizhou)
1 1
(1)
No change
Calcareoboea C.Y.
Wu ex H.W.Li
朱红苣苔属
- - Sunk into Petrocodon Wang et alꎬ
2011ꎻ Weber
et alꎬ 2011b
See text under
Petrocodon
Chirita Buch.
-Ham. ex D.Don
唇柱苣苔属
- - Split into 5 genera and
synomymisation with
Henckelia
Wang et alꎬ
2011ꎻ Weber
et alꎬ 2011a
See text under
Chirita
Chiritopsis W.T.
Wang
小花苣苔属
- - Sunk into Primulina Wang et alꎬ
2011ꎻ Weber
et alꎬ 2011a
See text under
Chirita
Cathayanthe Chun
扁蒴苣苔属
Didymocarpoideae
Trichosporeae
Didymocarpinae
S China (Hainan) 1 1
(1)
No change
Conandron Sieb.
& Zucc.
苦苣苔属
Didymocarpoideae
Trichosporeae
Didymocarpinae
E Chinaꎬ Taiwan re ̄
gion of Chinaꎬ S Ja ̄
pan
1 1 No change
Corallodiscus
Batalin
珊瑚苣苔属
Didymocarpoideae
Trichosporeae
Corallodisceae
Bhutanꎬ Chinaꎬ N &
NE Indiaꎬ Nepalꎬ
Thailand
3-5 3 No change
Cyrtandra J.R.
Forst. & G.Forst.
浆果苣苔属
Didymocarpoideae
Trichosporeae
Didymocarpinae
Nicobar Islands and
S Thailand through
Malesia incl. Taiwan
region of China and
the S Pacific to the
Hawaiian Islands
652-818 1
(1)
No change
84 广 西 植 物 36卷
续表2
Genus
(alphabet.)
(Chinese name)
Infrafamilial
position:
Subfamily
Tribe Subtribe
Distribution Speciesnumber
Species no.
in China
(endemic)
Tax. status Reference Remarks
Damrongia Kerr
丹氏苣苔属(新拟)
Didymocarpoideae
Trichosporeae
Loxocarpinae
China to Sumatra 10 1
(1)
Re ̄established for
particular species of
erstwhile Chiritaꎻ
inclusion of Boea clar ̄
keana Hemsl. and the
Asian species descri ̄
bed in Streptocarpus
Weber et alꎬ
2011a ꎻ
Puglisi et alꎬ
in press
See text under
Chirita and
Damrongia
Dayaoshania W.T.
Wang
瑶山苣苔属
- - Sunk into Oreocharis Möller et alꎬ
2011b
See text under
Oreocharis
Deinocheilos W.T.
Wang
全唇苣苔属
- - Sunk into Oreocharis Möller et alꎬ
2011b
See text under
Oreocharis
Didymocarpus
Wall.
长蒴苣苔属
Didymocarpoideae
Trichosporeae
Didymocarpinae
from N and NE Indi ̄
aꎬ Nepal and S Chi ̄
na southwards to the
Malay Peninsula and
N Sumatra
>70 ≈30
(≈24)
Some spp. transferred
to Petrocodon
Weber et alꎬ
2011b
See text under
Petrocodon
Didymostigma
W.T.Wang
双片苣苔属
Didymocarpoideae
Trichosporeae
Didymocarpinae
SE China (Guang ̄
dongꎬ Fujianꎬ
Guangxi)
3 3
(3)
No change
Dolicholoma
D.Fang & W.T.
Wang
长檐苣苔属
- - Sunk into Petrocodon Weber et alꎬ
2011b
See text under
Petrocodon
Dorcoceras Bunge
羚角苣苔属(新拟)
Didymocarpoideae
Trichosporeae
Loxocarpinae
Chinaꎬ Thailandꎬ
Cambodiaꎬ Vietnamꎬ
Philippines and
Indonesia
4 2
(1 or 2?)
Re ̄established for par ̄
ticular ( non ̄Australa ̄
sian) species of Boea
Puglisi et alꎬ
in press
See text under
Boea and Dorco ̄
ceras
Epithema Blume
盾座苣苔属
Didymocarpoideae
Epithemateae
Epitheminae
S Chinaꎬ Taiwan re ̄
gion of China
20 2 No change at genus
level
revised by
Bransgrove &
Middletonꎬ
2015
Glabrella Mich.
Möller & W.H.
Chen
光叶苣苔属
Didymocarpoideae
Trichosporeae
Didymocarpinae
S Chinaꎬ Taiwan re ̄
gion of China
3 3
(3)
New genus estblished
for 3 spp. of Briggsia
not to be included in
Oreocharis or Loxostig ̄
ma
Möller et alꎬ
2014ꎻ Wen et
alꎬ 2015aꎬb
See text under
Briggsia and
Oreocharis
Gyrocheilos
W.T.Wang
圆唇苣苔属
Didymocarpoideae
Trichosporeae
Didymocarpinae
S China ( Guangxiꎬ
Guangdongꎬ SE
Guizhou)ꎬ Vietnam
5 4 (3) No change
Gyrogyne
W.T.Wang
圆果苣苔属
Didymocarpoideae
Epithemateae
Loxoniinae (?)
S China ( W Guan ̄
gxi)
1 1
(1)
Position in
Epithemateae ̄
Loxoniinae uncertain
Probably extinct
Hemiboea
C.B.Clarke
半蒴苣苔属
Didymocarpoideae
Trichosporeae
Didymocarpinae
C & S Chinaꎬ Tai ̄
wan region of Chinaꎬ
N Vietnamꎬ S Japan
31 31
(30)
Inclusion of
Metabriggsia (2 spp.)
Weber et alꎬ
2011c
Inclusion does
not affect the
traditional con ̄
cept of HemiboeaHemiboeopsis W. T.
Wang
- - Sunk into Henckelia Weber et alꎬ
2011a
Henckelia Spreng.
南洋苣苔属 /
汉克丽亚花属
Didymocarpoideae
Trichosporeae
Didymocarpinae
From S China to Sri
Lanka
c. 55 23
(12)
Redefined to include
Chirita p.p.( excl. Mi ̄
crochirita and Primuli ̄
na) and Hemiboepsisꎬ
and to exclude
Codonoboaea
Weber et alꎬ
2011aꎻ Mid ̄
dleton et alꎬ
2013
See text under
Chirita and
Henckelia
Isometrum Craib
金盏苣苔属
- - Sunk into Oreocharis Möller et alꎬ
2011b
See text under
Oreocharis
Lagarosolen W.T.
Wang
细筒苣苔属
- - Sunk into Petrocodon Weber et alꎬ
2011b
See text under
Petrocodon
Leptoboea Benth.
细蒴苣苔属
Didymocarpoideae
Trichosporeae
Leptoboeinae
Bhutanꎬ N and NE
Indiaꎬ S China
( Yunnan )ꎬ Myan ̄
marꎬ Thailand
2-3 1 No change
941期 MÖLLER Michael et al. 得与失: 苦苣苔科新的属级界定与分类系统———中国该科植物之变迁
续表2
Genus
(alphabet.)
(Chinese name)
Infrafamilial
position:
Subfamily
Tribe Subtribe
Distribution Speciesnumber
Species no.
in China
(endemic)
Tax. status Reference Remarks
Litostigma Y.G.
Weiꎬ F.Wen &
Mich. Möller
凹柱苣苔属
Didymocarpoideae
Trichosporeae
Litostigminae
China (Guizhouꎬ
Yunnan)
2 2
(2)
Genus recently
established
Wei et alꎬ
2010
See text under
Litostigma
Loxostigma
C.B.Clarke
斜片 苣苔 属 (新
拟) /紫花苣苔属
Didymocarpoideae
Trichosporeae
Didymocarpinae
S China (Sichuanꎬ
Yunnanꎬ Guizhouꎬ
Guangxi)ꎬ N
Vietnam
11 11
(8)
Recently inclusion of
caulescent Briggsia
species
Möller et alꎬ
2014
See text under
Briggsia
Lysionotus D.Don
吊石苣苔属
Didymocarpoideae
Trichosporeae
Didymocarpinae
From N India and
Nepal eastwards
through N Thailandꎬ
N Vietnam and S
China to S Japan
28 26
(18)
No change
Metabriggsia W. T.
Wang
单座苣苔属
- - Included in Hemiboea Weber et alꎬ
2011c
Metapetrocosmea
W.T.Wang
盾叶苣苔属
Didymocarpoideae
Trichosporeae
Didymocarpinae
S China (Hainan) 1 1 No change
Microchirita ( C. B.
Clarke) Y.Z.Wang
钩序苣苔属
Didymocarpoideae
Trichosporeae
Didymocarpinae
From the Western
Ghats of India to the
foothills of the Hima ̄
layasꎬ through conti ̄
nental SE Asia to
Sumatraꎬ Borneo and
Java
≈18 2
(1)
Raised from Chirita
sect. Microchirita
to generic rank
Wang et alꎬ
2011ꎻ Weber
et alꎬ 2011a
See text under
Chirita
Middletonia
C. Puglisi
米氏苣苔属
Didymocarpoideae
Trichosporeae
Loxocarpinae
Indiaꎬ Bangladeshꎬ
Bhutanꎬ Chinaꎬ
Burmaꎬ Thailandꎬ
Laosꎬ Cambodiaꎬ
Vietnamꎬ Malaysia
4 1 Genus recently estab ̄
lished for four species
of Paraboea
Puglisi et alꎬ
in press
See text under
Middletonia and
Paraboea
Opithandra B.L.
Burtt
后蕊苣苔属
- - Sunk into Oreocharis Möller et alꎬ
2011b
See text under
Oreocharis
Oreocharis Benth.
马铃苣苔属
Didymocarpoideae
Trichosporeae
Didymocarpinae
Chinaꎬ Thailandꎬ
Vietnamꎬ Myanmarꎬ
Bhutanꎬ NE Indiaꎬ
Japan
>106 >102
(>98)
Expanded to include
Ancylostemon Craibꎬ
Bournea Oliv.ꎬ Brigg ̄
sia Craib p. p. ̄incl.
typeꎬ Dayaoshania W.
T. Wangꎬ Deinocheilos
W. T. Wangꎬ Isome
trum Craibꎬ Opithandra
B. L. Burttꎬ Parai
sometrum W. T. Wangꎬ
Thamnocharis W. T.
Wangꎬ and Tremacron
Craibꎻ Inclusion of
further ten spp.of
Briggsia
Möller et alꎬ
2011bꎻ
Möller et alꎬ
2014ꎻ Chen
et alꎬ 2014b
See text under
Briggsia and
Oreocharis
Ornithoboea Parish
ex C.B.Clarke
喜鹊苣苔属
Didymocarpoideae
Trichosporeae
Loxocarpinae
From S China and
Vietnam southwards
to N Penins. Malaysia
16 5
(1)
No change Revised by
Scott & Mid ̄
dletonꎬ 2014
Paraboea (C.B.
Clarke) Ridl.
蛛毛苣苔属
Didymocarpoideae
Trichosporeae
Loxocarpinae
Bhutanꎬ Chinaꎬ
Indonesiaꎬ Malaysiaꎬ
Myanmarꎬ Philippinesꎬ
Thailandꎬ Vietnam
132 26
(14)
Expanded by inclusion
of Phylloboea and
Trisepalumꎻ removal of
four species and place ̄
ment in the new
genus Middletonia
Puglisi et alꎬ
2011
Puglisi et alꎬ
in press
See text under
Paraboea and
Middletonia
Paraisometrum W.
T.Wang
弥勒苣苔属
- - Sunk into Oreocharis Möller et alꎬ
2011b
See text under
Oreocharis
Paralagarosolen Y.
G. Wei
方鼎苣苔属
- - Sunk into Petrocodon Wang et alꎬ
2011ꎻ Weber
et alꎬ 2011b
See text under
Petrocodon
05 广 西 植 物 36卷
续表2
Genus
(alphabet.)
(Chinese name)
Infrafamilial
position:
Subfamily
Tribe Subtribe
Distribution Speciesnumber
Species no.
in China
(endemic)
Tax. status Reference Remarks
Petrocodon Hance
石山苣苔属
Didymocarpoideae
Trichosporeae
Didymocarpinae
Chinaꎬ N Vietnamꎬ
NE Thailand
29 28
(27)
Expanded to include
Calcareoboea C. Y.Wu
ex H. W. Liꎬ Didymo ̄
carpus Wall. p.p.( ex ̄
cl. type )ꎬ Dolicholo ̄
ma D. Fang & W. T.
Wangꎬ Lagarosolen
W. T. Wangꎬ Parala ̄
garosolen Y. G. Weiꎬ
Tengia Chun and
Wentsaiboea D.Fang &
D.H.Qinꎬ p. p. ( excl.
type)
Wang et alꎬ
2011ꎻ Weber
et alꎬ 2011b
See text under
Petrocodon
Petrocosmea Oliv.
石蝴蝶属
Didymocarpoideae
Trichosporeae
Didymocarpinae
NE Indiaꎬ S Chinaꎬ
Myanmarꎬ Thailandꎬ
S Vietnam.
42 34
(34)
No change
Platystemma Wall.
堇叶苣苔属
Didymocarpoideae
Trichosporeae
Leptoboeinae
Nepalꎬ Bhutanꎬ N
Indiaꎬ SW China
1 1 No change
Primulina Hance
报春苣苔属
Didymocarpoideae
Trichosporeae
Didymocarpinae
Essentially southern
half of China and Vi ̄
etnam
>169 >154
(>151)
Enormous expansion of
the previously mono ̄
typic genus by inclu ̄
sion of Chirita sect.
Gibbosaccusꎬ Chiritop ̄
sisꎬ Deltocheilosꎬ and
Wentsaiboea p.p.(incl.
type)
Wang et alꎬ
2011ꎻ Weber
et alꎬ 2011a
See text under
Chirita
Pseudochirita W. T.
Wang
异裂苣苔属
Didymocarpoideae
Trichosporeae
Didymocarpinae
S China (C & W
Guangxi)ꎬ Vietnam
1 1
(1)
No change
Raphiocarpus Chun
漏斗苣苔属
Didymocarpoideae
Trichosporeae
Didymocarpinae
S China and N & C
Vietnam
13 8
(7)
No change since We ̄
ber 2004ꎬ but changes
to be expected
Rhabdothamnopsis
Hemsl.
长冠苣苔属
Didymocarpoideae
Trichosporeae
Loxocarpinae
S China 1 1
(1)
No change
Rhynchoglossum
Blume
尖舌苣苔属
Didymocarpoideae
Epithemateae
Loxotidinae
From India and S
China to New
Guineaꎬ one (to
3?) spp. in C
America
≈15 2
(1)
No change Recent descrip ̄
tion of 2 new and
morphologically
odd spp.from
Thailand ( Pat ̄
tharahirantricin
2014)Rhynchotechum
Blume
线柱苣苔属
Didymocarpoideae
Trichosporeae
Leptoboeinae
NE Indiaꎬ Nepalꎬ
Bhutanꎬ SW & S
Chinaꎬ SE Asia and
Malesia to New
Guinea
21 6
(2)
No change
Stauranthera
Benth.
十字苣苔属
Didymocarpoideae
Epithemateae
Loxoniinae
Malesiaꎬ S China ≈5 1 No change Revision in prep.
(Weberꎬ in
prep.)
Streptocarpusꎬ Asiat ̄
ic species
扭果花属 /海角苣
苔属
- - Sunk into Damrongia Puglisi et alꎬ
in press
See text under
Damrongia
Tengia Chun
世纬苣苔属
- - Sunk into Petrocodon Wang et alꎬ
2011ꎻ Weber
et alꎬ 2011b
See text under
Petrocodon
Thamnocharis W. T.
Wang
辐花苣苔属
- - Sunk into Oreocharis Möller et alꎬ
2011b
See text under
Oreocharis
Titanotrichum
Soler.
台闽苣苔属 /俄氏
草属
Gesnerioideae
Titanotricheae
SE Chinaꎬ Taiwan
region of Chinaꎬ S
Japan
1 1 Placed in subfam.
Gesnerioideae
Wang et alꎬ
2004aꎻ
Perret et alꎬ
2013ꎻ Weber
et alꎬ 2013
151期 MÖLLER Michael et al. 得与失: 苦苣苔科新的属级界定与分类系统———中国该科植物之变迁
续表2
Genus
(alphabet.)
(Chinese name)
Infrafamilial
position:
Subfamily
Tribe Subtribe
Distribution Speciesnumber
Species no.
in China
(endemic)
Tax. status Reference Remarks
Tremacron Craib
短檐苣苔属
- - Sunk into Oreocharis Möller et alꎬ
2011b
See text under
Oreocharis
Trisepalum C. B.
Clarke
唇萼苣苔属
- - Sunk into Paraboea Puglisi et alꎬ
2011
See text under
Paraboea
Wentsaiboea
D.Fang & D.H.Qin
文采苣苔属
- - Partly ( incl. type )
sunk into Primulinaꎬ
partly into Petrocodon
Weber et alꎬ
2011aꎬb
See text under
Chirita and Pet ̄
rocodon
Whytockia
W. W. Sm.
异叶苣苔属
Didymocarpoideae
Epithemateae
Monophyllaeinae
S Chinaꎬ Taiwan re ̄
gion of China
8 8
(8)
No change
Note: Presently accepted genera in bold face. Genera that have been synoymised since the publication of the “Flora of China” (Wang 1990ꎬ Wang et alꎬ 1998) are given
in square brackets and not bold. Infrafamilial position according to Weber et al (2013) .
With regard to subfam. Didymocarpoideaeꎬ both
tribesꎬ Epithemateae and Trichosporeaeꎬ are represen ̄
ted in China.
In tribe Epithemateaeꎬ which is apparently a rel ̄
ict tribe with morphologically rather odd and isolated
generaꎬ all four subtribes are represented in China.
From the seven genera two are missing in China: Mono ̄
phyllaea in subtribe Monophyllaeinae ( this thus con ̄
tains only Whytockiaꎬ with some six species in South
China and Taiwan region of China)ꎬ and Loxonia in
Loxoniinae (containing in China Stauranthera andꎬ with
some uncertaintyꎬ the ill ̄known and possibly extinct Gy ̄
rogyne). In both cases the genera are distributed in the
SE Asian tropics: Monophyllaea includes more than 40
speciesꎬ with distribution throughout Malesia (Sumatra
to New Guinea and S Thailand to Java)ꎬ Loxonia com ̄
prises three species restricted to western Malesia (Su ̄
matraꎬ Malay Peninsula and Borneo). Whytockia is par ̄
ticularly remarkable as morphological studies (Weberꎬ
1976ꎬ 1982) indicate that the genus has retained primi ̄
tive characters in relation to Monophyllaeaꎬ suggesting
that subtribe Monophyllaeinae has originated in the ex ̄
tratropics andꎬ after reaching the South East Asian trop ̄
icsꎬ radiated into the many species of Monophyllaea.
Stauranthera and Gyrogyne are represented with a single
species each in Chinaꎬ Rhynchoglossum and Epithema
with two species each.
Tribe Trichosporeaeꎬ with 10 subtribes tentatively
recognised by Weber et al (2013)ꎬ is represented by
five subtribes in China.
Among the more primitive alliancesꎬ subtribe Cor ̄
allodiscinaeꎬ with the single genus Corallodiscusꎬ is re ̄
markable. Its few species are rosette plants with scapose
inflorescences and tetrastaminate flowers.
Morphologically heterogeneous is subtribe Lepto ̄
boeinaeꎬ which is represented by almost all of its gene ̄
ra in China (the only lacking genus is the South Indian
Championiaꎬ the inclusion of which in Leptoboeinae
still requires confirmation). Leptoboeinae includes ro ̄
sette plants ( Beccarinda )ꎬ sub ̄shrubby caulescent
plants (Leptoboeaꎬ Boeicaꎬ Rhynchotechum) and unifo ̄
liate plants ( Platystemma ). Leptoboeaꎬ Boeica and
Rhynchotechum have (correctly!) thought to be closely
related in older classificationsꎬ but Burtt (1963) as ̄
sumed a close relationship of Rhynchotechum with Cyr ̄
tandraꎬ placing both in subfam. Cyrtandroideae tribe
Cyrtandreaeꎬ because the two genera have indehiscentꎬ
berry ̄like fruits in common. Apart from the changes in
taxonomic position as compared to former classifica ̄
tionsꎬ no changes in the definition of the genera have
been prompted by molecular systematic studies.
Another monogeneric subtribe is Litostigminae.
The genus Litostigma was recently established in 2010ꎬ
based on newly collected material from Guizhou and
Yunnan. The genus represents an important morphologi ̄
cal link between the basal lineages of tribe Trichospore ̄
ae with seeds without elaborate testa cell ornamentationꎬ
straight fruit and septicidal and loculicidal dehiscence
and derived lineages with predominantly loculicidal de ̄
hiscence and two stamens (Wei et alꎬ 2010).
25 广 西 植 物 36卷
Subtribe Loxocarpinae is the second largest sub ̄
tribe of Trichosporeae. In total it includes about a dozen
genera. Two of themꎬ Dorcoceras and Middletoniaꎬ have
been (re ̄)established very recently ( Puglisi et alꎬ in
press)ꎬ and both are represented in Chinaꎬ together
with Damrongiaꎬ Ornithoboeaꎬ Paraboea and Rhabdoth ̄
amnopsis. The latter is with a single species endemic to
Chinaꎬ while the remaining genera have their main dis ̄
tribution mostly further south. A characteristic feature of
the subtribe (but not present in all taxa) is the twisted
capsule. Both in this and the following subtribe many
changes in generic delimitations have been induced by
the recent molecular ̄systematic investigations. These are
listed in Table 2 and discussed in the following chapter.
Subtribe Didymocarpinae is by far the largest
subtribe of tribe Trichosporeae. It includes over 30 gene ̄
raꎬ with 24 represented in China. Six genera with one or
very few species (i.e. Allocheilosꎬ Allostigmaꎬ Briggsi ̄
opsisꎬ Cathayantheꎬ Didymostigmaꎬ Metapetrocosmea)
are endemic to this countryꎬ others have their main dis ̄
tribution elsewhere and reach with one or few species
the southernmost part of China (e. g.ꎬ Cyrtandraꎬ with
>800 species is the largest genus of Gesneriaceaeꎬ rea ̄
ches with a single species to South Taiwanꎬ China). Mi ̄
crochirita with a wide distribution from the Western
Ghats of Indiaꎬ the foothills of the Himalayasꎬ through
continental SE Asia into Sumatraꎬ Borneo and Javaꎬ has
two species in China (out of c. 18).
The subtribes of Trichosporeae not represented in
China are Jerdoniinae (only Jerdonia indicaꎬ S India)ꎬ
Tetraphyllinae (only Tetraphyllumꎬ 3 spp.ꎬ NE Indiaꎬ
Bangladeshꎬ Burmaꎬ Thailand)ꎬ Ramondinae (Haber ̄
leaꎬ Ramondaꎬ Jancaeaꎻ 5 spp.ꎬ SW and SE Europe)ꎬ
Streptocarpinae ( traditionally 9 generaꎬ recently all a ̄
malgamated in Streptocarpusꎬ Nishii et alꎬ 2015ꎻ 176
spp.ꎻ Africaꎬ Comoro Islands and Madagascar)ꎬ and
Didissandrinae (Didissandraꎬ Tribouniaꎻ c. 10 spp.ꎬ W
Malesiaꎬ Thailand). Altogetherꎬ these subtribes include
only one or few generaꎻ as to species numberꎬ Strepto ̄
carpinae is clearly the largest subtribe. The link from
the African to the Asiatic (including Chinese) species
via the “Asiatic species of Streptocarpus” proved unten ̄
able: the Asiatic species described in Streptocarpus be ̄
long to the purely Asian genus Damrongia (Puglisi et
alꎬ in press).
3 Taxonomic fate of the Chinese
genera of Gesneriaceae
In Table 2 all genera that have been used to ac ̄
commodate Chinese (including Taiwan) species are lis ̄
ted and briefly commented on. In some genera (that isꎬ
for instanceꎬ all genera of tribe Epithemateae) no chan ̄
ges in the taxonomic delineation have occurredꎬ in oth ̄
ers the inclusion of formerly distinct genera has little
bearing on their new definition (e. g. the inclusion of
Metabriggsia into Hemiboeaꎬ in which differences in the
ovary structure simply proved erroneousꎬ Weber et alꎬ
2011c). Howeverꎬ others have a rather complex taxo ̄
nomic history which require a more detailed explanation.
In the following the fate of the latter generaꎬ plus new or
newly established genera is briefly outlinedꎬ as far as
changes since the treatment of Wang et al (1990) and
Wang et al (1998) are concerned. De factoꎬ all changes
occurred from 2010 onwardsꎬ with the new genus
Litostigma (Wei et alꎬ 2010) as the first.
Boea. This was previously a widespread Asiatic ̄
Malesian genusꎬ with three species (B. clarkeanaꎬ B.
hygrometricaꎬ and B. philippensis) represented in south ̄
ern China. From the recent molecular ̄systematic studies
of Puglisi et al (in press)ꎬ Boea emerged as an essen ̄
tially Australasian genusꎬ with distribution in Eastern
Indonesiaꎬ Papua New Guineaꎬ the Solomon Islands and
Queensland (Australia). The genus Dorcoceras was re ̄
established to accommodate the remaining species dis ̄
tributed in Burma (1 sp.)ꎬ Cambodia (1 sp.) and Chi ̄
na (2 spp.). In additionꎬ Boea hygrometrica (endemic
to China) and B. philippensis ( Philippines and Viet ̄
nam) were included in Dorcoceras (the former represen ̄
ting the type species of Dorcoceras). Boea clarkeana (≡
Streptocarpus clarkeanus)ꎬ howeverꎬ was attributed to
the genus Damrongia (see there).
Briggsia. The genus is no longer relevantꎬ as the
alliance around the type species (B. longifolia) has
been synonymised with Oreocharis ( Möller et alꎬ
2011bꎬ 2014). In its traditional conceptꎬ Briggsia in ̄
351期 MÖLLER Michael et al. 得与失: 苦苣苔科新的属级界定与分类系统———中国该科植物之变迁
cluded acaulescent rosette plants as well as caulescent
plantsꎬ both with characteristic large and ventrally
pouched (“briggsioid”) flowers. Its three acaulescent
and strikingly glabrous species are now in the new genus
Glabrella and the three caulescent species are in Lox ̄
ostigma (Möller et alꎬ 2014ꎻ Wen et alꎬ 2015aꎬb).
The “briggsoid” flowers arose apparently independently
in different alliances (Möller et alꎬ 2011b). See also
notes for Oreocharis.
Chirita. Dramatic changes occurred in this tradi ̄
tional and large genus (with up to 140 species described
hereinꎬ placed in four sections: sect. Chiritaꎬ sect.
Liebigiaꎬ sect. Microchiritaꎬ and sect. Gibbosaccusꎻ
Woodꎬ 1974ꎻ Hilliardꎬ 2003)ꎬ ending in a synonymisa ̄
tion of the genus with Henckelia (Weber et alꎬ 2011a).
The character defining Chirita was traditionally seen in
the “chiritoid” stigma: a stigma with upper and lower
lobeꎬ but the upper largely or completely reducedꎬ and
the lower one expandend and often bipartite. The molec ̄
ular studies clearly revealed that this character is homo ̄
plasticꎬ having evolved independently in several alli ̄
ances of tribe Trichosporeae. As a consequenceꎬ Chirita
was split into five generaꎬ with the majority of Chirita
sect. Chirita (and the monotypic Hemiboeopsisꎬ China
and Laos) amalgamated with Henckelia sect. Henckelia
(from South India). The remaining species of Chirita
sect. Chirita were included in the revived genus Dam ̄
rongia (see below). Chirita sect. Liebigia was raised to
generic level (Liebigia Endl.ꎬ not represented in Chi ̄
na). Chirita sect. Microchirita was also raised to genus
level (Microchiritaꎬ with two species in China)ꎬ and
Chirita sect. Gibbosaccus wasꎬ together with Chiritopsis
and Wentsaiboea (including its type species and a later
published speciesꎬ W. luochengensisꎬ Liu et alꎬ 2010ꎻ
Xu et alꎬ 2012)ꎬ included)ꎬ included in the originally
monotypic genus Primulina (Wang et alꎬ 2011ꎻ Weber
et alꎬ 2011aꎻ Li & Xiaꎬ 2012). The latter genus is es ̄
sentially geographically restricted to Chinaꎬ with some
species extending or endemic to Vietnam.
Primulina has been expanded from a monotypic ge ̄
nus to one with 100 species at the time of its redefinition
(Weber et alꎬ 2011a) and over the last few years has be ̄
come a genus with over 160 species. The morphological
variation is relatively limited compared to other generaꎬ
and is based on diandrous zygomorphic flowers usually in ̄
fundibuliform but variable in size and colorationꎬ and a
vegetative habit with rhizomatous compact stems with
leaves in a basal rosetteꎬ with decussate phyllotaxy or in
whorls of three (rarely alternate). This character distin ̄
guishes this genus to a great length from Petrocodonꎬ
where the leaves are always alternate. Another possibly
distinguishing feature are the chromosome numbers which
are almost uniformly 2n = 36 in Primulina (>100 species
countedꎬ with one exception of a tetraploid numberꎬ
Christie et alꎬ 2012)ꎬ whereas it is 2n = 20 in the only
member of Petrocodon so far counted (P. hanceiꎬ Cao et
alꎬ 2003ꎻ Möller & Pullanꎬ 2015 onwards).
Damrongia. This is a genus re ̄established by We ̄
ber et al (2011a) for the accomodation of a couple of
species previously placed in Chirita. The species repre ̄
sent rosette plants with scapose inflorescencesꎬ infun ̄
dibuliform corollas with 2 stamensꎬ and a pistil with a
“chiritoid” stigma (see Chirita). It was thought to be
restricted to Thailand and the NW of Peninsular Malay ̄
siaꎬ but the Chinese Boea clarkeana also proved to be ̄
long to that genus (Puglisi et alꎬ in press). The same
applies for the Asian species described in Streptocarpus
on grounds of the twisted fruits ( Puglisi et alꎬ in
press).
Dorcoceras. Re ̄established genus segregated from
Boea by Puglisi et al (in press). See Boea above.
Glabrella. New genus established by Möller et al
(2014) for two species of Briggsiaꎬ which do not fit in ̄
to Oreocharis or Loxostigma. A third species was added
by Wen et al (2015aꎬb). See Briggsia and Oreocharis.
Henckelia. The genus Henckelia Spreng. was re ̄es ̄
tablished when the unwieldy and broadly circumscribed
Didymocarpus was changed and split into 3 smaller enti ̄
ties: Didymocarpus s. str.ꎬ Henckelia (with five sec ̄
tions: sect. Henckeliaꎬ S India and Sri Lankaꎬ sect.
Loxocarpusꎬ sect. Heteroboeaꎬ sect. Loxocarpusꎬ sect.
Didymanthus and sect. Glossadenia ) and Hovanella
(Weber & Burttꎬ 1998). The molecular data of Möller
et al (2009) indicated that the split was not sufficient
and that Henckelia had to be redefined. This was done
by Weber et al (2011a). The type section (Henckelia.
45 广 西 植 物 36卷
sect. Henckeliaꎬ S India and Sri Lanka) was amalgama ̄
ted with most of Chirita sect. Chirita and the monotypic
Hemiboepsis ( see under Chirita above)ꎬ sect. Loxo ̄
carpus has returned to generic level ( Weber et alꎬ
2011aꎻ Middleton et alꎬ 2013ꎻ Yaoꎬ 2012)ꎬ and the
last three sections have been included in the re ̄estab ̄
lished and largely expanded genus Codonoboea (Kiew &
Limꎬ 2011ꎻ Middleton et alꎬ 2013). Neither Loxocarpus
nor Codonoboea have species in China.
Litostigma. This new genus established by Wei et
al (2010) for two Chinese species form a separate sub ̄
tribe (Litostigminae) (see there).
Microchirita. This genus has been established by
raising Chirita sect. Microchirita to generic level (Wang
et alꎬ 2011ꎻ Weber et alꎬ 2011a). It can be character ̄
ized by inflorescences that usually appear in two or sev ̄
eral in a leaf axilꎬ often displaced onto the petioleꎬ of ̄
ten consisting of a short ̄stalked serial flower pair onlyꎬ
but this repeated several times. From the c. 25 species 2
occur in China: M. hamosa and M. prostrata (recently
described by Li & Xiaꎬ 2012).
Middletonia. A new genus established by Puglisi
et al (in press) for the accommodation of four species
separated from Paraboeaꎬ with one species distributed
in China (M. multiflora).
Oreocharis. While Oreocharis included some 27
species in its traditional delineation ( Wang et alꎬ
1998)ꎬ the number increased enormously through the
inclusion of Ancylostemonꎬ Bourneaꎬ Briggsia p. p.
(incl. type)ꎬ Dayaoshaniaꎬ Deinocheilosꎬ Isometrumꎬ
Opithandraꎬ Paraisometrumꎬ Thamnocharisꎬ and Trem ̄
acron (Möller et alꎬ 2011bꎻ Middleton et alꎬ 2013). As
the species of Briggsia transferred to Oreocharis includ ̄
ed the type species (B. longifolia)ꎬ the remaining spe ̄
cies were left without generic placement. In recent stud ̄
ies (Chen et alꎬ 2014bꎻ Möller et alꎬ 2014ꎻ Wen et alꎬ
2015aꎬb)ꎬ the rest were attributed to the following three
genera: the caulescent species were referred or returned
to Loxostigmaꎬ three acaulescent species with glabrous
leaves and stems were moved into a new genus Glabrel ̄
laꎬ and 11 acaulescent species were transferred to Oreo ̄
charis. New species were recently described (Liu et alꎬ
2012ꎻ Chen et al 2013ꎬ 2015ꎻ Tan et al 2013ꎬ 2015ꎻ
Rossini & Freitas 2014ꎻ Li & Li 2015ꎬ Yang et alꎬ
2015)ꎬ and Oreocharis comprises 106 speciesꎬ all repre ̄
senting acaulescent rosette plants with scapose inflores ̄
cencesꎬ but with considerable variability in the symme ̄
tryꎬ shape and coloration of the flowers. Oreocharis leio ̄
phylla and O. sinensis (making up the former genus
Bournea) have actinomorphic flowers. The corolla shape
in Oreocharis varies from tubularꎬ through funnel ̄
shapedꎬ campanulate to “briggsioid”ꎬ the stamen num ̄
ber is (1)ꎬ 2ꎬ 4 or (in actinomorphic flowers) 4 or 5 in
line with the number of petalsꎬ in some diandrous spe ̄
cies (previously constituting the genus Opithandra) the
posterior stamens are the fertile ones (while it is the re ̄
verse in most other diandrous Gesneriaceae)ꎬ and the
anthers may be fused or free and coherent at the tips or
at the faces. These floral characters have been formerly
used to define generaꎬ butꎬ as the molecular data showꎬ
the species relationships run across genera and these are
thus untenable. Like in Petrocodonꎬ the flowers combine
a range of corolla shapes and colours suggesting different
pollinator adaptationsꎬ sometimes in parallel.
Paraboea. This widespread and species ̄rich genus
was revised by Xu et al (2008)ꎬ who recognised 89
speciesꎬ c. 20 of them found in China. Changes since
then relate to the inclusion of the monotypic Phylloboea
(Myanmar) and Trisepalum (c. 13 spp.ꎬ 1 in China)
(Puglisi et alꎬ 2011). More recentlyꎬ Puglisi et al (in
press) removed four species from Paraboea and placed
them in the new genus Middletonia. In the new delinea ̄
tion the genus contains 132 species of which 26 occur in
Chinaꎬ of these 14 exclusively there (i.e. endemic).
Petrocodon. In its traditional conceptꎬ Petrocodon
was a genus with two Chinese speciesꎬ both with smallꎬ
white flowers and two fertile stamens. Based on molecu ̄
lar dataꎬ Wang et al (2011) added Tengia (with small
whiteꎬ pentamerous actinomorphic flowers and five sta ̄
mens)ꎬ Calcareoboea (with largeꎬ red and tubular flow ̄
ers)ꎬ and Paralagarosolen (with long ̄tubed hypocrater ̄
iform flowers). Shortly afterwardsꎬ Weber et al (2011b)
added the monotypic Dolicholoma ( with flowers with
narrow tube and subactinomorphic limb with longꎬ acute
lobes)ꎬ all species of Lagarosolen (with flowers similar
to those of Dolicholoma and Paralagarosolen)ꎬ one spe ̄
551期 MÖLLER Michael et al. 得与失: 苦苣苔科新的属级界定与分类系统———中国该科植物之变迁
cies of Wentsaiboea ( excl. type) and three species of
Didymocarpus (D. hanceiꎬ D. mollifoliusꎬ D. niveolano ̄
sus). This resulted in a genus size of around 20 species.
The recent description of new species by Wen et al
(2012)ꎬ Chen et al(2014a)ꎬ Hong et al (2014)ꎬ Xu
et al (2014) and Li & Wang (2015) increased the spe ̄
cies number to almost 30. The different flower shapes
and colours now assembled in Petrocodon suggest differ ̄
ent pollinator adaptationsꎬ including beeꎬ butterfly and
bird pollination. Until recentlyꎬ Petrocodon could be de ̄
fined by possessing 2 stamens or 5 (in the former genus
Tengia with actinomorphic flower). Using both morpho ̄
logical and molecular dataꎬ a species possessing 4 sta ̄
mens was described: P. hunanensis (Yu et alꎬ 2015).
This indicates (a) the great diversity that is still unex ̄
plored in Chinaꎬ and (b) that the use of molecular data
has apparently given the right signal to place the species
in the right genus.
4 Discussion and Outlook
4.1 You win someꎬ you lose some
The recent taxonomic changes have changed the
generic picture of Gesneriaceae in China considerably.
From the 56 genera listed in Wang et al (1998)ꎬ and
two genera established a few years later (Paralagaroso ̄
len: Weiꎬ 2004ꎻ Wentsaiboea: Fang & Qinꎬ 2004)ꎬ the
number has dropped to 45 (Table 1). Since 2010ꎬ and
essentially based on molecular dataꎬ seven new genera
have been added to Chinaꎬ either based on newly col ̄
lected material (Litostigma)ꎬ or from segregation of ex ̄
isting genera in China (Glabrellaꎬ Damrongiaꎬ Dorco ̄
cerasꎬ Middletoniaꎬ Microchiritaꎬ and Primulinaꎻ see
previous chapter)ꎬ or by a new circumscription of gene ̄
ra (Henckelia).
On the other handꎬ a considerable number of gene ̄
ra (22) are lost by synonymisation with other genera
(Ancylostemonꎬ Boeaꎬ Bourneaꎬ Briggsiaꎬ Calcareo ̄
boeaꎬ Chiritaꎬ Chiritopsisꎬ Dayaoshaniaꎬ Deinocheilosꎬ
Dolicholomaꎬ Hemiboeopsisꎬ Isometrumꎬ Lagarosolenꎬ
Metabriggsiaꎬ Opithandraꎬ Paraisometrumꎬ Paralagaro ̄
solenꎬ Tengiaꎬ Thamnocharisꎬ Tremacronꎬ Trisepalumꎬ
Wentsaiboea). Alsoꎬ the four Asiatic species of Strepto ̄
carpus (Hilliard & Burttꎬ 1971)ꎬ one of themꎬ S. clar ̄
keanus (Hemsl.) Hilliard and B.L.Burtt (≡Boea clar ̄
keana Hemsl.ꎬ as such in the Flora of China of Wang et
alꎬ 1990ꎻ Wang et alꎬ 1998)ꎬ from Chinaꎬ have found
a proper generic place (Damrongia ). Streptocarpusꎬ
thereforeꎬ can be definitely removed from the list of
genera extending into China.
By tribeꎬ there is no change in generic state in Ep ̄
ithemateaeꎬ as to Trichosporeae ̄Litostigminae there is
the addition of one genus (Litostigma)ꎬ in Trichospore ̄
ae ̄Loxocarpinae there are two fewer genera in China
(Boeaꎬ Trisepalum)ꎬ but three additional ones (Dam ̄
rongiaꎬ Dorcocerasꎬ Middletonia)ꎬ an overall net gain
of one. The greatest changes occurred in Trichosporeae ̄
Didymocarpinaeꎬ with a loss of 20 generaꎬ and a gain of
three ( Glabrellaꎬ Henckeliaꎬ Microchirita )ꎬ thus an
overall loss of 17 genera. This tribe is still not fully un ̄
derstood and further work is necessary to understand its
evolutionꎬ and further changes may be in the waitingꎬ
though these may be less drasticꎬ but fine ̄tuning.
The number of Gesneriaceae genera endemic to
China has decreased from 27 (Wang et alꎬ 1998ꎻ Wei
2004ꎻ Fang & Qinꎬ 2004) to 11. Ten of them included
only one to three species (Allocheilos [2]ꎬ Allostigma
[1]ꎬ Briggsiopsis [1]ꎬ Cathayanthe [1]ꎬ Didymostig ̄
ma [3]ꎬ Glabrella [ 3]ꎬ Gyrogyne [ 1]ꎬ Litostigma
[2]ꎬ Metapetrocosmea [1]ꎬ Rhabdothamnopsis [1])ꎬ a
drop from 22 previously. Of the remaining endemic gen ̄
eraꎬ almost a dozen have unusual morphologies com ̄
bined with isolated evolutionary positions(Möller et alꎬ
2011a).
Over the decadesꎬ there seems to be an overall
tendency to include small or monotypic genera (and the
large Dichrotrichum) that had been raised on the basis
of the presence of an unusual characteristicꎬ into larger
ones on grounds of more shared than distinguishing
characteristics. This is importantꎬ since single character
taxonomy is fraught with problems when picking the
‘wrong’ character a genus was not established for.
4.2 Morphological parallelisms and reversals
Burtt ( e. g. Burttꎬ 1963) pointed out that in
Gesneriaceae there are many exceptions and apparent
parallelisms in formꎬ and that the homology of similar
65 广 西 植 物 36卷
characters on which a genus is basedꎬ is sometimes dif ̄
ficult to ascertain (e. g. Burttꎬ 1968). The erstwhile ge ̄
nus Chirita is a prime exampleꎬ in which strong empha ̄
sis was given to the special type of stigma (“chiritoid
stigma ”). The molecular data have unambiguously
shown that the chiritoid stigma has evolved several times
independentlyꎬ and Chirita thus had to be split into five
genera (Weber et alꎬ 2011a) (see also below).
At the generic levelꎬ the number of fertile stamens
(4 or 2) played a significant roleꎬ especially in Chinese
Gesneriaceae. For exampleꎬDeinocheilos ( with 2 sta ̄
mens) has been kept separate from Tremacron (4 sta ̄
mens)ꎬ though the two genera are otherwise indistin ̄
guishable.
Floral symmetry is another problematic character of
importance both at the generic and at the tribal level.
Fritsch (1893 / 1894ꎬ 1908) established tribe Ramonde ̄
ae to include Corallodiscusꎬ Haberleaꎬ Petrocosmeaꎬ
Ramonda and Saintpaulia. His intention was to recog ̄
nise a group with similar vegetative habit ( flat rosette
plants) with septicidal capsule dehiscenceꎬ rather than
floral shape. Thusꎬ this tribe included four genera with
zygomorphic flowers (two of which have long floral tubes
and four stamens: Haberlea and Corallodiscusꎬ and two
with flat ̄faced corollas with 2 stamens: Petrocosmea and
Saintpaulia) ꎬ and one with actinomorphic flowers
( “Ramondia” = Ramonda). He placed another genus
with actinomorphic flowersꎬ Conandronꎬ in a separate
tribe. Burtt (1970) accepted this view and suggested
that actinomorphic genera were not relatedꎬ that zygo ̄
morphy was ancestral in the family and actinomorphic
genera the result of separate independent losses of flower
asymmetry. Wang et al (1990) and Wang et al (1992)
held the opposite view and considered actinomorphy as
ancestral. They redefined tribe Ramondeae to include
only genera with actinomorphic flowers: Bourneaꎬ Co ̄
nandronꎬ Ramondaꎬ Tengia and Thamnocharis. Molecu ̄
lar phylogenetic work has proven Burtt’s view to be cor ̄
rect since they showed that all actinomorphic genera
have independent origins (Möller et alꎬ 1999ꎬ 2009ꎬ
2011aꎻ Wang et alꎬ 2010).
Hidden homoplasies destabilize a classification sys ̄
tem that wants to reflect phylogenyꎬ and single character
taxonomy may lead to an inflation of small or monotypic
genera. With the rise of molecular phylogenetic methods
during the last 20 years or soꎬ we have become more a ̄
ware of the presence and levels of homoplasies among
morphological characters and how small and monotypic
genera are related or part of larger entities (Möller et
alꎬ 2011a). This has been shown for the Neotropical
Gesneriaceae as well as for the Afro ̄Malagasy Gesneri ̄
aceae of tribe Streptocarpinaeꎬ which have been reduced
from nine often small genera (7 with <3 species) to one
genus Streptocarpus (Nishii et alꎬ 2015). There are nu ̄
merous examples of morphological homoplasies that have
resulted in dramatic changes in classification for the
New World Gesneriaceae. The convergence of resupinate
flowers was recently discovered to be independently de ̄
rived in three lineages that resulted in a new generic
classification for members of the Columneinae (Clark &
Zimmerꎬ 2003ꎻ Clark et alꎬ 2006). The convergence of
hypocyrtoid or “pouched” flowers within Drymonia and
across several lineages of closely related genera has re ̄
sulted in a revised circumscription of genera in the Col ̄
umneinae (Clark et alꎬ 2012ꎻ Smith & Clark 2013ꎻ
Clark et alꎬ 2015). Recent phylogenetic results have
elucidated the convergence of radially symmetrical flow ̄
ers and that has resulted in the circumscription of more
narrowly defined genera in the Gloxiniinae (Smith et alꎬ
2004ꎻ Roalson et alꎬ 2005bꎻ Clark et alꎬ 2011).
4.3 Holistic approach to the taxonomy
Uncertainties in the correct genus assignment
abound. Photographs of Litostigma crystallina were pub ̄
lished under Petrocosmea crystallina ( Shui & Chen
2006) before its accommodation in a new genus (Wei et
alꎬ 2010). The recently described speciesꎬ Primulina
guangxiensis was initially placed with a superficially
similar species (Liu et alꎬ 2011)ꎬ but after molecular
studiesꎬ partly by the same authorsꎬ it was discovered
that it belonged in Petrocodon (assuming the integrity of
the molecular data) and was transferred to this genus
(Xu et alꎬ 2014).
In some cases incomplete knowledge of generic
concepts might be at issueꎬ in others perhaps the dis ̄
covery of species with new character combinationsꎬ such
as Petrocodon hunanensis with four stamensꎬ cause un ̄
751期 MÖLLER Michael et al. 得与失: 苦苣苔科新的属级界定与分类系统———中国该科植物之变迁
certainty. Howeverꎬ with the availability of molecular
data and an extensive data set of molecular sequences
on public databasesꎬ such as GenBankꎬ these uncer ̄
tainties can be addressed by molecular approaches.
Howeverꎬ where traditional taxonomic approaches
are concernedꎬ the utilisation and balancing of all char ̄
actersꎬ not just one or a fewꎬ has to be considered in
taxonomic decisions. This might have become more com ̄
plex with the new delineations of some generaꎬ and the
‘traditional’ characters of corolla shapeꎬ stigma or an ̄
droecium may be largely inadequate. Additional or hith ̄
erto undervalued characters may be required.
4.4 Open issues
At the suprageneric taxonomic level we present
classification on the basis of what is currently known.
The classification of tribe Trichosporeaeꎬ particularly
subtribe Didymocarpinaeꎬ is still to be completed. Be ̄
cause of the large number of representatives in this sub ̄
tribe residing in Chinaꎬ it can play a pivotal role in this
endeavour.
At the generic levelꎬ Raphiocarpus is still an unsat ̄
isfactorily known entity. The data so far indicate intri ̄
cate links to several genera including Loxostigma. The
inclusion of the caulescent ‘Briggsia’ into Loxostigma
(Möller et alꎬ 2014) has expanded its definition to in ̄
clude seeds without appendagesꎬ narrowing its gap to
Raphiocarpusꎬ but its link to this genus is unexplored.
The relationship between Didymocarpus and Gyro ̄
cheilos is still unresolved. Some Chinese species of Did ̄
ymocarpus were transferred to Petrocodon (Weber et alꎬ
2011b). In the phylogeny of Möller et al (2011a)ꎬ D.
cortusifolius fell as sister to Gyrocheilosꎬ apart from Did ̄
ymocarpus properꎬ and may belong in this genus. Li et
al (2015) published a molecular phylogeny on Chinese
species and found similar results to Möller et al
(2011a). Perhaps other basal rosette ̄forming ones from
sect. Heteroboea may follow as suggested by Weber &
Burtt (1998).
The new circumscriptions of Henckelia and Dam ̄
rongia are quite wide. They include acaulescent and
caulescent speciesꎬ and the latter species with straight
and twisted fruits. Such an assemblage seems at first
dissatisfyingꎬ but it has precedence in other genera. The
genus Streptocarpus was characterised for species posses ̄
sing twisted fruits. Though it includes a wide range of
morphological formsꎬ from unifoliatesꎬ plurifoliatesꎬ ro ̄
sulatesꎬ rosettesꎬ herbaceous and woody caulescent
forms (Hilliard & Burttꎬ 1971). It has now also been
extended to include species with non ̄twisted fruits
(Nishii et alꎬ 2015).
4.5 Outlook
The work summarised here is very much a work in
progress and far from complete. The finding of new gen ̄
era based on new collections (e. g. Litostigma)ꎬ and
morphotypes that expand the new circumcriptions (e. g.
Petrocodon hunanensis with four stamens)ꎬ are examples
which demonstrate that the full diversity in Gesnericeae
in China is not yet known and is an exciting prospect for
finding further ‘links’ through intensified and systemat ̄
ic fieldwork. Alsoꎬ the great number of new species de ̄
scribed in some generaꎬ especially Primulinaꎬ which
ballooned from around 100 when it was redefined in
2011ꎬ to currently >160 with many more species to be
describedꎬ illustrates the need for continued floristic
surveys and detailed fieldwork to fully document the di ̄
versity of Gesneriaceae in China. Systematic fieldwork
efforts combined with detailed herbarium studies and
molecular work has unearthed several genera over the
last few years from Thailand and Vietnam (e. g. Somra ̄
niaꎬ Middleton & Triboun 2012ꎻ Tribouniaꎬ Middleton
et alꎬ 2012ꎻ Billoliviaꎬ Middleton et alꎬ 2014ꎻ Chay ̄
amaritia Middleton et alꎬ 2015). Recent discoveries
outlined here clearly demonstrate the recent progress in
Gesneriaceae diversity of Chinaꎬ but more importantly
the future need to continue phylogenetic and taxonomic
studies for a long ̄term stable classification. Without a
doubtꎬ there is still much to be done.
Acknowledgements We thank the Chinese
Academy of Sciences Visiting Professorship scheme for
supporting part (MMO) of the research. We also thank
the Royal Botanic Garden Edinburghꎬ supported by the
Rural and Environment Science and Analytical Services
division (RESAS) in the Scottish Government.
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