You win some you lose someE updated generic delineations and classification of Gesneriaceae-implications for the family in China

Abstract

Over the last two decades molecular phylogenetic research on Gesneriaceae has greatly advanced our understanding 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 split，merged or newly defined. Additionally, new genera have been established based on newly collected material, illustrating on the one hand gaps in field work in China, and on the other hand the biological richness of the Gesneriaceae in China. Here, we summarize and present an overview of our work and the taxonomic consequences.

Full text

书书书
Guihaia Jan． 2016，36（1） ： 44－60 http：/ / journal． gxzw． gxib． cn
http：/ /www． guihaia－journal． com
DOI：10． 11931 / guihaia． gxzw201512015
MLLEＲ Michael，
韦毅刚，
温放，
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，2016，36（1） ：44－60
MLLEＲ M，WEI YG，WEN F，et al． You win some you lose some：updated generic delineations and classification of Gesneriaceae-implications for the fam-
ily in China［J］． Guihaia，2016，36（1） ：44－60
You win some you lose some：updated generic
delineations and classification of Gesneriaceae-
implications for the family in China
MLLEＲ Michael1*，WEI Yi-Gang2，WEN Fang2，CLAＲK John L． 3，WEBEＲ Anton4
（1． Ｒoyal Botanic Garden Edinburgh，Edinburgh EH3 5LＲ，UK；2． Guangxi Key Laboratory of Plant Conservation and Ｒestoration Ecology in Karst
Terrain，Guangxi Institute of Botany，Guilin 541006，China；3． Department of Biological Sciences，The University of Alabama，Tuscaloosa，
AL 35487，USA；4． Department of Botany and Biodiversity Ｒesearch，University of Vienna，Ｒennweg 14，A-1030 Vienna，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 fami-
ly that is thought to reflect the natural relationships of the taxa better than traditional morphological classifications． Dra-
matic taxonomic changes were implemented affecting the classification of Gesneriaceae in China． Many traditional genera
have been split，merged or newly defined． Additionally，new genera have been established based on newly collected ma-
terial，illustrating on the one hand gaps in field work in China，and on the other hand the biological richness of the
Gesneriaceae in China． Here，we summarize and present an overview of our work and the taxonomic consequences．
Key words：China，classification，Gesneriaceae，molecular phylogenies，systematics
CLC number：Q949． 4 Document code：AArticle 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
MLLEＲ Michael1*，韦毅刚2，温 放2，CLAＲK John L． 3，WEBEＲ Anton4
（1． 英国皇家爱丁堡植物园，爱丁堡，英国；2． 广西喀斯特植物保育与恢复生态学重点实验室，广西壮族自治区
中国科学院广西植物研究所，
广西 桂林 541006，中国；3． 生命科学部 阿拉巴马大学，塔斯卡卢萨，阿拉巴马州，美国；
4． 植物学与生物多样性研究部 维也纳大学，维也纳，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。该文提供了一个苦苣苔科植物新分类系统，
与传统的经典形态学分类系统比较，
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。众多传统意义上的属被分割、
合并或者重新定义，
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。此外，
基于最近采集的材料，
一些新属得以建立，
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 ） ； the National Natural Science Foundation of China
（31000258，31470306，31260038，31460159） ； the Key Laboratory of Plant Diversity and Biogeography of East Asia，Kunming Institute of Botany （KIB）
＆ the Chinese Academy of Sciences （2014CB954100）and Guangxi Natural Science Foundation （2015GXNSFBB139004 ＆ 2015GXNSFBA139105） ；
Guangxi Forest Science ＆ Technology Foundation （Gui Lin Ke Zi〔2014〕No． 27 ）．
Biography：MLLEＲ Michael （1960-） ，Male，German，Ph． D． ＆ Professor，focuses on the Old World Gesneriaceae，research interests on Phylogenetics ＆ Sys-
tematics，Finescale Biogeography，Character Evolution ＆ Evolutionary Development，Genome ＆ Chromosome Evolution，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；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。
在此，
针对目前已经完成的工作和分类结论，
对中国苦苣苔科植物的新分类系统进行了总结与概述，
并深入地讨
论了部分传统分类学上存在困惑的疑难属。
关键词：中国，分类，苦苣苔科，分子系统学，系统分类
Gesneriaceae are a medium-sized family in the or-
der Lamiales，with main distributions 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”，Wang et al
（1990）listed 413 species in 56 genera，in the later
English version （Wang et al，1998）the number in-
creased to 442 species，while the number of genera re-
mained the same：354 species and 25 genera （many of
them monotypic） ，respectively，were considered to be
endemic． Without a doubt，China is a significant centre
of diversity of the family，with the majority of taxa found
in China’s South and Southwest，in the provinces of
Yunnan，Guizhou，Guangdong and Guangxi Autono-
mous Ｒegion （Wang et al，1992）． The species density
decreases northwards and the furthest locality North are
the Yenshan Mts at 41°N．
In the last two decades，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 Gesneriaceae，in which an unproportionally high
number of taxonomic changes has been made． In the
following the partly dramatic changes of traditional ge-
neric concepts of Chinese Gesneriaceae are surveyed，
and the position of the Chinese Gesneriaceae in recent
classifications is shown and discussed．
1 Classifications of Gesneriaceae and
amendments at generic level
Traditionally，the family was subdivided into two
subfamilies，Gesnerioideae and Cyrtandroideae （e． g． ，
Bentham，1876；Fritsch，1893 － 94；Burtt，1963）． The
last formal classification based on morphology （and
partly cytogenetic data）was that of Burtt and Wiehler
（1995） ，who recognized three subfamilies： （ 1）Gesne-
rioideae （56 genera；predominantly Neotropical） ； （ 2）
Coronantheroideae （9 genera，southern hemisphere：
temperate South America to Australia） ； （ 3）Cyrtan-
droideae （= Didymocarpoideae；82 genera，chiefly Pal-
eotropical，including three genera in Europe）． The dis-
tribution of the c． 3 400 species is roughly equal be-
tween the Neotropics and Palaeotropics．
Traditional classifications were based on a few key
morphological features such as seed anatomy，post-ger-
mination characteristics of the seedlings，nectary struc-
ture and ovary position． Thus，the Neotropical Gesneri-
aceae were characterized by the presence of seed endo-
sperm，isocotyly （both cotyledons showing equal and
limited growth after germination） ，a nectary often con-
sisting of separate glands，and often partly or fully infe-
rior ovary position，while the Palaeotropical Gesneriace-
ae were characterized by the lack of endosperm，by con-
tinued growth of one cotyledon after germination （aniso-
cotyly；Fritsch，1904；Burtt，1963，1970；Jong，1970；
Nishii et al，2004；Mantegazza et al，2007） ，a basical-
ly ring-shaped nectary and superior ovary position． Sub-
fam． Coronantheroideae was characterized by isocotylous
seedlings，a nectary ‘adnate’to the ovary base，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） ，Epi-
scieae （21） ，Beslerieae （8） ，Napeantheae （1）and
Gesnerieae （2） ； ［2］Coronantheroideae：only Coronan-
thereae （9） ，［3］．
Cyrtandroideae：Klugieae （7） ，Didymocarpeae
（64 ） ，Trichosporeae （6） ，Cyrtandreae （3） ，Tit-
anotricheae （1）．
The first classification taking to some extent molec-
ular data into account，was that of Weber （2004 ）．
However，particulary for the Palaeotropical Gesneriace-
ae the molecular data situation was insufficient and no
formal classification was presented． Weber （2004 ） ，
therefore，proposed a provisional classification，recog-
nising four informal groups： （ 1）Coronantheroid Gesne-
54
1期MLLEＲ 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

riaceae； （ 2）Gesnerioid Gesneriaceae； （ 3）Epithema-
toid Gesneriaceae； （ 4）Didymocarpoid Gesneriaceae．
The latter and the largest group was subdivided into
morpho-geographical assemblages
：‘Basal Asiatic gene-
ra’，‘European genera’，‘African and Madagascan
genera’and‘Advanced Asiatic and Malesian genera’．
Especially by the work of the first author （MMO）
and associates，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 parallel，also considerable progress
was made with regard to Neotropical Gesneriaceae，in-
cluding the work of one of the authors （JLC）． In 2013 a
new formal classification was published that was entirely
based on molecular data （Weber et al，2013）．
By the inclusion of Sanango，which was repeatedly
shown to be sister to Gesneriaceae in previous molecular
studies，three subfamilies were recognised；Sanan-
goideae （only Sanango） ，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 Beslerieae，Napenatheae and Gesnerieae）
a fifth，and rather aberrant，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 hemisphere，but with a single
species （Titanotrichum oldhamii）in East Asia （E Chi-
na，Taiwan region of China，S Japan）． The former Cyr-
tandoideae，now referred to as Didymocarpoideae，were
subdivided into two tribes：Epithemateae and Tri-
chosporeae． The former is apparently an ancient assem-
blage of few and rather isolated genera that fall into four
subtribes，while the latter includes a large number of gen-
era and is obviously still in a stage of active evolution． In
this tribe provisionally 10 subtribes have been recognised，
the by far largest being subtribe Didymocarpinae．
Apart from a new infrafamilial classification of the
family，another important aspect of the molecular work
is the improvement in the understanding of generic de-
limitations． In fact，a wealth of taxonomic changes re-
sulted from the molecular studies，both with respect to
Neotropical and Palaeotropical taxa． On the one hand，
many genera have been reduced to synonymy （e． g．
Mller et al，2011b；Weber et al，2011b，c；Nishii et
al，2015；Puglisi et al，in press） ，on the other hand，
some genera have been revived （e． g． Centrosolenia：
Mora ＆ Clark 2016；Crantzia：Clark，2005，Clark et
al，2006；Damrongia：Weber et al，2011a；Glossolo-
ma：Clark et al，2006；Clark，2009；Liebigia：Weber
et al，2011a；Mandirola：Ｒoalson et al，2005a，b；
Boggan 2006；Seemannia：Ｒoalson et al，2005a，b；
Boggan 2006；Trichodrymonia：Mora ＆ Clark 2016 ）
or newly described （e． g． Billolivia，Middleton et al，
2014；Chautemsia：Araujo et al，2010；Christopheria：
Smith ＆ Clark 2013；Chayamaritia：Middleton et al，
2015；Glabrella：Mller et al，2014；Gloxinella：Ｒoal-
son et al，2005a，b，Boggan，2006；Gloxiniopsis：
Ｒoalson et al，2005a，b；Boggan，2006；Lesia：Smith
＆ Clark，2013；Litostigma：Wei et al，2010；
Pachychaulos：Smith ＆ Clark，2013；Pagothyra：
Smith ＆ Clark，2013；Somrania：Middleton ＆ Triboun
，2012；Sphaerorrhiza：Ｒoalson et al，2005a，b；Tri-
bounia：Middleton ＆ Mller，2012）． In many cases
the species content changed，partly dramatically （e．
g． ，Primulina：increment from 1 to ＞150 species；Al-
loplectus：reduction from c． 140 to 5 species：Clark et
al，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 given，with indication of the position of gen-
era presently recognised for China． Distribution is ei-
ther endemic or includes adjacent countries （some gen-
era 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 Scrophu-
lariaceae． It is characterised by a terminal racemose
inflorescence and the presence of masses of seedling-
like propagules in the upper part of the inflorescence
（Wang et al，2004b ）． Burtt （1963，1977 ）ear-
64 广 西 植 物 36 卷

marked Titanotrichum as a ‘genus anomalum’within Gesneriace-
Table 1 Classification of Gesneriaceae acc． 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． Weber，J． L． Clark ＆ Mich． Mller 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． Woo，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． Mller 2 / 38+ Whytockia W． W． Sm．
3． 1． 3． Subtribe Loxoniinae A． DC． 2（3）/ 9+ ？Gyrogyne W． T． Wang，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． Mller 1 / 1 1
3． 2． 02． Subtribe Corallodiscinae A． Weber ＆ Mich． Mller 1 / 3－5 Corallodiscus Batalin
3． 2． 03． Subtribe Tetraphyllinae A． Weber ＆ Mich． Mller 1 / 3
3． 2． 04． Subtribe Leptoboeinae C． B． Clarke 6/ 43 Beccarinda Kuntze，Boeica C． B． Clarke，Leptoboea
Benth． ，
Platystemma Wall． ，Ｒhynchotechum Blume
3． 2． 05． Subtribe Ｒamondinae DC． ex Meisn． 3（2）/ 5
3． 2． 06． Subtribe Litostigminae A． Weber ＆ Mich． Mller 1 / 2 Litostigma G． Wei，F． Wen ＆ Mich． Mller
3． 2． 07． Subtribe Streptocarpinae Ivanina 1 / 177
3． 2． 08． Subtribe Didissandrinae A． Weber ＆ Mich． Mller 2 / 10
3． 2． 09． Subtribe Loxocarpinae A． DC． 14 / 202+ Damrongia Kerr，Dorcoceras Bunge，Middletonia C．
Puglisi，Ornithoboea Parish ex C． B． Clarke，Para-
boea （C． B． Clarke ）Ｒidl． ，Ｒhabdothamnopsis
Hemsl．
3． 2． 10． Subtribe Didymocarpinae G． Don 32 1660－ Aeschynanthus Jack，Allocheilos W． T． Wang，Allostig-
ma
W． T． Wang，Anna
1830 Pellegr． ，Briggsiopsis K． Y． Pan，Cathayanthe Chun，
Conandron Sieb． ＆ Zucc． ，Cyrtandra J． Ｒ． Forst． ＆ G．
Forst． ，Didymocarpus Wall． ，Didymostigma W． T．
Wang，Glabrella Mich． Mller ＆ W． H． Chen，Gyro-
cheilos W． T． Wang，Hemiboea C． B． Clarke，Hencke-
lia Spreng． ，Loxostigma C． B． Clarke，Lysionotus D．
Don，Metapetrocosmea W． T． Wang，Microchirita （C．
B． Clarke）Yin Z． Wang，Oreocharis Benth． ，Petroc-
odon Hance，Petrocosmea Oliv． ，Primulina Hance，
Pseudochirita W． T． Wang，Ｒaphiocarpus Chun
Note：Bold face． Taxon represented in China；Underlined． Endemic to China；Spaced print． New genus for China； ？ represents it maybe extinct．
ae． Based on the aberrant morphology，Wang et al
（1990）established a separate tribe in subfam． Cyrtan-
droideae （now Didymocarpoideae）for its accomodation，
while Weber （2004）proposed to exclude it from Gesne-
riaceae． The molecular study of Wang et al （2004a）
confirmed its position within Gesneriaceae，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） ，therefore，included Titanotrichum in sub-
fam． Gesnerioideae as a monospecific tribe． Apart from
the geographical distribution （SE mainland China，Tai-
wan region of China，S Japan）which is difficult to ex-
plain for a member of subfam． Gesnerioideae，this taxo-
nomic placement is in line with the isocotylous seedlings
（the interpretation of Wang et al，2002 as anisocotylous
is erroneous）． Whether the distribution of Titanotrichum
74
1期MLLEＲ 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

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
seedlings it is similar to both groups，but other charac-
ters，including the curious propagules，indicate an iso-
lated position．
Table 2 List of present and past genera of Chinese Gesneriaceae．
Genus
（alphabet． ）
（Chinese name）
Infrafamilial
position：
Subfamily
Tribe Subtribe
Distribution Species
number
Species no．
in China
（endemic）Tax． status Ｒeference Ｒemark
Aeschynanthus Jack
芒毛苣苔属 Didymocarpoideae
Trichosporeae
Didymocarpinae
From S China，N ＆ S
India throughout
Malesia to New
Guinea and the
Solomon Islands
≈185 ≈34
（≈13）Emended by inclusion
of Micraeschynanthus
Middleton，
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 （Guizhou，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 Mller et al，
2011b；Chen
et al，2014b
See text under
Oreocharis
Anna Pellegr．
大苞苣苔属 Didymocarpoideae
Trichosporeae
Didymocarpinae
China，N Vietnam 4 4
（3）No change
Beccarinda Kuntze
横蒴苣苔属 Didymocarpoideae
Trichosporeae
Leptoboeinae
NE India，Burma，S
China，Vietnam，Su-
matra
≈7 5
（4）No change
Boea Comm． ex
Lam．
旋蒴苣苔属
10 － Ｒedefined；Chinese
spp． now in Dorcoceras
and Damrongia
Puglisi et al，
in press
See text under
Boea
Boeica C． B．
Clarke
短筒苣苔属
Didymocarpoideae
Trichosporeae
Leptoboeinae
Bhutan，S China，N
＆ NE India，Myan-
mar，N Vietnam，NW
Malaya
≈12 7
（3）No change
Bournea Oliv．
四数苣苔属 － － Sunk into Oreocharis Mller et al，
2011b
See text under
Oreocharis
Briggsia Craib
粗筒苣苔属 － － Partly sunk into Oreo-
charis，partly transferred
to Loxostigma，and two
spp． forming the new
genus Glabrella
Mller et al，
2011b；Chen
et al，2014b；
Mller et al，
2014
See text under
Briggsia and
Oreocharis
Briggsiopsis
K． Y． Pan
筒花苣苔属
Didymocarpoideae
Trichosporeae
Didymocarpinae
S China （C ＆ S Si-
chuan，NE Yunnan，
Guizhou）
1 1
（1）No change
Calcareoboea C． Y．
Wu ex H． W． Li
朱红苣苔属
－ － Sunk into Petrocodon Wang et al，
2011；Weber
et al，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 al，
2011；Weber
et al，2011a
See text under
Chirita
Chiritopsis W． T．
Wang
小花苣苔属
－ － Sunk into Primulina Wang et al，
2011；Weber
et al，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 China，Taiwan re-
gion of China，S Ja-
pan
1 1 No change
Corallodiscus
Batalin
珊瑚苣苔属
Didymocarpoideae
Trichosporeae
Corallodisceae
Bhutan，China，N ＆
NE India，Nepal，
Thailand
3－5 3 No change
84 广 西 植 物 36 卷

续表2
Genus
（alphabet． ）
（Chinese name）
Infrafamilial
position：
Subfamily
Tribe Subtribe
Distribution Species
number
Species no．
in China
（endemic）Tax． status Ｒeference Ｒemark
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
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 Chirita；
inclusion of Boea clar-
keana Hemsl． and the
Asian species descri-
bed in Streptocarpus
Weber et al，
2011a ；
Puglisi et al，
in press
See text under
Chirita and
Damrongia
Dayaoshania W． T．
Wang
瑶山苣苔属
－ － Sunk into Oreocharis Mller et al，
2011b
See text under
Oreocharis
Deinocheilos W． T．
Wang
全唇苣苔属
－ － Sunk into Oreocharis Mller et al，
2011b
See text under
Oreocharis
Didymocarpus
Wall．
长蒴苣苔属
Didymocarpoideae
Trichosporeae
Didymocarpinae
from N and NE Indi-
a，Nepal and S Chi-
na southwards to the
Malay Peninsula and
N Sumatra
＞70 ≈30
（≈24）Some spp． transferred
to Petrocodon
Weber et al，
2011b
See text under
Petrocodon
Didymostigma
W． T． Wang
双片苣苔属
Didymocarpoideae
Trichosporeae
Didymocarpinae
SE China （Guang-
dong，Fujian，
Guangxi）
3 3
（3）No change
Dolicholoma
D． Fang ＆ W． T．
Wang
长檐苣苔属
－ － Sunk into Petrocodon Weber et al，
2011b
See text under
Petrocodon
Dorcoceras Bunge
羚角苣苔属（新拟）Didymocarpoideae
Trichosporeae
Loxocarpinae
China，Thailand，
Cambodia，Vietnam，
Philippines and
Indonesia
4 2
（1 or 2？） Ｒe-established for par-
ticular （non-Australa-
sian）species of Boea
Puglisi et al，
in press
See text under
Boea and Dorco-
ceras
Epithema Blume
盾座苣苔属 Didymocarpoideae
Epithemateae
Epitheminae
S China，Taiwan re-
gion of China
20 2 No change at genus
level
revised by
Bransgrove ＆
Middleton，
2015
Glabrella Mich．
Mller ＆ W． H．
Chen
光叶苣苔属
Didymocarpoideae
Trichosporeae
Didymocarpinae
S China，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
Mller et al，
2014；Wen et
al，2015 a，b
See text under
Briggsia and
Oreocharis
Gyrocheilos
W． T． Wang
圆唇苣苔属
Didymocarpoideae
Trichosporeae
Didymocarpinae
S China （Guangxi，
Guangdong，SE
Guizhou） ，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 China，Tai-
wan region of China，
N Vietnam，S Japan
31 31
（30）Inclusion of
Metabriggsia （2
spp． ）
Weber et al，
2011c
Inclusion does
not affect the
traditional con-
cept of Hemiboea
Hemiboeopsis W． T．
Wang
－ － Sunk into Henckelia Weber et al，
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．
Microchirita and Prim-
ulina）and Hemiboep-
sis，and to exclude
Codonoboaea
Weber et al，
2011a；Mid-
dleton et al，
2013
See text under
Chirita and
Henckelia
Isometrum Craib
金盏苣苔属 － － Sunk into Oreocharis Mller et al，
2011b
See text under
Oreocharis
94
1期MLLEＲ 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 Species
number
Species no．
in China
（endemic）Tax． status Ｒeference Ｒemark
Lagarosolen W． T．
Wang
细筒苣苔属
－ － Sunk into Petrocodon Weber et al，
2011b
See text under
Petrocodon
Leptoboea Benth．
细蒴苣苔属 Didymocarpoideae
Trichosporeae
Leptoboeinae
Bhutan，N and NE
India，S China
（Yunnan ） ， Myan-
mar，Thailand
2－3 1 No change
Litostigma Y． G．
Wei，F． Wen ＆
Mich． Mller
凹柱苣苔属
Didymocarpoideae
Trichosporeae
Litostigminae
China （Guizhou，
Yunnan）2 2
（2）Genus recently
established
Wei et al，
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 （Sichuan，
Yunnan，Guizhou，
Guangxi） ，N
Vietnam
11 11
（8）Ｒecently inclusion of
caulescent Briggsia
species
Mller et al，
2014
See text under
Briggsia
Lysionotus D． Don
吊石苣苔属 Didymocarpoideae
Trichosporeae
Didymocarpinae
From N India and
Nepal eastwards
through N Thailand，
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 al，
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-
layas，through conti-
nental SE Asia to
Sumatra，Borneo and
Java
≈18 2
（1）Ｒaised from Chirita
sect． Microchirita
to generic rank
Wang et al，
2011；Weber
et al，2011a
See text under
Chirita
Middletonia
C． Puglisi
米氏苣苔属
Didymocarpoideae
Trichosporeae
Loxocarpinae
India，Bangladesh，
Bhutan，China，
Burma，Thailand，
Laos，Cambodia，
Vietnam，Malaysia
4 1 Genus recently estab-
lished for four species
of Paraboea
Puglisi et al，
in press
See text under
Middletonia and
Paraboea
Opithandra B． L．
Burtt
后蕊苣苔属
－ － Sunk into Oreocharis Mller et al，
2011b
See text under
Oreocharis
Oreocharis Benth．
马铃苣苔属 Didymocarpoideae
Trichosporeae
Didymocarpinae
China，Thailand，
Vietnam，Myanmar，
Bhutan，NE India，
Japan
＞105 ＞102
（＞98）Expanded to include
Ancylostemon Craib，
Bournea Oliv． ，Brigg-
sia Craib p． p． － incl．
type，Dayaoshania
W． T． Wang，Deino-
cheilos W． T． Wang，
Isome
trum Craib，Opithandra
B． L． Burtt，Parai-
sometrum W． T．
Wang，Thamnocharis
W． T．
Wang，and Tremacron
Craib；Inclusion of
further ten spp． of
Briggsia
Mller et al，
2011b；
Mller et al，
2014；Chen
et al，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
Vietbam southwards
to N Penins． Malaysia
16 5
（1）No change Ｒevised by
Scott ＆ Mid-
dleton，2014
Paraboea （C． B．
Clarke）Ｒidl．
蛛毛苣苔属
Didymocarpoideae
Trichosporeae
Loxocarpinae
Bhutan，China，
Indonesia，Malaysia，
Myanmar，Philippines，
Thailand，Vietnam
132 26
（14）Expanded by inclusion
of Phylloboea and
Trisepalum；removal of
four species and place-
ment in the new
genus Middletonia
Puglisi et al，
2011
Puglisi et al，
in press
See text under
Paraboea and
Middletonia
05 广 西 植 物 36 卷

续表2
Genus
（alphabet． ）
（Chinese name）
Infrafamilial
position：
Subfamily
Tribe Subtribe
Distribution Species
number
Species no．
in China
（endemic）Tax． status Ｒeference Ｒemark
Paraisometrum W．
T． Wang
弥勒苣苔属
－ － Sunk into Oreocharis Mller et al，
2011b
See text under
Oreocharis
Paralagarosolen Y．
G． Wei
方鼎苣苔属
－ － Sunk into Petrocodon Wang et al，
2011；Weber
et al，2011b
See text under
Petrocodon
Petrocodon Hance
石山苣苔属 Didymocarpoideae
Trichosporeae
Didymocarpinae
China，N Vietnam，
NE Thailand
29 28
（27）Expanded to include
Calcareoboea C． Y． Wu
ex H． W． Li，Didymo-
carpus Wall． p． p． -ex-
cl． type，Dolicholoma
D． Fang ＆ W． T．
Wang，Lagarosolen
W． T．
Wang，Paralagaroso-
len Y． G． Wei，Tengia
Chun and Wentsaiboea
D． Fang ＆ D． H． Qin，
p． p． （excl． type）
Wang et al，
2011；Weber
et al，2011b
See text under
Petrocodon
Petrocosmea Oliv．
石蝴蝶属 Didymocarpoideae
Trichosporeae
Didymocarpinae
NE India，S China，
Myanmar，Thailand，
S Vietnam．
42 34
（34）No change
Platystemma Wall．
堇叶苣苔属 Didymocarpoideae
Trichosporeae
Leptoboeinae
Nepal，Bhutan，N
India，SW China
1 1 No change
Primulina Hance
报春苣苔属 Didymocarpoideae
Trichosporeae
Didymocarpinae
Essentially southern
half of China and Vi-
etnam
＞168 ＞154
（＞151）Enormous expansion of
the previously mono-
typic genus by inclu-
sion of Chirita sect．
Gibbosaccus，Chiritop-
sis，Deltocheilos，and
Wentsaiboea p． p． -
incl．
type
Wang et al，
2011；Weber
et al，2011a
See text under
Chirita
Pseudochirita W．
T． Wang
异裂苣苔属
Didymocarpoideae
Trichosporeae
Didymocarpinae
S China （C ＆ W
Guangxi） ，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 2004，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
Guinea，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 India，Nepal，
Bhutan，SW ＆ S
China，SE Asia and
Malesia to New
Guinea
21 6
（2）No change
Stauranthera
Benth．
十字苣苔属
Didymocarpoideae
Epithemateae
Loxoniinae
Malesia，S China ≈5 1 No change Ｒevision in
prep． （Weber，
in
prep． ）
Streptocarpus，Asiat-
ic species
扭果花 属/海 角苣
苔属
－ － Sunk into Damrongia Puglisi et al，
in press
See text under
Damrongia
Tengia Chun
世纬苣苔属 － － Sunk into Petrocodon Wang et al，
2011；Weber
et al，2011b
See text under
Petrocodon
Thamnocharis W． T．
Wang
辐花苣苔属
－ － Sunk into Oreocharis Mller et al，
2011b
See text under
Oreocharis
15
1期MLLEＲ 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 Species
number
Species no．
in China
（endemic）Tax． status Ｒeference Ｒemark
Titanotrichum
Soler．
台闽苣 苔属 /俄氏
草属
Gesnerioideae
Titanotricheae
SE China，Taiwan
region of China，S
Japan
1 1 Placed in subfam．
Gesnerioideae
C． N． Wang
et al，2004a；
Perret et al，
2013；Weber
et al，2013
Tremacron Craib
短檐苣苔属 － － Sunk into Oreocharis Mller et al，
2011b
See text under
Oreocharis
Trisepalum C． B．
Clarke
唇萼苣苔属
－ － Sunk into Paraboea Puglisi et al，
2011
See text under
Paraboea
Wentsaiboea
D． Fang ＆ D． H． Qin
文采苣苔属
－ － Partly （incl． type ）
sunk into Primulina，
partly into Petrocodon
Weber et al，
2011a，b
See text under
Chirita and Pet-
rocodon
Whytockia
W． W． Sm．
异叶苣苔属
Didymocarpoideae
Epithemateae
Monophyllaeinae
S China，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 1990，Wang et al，1998 ）are given
in square brackets and not bold． Infrafamilial position according to Weber et al，（ 2013）．
With regard to subfam． Didymocarpoideae，both
tribes，Epithemateae and Trichosporeae，are represen-
ted in China．
In tribe Epithemateae，which is apparently a rel-
ict tribe with morphologically rather odd and isolated
genera，all four subtribes are represented in China．
From the seven genera two are missing in Cina：Mono-
phyllaea in subtribe Monophyllaeinae （this thus con-
tains only Whytockia，with some six species in South
China and Taiwan region of China） ，and Loxonia in
Loxoniinae （containing in China Stauranthera and-with
some uncertainty the ill-known and possibly extinct Gyr-
ogyne）． In both cases the genera are distributed in the
SE Asian tropics：Monophyllaea includes more than 40
species，with distribution throughout Malesia （Sumatra
to New Guinea and S Thailand to Java） ，Loxonia com-
prises three species restricted to western Malesia （Su-
matra，Malay Peninsula and Borneo）．Whytockia is par-
ticularly remarkable as morphological studies （Weber，
1976，1982）indicate that the genus has retained primi-
tive characters in relation to Monophyllaea，suggesting
that subtribe Monophyllaeinae has originated in the ex-
tratropics and，after reaching the South East Asian trop-
ics，radiated into the many species of Monophyllaea．
Stauranthera and Gyrogyne are represented with a single
species each in China，Ｒhynchoglossum and Epithema
with two species each．
Tribe Trichosporeae，with 10 subtribes tentatively
recognised by Weber et al （2013） ，is represented by
five subtribes in China．
Among the more primitive alliances，subtribe Cor-
allodiscinae，with the single genus Corallodiscus，is re-
markable． Its few species are rosette plants with scapose
inflorescences and tetrastaminate flowers．
Morphologically heterogeneous is subtribe Lepto-
boeinae，which is represented by almost all of its genera
in China （the only lacking genus is the South Indian
Championia，the inclusion of which in Leptoboeinae
still requires confirmation）． Leptoboeinae includes ro-
sette plants （Beccarinda ） ， sub-shrubby caulescent
plants （Leptoboea，Boeica，Ｒhynchotechum）and unifo-
liate plants （Platystemma ）．Leptoboea，Boeica and
Ｒhynchotechum have （correctly！） thought to be closely
related in older classifications，but Burtt （1963）as-
sumed a close relationship of Ｒhynchotechum with Cyr-
tandra（“subfam． Cyrtandroideae tribe Cyrtandreae”） ，
because the two genera have indehiscent，berry-like
fruits in common． Apart from the changes in taxonomic
position as compared to former classifications，no chan-
ges 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 2010，
based on newly collected material from Guizhou and
Yunnan． The genus represents an important morphologi-
cal link between the basal lineages of tribe Trichospore-
25 广 西 植 物 36 卷

ae with seeds without elaborate testa cell ornamentation，
straight fruit and septicidal and loculicidal dehiscence
and derived lineages with predominantly loculicidal de-
hiscence and two stamens （Wei et al，2010）．
Subtribe Loxocarpinae is the second largest sub-
tribe of Trichosporeae． In total it includes about a dozen
genera． Two of them，Dorcoceras and Middletonia，have
been （re-）established very recently and both are repre-
sented in China，together with Damrongia，Ornitho-
boea，Paraboea and Ｒhabdothamnopsis． The latter is
with a single species endemic to China，while the re-
maining genera have their main distribution mostly fur-
ther 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 molecu-
lar-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 gen-
era，with 24 represented in China． Six genera with one
or very few species （i． e． Allocheilos，Allostigma，
Briggsiopsis，Cathayanthe，Didymostigma，Metapetro-
cosmea）are endemic to this country，others have their
main distribution elsewhere and reach with one or few
species the southernmost part of China （e． g． ，Cyrtan-
dra，with＞ 800 species is the largest genus of Gesneri-
aceae，reaches with a single species to South Taiwan，
China）．Microchirita with a wide distribution from the
Western Ghats of India，the foothills of the Himalayas，
through continental SE Asia into Sumatra，Borneo and
Java，has two species in China （out of c． 18）．
The subtribes of Trichosporeae not represented in
China are Jerdoniinae （only Jerdonia indica，S India） ，
Tetraphyllinae （only Tetraphyllum，3 spp． ，NE India，
Bangladesh，Burma，Thailand） ，Ｒamondinae （Haber-
lea，Ｒamonda，Jancaea；5 spp． ，SW and SE Europe） ，
Streptocarpinae （traditionally 9 genera，recently all amal-
gamated in Streptocarpus，Nishii et al，2015；177 spp． ；
Africa，Comoro Islands and Madagascar） ，and Didissan-
drinae （Didissandra，Tribounia；c． 10 spp． ，W Malesia，
Thailand）． Altogether，these subtribes include only one
or few genera；as to species number，Streptocarpinae is
clearly the largest subtribe． The link from the African to
the Asiatic （including Chinese）species via the “Asiatic
species of Streptocarpus”proved untenable：the Asiatic
species described in Streptocarpus belong to the purely A-
sian genus Damrongia （Puglisi et al，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
listed and briefly commented on． In some genera （that
is，for instance，all genera of tribe Epithemateae）no
changes in the taxonomic delineation have occurred，in
others the inclusion of formerly distinct genera has little
bearing on their new definition （e． g． the inclusion of
Metabriggsia into Hemiboea，in which differences in
the ovary structure simply proved erroneous，Weber et
al，2011c） ，but others have a rather complex tax-
onomic history which require a more detailed explana-
tion． In the following the fate of the latter genera，plus
new or newly established genera is briefly outlined，as far
as changes since the treatment of Wang et al （1990）and
Wang et al （1998）are concerned． De facto，all changes
occurred from 2010 onwards，with the new genus Litostig-
ma （Wei et al，2010）as the first．
Boea． This was previously a widespread Asiatic-
Malesian genus，with three species （B． clarkeana，B．
hygrometrica，and B． philippensis ）represented in
southern China． From the recent molecular-systematic
studies of Puglisi et al，（ in press） ，Boea emerged as
an essentially Australasian genus，with distribution in
Eastern Indonesia，Papua New Guinea，the Solomon
Islands and Queensland （Australia ）． The genus
Dorcoceras was re-established to accommodate the re-
maining species distributed in Burma （1 sp． ） ，Cam-
bodia （1 sp． ）and China （2 spp． ）． In addition，Boea
hygrometrica （endemic to China）and B． philippensis
（Philippines and Vietnam）were included in Dorco-
ceras （the former representing the type species of
Dorcoceras）．Boea clarkeana （≡Streptocarpus clar-
keanus） ，however，was attributed to the genus Dam-
rongia （see there）．
Briggsia． The genus is no longer relevant，as the
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1期MLLEＲ 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

alliance around the type species （B． longifolia）has
been synonymised with Oreocharis （Mller et al，
2011b，2014）． In its traditional concept，Briggsia in-
cluded acaulescent rosette plants as well as caulescent
plants，both with characteristic large and ventrally
pouched（“briggsioid”）flowers． Its three acaulescent-
and strikingly glabrous-species are now in the new ge-
nus Glabrella and the three caulescent species are in
Loxostigma （Mller et al，2014；Wen et al，2015a，
b）． The “briggsoid”flowers arose apparently inde-
pendently in different alliances （Mller et al，2011b）．
See also notes for Oreocharis．
Chirita． Dramatic changes occurred in this tradi-
tional and large genus （with up to 140 species de-
scribed herein，placed in four sections：sect． Chirita，
sect． Liebigia，sect． Microchirita，and sect． Gibbosac-
cus；Wood，1974；Hilliard，2003 ） ，ending in a syn-
onymisation of the genus with Henckelia （Weber et al，
2011a）． The character defining Chirita was traditional-
ly seen in the “chiritoid”stigma：a stigma with upper
and lower lobe，but the upper largely or completely re-
duced，and the lower one expandend and often bipar-
tite． The molecular studies clearly revealed that this
character is homoplastic，having evolved independently
in several alliances of tribe Trichosporeae． As a conse-
quence，Chirita was split into five genera，with the
majority of Chirita sect． Chirita （and the monotypic
Hemiboeopsis，China and Laos ）amalgamated with
Henckelia sect． Henckelia （from South India）． The re-
maining species of Chirita sect． Chirita were included
in the revived genus Damrongia （see below）．Chirita
sect． Liebigia was raised to generic level （Liebigia
Endl． ，not represented in China ）．Chirita sect． Mi-
crochirita was also raised to genus level （Microchirita，
with two species in China） ，and Chirita sect． Gib-
bosaccus was，together with Chiritopsis and Wentsaiboea
（including its type species and a later published species，
W． luochengensis，Liu et al，2010；Xu et al，2012） ，in-
cluded） ，included in the originally monotypic genus
Primulina （Wang et al，2011；Weber et al，2011a；Li ＆
Xia 2012）． The latter genus is essentially geographically
restricted to China，now with ＞150 species，with some
species extending or endemic to Vietnam．
The newly defined Henckelia has a broad circum-
scription at present，and is currently under further stud-
ies． Primulina has been expanded from a monotypic ge-
nus to one with 100 species at the time of its redefinition
（Weber et al，2011a）and over the last few years has be-
come a genus with over 150 species． The morphological
variation is relatively limited compared to other genera，
and is based on diandrous zygomorphic flowers usually in-
fundibuliform but variable in size and coloration，and a
vegetative habit with rhizomatous compacted stems with
leaves in a basal rosette，with decussate phyllotaxy or in
whorls of three （rarely alternate）． This character distin-
guishes this genus to a great length from Petrocodon，
where the leaves are always alternate． Another possibly
distinguishing feature are the chromosome numbers which
are almost uniformly 2n = 36 in Primulina （＞100 species
counted，with one exception of a tetraploid number，
Christie et al，2012） ，whereas it is 2n = 20 in the only
member of Petrocodon so far counted （P． hancei，Cao et
al，2003；Mller ＆ Pullan，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 inflorescences，infun-
dibuliform corollas with 2 stamens，and a pistil with a
“chiritoid”stigma （see Chirita）． It was thought to be
restricted to Thailand and the NW of Peninsular Malay-
sia，but the Chinese Boea clarkeana also proved to be-
long to that genus （Puglisi et al，in press）． The same
applies for the Asian species described in Streptocarpus
on grounds of the twisted fruits （Puglisi et al，in press）．
Dorcoceras． Ｒe-established genus segregated from
Boea by Puglisi et al （in press）． See Boea above．
Glabrella． New genus established by Mller et al
（2014）for two species of Briggsia，which do not fit in-
to Oreocharis or Loxostigma． A third species was added
by Wen et al （2015a，b）． See Briggsia and Oreocharis．
Henckelia． The genus Henckelia Spreng． was re-
established when the unwieldy and broadly circum-
scribed Didymocarpus was changed and split into 3
smaller entities：Didymocarpus s． str． ，Henckelia （with
five sections：sect． Henckelia，S India and Sri Lanka，
sect． Loxocarpus，sect． Heteroboea，sect． Loxocarpus，
45 广 西 植 物 36 卷

sect． Didymanthus and sect． Glossadenia）and Hovanel-
la，Weber ＆ Burtt 1998）． The molecular data of Mller
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．
sect． Henckelia，S India and Sri Lanka）was amalgamt-
ed with most of Chirita sect． Chirita and the monotypic
Hemiboepsis （see under Chirita above） ，sect． Loxo-
carpus has returned to generic level （Weber et al，
2011a；Middleton et al，2013；Yao，2012） ，and the
last three sections have been included in the re-estab-
lished and largely expanded genus Codonoboea （Kiew ＆
Lim 2011；Middleton et al，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 al，2011；Weber et al，2011a）． It can be character-
ized by inflorescences that usually appear in two or sev-
eral in a leaf axil，often displaced onto the petiole，of-
ten consisting of a short-stalked serial flower pair only，
but this repeated several times． From the c． 25 species
2 occur in China：M． hamosa and M． prostrata （re-
cently described by Li ＆ Xia，2012）．
Middletonia． A new genus established by Puglisi
et al，（ in press）for the accommodation of four species
separated from Paraboea，with one species distributed
in China （M． multiflora）．
Oreocharis．While Oreocharis included some 27
species in its traditional delineation （Wang et al，
1998） ，the number increased enormously through the
inclusion of Ancylostemon，Bournea，Briggsia p． p．
（incl． type） ，Dayaoshania，Deinocheilos，Isometrum，
Paraisometrum，Thamnocharis，and Tremacron （Mller
et al，2011b；Middleton et al，2013）． As the species of
Briggsia transferred to Oreocharis included the type spe-
cies （B． longifolia） ，the remaining species were left
without generic placement． In recent studies （Chen et
al，2014b；Mller et al，2014；Wen et al，2015a，b） ，
the rest were attributed to the following three genera：
the caulescent species were referred or returned to Lox-
ostigma，three acaulescent species with glabrous leaves
and stems were moved into a new genus Glabrella Mich．
Mller ＆ W． H． Chen，and 11 acaulescent species were
transferred to Oreocharis． New species were recently de-
scribed （Liu et al，2012；Chen et al 2013，2015；Tan
et al 2013，2015；Ｒossini and Freitas 2014；Li and Li
2015，Yang et al，2015） ，and Oreocharis now compri-
ses now 106 species，all representing acaulescent rosette
plants with scapose inflorescences，but with considera-
ble variability in the symmetry，shape and coloration of
the flowers． Oreocharis leiophylla and O． sinensis （mak-
ing up the former genus Bournea）have actinomorphic
flowers． The corolla shape in Oreocharis varies from tu-
bular，through funnel-shaped，campanulate to
“briggsioid”，the stamen number is （1） ，2，4 or （in
actinomorphic flowers）4 or 5 in line with the number of
petals，in some diandrous species （previously constitu-
ting the genus Opithandra）the posterior stamens are the
fertile ones （while it is the reverse in most other dian-
drous Gesneriaceae） ，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 genera，
but，as the molecular data show，the species relation-
ships run across genera and these are thus untenable．
Like in Petrocodon，the flowers combine a range of co-
rolla shapes and colours suggesting different pollinator
adaptations sometimes in parallel．
Paraboea． This widespread and species-rich genus
was revised by Xu et al （2008） ，who recognised 89
species，c． 20 of them found in China． Changes since
then relate to the inclusion of the monotypic Phylloboea
（Myanmar）and Trisepalum （c． 13 spp． ，1 in China）
（Puglisi et al，2011）． More recently，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
China，of these 14 exclusively there （i． e． endemic）．
Petrocodon．In its traditional concept，Petrocodon
was a genus with two Chinese species，both with small，
white flowers and two fertile stamens． Based on molecu-
lar data，Wang et al （2011）added Tengia （with small
white，pentamerous actinomorphic flowers and five sta-
mens） ，Calcareoboea （with large，red and tubular flow-
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1期MLLEＲ 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

ers） ，and Paralagarosolen （with long-tubed hypocrater-
iform flowers）． Shortly later，Weber et al （2011b ）
added the monotypic Dolicholoma （with flowers with
narrow tube and subactinomorphic limb with long，acute
lobes） ，all species of Lagarosolen （with flowers similar
to those of Dolicholoma and Paralagarosolen） ，one spe-
cies of Wentsaiboea （without type）and three species of
Didymocarpus （D． hancei，D． mollifolius，D． niveola-
nosus）． This resulted in a genus size of around 20 spe-
cies． The recent description of new species by Wen et al
（2012） ，Chen et al（2014a） ，Hong et al （2014） ，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 adaptations，including bee，butterfly and
bird pollination． Until recently，Petrocodon could be
defined by possessing 2 stamens or 5 （in the former ge-
nus Tengia with actinomorphic flower）． Using both mor-
phological and molecular data，a species possessing 4
stamens was described：P． hunanensis （Yu et al，
2015）． This indicates （a）the great diversity that is
still unexplored in China，and （b）that the use of mo-
lecular data has apparently given the right signal to
place the species in the right genus．
4 Discussion and Outlook
4． 1 You win some，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） ，and
two genera established a few years later （Paralagaroso-
len：Wei，2004；Wentsaiboea：Fang ＆ Qin，2004） ，the
number has dropped to 45 （Table 1）． Since 2010，and
essentially based on molecular data，seven new genera
have been added to China，either based on newly col-
lected material （Litostigma） ，or from segregation of ex-
isting genera in China （Glabrella，Damrongia，Dorco-
ceras，Middletonia，Microchirita，and Primulina；see
previous chapter） ，or by a new circumscription of gene-
ra （Henckelia）．
On the other hand，a considerable number of gene-
ra （22）are lost by synonymisation with other genera
（Ancylostemon，Boea，Bournea，Briggsia，Calcareo-
boea，Chirita，Chiritopsis，Dayaoshania，Deinocheilos，
Dolicholoma，Hemiboeopsis，Isometrum，Lagarosolen，
Metabriggsia，Opithandra，Paraisometrum，Paralagaro-
solen，Tengia，Thamnocharis，Tremacron，Trisepalum，
Wentsaiboea）． Also，the four Asiatic species of Strepto-
carpus （Hilliard and Burtt 1971） ，one of them，S．clar-
keanus （Hemsl． ）Hilliard and B． L． Burtt （≡Boea clar-
keana Hemsl． ，as such in the Flora of China of Wang et
al，1990；Wang et al，1998） ，from China，have found
a proper generic place （Damrongia ）．Streptocarpus，
therefore，can be definitely removed from the list of
genera extending into China．
By tribe，there is no change in generic state in Ep-
ithemateae，as to Trichosporeae-Litostigminae there is
the addition of one genus （Litostigma） ，in Trichospore-
ae-Loxocarpinae there are two fewer genera in China
（Boea，Trisepalum） ，but three additional ones （Dam-
rongia，Dorcoceras，Middletonia） ，an overall net gain
of one． The greatest changes occurred in Trichosporeae-
Didymocarpinae，with a loss of 20 genera，and a gain of
three （Glabrella，Henckelia，Microchirita） ，thus an o-
verall loss of 17 genera． This tribe is still not fully un-
derstood and further work is necessary to understand its
evolution，and further changes may be in the waiting，
though these may be less drastic，but fine-tuning．
The number of endemic genera has decreased from
27 （Wang et al，1998；Wei 2004；Fang ＆ Qin，2004）
to 11． Ten of them include only one to three species
（Allocheilos ［2］，Allostigma ［1］，Briggsiopsis ［1］，
Cathayanthe ［1］，Didymostigma ［3］，Glabrella ［3］，
Gyrogyne ［1］，Litostigma ［2］，Metapetrocosmea ［1］，
Ｒhabdothamnopsis ［1］） ，a drop from 22 previously．
The remaining almost a dozen small genera have often
unusual morphologies combined with isolated evolution-
ary positions．
Over the decades，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 characteristic，into larger
ones on grounds of more shared than distinguishing
characteristics． This is important，since single character
taxonomy is fraught with problems when picking the
65 广 西 植 物 36 卷

‘wrong’character a genus was not established for．
4． 2 Morphological parallelisms and reversals
Burtt （e． g． Burtt，1963 ）pointed out that in
Gesneriaceae there are many exceptions and apparent
parallelisms in form，and that the homology of similar
characters on which a genus is based，is sometimes dif-
ficult to ascertain （e． g． Burtt，1968）． The erstwhile
genus Chirita is a prime example，in which strong em-
phasis was given to the special type of stigma （“chiri-
toid stigma”）． The molecular data have unambiguously
shown that the chiriotoid stigma has evolved several
times independently，and Chirita thus had to be split
into five genera （Weber et al，2011a） （ see also be-
low）．
At the generic level，the number of fertile stamens
（4 or 2）played a significant role，especially in Chinese
Gesneriaceae． For example，Deinocheilos （with 2 sta-
mens）has been kept separate from Tremacron （4 sta-
mens） ，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 /94，1908）established tribe Ｒamondeae
to include Corallodiscus，Haberlea，Petrocosmea，
Ｒamonda，and Saintpaulia． His intention was to recog-
nise a group with similar vegetative habit （flat rosette
plants）with septicidal capsule dehiscence，rather than
floral shape． Thus，this tribe includes four genera with
zygomorphic flowers，two of which have long floral tubes
and four stamens （Haberlea and Corallodiscus） ，two
with flat-faced （truncate）zygomorphic corollas with 2
stamens （Petrocosmea and Saintpaulia）and one actino-
morphic genus Ｒamondia （=Ｒamonda）． He placed an-
other genus with actinomorphic flowers，Conandron，in
a separate tribe． Burtt （1970）accepted this view and
suggested that actinomorphic genera were not related，
that zygomorphy was ancestral in the family and actino-
morphic genera the result of separate independent losses
of flower asymmetry． Wang et al （1990）and Wang et
al （1992）with the reverse view of an ancestral actino-
morphy and derived zygomorphy redefined tribe Ｒamon-
deae to include only actinomorphic genera，Bournea，
Conandron，Ｒamonda，Tengia，and Thamnocharis．
Molecular phylogenetic work has proven Burtt’s view to
be correct since they showed that all actinomorphic gen-
era have independent origins （Mller et al，1999，
2009，2011a；Wang et al，2010）．
Hidden homoplasies destabilize a classification sys-
tem that wants to reflect phylogeny，and single character
taxonomy may lead to an inflation of the number of small
or monotypic genera． With the rise of molecular phylo-
genetic methodsduring the last 20 years or so，we have
become more aware of the presence and levels of homo-
plasies among morphological characters and how small
and monotypic genera are related or part of larger enti-
ties （Mller et al，2011a）． This has been shown for the
Neotropical Gesneriaceae as well as for the Afro-Mala-
gasi Gesneriaceae of tribe Streptocarpinae，which have
been reduced from nine often small genera （7 with ＜3
species）to one genus Streptocarpus （Nishii et al，
2015）． There are numerous examples of morphological
homoplasies that have resulted in dramatic changes in
classification for the New World Gesneriaceae． The con-
vergence of resupinate flowers was recently discovered to
be independently derived in three lineages that resulted
in a new generic classification for members of the Col-
umneinae （Clark et al，2006；Clark ＆ Zimmer 2003）．
The convergence of hypocyrtoid or “pouched”flowers
within Drymonia and across several lineages of closely
related genera has resulted in a revised circumscription
of genera in the Columneinae （Clark et al，2015；Clark
et al，2012；Smith ＆ Clark 2013． Ｒecent phylogenetic
results that have elucidated the convergence of radially
symmetrical flowers and that has resulted in the circum-
scription of more narrowly defined genera in the Gloxini-
inae （Smith et al，2004；Ｒoalson et al，2005b；Clark
et al，2011）．
4． 3 Holistic approach to the taxonomy
Uncertainties in the correct genus assignment a-
bound． Photographs of Litostigma crystallina were pub-
lished under Petrocosmea crystallina （Shui ＆ Chen
2006）before its accommodation in a new genus （Wei et
al，2010）． The recently described species，Primulina
guangxiensis Yan Liu and W． B． Xu was initially
placed with a superficially similar species （Liu et al，
2011） ，but after molecular studies，partly by the same
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1期MLLEＲ 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

authors，it was discovered that it belonged in Petrocodon
（assuming the integrity of the molecular data）and was
transferred to this genus （Xu et al，2014）．
In some cases incomplete knowledge of generic
concepts might be at issue，in others perhaps the dis-
covery of species with new character combinations，such
as Petrocodon hunanensis with four stamens，cause un-
certainty． However，with the availability of molecular
data and an extensive data set of molecular sequences
on public databases，such as GenBank，these uncer-
tainties can be addressed by molecular approaches．
However，where traditional taxonomic approaches
are concerned，the utilisation and balancing of all char-
acters，not just one or a few，has to be considered in
taxonomic decisions． This might have become more
complex with the new delineations of some genera，and
the‘traditional’characters of corolla shape，stigma or
androecium may be largely inadequate． Additional or
hitherto undervalued characters may be required．
4． 4 Open issues
At the higher taxonomic level we present classifica-
tion on the basis of whatis currently known． The classi-
fication of tribe Trichosporeae，particularly subtribe
Didymocarpinae，is still to be completed． Because of
the large number of representatives in this subtribe resi-
ding in China，it can play a pivotal role in this endeav-
our．
At the generic level，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
（Mller et al，2014）has expanded its definition to in-
clude seeds without appendages，narrowing its gap to
Ｒaphiocarpus，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 al，
2011b）． In the phylogeny of Mller et al （2011a） ，D．
cortusifolius fell as sister to Gyrocheilos，apart from Did-
ymocarpus proper，and may belong in this genus． Li et
al （2015）published a molecular phylogeny on Chinese
species and found similar results to Mller 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 species，and the latter species with straight
and twisted fruits． Such an assemblage seems at first
dissatisfying，but it has precedence in other genera．
The genus Streptocarpus was characterised for species
possessing twisted fruits． Though it includes a wide
range of morphological forms，from unifoliates，plurifoli-
ates，rosulates，rosettes，herbaceous and woody caules-
cent forms （Hilliard ＆ Burtt，1971）． It has now also
been extended to include species with non-twisted fruits
（Nishii et al，2015）．
4． 5 Outlook
The work summarised here is very much a work in
progress and far from completed． The finding of new
genera on new collections （e． g． Litostigma） ，and mor-
photypes that expand the new circumcriptions （e． g．
Petrocodon hunanensis with four stamens） ，are examples
which demonstrate that the full diversity in Gesnericeae
in China are not yet known and it is an exciting prospect
for finding further ‘links’through intensified and sys-
tematic taxon sampling． Also，the great number of new
species described in some genera，especially Primulina，
which ballooned from around 100 when it was redefined
in 2011，to now ＞150 with many more species to be de-
scribed，illustrates the need for continued floristic sur-
veys and detailed fieldwork to fully document the diver-
sity of Gesneriaceae in China． Systematic fieldwork ef-
forts combined with detailed herbarium studies and mo-
lecular work has unearthed several genera over the last
few years from Thailand and Vietnam （e． g． Somrania，
Middleton ＆ Triboun 2012；Tribounia，Middleton et al，
2012；Billolivia，Middleton et al，2014；Chayamaritia
Middleton et al，2015）． Ｒecent discoveries outlined
here clearly demonstrate the recent progress in Gesneri-
aceae diversity of China，but more importantly the fu-
ture need to continue phylogenetic and taxonomic stud-
ies for a long-term stable classification． Without a
doubt，there is still much to be done．
Acknowledgements We thank the Chinese A-
cademy of Sciences Visiting Professorship scheme for
85 广 西 植 物 36 卷

supporting our research． We also thank the Ｒoyal Bo-
tanic Garden Edinburgh，supported by the Ｒural and
Environment Science and Analytical Services division
（ＲESAS）in the Scottish Government．
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1期 吴佩纹等：蓝花丹结实率低的传粉生物学和繁育系统初探