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

  • Guangxi Institute of Botany, CAS
  • Lawrenceville School


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.
Guihaia Jan. 2016361) : 44-60 http/ / journal. gxzw. gxib. cn
http/ /www. guihaia-journal. com
DOI10. 11931 / guihaia. gxzw201512015
MLLER Michael
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2016361) :44-60
MLLER MWEI YGWEN Fet al. You win some you lose someupdated generic delineations and classification of Gesneriaceae-implications for the fam-
ily in ChinaJ. Guihaia2016361) :44-60
You win some you lose someupdated generic
delineations and classification of Gesneriaceae-
implications for the family in China
MLLER Michael1*WEI Yi-Gang2WEN Fang2CLARK John L. 3WEBER Anton4
1. Royal Botanic Garden EdinburghEdinburgh EH3 5LRUK2. Guangxi Key Laboratory of Plant Conservation and Restoration Ecology in Karst
TerrainGuangxi Institute of BotanyGuilin 541006China3. Department of Biological SciencesThe University of AlabamaTuscaloosa
AL 35487USA4. Department of Botany and Biodiversity ResearchUniversity of ViennaRennweg 14A-1030 ViennaAustria
AbstractOver 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 splitmerged or newly defined. Additionallynew genera have been established based on newly collected ma-
terialillustrating on the one hand gaps in field work in Chinaand on the other hand the biological richness of the
Gesneriaceae in China. Herewe summarize and present an overview of our work and the taxonomic consequences.
Key wordsChinaclassificationGesneriaceaemolecular phylogeniessystematics
CLC numberQ949. 4 Document codeAArticle ID1000-3142201601-0044-17
MLLER Michael1*韦毅刚2温 放2CLARK John L. 3WEBER 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
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
Received date2015-12-21 Accepted date2016-01-07
Foundation itemthe Chinese Academy of Sciences Visiting Professorship Scheme 2011T1S20 ) ; the National Natural Science Foundation of China
31000258314703063126003831460159) ; the Key Laboratory of Plant Diversity and Biogeography of East AsiaKunming Institute of Botany KIB
& the Chinese Academy of Sciences 2014CB954100and Guangxi Natural Science Foundation 2015GXNSFBB139004 & 2015GXNSFBA139105) ;
Guangxi Forest Science & Technology Foundation Gui Lin Ke Zi2014No. 27
BiographyMLLER Michael 1960-) ,MaleGermanPh. D. & Professorfocuses on the Old World Gesneriaceaeresearch interests on Phylogenetics & Sys-
tematicsFinescale BiogeographyCharacter Evolution & Evolutionary DevelopmentGenome & Chromosome Evolutionetc. E-mailm. moeller@ rbge. ac. uk.
* Corresponding author
Gesneriaceae are a medium-sized family in the or-
der Lamialeswith 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
Chineseversion of theFlora of ChinaWang et al
1990listed 413 species in 56 generain the later
English version Wang et al1998the number in-
creased to 442 specieswhile the number of genera re-
mained the same354 species and 25 genera many of
them monotypic) ,respectivelywere considered to be
endemic. Without a doubtChina is a significant centre
of diversity of the familywith the majority of taxa found
in Chinas South and Southwestin the provinces of
YunnanGuizhouGuangdong and Guangxi Autono-
mous Region Wang et al1992 The species density
decreases northwards and the furthest locality North are
the Yenshan Mts at 41°N.
In the last two decadesthe 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 Gesneriaceaein 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
Traditionallythe family was subdivided into two
subfamiliesGesnerioideae and Cyrtandroideae e. g.
Bentham1876Fritsch1893 - 94Burtt1963. The
last formal classification based on morphology and
partly cytogenetic datawas that of Burtt and Wiehler
1995) ,who recognized three subfamilies 1Gesne-
rioideae 56 generapredominantly Neotropical 2
Coronantheroideae 9 generasouthern hemisphere
temperate South America to Australia) ; 3Cyrtan-
droideae = Didymocarpoideae82 generachiefly Pal-
eotropicalincluding 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 anatomypost-ger-
mination characteristics of the seedlingsnectary struc-
ture and ovary position. Thusthe Neotropical Gesneri-
aceae were characterized by the presence of seed endo-
spermisocotyly both cotyledons showing equal and
limited growth after germination) ,a nectary often con-
sisting of separate glandsand often partly or fully infe-
rior ovary positionwhile the Palaeotropical Gesneriace-
ae were characterized by the lack of endospermby con-
tinued growth of one cotyledon after germination aniso-
Nishii et al2004Mantegazza et al2007) ,a basical-
ly ring-shaped nectary and superior ovary position. Sub-
fam. Coronantheroideae was characterized by isocotylous
seedlingsa nectary adnateto the ovary baseand
superior ovary position.
Burtt & Wiehler 1995subdivided the three sub-
families into the following tribes number of genera in
brackets) : 1GesnerioideaeGloxinieae 23) ,Epi-
scieae 21) ,Beslerieae 8) ,Napeantheae 1and
Gesnerieae 2) ; 2Coronantheroideaeonly Coronan-
thereae 9) ,3
CyrtandroideaeKlugieae 7) ,Didymocarpeae
64 ) ,Trichosporeae 6) ,Cyrtandreae 3) ,Tit-
anotricheae 1
The first classification taking to some extent molec-
ular data into accountwas that of Weber 2004
Howeverparticulary for the Palaeotropical Gesneriace-
ae the molecular data situation was insufficient and no
formal classification was presented. Weber 2004 ) ,
thereforeproposed a provisional classificationrecog-
nising four informal groups 1Coronantheroid Gesne-
1M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 2Gesnerioid Gesneriaceae 3Epithema-
toid Gesneriaceae 4Didymocarpoid Gesneriaceae.
The latter and the largest group was subdivided into
morpho-geographical assemblages
Basal Asiatic gene-
raEuropean generaAfrican and Madagascan
generaandAdvanced Asiatic and Malesian genera
Especially by the work of the first author MMO
and associatesin 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 parallelalso considerable progress
was made with regard to Neotropical Gesneriaceaein-
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 al2013
By the inclusion of Sanangowhich was repeatedly
shown to be sister to Gesneriaceae in previous molecular
studiesthree subfamilies were recognisedSanan-
goideae only Sanango) ,Gesnerioideae and Didymo-
carpoideae. Wiehlers subfam. Coronantheroideae was
included as a tribe Coronanthereaein Gesnerioideae.
Apart from the traditional Gesnerioideae subdivided in-
to the tribes BeslerieaeNapenatheae and Gesnerieae
a fifthand rather aberranttribe was included in
Gesnerioideaethe monospecific Titanotricheae from E
Asia. This is the only tribe that does not occur in the
Neotropics or the southern hemispherebut with a single
species Titanotrichum oldhamiiin East Asia E Chi-
naTaiwan region of ChinaS Japan The former Cyr-
tandoideaenow referred to as Didymocarpoideaewere
subdivided into two tribesEpithemateae and Tri-
chosporeae. The former is apparently an ancient assem-
blage of few and rather isolated genera that fall into four
subtribeswhile 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
familyanother important aspect of the molecular work
is the improvement in the understanding of generic de-
limitations. In facta wealth of taxonomic changes re-
sulted from the molecular studiesboth with respect to
Neotropical and Palaeotropical taxa. On the one hand
many genera have been reduced to synonymy e. g.
Mller et al2011bWeber et al2011bcNishii et
al2015Puglisi et alin press) ,on the other hand
some genera have been revived e. g. Centrosolenia
Mora & Clark 2016CrantziaClark2005Clark et
al2006DamrongiaWeber et al2011aGlossolo-
maClark et al2006Clark2009LiebigiaWeber
et al2011aMandirolaRoalson et al2005ab
Boggan 2006SeemanniaRoalson et al2005ab
Boggan 2006TrichodrymoniaMora Clark 2016
or newly described e. g BilloliviaMiddleton et al
2014ChautemsiaAraujo et al2010Christopheria
Smith & Clark 2013ChayamaritiaMiddleton et al
2015GlabrellaMller et al2014GloxinellaRoal-
son et al2005abBoggan2006Gloxiniopsis
Roalson et al2005abBoggan2006LesiaSmith
& Clark2013LitostigmaWei et al2010
PachychaulosSmith & Clark2013Pagothyra
Smith & Clark2013SomraniaMiddleton & Triboun
2012SphaerorrhizaRoalson et al2005abTri-
bouniaMiddleton & Mller2012 In many cases
the species content changedpartly dramatically e.
g. Primulinaincrement from 1 to >150 speciesAl-
loplectusreduction from c. 140 to 5 speciesClark et
2 Position of Chinese Generiaceae in
the classification of Weber et al2013
In Table 1 the classification of Weber et al
2013is givenwith 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 al2004b Burtt 19631977 ear-
64 广 西 植 物 36
marked Titanotrichum as a genus anomalumwithin Gesneriace-
Table 1 Classification of Gesneriaceae acc. to Weber et al 2013and taxa represented in China
Rank No. of genera / species no. /genera represented in China
Gesneriaceae Rich. & Juss. in DC.
1. Subfam. Sanangoideae A. WeberJ. 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. WooJ. 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. 23/ 9+ Gyrogyne W. T. WangStauranthera 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 KuntzeBoeica C. B. ClarkeLeptoboea
Platystemma Wall. Rhynchotechum Blume
3. 2. 05. Subtribe Ramondinae DC. ex Meisn. 32/ 5
3. 2. 06. Subtribe Litostigminae A. Weber & Mich. Mller 1 / 2 Litostigma G. WeiF. 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 KerrDorcoceras BungeMiddletonia C.
PuglisiOrnithoboea Parish ex C. B. ClarkePara-
boea C. B. Clarke Ridl. habdothamnopsis
3. 2. 10. Subtribe Didymocarpinae G. Don 32 1660- Aeschynanthus JackAllocheilos W. T. WangAllostig-
W. T. WangAnna
1830 Pellegr. Briggsiopsis K. Y. PanCathayanthe Chun
Conandron Sieb. & Zucc. Cyrtandra J. R. Forst. & G.
Forst. Didymocarpus Wall. Didymostigma W. T.
WangGlabrella Mich. Mller & W. H. ChenGyro-
cheilos W. T. WangHemiboea C. B. ClarkeHencke-
lia Spreng. Loxostigma C. B. ClarkeLysionotus D
DonMetapetrocosmea W. T. WangMicrochirita C
B. ClarkeYin Z. WangOreocharis Benth. Petroc-
odon HancePetrocosmea Oliv. Primulina Hance
Pseudochirita W. T. WangRaphiocarpus Chun
NoteBold face. Taxon represented in ChinaUnderlined. Endemic to ChinaSpaced print. New genus for China represents it maybe extinct.
ae. Based on the aberrant morphologyWang et al
1990established a separate tribe in subfam. Cyrtan-
droideae now Didymocarpoideaefor its accomodation
while Weber 2004proposed to exclude it from Gesne-
riaceae. The molecular study of Wang et al 2004a
confirmed its position within Gesneriaceaebut yielded
the surprising result that it is more closely allied to the
New World than to the Old World Gesneriaceae. Weber
et al 2013) ,thereforeincluded Titanotrichum in sub-
fam. Gesnerioideae as a monospecific tribe. Apart from
the geographical distribution SE mainland ChinaTai-
wan region of ChinaS Japanwhich is difficult to ex-
plain for a member of subfam. Gesnerioideaethis taxo-
nomic placement is in line with the isocotylous seedlings
the interpretation of Wang et al2002 as anisocotylous
is erroneous Whether the distribution of Titanotrichum
1M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 groupsbut other charac-
tersincluding the curious propagulesindicate an iso-
lated position.
Table 2 List of present and past genera of Chinese Gesneriaceae.
Chinese name
Tribe Subtribe
Distribution Species
Species no.
in China
endemicTax. status Reference Remark
Aeschynanthus Jack
芒毛苣苔属 Didymocarpoideae
From S ChinaN & S
India throughout
Malesia to New
Guinea and the
Solomon Islands
185 34
13Emended by inclusion
of Micraeschynanthus
The inclusion of
the monotypic
genus Micraeschy-
nanthus Malay
little bearing on
the definition of
W. T. Wang
S China GuizhouE
Yunnan2 2
2No change
Allostigma W. T.
S China 1 1
1No change See text under
Sunk into Oreocharis Mller et al
et al2014b
See text under
Anna Pellegr.
大苞苣苔属 Didymocarpoideae
ChinaN Vietnam 4 4
3No change
Beccarinda Kuntze
横蒴苣苔属 Didymocarpoideae
NE IndiaBurmaS
7 5
4No change
Boea Comm. ex
10 - RedefinedChinese
spp. now in Dorcoceras
and Damrongia
Puglisi et al
in press
See text under
Boeica C. B.
BhutanS ChinaN
& NE IndiaMyan-
marN VietnamNW
12 7
3No change
Bournea Oliv
四数苣苔属 Sunk into Oreocharis Mller et al
See text under
Briggsia Craib
粗筒苣苔属 Partly sunk into Oreo-
charispartly transferred
to Loxostigmaand two
spp. forming the new
genus Glabrella
Mller et al
et al2014b
Mller et al
See text under
Briggsia and
K. Y. Pan
S China C & S Si-
chuanNE Yunnan
1 1
1No change
Calcareoboea C. Y.
Wu ex H. W. Li
Sunk into Petrocodon Wang et al
et al2011b
See text under
Chirita Buch.
-Ham. ex D. Don
Split into 5 genera and
synomymisation with
Wang et al
et al2011a
See text under
Chiritopsis W. T.
Sunk into Primulina Wang et al
et al2011a
See text under
Cathayanthe Chun
扁蒴苣苔属 Didymocarpoideae
S China Hainan1 1
1No change
Conandron Sieb.
& Zucc.
E ChinaTaiwan re-
gion of ChinaS Ja-
1 1 No change
BhutanChinaN &
NE IndiaNepal
3-5 3 No change
84 广 西 植 物 36
Chinese name
Tribe Subtribe
Distribution Species
Species no.
in China
endemicTax. status Reference Remark
Cyrtandra J. R.
Forst. G. Forst.
Nicobar Islands and
S Thailand through
Malesia incl. Taiwan
region of China and
the S Pacific to the
Hawaiian Islands
652-818 1
1No change
Damrongia Kerr
China to Sumatra 10 1
1Re-established for
particular species of
erstwhile Chirita
inclusion of Boea clar-
keana Hemsl. and the
Asian species descri-
bed in Streptocarpus
Weber et al
Puglisi et al
in press
See text under
Chirita and
Dayaoshania W. T.
Sunk into Oreocharis Mller et al
See text under
Deinocheilos W. T.
Sunk into Oreocharis Mller et al
See text under
from N and NE Indi-
aNepal and S Chi-
na southwards to the
Malay Peninsula and
N Sumatra
>70 30
24Some spp. transferred
to Petrocodon
Weber et al
See text under
W. T. Wang
SE China Guang-
3 3
3No change
D. Fang W. T.
Sunk into Petrocodon Weber et al
See text under
Dorcoceras Bunge
Philippines and
4 2
1 or 2?) Re-established for par-
ticular non-Australa-
sianspecies of Boea
Puglisi et al
in press
See text under
Boea and Dorco-
Epithema Blume
S ChinaTaiwan re-
gion of China
20 2 No change at genus
revised by
Bransgrove &
Glabrella Mich.
Mller & W. H
S ChinaTaiwan re-
gion of China
3 3
3New genus estblished
for 3 spp. of Briggsia
not to be included in
Oreocharis or Loxostig-
Mller et al
2014Wen et
al2015 ab
See text under
Briggsia and
W. T. Wang
S China Guangxi
Guizhou) ,Vietnam
5 4 3No change
W. T. Wang
Loxoniinae ( ?)
S China W Guan-
gxi1 1
1Position in
Loxoniinae uncertain
Probably extinct
C. B. Clarke
C & S ChinaTai-
wan region of China
N VietnamS Japan
31 31
30Inclusion of
Metabriggsia 2
Weber et al
Inclusion does
not affect the
traditional con-
cept of Hemiboea
Hemiboeopsis W. T.
Sunk into Henckelia Weber et al
Henckelia Spreng.
From S China to Sri
c. 55 23
12Redefined to include
Chirita p. p. excl.
Microchirita and Prim-
ulinaand Hemiboep-
sisand to exclude
Weber et al
dleton et al
See text under
Chirita and
Isometrum Craib
金盏苣苔 Sunk into Oreocharis Mller et al
See text under
1M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
Chinese name
Tribe Subtribe
Distribution Species
Species no.
in China
endemicTax. status Reference Remark
Lagarosolen W. T.
Sunk into Petrocodon Weber et al
See text under
Leptoboea Benth.
细蒴苣苔属 Didymocarpoideae
BhutanN and NE
IndiaS China
Yunnan ) , Myan-
2-3 1 No change
Litostigma Y. G.
WeiF. Wen &
Mich. Mller
China Guizhou
Yunnan2 2
2Genus recently
Wei et al
See text under
C. B. Clarke
斜 片 苣 苔 属
S China Sichuan
Guangxi) ,N
11 11
8Recently inclusion of
caulescent Briggsia
Mller et al
See text under
Lysionotus D. Don
吊石苣苔属 Didymocarpoideae
From N India and
Nepal eastwards
through N Thailand
N Vietnam and S
China to S Japan
28 26
18No change
Metabriggsia W. T.
Included in Hemiboea Weber et al
W. T. Wang
S China Hainan1 1 No change
Microchirita C.
B. Clarke Y. Z.
From the Western
Ghats of India to the
foothills of the Hima-
layasthrough conti-
nental SE Asia to
SumatraBorneo and
18 2
1Raised from Chirita
sect. Microchirita
to generic rank
Wang et al
et al2011a
See text under
C. Puglisi
4 1 Genus recently estab-
lished for four species
of Paraboea
Puglisi et al
in press
See text under
Middletonia and
Opithandra B. L.
Sunk into Oreocharis Mller et al
See text under
Oreocharis Benth.
马铃苣苔属 Didymocarpoideae
BhutanNE India
>105 >102
>98Expanded to include
Ancylostemon Craib
Bournea Oliv. Brigg-
sia Craib p. p. - incl.
W. T. WangDeino-
cheilos W. T. Wang
trum CraibOpithandra
B. L. BurttParai-
sometrum W. T.
W. T.
Wangand Tremacron
CraibInclusion of
further ten spp. of
Mller et al
Mller et al
et al2014b
See text under
Briggsia and
Ornithoboea Parish
ex C. B. Clarke
From S China and
Vietbam southwards
to N Penins. Malaysia
16 5
1No change Revised by
Scott & Mid-
Paraboea C. B.
132 26
14Expanded by inclusion
of Phylloboea and
Trisepalumremoval of
four species and place-
ment in the new
genus Middletonia
Puglisi et al
Puglisi et al
in press
See text under
Paraboea and
05 广 西 植 物 36
Chinese name
Tribe Subtribe
Distribution Species
Species no.
in China
endemicTax. status Reference Remark
Paraisometrum W.
T. Wang
Sunk into Oreocharis Mller et al
See text under
Paralagarosolen Y.
G. Wei
Sunk into Petrocodon Wang et al
et al2011b
See text under
Petrocodon Hance
ChinaN Vietnam
NE Thailand
29 28
27Expanded to include
Calcareoboea C. Y. Wu
ex H. W. LiDidymo-
carpus Wall. p. p. -ex-
cl. typeDolicholoma
D. Fang W. T.
W. T.
len Y. G. WeiTengia
Chun and Wentsaiboea
D. Fang D. H. Qin
p. p. excl. type
Wang et al
et al2011b
See text under
Petrocosmea Oliv.
石蝴蝶属 Didymocarpoideae
NE IndiaS China
S Vietnam.
42 34
34No change
Platystemma Wall.
IndiaSW China
1 1 No change
Primulina Hance
Essentially southern
half of China and Vi-
>168 >154
>151Enormous expansion of
the previously mono-
typic genus by inclu-
sion of Chirita sect.
Wentsaiboea p. p. -
Wang et al
et al2011a
See text under
Pseudochirita W.
T. Wang
S China C & W
Guangxi) ,Vietnam
1 1
1No change
Raphiocarpus Chun
S China and N & C
13 8
7No change since We-
ber 2004but changes
to be expected
S China 1 1
1No change
From India and S
China to New
Guineaone to
3?) spp. in C
15 2
1No change Recent descrip-
tion of 2 new and
odd spp. from
Thailand Pat-
NE IndiaNepal
BhutanSW & S
ChinaSE Asia and
Malesia to New
21 6
2No change
MalesiaS China 5 1 No change Revision in
prep. Weber
ic species
Sunk into Damrongia Puglisi et al
in press
See text under
Tengia Chun
世纬苣苔 Sunk into Petrocodon Wang et al
et al2011b
See text under
Thamnocharis W. T.
Sunk into Oreocharis Mller et al
See text under
1M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
Chinese name
Tribe Subtribe
Distribution Species
Species no.
in China
endemicTax. status Reference Remark
SE ChinaTaiwan
region of ChinaS
1 1 Placed in subfam.
C. N. Wang
et al2004a
Perret et al
et al2013
Tremacron Craib
短檐苣苔属 Sunk into Oreocharis Mller et al
See text under
Trisepalum C. B.
Sunk into Paraboea Puglisi et al
See text under
D. Fang & D. H. Qin
Partly incl. type
sunk into Primulina
partly into Petrocodon
Weber et al
See text under
Chirita and Pet-
W. W. Sm.
S ChinaTaiwan re-
gion of China
8 8
8No change
NotePresently accepted genera in bold face. Genera that have been synoymised since the publication of theFlora of ChinaWang 1990Wang et al1998 are given
in square brackets and not bold. Infrafamilial position according to Weber et al,( 2013
With regard to subfam. Didymocarpoideaeboth
tribesEpithemateae and Trichosporeaeare represen-
ted in China.
In tribe Epithemateaewhich is apparently a rel-
ict tribe with morphologically rather odd and isolated
generaall four subtribes are represented in China.
From the seven genera two are missing in CinaMono-
phyllaea in subtribe Monophyllaeinae this thus con-
tains only Whytockiawith 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 tropicsMonophyllaea includes more than 40
specieswith distribution throughout Malesia Sumatra
to New Guinea and S Thailand to Java) ,Loxonia com-
prises three species restricted to western Malesia Su-
matraMalay Peninsula and BorneoWhytockia is par-
ticularly remarkable as morphological studies Weber
19761982indicate that the genus has retained primi-
tive characters in relation to Monophyllaeasuggesting
that subtribe Monophyllaeinae has originated in the ex-
tratropics andafter reaching the South East Asian trop-
icsradiated into the many species of Monophyllaea
Stauranthera and Gyrogyne are represented with a single
species each in ChinaRhynchoglossum and Epithema
with two species each.
Tribe Trichosporeaewith 10 subtribes tentatively
recognised by Weber et al 2013) ,is represented by
five subtribes in China.
Among the more primitive alliancessubtribe Cor-
allodiscinaewith the single genus Corallodiscusis re-
markable. Its few species are rosette plants with scapose
inflorescences and tetrastaminate flowers.
Morphologically heterogeneous is subtribe Lepto-
boeinaewhich is represented by almost all of its genera
in China the only lacking genus is the South Indian
Championiathe inclusion of which in Leptoboeinae
still requires confirmation Leptoboeinae includes ro-
sette plants Beccarinda ) , sub-shrubby caulescent
plants LeptoboeaBoeicaRhynchotechumand unifo-
liate plants Platystemma LeptoboeaBoeica and
Rhynchotechum have correctlythought to be closely
related in older classificationsbut Burtt 1963as-
sumed a close relationship of Rhynchotechum with Cyr-
tandrasubfam. Cyrtandroideae tribe Cyrtandreae) ,
because the two genera have indehiscentberry-like
fruits in common. Apart from the changes in taxonomic
position as compared to former classificationsno 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
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 al2010
Subtribe Loxocarpinae is the second largest sub-
tribe of Trichosporeae. In total it includes about a dozen
genera. Two of themDorcoceras and Middletoniahave
been re-established very recently and both are repre-
sented in Chinatogether with DamrongiaOrnitho-
boeaParaboea and Rhabdothamnopsis The latter is
with a single species endemic to Chinawhile the re-
maining genera have their main distribution mostly fur-
ther south. A characteristic feature of the subtribe but
not present in all taxais 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-
erawith 24 represented in China. Six genera with one
or very few species i. e AllocheilosAllostigma
cosmeaare endemic to this countryothers have their
main distribution elsewhere and reach with one or few
species the southernmost part of China e. gCyrtan-
drawith> 800 species is the largest genus of Gesneri-
aceaereaches with a single species to South Taiwan
ChinaMicrochirita with a wide distribution from the
Western Ghats of Indiathe foothills of the Himalayas
through continental SE Asia into SumatraBorneo and
Javahas two species in China out of c. 18
The subtribes of Trichosporeae not represented in
China are Jerdoniinae only Jerdonia indicaS India) ,
Tetraphyllinae only Tetraphyllum3 spp. NE India
BangladeshBurmaThailand) ,amondinae Haber-
leaRamondaJancaea5 spp. SW and SE Europe) ,
Streptocarpinae traditionally 9 generarecently all amal-
gamated in StreptocarpusNishii et al2015177 spp.
AfricaComoro Islands and Madagascar) ,and Didissan-
drinae DidissandraTribouniac. 10 spp. W Malesia
Thailand. Altogetherthese subtribes include only one
or few generaas to species numberStreptocarpinae is
clearly the largest subtribe. The link from the African to
the Asiatic including Chinesespecies via the Asiatic
species of Streptocarpusproved untenablethe Asiatic
species described in Streptocarpus belong to the purely A-
sian genus Damrongia Puglisi et alin 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
isfor instanceall genera of tribe Epithemateaeno
changes in the taxonomic delineation have occurredin
others the inclusion of formerly distinct genera has little
bearing on their new definition e. g the inclusion of
Metabriggsia into Hemiboeain which differences in
the ovary structure simply proved erroneousWeber et
al2011c) ,but others have a rather complex tax-
onomic history which require a more detailed explana-
tion. In the following the fate of the latter generaplus
new or newly established genera is briefly outlinedas far
as changes since the treatment of Wang et al 1990and
Wang et al 1998are concerned. De factoall changes
occurred from 2010 onwardswith the new genus Litostig-
ma Wei et al2010as the first.
Boea This was previously a widespread Asiatic-
Malesian genuswith three species B. clarkeanaB
hygrometricaand B. philippensis represented in
southern China. From the recent molecular-systematic
studies of Puglisi et al,( in press) ,Boea emerged as
an essentially Australasian genuswith distribution in
Eastern IndonesiaPapua New Guineathe 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 additionBoea
hygrometrica endemic to Chinaand B. philippensis
Philippines and Vietnamwere included in Dorco-
ceras the former representing the type species of
DorcocerasBoea clarkeana Streptocarpus clar-
keanus) ,howeverwas attributed to the genus Dam-
rongia see there
Briggsia The genus is no longer relevantas the
1M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. longifoliahas
been synonymised with Oreocharis Mller et al
2011b2014 In its traditional conceptBriggsia in-
cluded acaulescent rosette plants as well as caulescent
plantsboth with characteristic large and ventrally
pouchedbriggsioidflowers. 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 al2014Wen et al2015a
b The briggsoidflowers arose apparently inde-
pendently in different alliances Mller et al2011b
See also notes for Oreocharis
Chirita Dramatic changes occurred in this tradi-
tional and large genus with up to 140 species de-
scribed hereinplaced in four sectionssectChirita
sect. LiebigiasectMicrochiritaand sect. Gibbosac-
cusWood1974Hilliard2003 ) ,ending in a syn-
onymisation of the genus with Henckelia Weber et al
2011a The character defining Chirita was traditional-
ly seen in the chiritoidstigmaa stigma with upper
and lower lobebut the upper largely or completely re-
ducedand the lower one expandend and often bipar-
tite. The molecular studies clearly revealed that this
character is homoplastichaving evolved independently
in several alliances of tribe Trichosporeae. As a conse-
quenceChirita was split into five generawith the
majority of Chirita sect. Chirita and the monotypic
HemiboeopsisChina 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 belowChirita
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 wastogether with Chiritopsis and Wentsaiboea
including its type species and a later published species
W. luochengensisLiu et al2010Xu et al2012) ,in-
cluded) ,included in the originally monotypic genus
Primulina Wang et al2011Weber et al2011aLi &
Xia 2012 The latter genus is essentially geographically
restricted to Chinanow with >150 specieswith some
species extending or endemic to Vietnam.
The newly defined Henckelia has a broad circum-
scription at presentand 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 al2011aand 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 colorationand a
vegetative habit with rhizomatous compacted stems with
leaves in a basal rosettewith 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
countedwith one exception of a tetraploid number
Christie et al2012) ,whereas it is 2n = 20 in the only
member of Petrocodon so far counted P. hanceiCao et
al2003Mller & Pullan2015 onwards
Damrongia. This is a genus re-established by We-
ber et al 2011afor the accomodation of a couple of
species previously placed in Chirita The species repre-
sent rosette plants with scapose inflorescencesinfun-
dibuliform corollas with 2 stamensand a pistil with a
chiritoidstigma see Chirita It was thought to be
restricted to Thailand and the NW of Peninsular Malay-
siabut the Chinese Boea clarkeana also proved to be-
long to that genus Puglisi et alin press The same
applies for the Asian species described in Streptocarpus
on grounds of the twisted fruits Puglisi et alin 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
2014for two species of Briggsiawhich do not fit in-
to Oreocharis or Loxostigma A third species was added
by Wen et al 2015ab 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 entitiesDidymocarpus s. str. Henckelia with
five sectionssect. HenckeliaS India and Sri Lanka
sect. Loxocarpussect. Heteroboeasect. Loxocarpus
45 广 西 植 物 36
sect. Didymanthus and sect. Glossadeniaand Hovanel-
laWeber & Burtt 1998 The molecular data of Mller
et al 2009indicated that the split was not sufficient
and that Henckelia had to be redefined. This was done
by Weber et al2011a The type section Henckelia
sect. HenckeliaS India and Sri Lankawas 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
2011aMiddleton et al2013Yao2012) ,and the
last three sections have been included in the re-estab-
lished and largely expanded genus Codonoboea Kiew &
Lim 2011Middleton et al2013 Neither Loxocarpus
nor Codonoboea have species in China.
Litostigma This new genus established by Wei et
al2010for 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 al2011Weber et al2011a It can be character-
ized by inflorescences that usually appear in two or sev-
eral in a leaf axiloften displaced onto the petioleof-
ten consisting of a short-stalked serial flower pair only
but this repeated several times. From the c. 25 species
2 occur in ChinaM. hamosa and M. prostrata re-
cently described by Li & Xia2012
Middletonia A new genus established by Puglisi
et alin pressfor the accommodation of four species
separated from Paraboeawith one species distributed
in China M. multiflora
OreocharisWhile Oreocharis included some 27
species in its traditional delineation Wang et al
1998) ,the number increased enormously through the
inclusion of AncylostemonBourneaBriggsia p. p.
incl. type) ,DayaoshaniaDeinocheilosIsometrum
ParaisometrumThamnocharisand Tremacron Mller
et al2011bMiddleton et al2013 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
al2014bMller et al2014Wen et al2015ab) ,
the rest were attributed to the following three genera
the caulescent species were referred or returned to Lox-
ostigmathree acaulescent species with glabrous leaves
and stems were moved into a new genus Glabrella Mich.
Mller W. H. Chenand 11 acaulescent species were
transferred to Oreocharis New species were recently de-
scribed Liu et al2012Chen et al 20132015Tan
et al 20132015Rossini and Freitas 2014Li and Li
2015Yang et al2015) ,and Oreocharis now compri-
ses now 106 speciesall representing acaulescent rosette
plants with scapose inflorescencesbut with considera-
ble variability in the symmetryshape and coloration of
the flowers. Oreocharis leiophylla and O. sinensis mak-
ing up the former genus Bourneahave actinomorphic
flowers. The corolla shape in Oreocharis varies from tu-
bularthrough funnel-shapedcampanulate to
briggsioidthe stamen number is 1) ,24 or in
actinomorphic flowers4 or 5 in line with the number of
petalsin some diandrous species previously constitu-
ting the genus Opithandrathe 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
butas the molecular data showthe species relation-
ships run across genera and these are thus untenable.
Like in Petrocodonthe 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
speciesc 20 of them found in China. Changes since
then relate to the inclusion of the monotypic Phylloboea
Myanmarand Trisepalum c. 13 spp. 1 in China
Puglisi et al2011 More recentlyPuglisi et alin
pressremoved 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
Chinaof these 14 exclusively there i. e endemic
PetrocodonIn its traditional conceptPetrocodon
was a genus with two Chinese speciesboth with small
white flowers and two fertile stamens. Based on molecu-
lar dataWang et al 2011added Tengia with small
whitepentamerous actinomorphic flowers and five sta-
mens) ,Calcareoboea with largered and tubular flow-
1M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 laterWeber et al 2011b
added the monotypic Dolicholoma with flowers with
narrow tube and subactinomorphic limb with longacute
lobes) ,all species of Lagarosolen with flowers similar
to those of Dolicholoma and Paralagarosolen) ,one spe-
cies of Wentsaiboea without typeand three species of
Didymocarpus D. hanceiD. mollifoliusD. 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 al2014a) ,Hong et al 2014) ,Xu
et al 2014and Li & Wang 2015increased the spe-
cies number to almost 30. The different flower shapes
and colours now assembled in Petrocodon suggest differ-
ent pollinator adaptationsincluding beebutterfly and
bird pollination. Until recentlyPetrocodon could be
defined by possessing 2 stamens or 5 in the former ge-
nus Tengia with actinomorphic flower Using both mor-
phological and molecular dataa species possessing 4
stamens was describedP. hunanensis Yu et al
2015 This indicates athe great diversity that is
still unexplored in Chinaand bthat 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 someyou 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-
lenWei2004WentsaiboeaFang & Qin2004) ,the
number has dropped to 45 Table 1 Since 2010and
essentially based on molecular dataseven new genera
have been added to Chinaeither based on newly col-
lected material Litostigma) ,or from segregation of ex-
isting genera in China GlabrellaDamrongiaDorco-
cerasMiddletoniaMicrochiritaand Primulinasee
previous chapter) ,or by a new circumscription of gene-
ra Henckelia
On the other handa considerable number of gene-
ra 22are lost by synonymisation with other genera
Wentsaiboea. Alsothe four Asiatic species of Strepto-
carpus Hilliard and Burtt 1971) ,one of themSclar-
keanus Hemsl. Hilliard and B. L. Burtt Boea clar-
keana Hemsl. as such in the Flora of China of Wang et
al1990Wang et al1998) ,from Chinahave found
a proper generic place Damrongia Streptocarpus
thereforecan be definitely removed from the list of
genera extending into China.
By tribethere is no change in generic state in Ep-
ithemateaeas to Trichosporeae-Litostigminae there is
the addition of one genus Litostigma) ,in Trichospore-
ae-Loxocarpinae there are two fewer genera in China
BoeaTrisepalum) ,but three additional ones Dam-
rongiaDorcocerasMiddletonia) ,an overall net gain
of one. The greatest changes occurred in Trichosporeae-
Didymocarpinaewith a loss of 20 generaand a gain of
three GlabrellaHenckeliaMicrochirita) ,thus an o-
verall loss of 17 genera. This tribe is still not fully un-
derstood and further work is necessary to understand its
evolutionand further changes may be in the waiting
though these may be less drasticbut fine-tuning.
The number of endemic genera has decreased from
27 Wang et al1998Wei 2004Fang & Qin2004
to 11. Ten of them include only one to three species
Allocheilos 2Allostigma 1Briggsiopsis 1
Cathayanthe 1Didymostigma 3Glabrella 3
Gyrogyne 1Litostigma 2Metapetrocosmea 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 decadesthere seems to be an overall
tendency to include small or monotypic genera and the
large Dichrotrichumthat had been raised on the basis
of the presence of an unusual characteristicinto larger
ones on grounds of more shared than distinguishing
characteristics. This is importantsince single character
taxonomy is fraught with problems when picking the
65 广 西 植 物 36
wrongcharacter a genus was not established for.
4. 2 Morphological parallelisms and reversals
Burtt e. g. Burtt1963 pointed out that in
Gesneriaceae there are many exceptions and apparent
parallelisms in formand that the homology of similar
characters on which a genus is basedis sometimes dif-
ficult to ascertain e. g. Burtt1968 The erstwhile
genus Chirita is a prime examplein 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 independentlyand Chirita thus had to be split
into five genera Weber et al2011a see also be-
At the generic levelthe number of fertile stamens
4 or 2played a significant roleespecially in Chinese
Gesneriaceae. For exampleDeinocheilos with 2 sta-
menshas been kept separate from Tremacron 4 sta-
mens) ,though the two genera are otherwise indistin-
Floral symmetry is another problematic character of
importance both at the generic and at the tribal level.
Fritsch 1893 /941908established tribe Ramondeae
to include CorallodiscusHaberleaPetrocosmea
Ramondaand Saintpaulia His intention was to recog-
nise a group with similar vegetative habit flat rosette
plantswith septicidal capsule dehiscencerather than
floral shape. Thusthis tribe includes four genera with
zygomorphic flowerstwo of which have long floral tubes
and four stamens Haberlea and Corallodiscus) ,two
with flat-faced truncatezygomorphic corollas with 2
stamens Petrocosmea and Saintpauliaand one actino-
morphic genus Ramondia =amonda He placed an-
other genus with actinomorphic flowersConandronin
a separate tribe. Burtt 1970accepted 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 1990and Wang et
al 1992with the reverse view of an ancestral actino-
morphy and derived zygomorphy redefined tribe Ramon-
deae to include only actinomorphic generaBournea
ConandronRamondaTengiaand Thamnocharis
Molecular phylogenetic work has proven Burtts view to
be correct since they showed that all actinomorphic gen-
era have independent origins Mller et al1999
20092011aWang et al2010
Hidden homoplasies destabilize a classification sys-
tem that wants to reflect phylogenyand 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 sowe 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 al2011a This has been shown for the
Neotropical Gesneriaceae as well as for the Afro-Mala-
gasi Gesneriaceae of tribe Streptocarpinaewhich have
been reduced from nine often small genera 7 with <3
speciesto 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 al2006Clark & Zimmer 2003
The convergence of hypocyrtoid or pouchedflowers
within Drymonia and across several lineages of closely
related genera has resulted in a revised circumscription
of genera in the Columneinae Clark et al2015Clark
et al2012Smith & 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 al2004Roalson et al2005bClark
et al2011
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
2006before its accommodation in a new genus Wei et
al2010 The recently described speciesPrimulina
guangxiensis Yan Liu and W. B. Xu was initially
placed with a superficially similar species Liu et al
2011) ,but after molecular studiespartly by the same
1M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
authorsit was discovered that it belonged in Petrocodon
assuming the integrity of the molecular dataand was
transferred to this genus Xu et al2014
In some cases incomplete knowledge of generic
concepts might be at issuein others perhaps the dis-
covery of species with new character combinationssuch
as Petrocodon hunanensis with four stamenscause un-
certainty. Howeverwith the availability of molecular
data and an extensive data set of molecular sequences
on public databasessuch as GenBankthese uncer-
tainties can be addressed by molecular approaches.
Howeverwhere traditional taxonomic approaches
are concernedthe utilisation and balancing of all char-
actersnot just one or a fewhas to be considered in
taxonomic decisions. This might have become more
complex with the new delineations of some generaand
thetraditionalcharacters of corolla shapestigma 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 Trichosporeaeparticularly subtribe
Didymocarpinaeis still to be completed. Because of
the large number of representatives in this subtribe resi-
ding in Chinait can play a pivotal role in this endeav-
At the generic levelRaphiocarpus 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 Briggsiainto Loxostigma
Mller et al2014has expanded its definition to in-
clude seeds without appendagesnarrowing its gap to
Raphiocarpusbut 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 Gyrocheilosapart from Did-
ymocarpus properand may belong in this genus. Li et
al 2015published 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 speciesand the latter species with straight
and twisted fruits. Such an assemblage seems at first
dissatisfyingbut it has precedence in other genera.
The genus Streptocarpus was characterised for species
possessing twisted fruits. Though it includes a wide
range of morphological formsfrom unifoliatesplurifoli-
atesrosulatesrosettesherbaceous and woody caules-
cent forms Hilliard & Burtt1971 It has now also
been extended to include species with non-twisted fruits
Nishii et al2015
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 linksthrough intensified and sys-
tematic taxon sampling. Alsothe great number of new
species described in some generaespecially Primulina
which ballooned from around 100 when it was redefined
in 2011to now >150 with many more species to be de-
scribedillustrates 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 2012TribouniaMiddleton et al
2012BilloliviaMiddleton et al2014Chayamaritia
Middleton et al2015 ecent discoveries outlined
here clearly demonstrate the recent progress in Gesneri-
aceae diversity of Chinabut more importantly the fu-
ture need to continue phylogenetic and taxonomic stud-
ies for a long-term stable classification. Without a
doubtthere 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
supporting our research. We also thank the Royal Bo-
tanic Garden Edinburghsupported by the Rural and
Environment Science and Analytical Services division
RESASin the Scottish Government.
calcicolaA new genus and species of Gloxinieae Gesneriace-
aefrom Minas GeraisBrazilJ. Taxon59203-208.
BENTHAM G1876. Gesneriaceae. Bentham GHooker JD
eds Genera plantarum 2 2) [M. LondonReeve and
Co. Williams and Norgate990-1 025.
BOGGAN JK2006. What happened to Gloxinia? [J. Gesneri-
ads563) : 26-32.
BRANSGROVE KMIDDLETON DJ2015. A revision of Epithe-
ma Gesneriaceae) [J Gard Bull Singapore671) : 159 -
BURTT BL1963. Studies in the Gesneriaceae of the Old World.
XXIV. Tentative keys to the tribes and generaJ Notes Roy
Bot Gard Edinburgh24205-220.
BURTT BL1968. Studies in the Gesneriaceae of the Old World
XXIX. A reconsideration of generic limits in tribe Trichosporeae
J Notes Roy Bot Gard Edinburgh29219-225.
BURTT BL1970. Studies in the Gesneriaceae of the Old World.
XXXI. Some aspects of functional evolution
J Notes Roy Bot
Gard Edinburgh301-10.
BURTT BL1977. Classification above the genusas exemplified
by Gesneriaceaewith parallels from other groupsJ Pl Syst
EvolSuppl. I97-109.
BURTT BLWIEHLER H1995. Classification of the family
GesneriaceaeJ Gesneriana 11-4.
CAO LMCAO MTANG XLet al2003. Chromosome numbers
of 4 species in the Gesneriaceae from GuangxiJ. Guihaia23
4) : 331-333.
CHEN WHSHUI YMMLLER Met al2013. Oreocharis jin-
pingensis Gesneriaceae ) ,a new species from the Xilong
Mountains in southwest ChinaJ Ann Bot Fenn50312 -
CHEN WHMLLER MSHUI YMet al2014a. Three new
species of Petrocodon Gesneriaceae) ,endemic to the limestone
areas of Southwest Chinaand preliminary insights into the di-
versification patterns of the genusJ Syst Bot391) : 316
CHEN WHSHUI YMMLLER M2014b. Two new combina-
tions in Oreocharis Gesneriaceaefrom ChinaJ. Candollea
692) : 179-182.
CHEN WHMLLER MCHUN RZet al2015. Oreocharis syner-
giaa new species of Gesneriaceae from Northwestern Yunnan
ChinaJ. Phytotaxa2331) : 90-93.
CHRISTIE FBARBER SMLLER M2012. New chromosome
counts in Old World GesneriaceaeNumbers for species hitherto
regarded as Chiritaand their systematic and evolutionary signif-
icanceJ Edinburgh J Bot692) : 323-345.
CLARK JL2005. A monograph of Alloplectus Gesneriaceae) [J
CLARK JLZIMMER EA2003. A preliminary phylogeny of Allo-
plectus Gesneriaceae) : implications for the evolution of flower
resupinationJ Syst Bot28365-375.
netic relationships and generic boundaries in the Episcieae
Gesneriaceaeinferred from nuclearchloroplastand morpho-
logical dataJ. Taxon55313-336.
CLARK JL2009. Systematics of Glossoloma Gesneriaceae) [J
Syst Bot Monogr891-126.
pendent origin of radial floral symmetry in the Gloxinieae
GesnerioideaeGesneriaceaeis supported by the rediscovery
of Phinaea pulchella in CubaJ Syst Bot36757-767.
CLARK JLFUNKE MMDUFFY AMet al2012. Phylogeny of a
Neotropical clade in the Gesneriaceaemore tales of convergent
evolutionJ Int J Plant Sci173894-916.
anti-bee pollination mechanisms in the Neotropical plant genus
Drymonia Gesneriaceae) [J Evol Ecol29355-377.
FANG DQIN DH2004. Wentsaiboea D. Fang and D. H. Qina
new genus of the Gesneriaceae from GuangxiChinaJ. Acta
Phytotax Sin42533-536.
FRITSCH K1893 1894. Gesneriaceae M/ /ENGLER A
PRANTL K eds Die natürlichen Pflanzenfamilien IV / 3B.
LeipzigW. Engelmann133-144 1893) ; 145-185 1894
FRITSCH K1904. Die Keimpflanzen der Gesneriaceen mit besonder-
er Berücksichtigung von Streptocarpusnebst vergleichender Studi-
en über die Morphologie dieser FamilieM. JenaG. Fischer.
eds Die natürlichen PflanzenfamilienNachtrag III.
Nachtrge zu Teil IVAbteilung 3B. LeipzigW. Engelmann
HILLIARD OM2003. A revision of Chirita sect. Liebigia Gesne-
riaceae) [J Edinburgh J Bot603) : 361-387.
HILLIARD OMBURTTBL1971. StreptocarpusAn African plant
studyM. PietermaritzburgNatal University Press.
HONG XZHOU SBWEN F2014. Petrocodon villosus Gesneri-
aceae) ,a new species from GuangxiChinaJ. Blumea59
JONG K1970. Developmental aspects of vegetative morphology of
StreptocarpusD. EdinburghUniversity of Edinburgh
KIEW RLIM CL2011. Names and new combinations for Penin-
sular Malaysian species of Codonoboea Ridl. Gesneriaceae
J Gard Bull Singapore622) : 253-276.
LI HQWANG YQ2015. Petrocodon confertiflorus Gesneriace-
ae) ,a new species from GuangdongChinaJ Ann Bot
Fenn521-2) :27-32.
LI JMXIA Z2012. A new species of Microchirita Gesneriace-
aefrom YunnanChinaJ J Syst Evol506) : 576.
LI JMLI ZM2015. Oreocharis brachypodus Gesneriaceae) ,a new
taxon from GuizhouChinaJ. Phytotaxa204 4) : 296-299.
LI JMSUN WJCHANG Yet al2015. Systematic position of
Gyrocheilos and some odd species of Didymocarpus Gesneriace-
aeinferred from molecular datawith reference to pollen and
other morphological charactersJ J Syst Evoldoi10. 1111 /
jse. 12169.
LIU YXU WBPAN B2010. Wentsaiboea tiandengensis sp. nov.
and W. luochengensis sp. nov. Gesneriaceaefrom Karst caves
in Guangxisouthern ChinaJ Nord J Bot28739-745.
LIU YXU WBHUANG YS2011. Primulina guangxiensis sp.
nov. Gesneriaceaefrom a karst cave in GuangxiChinaJ
Nord J Bot29682-686.
LIU YXU WBHUANG YSet al2012. Oreocharis dayaos-
hanioidesa rare new species of Gesneriaceae from eastern
GuangxiChinaJ Bot Stud53393-399.
1M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
cotyly and meristem initiation in an unorthodox plantStrepto-
carpus rexii Gesneriaceae) [J. Planta2253) : 653-663.
MIDDLETON DJMLLER M2012. Tribouniaa new genus of
Gesneriaceae from ThailandJ. Taxon616) : 1 286-1 295.
MIDDLETON DJTRIBOUN P2012Somraniaa new genus of
Gesneriaceae from ThailandJ Thai Forest Bull Bot. ) ,40
MIDDLETON DJWEBER AYAO Tet al2013. The current
status of the species hitherto assigned to Henckelia Gesneriace-
ae) [J Edinburgh J Bot703) : 385-404.
MIDDLETON DJATKINS HLUU HTet al2014. Billoliviaa
new genus of Gesneriaceae from Vietnam with five new species
J. Phytotaxa1614) : 241-269.
amaritia GesneriaceaeDidymocarpoideae) ,a new genus from
Southeast AsiaJ Pl Syst Evol3017) : 1 947-1 966.
MLLER MCLOKIE MCUBAS Pet al1999. Integrating mo-
lecular phylogenies and developmental geneticsa Gesneriaceae
GORNALL RJ eds Molecular systematics and plant evolu-
tion. LondonTaylor and Francis375-402.
MLLER MPFOSSER MJANG CGet al2009. A preliminary
phylogeny of the " Didymocarpoid Gesneriaceae" based on three
molecular data setsincongruence with available tribal classifi-
cationsJ Am J Bot965) : 989-1 010.
MLLER MFORREST AWEI YGet al2011a. A molecular
phylogenetic assessment of the advanced Asiatic and Malesian
didymocarpoid Gesneriaceae with focus on non - monophyletic
and monotypic generaJ Pl Syst Evol2923-4) : 223-248.
delineation for Oreocharis incorporating an additional ten genera
of Chinese GesneriaceaeJ. Phytotaxa231-36.
MLLER MCHEN WHSHUI YMet al2014. A new genus of
Gesneriaceae in China and the transfer of Briggsia species to
other generaJ Gard Bull Singapore662) : 195-205.
MLLER MPULLAN M2015onwards. RBGE WebCyte2-An
updated Gesneriaceae cytology databaseEB / OL. http/ / el-
mer. rbge. org. uk/Webcyte.
MORAMMJL CLARK2016. Molecular phylogeny of the neo-
tropical genus Paradrymonia Gesneriaceae) ,reexamination of
generic concepts and the resurrection of Trichodrymonia and
CentrosoleniaJ Syst Botin press.
NISHII KKUWABARA ANAGATA T2004. Characterization of
anisocotylous leaf formation in Streptocarpus wendlandii Gesne-
riaceae) : significance of plant growth regulatorsJ Ann Bot
NISHII KHUGHES MBRIGGS Met al2015. Streptocarpus re-
defined to include all Afro-Malagasy GesneriaceaeMolecular
phylogenies prove congruent with geography and cytology and
uncovers remarkable morphological homoplasiesJ. Taxonin
PATTHARAHIRANTRIC N2014. The genus Rhynchoglossum Blume
Gesneriaceaein ThailandJ Thai For Bull4224-34.
ral and spatial origin of Gesneriaceae in the New World inferred
from plastid DNA sequencesJ Bot J Linn Soc17161-79.
sights into the relationships between ParaboeaTrisepalumand
Phylloboea Gesneriaceaeand their taxonomic consequences
J. Taxon606) : 1 693-1 702.
PUGLISI CYAO TLMILNE Ret al. New generic recircum-
scription of the Loxocarpinae Gesneriaceae) ,as inferred by
phylogenetic and morphological dataJ. Taxonin press.
ROALSON EHBOGGAN JKSKOG LE2005a. Reorganization of
tribal and generic boundaries in the Gloxinieae Gesneriaceae
Gesnerioideaeand the description of a new tribe in the Gesne-
rioideaeSphaerorrhizeaeJ. Selbyana25225-238.
ling Gloxinieae Gesneriaceae I. Phylogenetic patterns and
generic boundaries inferred from nuclearchloroplastand mor-
phological cladistic datasetsJ. Taxon54389-410.
ROSSINI JFREITAS J2014. Oreocharis yunnanensisa new
name for the illegitimate Oreocharis glandulosa Gesneriaceae
from ChinaJ. Phytotaxa163 3) : 180 .
SCOTT SMMIDDLETON DJ2014. A revision of Ornithoboea
Gesneriaceae) [J Gard Bull Singapore661) : 73-119.
SHUI YMCHEN WH2006. Seed plants of the karst region in
ChinaM Monmouth JunctionUSAScience Press USA Inc.
SMITH JFCLARK JL2013. Molecular phylogenetic analyses re-
veal undiscovered monospecific genera in Episcieae Gesneri-
aceae Syst Bot382) : 451-463.
netic analysis within tribes Gloxinieae and Gesnerieae Gesneri-
oideaeGesneriaceae) [J Syst Bot29947-958.
SOLEREDER H1909. ber die Gattung RehmanniaJ. Ber
Deutsch Bot Ges27390-404.
TAN YHLI JWPAN Bet al2013. Oreocharis glandulosaa
new species of Gesneriaceae from southern YunnanChinaJ
TAN YHLI JWYIN JT2015. Oreocharis tsaiia new species of
Gesneriaceae from southern YunnanChinaJ. Phytotaxa195
2) : 188-192.
WANG CNMLLER MCRONK QCB2004a. Phylogenetic posi-
tion of Titanotrichum oldhamii Gesneriaceaeinferred from
four different gene regionsJ Syst Bot292) : 407-418.
WANG CNMLLER MCRONK QCB2004b. Altered expres-
sion of GFLOthe Gesneriaceae homologue of FLORICAULA /
LEAFYis associated with the transition to bulbil formation in
Titanotrichum oldhamiiJ Dev Gen Evol214122-127.
WANG WTPAN KYLI ZY1990. GesneriaceaeM/ / WANG
WT ed Flora Reipublicae Popularis Sinicae. Vol. 69. Bei-
jingScience Press125-581.
WANG WTPAN KYLI ZY1992. Keys to the Gesneriaceae of
ChinaJ Edinburgh J Bot495 - 74. Translated from Chi-
nese by W. T. Wangwith introduction and notes by B. L.
WANG WTPAN KYLI ZYet al1998. GesneriaceaeM/ /
WU ZYRAVEN PH eds. Flora of China. Vol. 18. Beijing
Science PressSt. LouisMissouriMissouri Botanical Garden
WANG YZLI ZYPAN KYet al2002. Pattern and significance
of seedling development in Titanotrichum oldhamii Gesneriace-
ae) [J Acta Bot Sin44903-907.
WANG YZLIANG RHWANG BHet al2010. Origin and phy-
logenetic relationships of the Old World Gesneriaceae with acti-
nomorphic flowers inferred from ITS and trnL-trnF sequences
J. Taxon594) : 1 044-1 052.
WANG YZMAO RBLIU Yet al2011. Phylogenetic recon-
struction of Chirita and allies Gesneriaceaewith taxonomic
treatmentsJ J Syst Evol4950-64.
下转第113 Continue on page 113
06 广 西 植 物 36
J中国药学杂志4614) :1 049-1 052.
YANG YJXIE SQMENG ZG2012. Pollination ecology of Coptis
teeta Wall. an endangered medicinal plant J Acta Bot Bore-
al-Occident Sin327 1 372 1 375. 杨艳娟
2012. 濒危药用植物云南黄连传粉生态学研究J西
北植物学报327) :1 372-1 375.
ZHANG JYANG FJLIU Set al2009. Breeding system study in
astragalusm embranaceus FicshBunge J Ginseng Res
3) :17-20.
2009. 膜荚黄芪繁殖系
统的研究J人参研究3) :17-20.
ZHANG RBDOU QLHE Pet al2006. Study on the breeding
system of the endangered plant Euonymus chloranthoides Yang
J. Guihaia263 308 -312. 张仁波
2006. 濒危植物缙云卫矛繁育系统研究J广西植物26
3) :308-312.
ZHAO S张硕) ,GAO SP高素萍) ,2014. Distyly and selfing in-
compatibility of Plumbago auriculata J. Guihaia346 747
高素萍2014. 蓝花丹的花部形态二态性及自
交不亲和特性J广西植物346) : 747-753.
ZHANG WBJIN ZX2009. Floral syndrome and breeding system
of endangered plantsinocalycanthus chinensis J J Zhejiang
Univ Med Sci362) : 204-210.
金则新2009. 濒危
362) :204-210.
ZHANG XHZHANG DXYIN DSet al2009. Study on vitality
measurement and storing methods of Asiatic Lily pollen J
Northern Hortic2) :172-175.
2009. 合花粉生活力测定和贮藏方法研究J北方园
2) : 172-175.
ZHANG ZL2003. Plant physiology experimental guidance M
BeijingHigher Education Press224-225.
ZHAO ZH2012. A preliminary study of Plumbago auriculata bio-
logical characteristicsD. YaanSichuan Agric Univ. 赵志
2012. 蓝花丹生物学特性初步研究D雅安四川农
ZHU HMA RJ2009. Preliminary studies on breeding system of
invasive weed Ipomoea cairica L. J J NW A & F Univ37
5) :211-216.
马瑞君2009. 入侵杂草五爪金龙有性
繁育系统的初步研究J西北农林科技大学学报375) :
ZHOU HJTANG LMA Xet al2003. A preliminary study of soil
Radix pollination biology and mating systemsJ J Beijing For
Univ395 669673.
2003. 土麦
395) :669-673.
上接第60 Continue from page 60
WEBER A1976. Beitrge zur morphologie und systematik der Klu-
gieae und Loxonieae Gesneriaceae. IIIWhytockia als mor-
phologische und phylogenetische Ausgangsform von Monophyl-
laeaJ Beitr Biol Pflanzen52183-205.
WEBER A1982. Contributions to the morphology and systematics
of Klugieae and Loxonieae Gesneriaceae IX. The genus
WhytockiaJ Notes Roy Bot Gard Edinburgh40113-121.
WEBER ABURTT BL1998. Remodelling of Didymocarpus and
associated genera Gesneriaceae) [M Beitr Biol Pflanzen
JW eds The families and genera of vascular plants. Vol. 7.
Flowering plants. Dicotyledons. Lamiales except Acanthaceae
including Avicenniaceae. Berlin/HeidelbergSpringer63-158.
WEBER AMiddleton DJForrest Aet al2011a. Molecular sys-
tematics and remodelling of Chirita and associated genera
Gesneriaceae) [J. Taxon60767-790.
WEBER AWEI YGPUGLISI Cet al2011b. A new definition of
the genus Petrocodon Gesneriaceae) [J. Phytotaxa2349-67.
WEBER AWEI YGSONTAG Set al2011c. Inclusion of
Metabriggsia into Hemiboea Gesneriaceae) [J. Phytotaxa23
WEBER ACLARK JLMLLER M2013. A new formal classifi-
cation of GesneriaceaeJ. Selbyana31268-94.
WEI YG2004. Paralagarosolen Y. G. Weia new genus of the
Gesneriaceae from GuangxiChinaJ Acta Phytotax Sin42
WEI YGWEN FCHEN WHet al2010. Litostigmaa new genus
from Chinaa morphological link between basal and derived Didy-
mocarpoid GesneriaceaeJ Edinburgh J Bot671) : 161-184.
WEN FLIANG GYWEI YG2012. Petrocodon lancifolius
Gesneriaceae) ,a new species endemic to a central subtropical
zone of Guizhou ProvinceChinaJ. Phytotaxa4945-49.
WEN FMACIEJEWSKI SHE XQet al2015a. Briggsia leio-
phyllaa new species of Gesneriaceae from southern Guizhou
ChinaJ. Phytotaxa202 1) : 51-56.
WENFWEI YGMLLER M2015b. Glabrella leiophylla
Gesneriaceae) ,a new combination for a former Briggsia spe-
cies from GuizhouChinaJ. Phytotaxa218 2) : 193-194.
Wood D1974. A revision of Chirita Gesneriaceae) [J. Notes
Roy Bot Gard Edinburgh33123-205.
XU WBMENG TZHANG Qet al2014. Petrocodon Gesneri-
aceaein the limestone karsts of GuangxiChinaThree new
species and a new combination based on morphological and mo-
lecular evidenceJ Syst Bot393) : 965-974.
XU WBZHANG QWEN Fet al2012. Nine new combinations
and one new name of Primulina Gesneriaceaefrom South Chi-
naJ. Phytotaxa641 -8
XU ZRBURTT BLSKOG LEet al2008. A revision of Para-
boea Gesneriaceae) [J Edinburgh J Bot652) : 161-347.
YANG CZCAI DLWEN F2015. Oreocharis striata Gesneri-
aceae) ,a new species from FujianChinaJ Ann Bot Fenn
YAO TL2012. A taxonomic revision of Loxocarpus Gesneriaceae
D MSc thesisFaculty of ScienceUniversiti MalayaKuala
YUXLLIMZHOU JJet al2015. Petrocodon hunanensis
Gesneriaceae) ,a new species identified by both morphological
and molecular evidence from limestone area in HunanChina
J. Phytotaxa1951) : 65-72.
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
... The genus Didymocarpus comprises over 100 species, widely distributed from the Himalayas to the Malay Peninsula (Nong et al., 2021;Souvannakhoummane et al., 2019). China and India are the highest centers of diversity of this genus, with 34 species and five varieties, 23 species and two varieties, respectively (Michael et al., 2016;Möller et al., 2017). Didymocarpus species are lithophytic perennials characterized by ovate to ovate-cordate lamina, predominantly glandular and hairy, oblique limbs on tube-like corollas, a couple of fertile staminate having three staminodes, a capitate stigma, an orthocarpic ovary and a bivalve capsule which dehisces in a loculicidal direction (Weber et al., 2000). ...
Ethnopharmacological relevance Medicinal plants have been extensively used to treat various illnesses since the dawn of civilization. The genus Didymorcapus Wall. comprises 100 species widely distributed in the tropical regions of Asia, with a few found scattered in Africa and Australia. Species in this genus have long been used in folk medicine to treat various illnesses, including wounds, kidney stones, inflammations, asthma, flu, eczema, dysentery, fractures, colic etc. Some species have applications as weight loss agents, laxatives, and protective medication after childbirth. Aim To provide comprehensive information on the current knowledge of the ethnobotanical uses, phytochemical compounds, pharmacological applications, and toxicology of genus Didymocarpus to reveal its therapeutic potential, offering insights into future research opportunities. Materials and methods Data were systematically obtained from books and online databases such as PubMed, Web of Science, Scopus, Sci Finder, Google Scholar, Science direct, ACS Publications, Elsevier, Wiley Online Library. Results Seventeen Didymocarpus species have applications in traditional medicine in different Asian countries. A total of 166 compounds have been isolated from the genus Didymocarpus including terpenoids, flavonoids, phenolic compounds, fatty acids, chalcones, steroids, and others. Among these constituents, terpenoids, flavonoids, chalcones, and phenolics are the significant contributors to pharmacological activities of the genus Didymocarpus, possessing wide-reaching biological activities both in vivo and in vitro. The crude extracts and isolated phytochemical compounds from this genus have been shown to exhibit various pharmacological activities, including antiurolithiatic, nephro-protective, antimicrobial, anticancer, antidiabetic, cytotoxic, wound healing, and antioxidant activities. Conclusions Traditional uses and scientific evaluation of Didymocarpus indicate that Didymocarpus pedicellata is one of the most widely used species in some parts of the world. Although substantial progress on the chemical and pharmacological properties of Didymocarpus species has been made, further studies on the pharmacology and toxicology of these species are needed to ensure safety, efficacy, and quality. Also, further research on the structure-activity relationship of some of the isolated phytocompounds may improve their biological potency and scientific exploitation of traditional uses of the Didymocarpus taxa.
... Petrocodon Hance (Gesneriaceae), is a small genus growing in mountainous karst habitats of S China, N Vietnam and NE Thailand (Hance 1883;Wang et al. 1998;Li and Wang 2004;Wei et al. 2010). Based on the latest molecular phylogenetic analyses, Petrocodon has been expanded to now include more than 30 species (Wang et al. 2011;Weber et al. 2011;M€ oller et al. 2016;IPNI 2019), and include all species previously referred to the genera Calcareoboea C. Y. Wu ex H. W. The DNA sequence of the chloroplast genome can be used as a super barcode or a resource for research in phylogeograhy, genetic diversity and evolution. For Petrocodon, however, no complete chloroplast sequence has been published to date. ...
Full-text available
The complete chloroplast genome of Petrocodon jingxiensis (Yan Liu, H.S. Gao, and W.B. Xu) A. Weber and Mich. Möller was determined in this study. The cp genome was 153,056 bp in length including two inverted repeats (IRs) of 25,405 bp, which were separated by LSC and SSC of 84,154 bp and 18,092 bp, respectively. The GC content was 41.9%. The genome encoded 112 functional genes, including 79 protein-coding genes, 29 tRNA genes, and four rRNA genes. This plastid genome is the first report for the Petrocodon in Gesneriaceae which will be useful data for developing markers for further studies on resolving the relationship within the genus.
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Gesneriaceae is a pantropical family of plants that, thanks to their lithophytic and epiphytic growth forms, have developed different strategies for overcoming water scarcity. Desiccation tolerance or “resurrection” ability is one of them: a rare phenomenon among angiosperms that involves surviving with very little relative water content in their tissues until water is again available. Physiological responses of desiccation tolerance are also activated during freezing temperatures, a stress that many of the resurrection gesneriads suffer due to their mountainous habitat. Therefore, research on desiccation- and freezing-tolerant gesneriads is a great opportunity for crop improvement, and some of them have become reference resurrection angiosperms (Dorcoceras hygrometrica, Haberlea rhodopensis and Ramonda myconi). However, their difficult indoor cultivation and outdoor accessibility are major obstacles for their study. Therefore, this review aims to identify phylogenetic, geoclimatic, habitat, and morphological features in order to propose new tentative resurrection gesneriads as a way of making them more reachable to the scientific community. Additionally, shared and species-specific physiological responses to desiccation and freezing stress have been gathered as a stress response metabolic basis of the family.
Remediation using plant-microbe interactions is an efficient and cost-effective technique. However, few studies have reported their effects in phosphate mining wastelands. In this study, two native phosphate-solubilizing microorganisms (PSM) (Trichoderma asperellum LZ1 and Serratia sp. LX2) and two native plants (Lolium perenne L. and Lactuca versicolor) were used to remediate excess phosphorus (P) contamination in phosphate mining wasteland soil. In pot experiments, PSM was found to increase P availability by reducing soil pH. The inoculation of strain LZ1 and LX2 significantly increased the total biomass of L. perenne L. by 41.73%, 6.27%, and 49.26%, respectively. Similarly, L. versicolor biomass also increased by 17.51%, 11.67%, and 41.16%, respectively. In addition, the root, shoot length, and chlorophyll content of L. perenne L. and L. versicolor were also significantly higher than those of the treatments inoculated without PSM. In PSM-inoculated soils, the P accumulation in L. perenne L. and L. versicolor was 11.71 and 5.54 mg/pot, respectively. The findings in this study showed that T. asperellum LZ1 and Serratia sp. LX2 could effectively promote the growth of L. perenne L. and L. versicolor, and increase the P accumulation in plants, which would be beneficial to the phytoremediation of excess P contamination in phosphate mining wasteland.
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A new species of Oreocharis Benth., O. rubrostriata F.Wen & L.E.Yang, is described and illustrated from Rongshui County, Guangxi Zhuangzu Autonomous Region, China, and a proposed conservation assessment rates the new species as Critically Endangered (CR).
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