全 文 ：单种属弥勒苣苔属系统位置研究: 基于分子和细胞学数据*
谭摇 英1,2, 王摇 智1,3, 隋学艺1,4, 胡光万1,
MOTLEY Timothy5,6, 龙春林1,6**
(1 中国科学院昆明植物研究所, 云南 昆明摇 650201; 2 中国科学院研究生院, 北京摇 100049; 3 西南林业大学林学院,
云南 昆明摇 650224; 4 云南农业大学农学院, 云南 昆明摇 650201; 5 奥多明尼奥大学生物学部,
美国 维吉尼亚州 诺福克摇 23529鄄0266; 6 中央民族大学生命环境科学学院, 北京摇 100081)
摘要: 弥勒苣苔属是苦苣苔科的单种属, 仅分布于中国西南部。 为探讨弥勒苣苔在苦苣苔亚科中的系统位
置, 我们选择了苦苣苔亚科 116 个类群, 外类群为苦苣苔亚科以外的 7 个物种。 用最大简约法 (MP) 和
贝叶斯分析 (BI), 对以上类群的核基因 ITS 以及两个叶绿体基因 trnL鄄F、 atpB鄄rbcL 数据进行了独立和联
合分析。 在三个片段联合分析的结果中, 弥勒苣苔与马铃苣苔属、 后蕊苣苔属、 金盏苣苔属、 直瓣苣苔属
以及川鄂粗筒苣苔构成一个强烈支持的分枝。 MP树中, 此分枝为并系, 而在 BI分析中, 弥勒苣苔与川鄂
粗筒苣苔、 直瓣苣苔属互为姐妹类群。 同时, 第一次报道了弥勒苣苔的染色体数目 (2n = 34)。 根据前人
报道, 马铃苣苔属、 后蕊苣苔属、 粗筒苣苔属和直瓣苣苔属的染色体数目同为 2n=34, 这进一步支持我们
的分子系统发育分析。
关键词: atpB鄄rbcL; 苦苣苔科; ITS; 弥勒苣苔属; 系统学; trnL鄄F
中图分类号: Q 942, Q 949摇 摇 摇 摇 摇 文献标识码: A摇 摇 摇 摇 摇 文章编号: 2095-0845(2011)05-465-12
The Systematic Placement of the Monotypic Genus Paraisometrum
(Gesneriaceae) Based on Molecular and Cytological Data*
TAN Ying1,2, WANG Zhi1,3, SUI Xue鄄Yi1,4, HU Guang鄄Wan1,
MOTLEY Timothy5,6, LONG Chun鄄Lin1,6**
(1 Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650201, China; 2 Graduate University of Chinese
Academy of Sciences, Beijing 100049, China; 3 School of Forestry, Southwest Forestry University, Kunming
650224, China; 4 College of Agronomy, Yunnan Agricultural University, Kunming 650201, China;
5 Department of Biological Sciences, Old Dominion University, Norfolk, Virginia 23529鄄0266, USA;
6 College of Life and Environmental Sciences, Minzu University of China, Beijing 100081, China)
Abstract: Paraisometrum, is a monotypic genus endemic to Southwest China, with an uncertain systematic position.
To resolve the phylogenetic relationships of Paraisometrum within subfamily Cyrtandroideae, we performed phyloge鄄
netic analyses of 116 taxa placed in the subfamily Cyrtandroideae and 7 outgroup species selected from other Gesneri鄄
aceae subfamilies. The independent and combined analyses were performed on three DNA datasets: the nuclear in鄄
ternal transcribed spacer regions ( ITS) and two chloroplast regions ( trnL鄄F, atpB鄄rbcL). Maximum Parsimony
(MP) and Bayesian Inference (BI) were used for tree construction. The combined three gene dataset strongly sup鄄
植 物 分 类 与 资 源 学 报摇 2011, 33 (5): 465 ~ 476
Plant Diversity and Resources摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 DOI: 10. 3724 / SP. J. 1143. 2011. 11075
*
**
Foundation items: The Chinese Academy of Sciences (KSCX2鄄YW鄄Z鄄0925 & O92441112F), the Ministry of Education of China through
its 985 and 111 projects (MUC 98503鄄001006 & B08044), the National Science Foundation of China (31070288), the Japan
Society for the Promotion of Science ( JSPS / AP / 109080 ), and the Ministry of Science and Technology of China
(2008FY110400鄄2鄄2)
Author for correspondence; E鄄mail: long@ mail. kib. ac. cn
Received date: 2011-05-10, Accepted date: 2011-06-16
作者简介: 谭英 (1986-) 女, 在读硕士研究生, 主要从事保护生物学研究。 E鄄mail: tanying@ mail. kib. ac. cn
port Paraisometrum in a clade containing Oreocharis, Opithandra, Isometrum, Ancylostemon, and Briggsia ros鄄
thomii. In the MP tree this lineage is an unresolved polytomy, but BI resolves Paraisometrum as sister to B. rosthomii
and the Ancylostemon clade. We did the first chromosome counts of Paraisometrum (2n=34). The diploid number
34 has also been reported for Oreocharis, Opithandra, Briggsia and Ancylostemon, which lends further support to the
placement of Paraisometrum in this lineage.
Key words: atpB鄄rbcL; Gesneriaceae; ITS; Paraisometrum; Phylogeny; trnL鄄F
摇 Paraisometrum W. T. Wang is a monotypic ge鄄
nus, containing P. mileense W. T. Wang, which is
endemic to Southwest China ( Weitzman et al.,
1997) in the Yunnan Province ( Shui, 2007a, b)
and Guangxi Region (Xu et al., 2009). The holo鄄
type of this plant was collected by French missionary
Ducloux F. in Mi鄄le County, Yunnan Province in
1906 and remained unnamed and stored in Paris for
nearly 100 years. In 1997, when Weitzman et al.
wrote an article on Gesneriaceae in the corpus for
Flora of China, Prof. W. T. Wang checked Du鄄
cloux忆 s specimens in Paris collected from Mi鄄le
County and concluded that it represented a new ge鄄
nus in Gesneriaceae and named it Paraisometrum
mileense (Weitzman et al., 1997). It was believed
extinct until 2006. In 2006, the species was redis鄄
covered in the Shilin County, Yunnan Province by Dr.
Yuming Shui, a botanist from the Kunming Institute
of Botany, Chinese Academy of Sciences ( Shui,
2007a, b). According to the IUCN Species Pro鄄
gramme, Paraisometrum mileense can be considered
an endangered species because of its narrow geo鄄
graphic range and small population size.
The Chinese Gesneriaceae, based on morpho鄄
logical characteristics, have been divided into six
tribes: Ramondeae, Didymocarpeae, Trichosporeae,
Cyrtandreae, Epithemateae and Titanotricheae (Wang
et al., 1992). Wang et al. (1998) placed P. mile鄄
ense in the Didymocarpeae, and stated that it is most
similar to the genus Isometrum Craib because they
both have four fertile stamens and a funnelform
tube, longer than the lobes. More recently, molecu鄄
lar analyses of subfamily Didymocarpoideae have
been conducted by Mayer et al. (2003) and M觟ller
et al. (2009) based on chloroplast DNA (cpDNA)
or combined ITS and cpDNA sequences, respective鄄
ly. Neither study included samples of P. mileense,
which left its phylogenetic position unresolved. The
rediscovery of a population of P. mileense have made
it possible determine the phylogenetic position of this
species to test the morphological hypothsis of Wang
et al. (1998).
To investigate the systematic position of Parai鄄
sometrum, we used DNA sequences for the nuclear
internal transcribed spacer regions ( ITS), cpDNA
trnL鄄F intron鄄spacer region ( trnL鄄F) and atpB鄄rbcL
spacer ( atpB鄄rbcL ). In addition, we report the
chromosome numbers of Paraisometrum for the first
time and discuss characters among closely related
genera.
1摇 Materials and methods
1. 1摇 Taxon sampling
To produce a reliable phylogeny of the subfami鄄
ly Cyrtandroideae the ingroup comprised 116 taxa,
representing 115 species in 46 genera and included
4 members of Gesnerioid and 3 members of Coronan鄄
theroid as outgroups ( Appendix). These outgroup
taxa were chosen based on results of recent studies of
Gesneriaceae (Zimmer et al., 2002; Perret et al.,
2003; M觟ller et al., 2009). Sequences used in this
study were from GenBank and supplemented with 17
new sequences (5 individuals of trnL鄄F, 6 atpB鄄
rbcL, 6 ITS). For the ingroup we followed the taxo鄄
nomic classification of Weber (2004). 13 species
(5 genera) belonged to Epithematoid Gesneriaceae
and 103 species (42 genera) came from Didymo鄄
carpoid Gesneriaceae (Appendix).
1. 2摇 Molecular methods
Materials for DNA extraction were from various
664摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 植 物 分 类 与 资 源 学 报摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 第 33 卷
forms and diverse sources, including fresh leaves
from research collections, silica鄄dried leaves from
our field collections. Genomic DNA was isolated
with a modified CTAB protocol (Doyle and Doyle,
1987).
Amplifications of ITS were carried out following
the PCR profile described by M觟ller and Cronk
(1997), using forward primer ‘ ITS 5P爷 and re鄄
verse primer ‘ ITS 8P爷. The trnL鄄F intron鄄spacer
region ( trnL鄄F) and atpB鄄rbcL spacer (atpB鄄rbcL)
were amplified following Mayer et al. (2003) using
primers ‘ c爷 and ‘ f爷 for trnL鄄F ( Taberlet et al.,
1991) and primers ‘JF31爷 and ‘JF5爷 for atpB鄄rbcL
(Samuel et al., 1997). PCR was performed using a
DNAEngine襆 peltier thermal cycler ( Bio鄄RAD,
Inc. ) . The PCR products were purified using a mul鄄
tifunction DNA purification KIT ( BioTeke Inc. ).
The purified PCR products were cycle鄄sequenced
using an ABI Prism BigDye Terminator Cycle Kit fol鄄
lowing standard kit protocols (Perkin鄄Elmer Applied
Biosystems) and run on a ABI鄄PRISM3730 se鄄
quencher. Forward and reverse sequences were as鄄
sembled using SeqMan of DNAStar ( DNASTAR,
1999), visually aligned with ClustalX v. 1. 83
(Thompson et al., 1998) followed by manual cor鄄
rection in BioEdit (Hall, 1999). Sequence align鄄
ments and PAUP / Nexus鄄formatted files for individu鄄
al regions and combined analyses are available from
the authors upon request.
1. 3摇 Phylogenetic analysis
Phylogenetic analyses were performed using Maxi鄄
mum Parsimony (MP) and Bayesian Inference (BI) u鄄
sing the protocols of Rannala and Yang (1996) on the
individual ITS, atpB鄄rbcL, trnL鄄F, combined cpDNA
and combined three gene datasets respectively. Maxi鄄
mum Parsimony analysis was performed with PAUP*
version 4.0b10 (Swofford, 2002). Gaps were trea鄄
ted as missing data, and characters were assumed to
be unordered. Heuristic searches were performed
employing 1 000 random replicates, holding one tree
at each step during stepwise addition, using tree鄄bi鄄
section鄄reconnection (TBR) branch鄄swapping algo鄄
rithm, MulTrees in effect, and steepest descent off.
Support for the nodes resolved in the strict consensus
of the MP trees was calculated by bootstrap analyses
( Felsenstein, 1985 ) with 1 000 heuristic鄄search
replicates as described above. Bayesian analysis was
performed using MrBayes v. 3. 1. 2 ( Huelsenbeck
and Ronquist, 2003) for all the datasets. The most
appropriate molecular model for each dataset was de鄄
termined with jModelTest 0. 1. 1 (Guindon and Gas鄄
cuel, 2003; Posada, 2008). The model GTR+I+G
was selected. The Markov chain Monte Carlo (MC鄄
MC) algorithm was run for 10 000 000 generations
with one cold and three heated chains in two inde鄄
pendent parallel analyses. Trees were sampled every
100 generations from the chain. The first 20% trees
were discarded as burn鄄in (generations prior to sta鄄
tionary of likelihood values).
1. 4摇 Cytological protocols
Root tips of Paraisometrum mileense were pre鄄
treated in 0. 002 mol·L-1 8鄄hydroxyquinoline for 4 to
5 h at room temperature, and then fixed in Carnoy忆s
I (3 ethanol 颐 1 glacial acetic acid) for 30 min at
5益 . After hydrolysis for 1 min in 1 mol·L-1 HCl at
60益, followed by washing through several changes
of distilled water, the root tips were dyed in carbol
fuchsin and then squashed. The chromosomes were
examined using Axioskop 40 made by ZEISS and
photographed using JVC TK鄄C1480BEC color video
camera.
2摇 Results
2. 1摇 Phylogenetic analysis
The main clades found in the parsimony analy鄄
sies of the individual ITS, trnL鄄F, atpB鄄rbcL and
combined cpDNA datasets (not shown) were broadly
congruent with, although less well resolved than,
the consensus trees from the combined three gene
dataset. This reflects the fewer of informative cha鄄
racters in the separate analyses. The results of the
combined three gene analyses are presented here.
Of the aligned 2990 characters (ITS: 791 char鄄
acters, trnL鄄F: 1189 characters, atpB鄄rbcL: 1010
7645 期摇 摇 摇 TAN Ying et al. : The Systematic Placement of the Monotypic Genus Paraisometrum (Gesneriaceae) …摇 摇 摇 摇
characters) analyzed, 1100 were constant, 549 un鄄
informative, and 1341 parsimony informative. Maxi鄄
mum parsimony analysis resulted in 4 most parsimo鄄
nious trees of 7058 steps, with a CI of 0. 44, a RI of
0. 68, and a RC of 0. 30. A strict consensus tree of
the combined dataset resolved 92 nodes with boot鄄
strap support (BS) 逸50% (Fig. 1). The phylo鄄
gram from the Bayesian analysis of the combined
dataset resolved 113 nodes, 95 of which had posteri鄄
or probabilities (PP) 逸0. 90 (Fig. 2). Support for
deep nodes was weak, but the support for the indi鄄
vidual major clades was strong.
The MP strict consensus tree had a similar tree
topology to that of Bayesian consensus tree. Subfam鄄
ily Cyrtandroideae is monophyletic and sister to sub鄄
family Gesnerioideae ( BS = 97% ; PP = 100% ).
The tribe Epithemateae is sister to the remaining old
world Gesneriaceae with strong support ( BS =
100% ; PP = 100% ). Genus Corallodiscus form a
clade (BS= 87% ; PP = 100% ) and is sister to the
remaining taxa (BS=87% ; PP= 100% ). Basal A鄄
siatic genera, Rhynchotechum, Platystemma, and
European genera, Jancaea, Ramonda, Haberlea,
constitute a clade (BS = 93% ), which is separated
from the African and remaining Asiatic genera with
moderate support (BS=87% ; PP=100% ). Among
the remaining taxa, Briggisa, Chirita, and Strepto鄄
carpus are not monophyletic. Paraisometrum is in a
well supported clade ( BS = 100% , PP = 100% )
with Briggsia rosthornii, Isometrum lungshengense,
Opithandra primuloides, Oreocharis auricula and
three species of Ancylostemon. The three species of
Ancylostemon form a monophyletic group with strong
support ( BS = 100% ; PP = 100% ). The BI tree
provides more resolution in the Paraisometrum
clade, Paraisometrum is sister to Ancylostemon and
Briggsia rosthornii (PP = 99% ) in a polytomy with
the other three genera of the lineagewhile the MP
tree does not resolve any of the relationships among
the genera in the clade.
2. 2摇 Cytology
Paraisometrum mileense has a diploid number of
2n=34, and the chromosome size is small, <2 滋m
(Fig. 3).
3摇 Discussion
3. 1摇 Phylogenetic relationships
In our study, Old World Gesneriaceae can be
clearly divided into two tribal groups that are congru鄄
ent with previous studies (Smith et al., 1997; Wang
and Li, 2002; Mayer et al., 2003; M觟ller et al.,
2009; Wang et al., 2010). Weber (2004) separa鄄
ted subfamily Cyrtandroideae into the small tribe Ep鄄
ithemateae ( Epithematoid Gesneriaceae ) and the
large tribe Didymocarpeae in which the members of
Cyrtandreae and Trichosporeae were polyphyletically
nested at various places ( Didymocarpoid Gesneri鄄
aceae). On the basis of morphological and geo鄄
graphical features, the Didymocarpoid Gesneriaceae
can be divided into following groups: Basal Asiatic
and European genera, African genera, Advanced A鄄
siatic genera (including genera with twisted capsular
fruits, straight capsular fruits, or indehiscent fruits).
The Asiatic and European, and African groups are
well defined and supported according to the phyloge鄄
netic result made by M觟ller et al. (2009). In their
study, the genus Briggsia is also not monophyletic,
and Oreocharis, one clade of Briggsia and Ancyloste鄄
mon compose a clade. Our results show that Oreo鄄
charis, Opithandra, Isometrum, Ancylostemon,
Briggsia rosthornii and Paraisometrum constitute a
clade in the Advanced Asiatic group, and suggest
Paraisometrum may be mostly related to Ancyloste鄄
mon and Briggesia rosthornii than other genera.
3. 2摇 Morphology
The 5 genera in the clade with Paraisometrum
have some shared floral characters, supporting their re鄄
lationship. These genera have campanulate calyx, ad
basin 5鄄sectus, bilabiate corollas, corolla lobes much
shorter than the corolla tube, and 4 fertile stamens,
stigma number varies, Isometrum, Briggsia, and Ancy鄄
lostemon have 2 stigmas, whereas Opithandra and Ore鄄
charis can have either 1 or 2 stigmas, and Paraisome鄄
trum always has a single stigma (Wang et al., 1998).
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Fig. 1摇 Strict consensus tree of 4 most parsimonious trees based on the combined ITS, trnL鄄F and atpB鄄rbcL data.
Numbers above branches are bootstrap support values 逸50% . The tribes are labeled with bars
9645 期摇 摇 摇 TAN Ying et al. : The Systematic Placement of the Monotypic Genus Paraisometrum (Gesneriaceae) …摇 摇 摇 摇
Fig. 2摇 Bayesian phylogram based on the combined ITS, trnL鄄F and atpB鄄rbcL data. Numbers above branches are posterior
probability values (only values 逸0. 90 are shown). The tribes are labeled with bars
074摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 植 物 分 类 与 资 源 学 报摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 第 33 卷
Fig. 3摇 Karyomorphology of Paraisometrum mileense. A. Interphase nucleus (伊1000); B. Mitotic prophase (伊1000);
C. Mitotic metaphase (伊1000); D. Mitotic anaphase (伊1000). The bar is 1 滋m long
Burtt (1962) and Pan (1986) held the opinion that
based on their observations that Isometrum Craib and
Ancylostemon Craib were closely related, which our
results support. Wang et al. (1998) hypothesized
that Paraisometrum was sister to Isometrum. Our
analysis confirmed that these two genera are closely
related in the same lineage, but Paraisometrum like鄄
ly has closer affinities to Ancylostemon and Briggsia
rosthornii. Different from other 3 genera, their aba鄄
xial lip is longer than adaxial lip.
3. 3摇 Distribution
Briggsia rosthornii is distributed in Sichuan,
Guizhou, and Hubei provinces of Southwest and
South鄄Central China, Opithandra is scattered throug鄄
hout Western China, except for O. primuloides, a
species restricted to Japan. The distributions of Ore鄄
ocharis and Ancylostemon range from Southwestern to
Central and Southeastern China (Li, 1996; Wang et
al., 1998; Li and Wang, 2004). Isometrum is en鄄
demic to China, and centered in Sichuan, where
82% of the species in the genus occur (Pan, 1986).
3. 4摇 Cytology
Chromosome ploidy levels and base numbers
can provide insight into plant evolution, because
sister groups often share chromosome numbers or
base number (Turner et al., 1961; Ehrendorfer et
al., 1968; Walker, 1972; Raven, 1975; Grant,
1982). However, chromosome counts at the spe鄄
cies鄄level across the family have been made for only
about 18% of the taxa in the Gesneriaceae, and the
number of genera without chromosome counts is even
greater in the subfamily Cyrtandroideae than subfami鄄
ly Gesnerioideae (M觟ller and Kiehn, 2004). In the
Epithemateae, the chromosome counts have high
variation, with the basic numbers in this group ran鄄
ging from n = 8 -12, (Ratter, 1975; Skog, 1984;
Wang et al., 1998). It is more complicated in the
Didymocarpoid Gesneriaceae, where basic numbers
are n=4, 8, 9, 10, 11, 12, 13, 14, 15, 16, 17,
18 (Ratter and Prentice, 1967; Ratter, 1975; Kie鄄
hn and Weber, 1998; Rashid et al., 2001; M觟ller
and Kiehn, 2004). In previous studies, the genera
1745 期摇 摇 摇 TAN Ying et al. : The Systematic Placement of the Monotypic Genus Paraisometrum (Gesneriaceae) …摇 摇 摇 摇
Oreocharis, Opithandra, and Ancylostemon have the
same chromosome number 2n= 34, Briggsia has the
chromosome number 2n = 34 or 2n = 68 ( Ratter,
1963; Ratter and Prentice, 1964; Wang and Gu,
1999; Zhou et al., 2004). Paraisometrum also has
a chromosome complement of 2n = 34, which is an鄄
other character uniting members of this clade. The
chromosome is too small to do karyotype analysis, or
we can do further comparison of these species. Al鄄
though the data are not yet available, we predict that
Isometrum to have a karyotype of 2n=34.
Our molecular analyses place the rare monotypic
genus, Paraisometrum, among the Didymocarpoid
genera, subfamily Cyrtandroideae of Gesneriaceae.
Paraisometrum belongs in a strongly supported clade
with Oreocharis, Opithandra, Isometrum, Ancyloste鄄
mon, and Briggsia rosthornii among and nested in an
informal group the Advanced Asiatic group. Bayesi鄄
an analyses support Paraisometrum, as sister to An鄄
cylostemon and Briggsia rosthornii. Because Briggsia
rosthornii is a single species (and not the type) in a
polyphyletic genus further study and taxonomic revi鄄
sion will be required on this clade. So at this time
the decision on whether Briggsia rosthornii should be
considered another monotypic genus or whether it
and / or Paraisometrum mileense should be combined
into an existing genus remain unanswered.
Acknowledgement: We thank Prof. Lei Shi from the Botan鄄
ical Garden of Institute of Botany, CAS, for providing some
plant materials, Mr. Yu Guo for laboratory assistance, and
Dr. Yanchun Li, Dr. Yuxiao Zhang, Dr. Wenbin Yu, Dr.
Bo Long and Dr. Jiahui Chen for advice on data analysis.
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Appendix: GenBank accession numbers for the sequences used in the phylogenetic analysis
Taxon Locality Voucher
GenBank accession No.
atpB鄄rbcL trnL鄄F ITS1 / ITS2
Outgroup taxon
Gesnerioid
Achimenes admirabilis Wiehler AJ439982 AJ439827 AY182173
Sinningia cardinalis (Lehm. ) H. E. Moore AJ490931 AJ492318 —
S. schiffneri Fritsch AJ439900 AJ439745 —
Gesneria humilis L. AJ439976 AJ439821 AY047051
Coronantheroid
Asteranthera ovata (Cav. ) Hanst FJ501371 FJ501427 EF445669
Fidldia australis Cunn. AY423112 AY423130 EF445687
Rhabdothamnux solandri Cunn. FJ501370 FJ501426 EF445700
Ingroup taxon
Didymocarpoid
Acanthonema strigosum Hook. f. — FJ501454 FJ501306
Aeschynanthus austroyunnanensis W. T. Wang FJ501396 FJ501500 AF349218 / AF349299
A. bracteatus Wall. ex DC. — FJ501501 AF349203 / AF349284
A. hildebrandtii Hemsl. — AY047099 AY047040
A. longiflorus (Blume) DC. AJ490920 AJ492307 FJ501333
Agalmyla biflora (Elmer) O.M.Hilliard & B.L.Burtt FJ501421 FJ501541 FJ501361
A. parasitica (Lam. ) Kuntze FJ501420 FJ501539 —
Ancylostemon aureus (Franch. ) B. L. Burtt FJ501398 FJ501505 FJ501336
A. convexus Craib — FJ501506 FJ501337
Anna mollifolia (W.T.Wang) W.T.Wang & K.Y.Pan — FJ501543 AF055050 / AF055051
A. submontana Pellegr. FJ501422 FJ501542 FJ501362
Boea hygrometrica (Bunge) R. Br. — FJ501476 FJ501319
B. magellanica Lam. — FJ501478 FJ501321
Briggsia longipes (Hemsl. Ex Oliv. ) Craib FJ501423 FJ501545 AF055052 / AF055053
B. mihieri Craib — FJ501544 FJ501363
B. rosthornii (Diels) B. L. Burtt FJ501425 FJ501547 FJ501365
Calcareoboea coccinea C. Y. Wu ex H. W. Li FJ501406 FJ501516 FJ501365
Chirita asperifolia (Blume) B. L. Burtt FJ501419 FJ501538 FJ501359
C. caliginosa C. B. Clarke FJ501391 FJ501488 FJ501325
C. gemella D. Wood FJ501408 FJ501523 FJ501345
C. hamosa R. Br. FJ501392 FJ501489 —
C. lacunosa (Hook f. ) B. L. Burtt FJ501384 FJ501458 FJ501308
C. lavandulacea Stapf. FJ501390 FJ501497 FJ501324
C. longgangensis W. T. Wang AJ490903 AJ492290 FJ501347
C. pinnata W. T. Wang — FJ501526 FJ501349
C. pinnatifida (Hand. 鄄Mazz. ) B. L. Burtt — FJ501527 FJ501350
C. pumila D. Don FJ501393 FJ501491 FJ501327
*Chirita sp.
China, Yunnan Prov.,
Jiaozixueshan Mountain
HGWB鄄754
(KUN) HQ327455 HQ327447 HQ327462
C. sinensis Lindl. FJ501409 FJ501524 FJ501348
C. spadiciformis W. T. Wang AJ490904 AJ492291 FJ501346
C. urticifolia Buch. 鄄Ham. Ex D. Don — FJ501492 FJ501328
C. walkei Gardner — FJ501490 FJ501326
Colpogyne betsiliensis B. L. Burtt — FJ501445 FJ501302
Conandron ramondioides Sieb. & Zucc. FJ501405 FJ501515 FJ501340
Corallodiscus conchifolia Batalin FJ501375 FJ501433 —
C. lanuginosus (Wall. Ex R. Br) B. L. Burtt FJ501374 FJ501432 —
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Continue
Taxon Locality Voucher
GenBank accession No.
atpB鄄rbcL trnL鄄F ITS1 / ITS2
C. lanuginosus (Wall. Ex R. Br. ) Burtt China, Yunnan,Nujiang
WZ2009CM03
(KUN) HQ327458 HQ327450 HQ327465
Cyrtandra cupulata Ridl FJ50141 FJ501532 AY818826 / AY818861
C. glabra Banks ex Gaertn. AY423119 AY423136 FJ501353
C. longifolia (Wawra) Hillebr. Ex C. B. Clarke FJ501413 FJ501531 AY818846 / AY818881
C. pendula Blume FJ501412 FJ501530 FJ501354
C. platyphylla A. Gray FJ501410 FJ591528 —
C. sessilis H. St. JohnKapua et al. s. n. FJ501411 FJ501529 —
Didissandra frutescens (Jack) C. B. Clarke U91313 FJ501521 —
Didymocarpus aromaticus Wall. ex D. Don FJ501402 FJ501511 —
D. citrinus Ridul. AJ490906 AJ492293 DQ912669
D. cordatus Wall. ex DC. — AJ492294 DQ912673
D. podocarpus C. B. Clarke FJ501404 FJ501514 DQ912688
D. purpureobraceatus W. W. Sm FJ501401 FJ501510 —
D. stenanthos C. B. Clarke FJ501403 FJ501512 DQ912687
Emarhendia bettiana (M. R. Hend) Kiew AJ490908 AJ492295 —
Haberlea rhodopensis Friv. AJ490909 AJ492296 —
Hemiboea bicornuta (Hayata) Ohwi FJ501416 FJ501534 FJ501356
H. cavaleriei H. Lev. FJ501415 FJ501533 FJ501355
*H. gracilis Franch
China, Hunan,
Zhangjiajie
WZ2010ZJJ01
(KUN) HQ327453 HQ327445 HQ327461
H. subcapitata C. B. Clarke FJ501417 FJ501535 FJ501357
Henckelia albomarginata (Hemsl. ) A. Weber AJ490910 AJ492297 —
H. humboldtiana (Gardner) A. Weber & B. L.
Burtt FJ501389 FJ501485 —
Isometrum lungshengense ( W. T. Wang) W. T.
Wang & K. Y. Pan — GU350690 GU350659
Jancaea heldreichii Boiss. FJ501378 FJ501439 —
Kaisupeea herbacea (C. B. Clarke) B. L. Burtt FJ501385 FJ501459 FJ501309
Loxostigma fimbrisepalum K. Y. Pan FJ501399 FJ501507 —
L. griffithii (Wight) C. B. Clarke FJ501400 FJ501508 FJ501338
Lysionotus chingii Chun ex W. T. Wang — FJ501498 FJ501332
L. forrestii W. W. Sm. FJ501394 FJ501495 AF349152 / AF349233
*L. microphyllus var. microphyllus W. T. Wang
China, Hunan,
Zhangjiajie
WZ2010ZJJ03
(KUN) HQ327456 — HQ327463
L. pauciflorus Maxim. FJ501395 FJ501497 FJ501331
Opithandra primuloider (Miq. ) B. L. Burtt FJ501424 FJ501546 FJ501364
Oreocharis auricular (S. Moore) C. B. Clarke — FJ501482 FJ501323
Ornithoboea arachnoidea (Diels) Craib FJ501387 FJ501461 FJ501312
O. wildeana Craib — FJ501462 FJ501313
Paraboea acutifolia (Ridl. ) B. L. Burtt — FJ501464 FJ501314
P. capitata Ridl. var. capitata AJ490911 AJ492298 FJ501315
P. rufescens (Franch. ) B. L. Burtt FJ501388 FJ501469 FJ501316
P. rufescens (Franch. ) B. L. Burtt var. umbellate
(Drake) K. Y. Pan — FJ501470 FJ501317
*Paraisometrum mileense W. T. Wang
China, Yunnan,
Shilin county
TY2009SL01
(KUN) HQ327452 HQ327444 HQ327460
Petrocodon dealbatus Hance FJ501418 FJ501538 FJ501358
Petrocosmea kerrii Craib FJ501397 FJ501502 FJ501334
5745 期摇 摇 摇 TAN Ying et al. : The Systematic Placement of the Monotypic Genus Paraisometrum (Gesneriaceae) …摇 摇 摇 摇
Continue
Taxon Locality Voucher
GenBank accession No.
atpB鄄rbcL trnL鄄F ITS1 / ITS2
P. nervosa Craib AJ4909012 AJ492299 FJ501335
Platystemma violoides Wall. FJ501382 FJ501443 —
Primulina tabacum Hance AJ490913 AJ492300 FJ501352
Ramonda myconi (L. ) Rchb. AJ490914 AJ492301 —
Raphiocarpus begoniifolius (H. Lev. ) B. L. Burtt — FJ501517 FJ501342
R. petelotii (Pellegr. ) B. L. Burtt — FJ501518 FJ501343
Rhabdothamnopsis sinensis Hemsl. AJ490915 AJ492302 FJ501310
Rhynchotechum discolor (Maxim. ) B. L. Burtt FJ501376 FJ501436 —
R. parciflorum Blume FJ501377 FJ501437 —
Saintpaulia tongwensis B. L. Burtt — FJ501446 FJ501303
S. velutina B. L. Burtt AJ490916 AJ492303 FJ501304
Schizoboea kamerunensis K. Fritsch (B. L. Burtt) — FJ501453 FJ501305
Spelaeanthus chinii Kiew — FJ501457 FJ501307
Streptocarpus andohahelensis Humbert — FJ501449 AF316903
S. beampingaratrensis Humbert var.
beampingaratrensis — FJ501448 AF316905
S. dunnii Hook. f. — FJ501456 AF316951
S. hilsenbergii R. Br. — FJ501450 AF316907
S. holstii Engl. AJ490917 AJ492304 AF316917
S. ibityensis Humbert — FJ501455 AF316926
S. orientalis Craib — FJ501444 AF316929
S. papangae Humbert — FJ501444 AF316929
S. rexii Lindl. AJ490918 AJ492305 AF316979
S. saxorum Engl. FJ501383 FJ501447 AF316914
Trisepalum speciosum (Ridl. ) B. L. Burtt AJ490919 AJ492306 —
Epithematoid
Epithema membranaceum (King) Kiew AJ490887 AJ492274 —
E. saxatile Blume Weber & Anthonysamy AJ490888 AJ492275 —
E. taiwanense S. S. Yin AJ490889 AJ492276 —
E. tenue C. B. Clarke AJ490890 AJ492277 —
Monophyllaea elongate B. L. Burtt AJ490892 AJ492279 —
M. glauca C. B. Clarke AJ490893 AJ492280 —
M. hiricalyx Franch. AJ490894 AJ492269 —
M. horfieldii R. Br U91315 AJ492269 —
Rhynchoglossum azureum (Schltdl. ) AJ490895 AJ492282 —
R. notonianum (Wall. ) B. L. Burtt AJ490896 AJ492283 —
R. obliquum Blume AJ490897 AJ492284 —
Stauranthera grandiflora Benth. AJ490900 AJ492287 —
Whytockia sasakii (Hayata) B. L. Burtt AJ490901 AJ492288 —
W. tsiangiana (Hand. 鄄Mazz) A. Weber AJ490902 AJ492289 —
*Sequences are newly submitted to GenBank. Voucher information of sequences obtained from GenBank is not provided here. Dashed (—)
indicate that no sequence was obtained
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