全 文 ：书广 西 植 物 Guihaia Apr． 2015，35(2):143 － 150 http:/ / journal． gxzw． gxib． cn
DOI:10． 11931 /guihaia． gxzw201402007
黎乾坤，张强，邓涛，等． 广西报春苣苔属一新种及其系统位置［J］． 广西植物，2015，35(2):143 －150
Li QK，Zhang Q，Deng T，et al． Primulina bobaiensis，a new species of Gesnericaeae from Guangxi，China and its phylogenetic placement revealed by the chlo-
roplast matK gene［J］． Guihaia，2015，35(2):143 －150
Primulina bobaiensis，a new species of Gesnericaeae
from Guangxi，China and its phylogenetic placement
revealed by the chloroplast matK gene
LI Qian-Kun1，ZHANG Qiang2，DENG Tao2，PAN Bo2，
HUANG Yu-Song2，LI Wen-Lan1，3*
(1． College of Chemistry and Bioengineering，Guilin University of Technology，Guilin 541004，China;2． Guangxi
Institute of Botany，Guangxi Zhuang Autonomous Ｒegion and the Chinese Academy of Sciences，Guilin 541006，
China;3． College of Life Sciences ＆ Technology of Guangxi University，Nanning 530005，China )
Abstract:Primulina(Gesneriaceae)is an enigmatic group of lithophytic plants，which encompass over 160 species mainly
distributed in the limestone region of southern China and nothern Vietnam． A majority of species of the genus have very nar-
row distribution，which have been found in only one or a few proximate localities． It seems that this genus could even pos-
sess much more diversity than presently known as lots of new species have recently been and are being discribed，particular-
ly from its distributional center． In the presnt study，a new species of Gesneriaceae，P． bobaiensis from Guangxi，China is
described and illustrated． We intuitively judged the higher taxonomic unit，i． e． genus，that the assumed new species should
belong to;compared the morphologies of this assumed new species with all other recognized species previously discribed in
the genus and deduced possible close relatives of the new species based on morphological similarity． Then we tested its af-
finities to other sampled species from the same large region and the divergence of DNA sequences between the assumed new
species and its closest relatives． Morphologically，this new species is assumed to belong to the genus Primulina and can be
easily distinguished from all other Gesneriaceae species by its special floral appearance，e． g． lower lip is much longer than
upper lip and it has spotted marks on the inside of the upper lip(the majority of other species have striped marks on the in-
side(s)of the lower and /or upper lips)． Subsequently，the taxonomic treatment of being attibuted to Primulina and the
monophyly of the recircumscribed genus Primulina s． l． are also corroborated by the molecular evidence based on the plastid
matK gene． Within the genus the new species is suggested to be the sister of a lineage comprising P． swinglei and P． laxi-
flora． Further examination of the matK sequences indicated 5 and 11 base differences of P． bobaiensis separating from P．
swinglei and P． laxiflora，respectively，implying potential long period of isolation from these species or high substitution
rate of the matK gene in these taxa． Moreover，the new species is restricted on Danxia rocks while its closest relatives as
well as most of its congeners are confined on limestone rocks，suggesting possible specific edaphic adaptation and isolation of
the new species． Further，the high endemism with low abundance and usually morphological and molecular distinctiveness
of this and many other species of Primulina would also superimpose the importance of conservation of these rare species．
Key words:Gesneriaceae;Primulina;new species;phylogeny;matK
CLC number:Q949 Document code:A Article ID:1000-3142(2015)02-0143-08
收稿日期:2014-04-23 修回日期:2014-06-27
基金项目:广西自然科学基金(2012GXNSFBA053067);广西林业科技项目(桂林科字［2014］第 27号);广西植物功能物质研究与利用重点实验室主
任基金(ZＲJJ2013-9);广西植物研究所基本业务费项目(Guizhiye 12011);广西植物研究所博士启动基金(Guizhiye 11006)。
作者简介:黎乾坤(1986-)，男，广西梧州人，硕士研究生，从事药用植物资源学研究，(E-mail)459672720@ qq． com。
* 通讯作者:李文兰，博士，从事植物分子生物学研究，(E-mail)liwenlan@ whu． edu． cn。
广西报春苣苔属一新种及其系统位置
黎乾坤1，张 强2，邓 涛2，盘 波2，黄俞淞2，李文兰1，3*
(1． 桂林理工大学 化学与生物工程学院，广西 桂林 541006;2． 广西壮族自治区
中 国 科 学 院
广西植物研究所，
广西 桂林 541006;3． 广西大学 生命科学与技术学院，南宁 530005 )
摘 要:报春苣苔是一类生长在石灰岩基质上的多年生草本植物，目前已知的种类160余种，主要分布在我国南
部和越南北部的石灰岩地区。该属植物许多种类分布极其狭域，往往仅零星分布在某个或少数几个地点。近年
来，该属植物许多新种类不断地被发现和报道，预示着该属植物可能有更加丰富的多样性。该研究报道了在广
西博白县发现的该属植物一新种。从形态上判断，该新种属于广义报春苣苔属，其具有的一些特别的形态特征，
如花冠下唇比上唇长得多，上唇内部具有两个近圆形斑点(其它多数种类具有条形斑点)，而易于与本属其它物
种区分。而且，基于叶绿体 matK基因序列重建的系统发育关系表明，该新种的确属于报春苣苔属且广义报春苣
苔属是单系类群。该新种与钟冠报春(P． swinglei)和疏花报春(P． laxiflora)最为近缘，其 matK基因与后两者分
别有 5个和 11个碱基差异，暗示这些物种间经历了较长时间的独立演化或 matK基因在这些物种中具有较高的
碱基替换速率。另外，该新种只生长在丹霞岩石上，而其近缘种以及同属大多数其它种类都生长在石灰岩基质
上，暗示该新种获得了特殊的适应能力以及与其它近缘种间存在生态隔离。该新种以及近年来本属许多其它新
种类的发现，进一步增加了对该属植物多样性的认识，以及该属植物的演化与多样性分化之谜。这些物种所具
有的高度区域或局域特有性和稀有性，以及形态上的特殊性和遗传上的独立性，进一步强化了对该属植物开展
保护的重要性。
关键词:苦苣苔科;报春苣苔属;新物种;系统发育;matK基因
The recently recircumscribed genus Primulina
Hance (Gesneriaceae)contains more than 130 species
with large morphological diversities (Weber et al．，
2011)． It has grown to be one of the species-richest
genus of Gensneriaceae in South China and its sur-
rounding regions，one of the distributional centres of the
family across the world，with 59 genera and over 460
discribed species found in China alone (Li et al．，
2004;Wei et al．，2010)． Primulina is still expanding
rapidly with lots of new species and varieties that have
been and are being discribed，mainly from the distrbu-
tional centre of the limestone regions of South China，
especially from Guangxi Zhuang Autonomous Ｒegion，
where dozens of new species have been discribed in the
last several years (Xu et al．，2008;Li et al．，2009;Li
et al．，2010;Xu et al．，2010;Huang et al．，2011;Wu
et al．，2011;Xu et al．，2011a，2011b;Hong et al．，
2012;Huang et al．，2012;Wu et al．，2012;Xu et al．，
2012;Chung et al．，2013;Wen et al．，2013;Zhao et
al．，2013)．
In the present study，we found and discribed a
Gesneriaceae population with special floral appearance
from the generic distributional centre of Guangxi，Chi-
na． The new population was found growing on Danxia
rocks，but not on limestones，habitat occupied by most
others of its congeners． According to its floral structure
and chararaters，this new population can be easily at-
tributed to the recently redefined genus Primulina and
is assumed to represent a new species in the genus after
carefully checking the morphologies against all the rec-
ognized congeners，including specimens and those dis-
cribed in literatures (Wang et al．，1990;1999，Li et
al．，2004;Wei et al．，2010)． Whereas，depending on
morphological comparisons alone，it is difficult to assess
which known species the new species is closely related
to due to large morphological diversities and complicat-
ed morphological evolution in Primulina (Weber et
al．，2011)．
Traditional taxonomy has been proved to largely
contradict with the molecular systematics in Gesneri-
aceae and the majority of the traditional genera have
been revealed as non-monophyly (e． g． Wang et al．，
2011;Weber et al．，2011)． Further，numerous mono-
typic genera with distinct morphology have been widely
proved to be embedded deeply within other genera and
their generic status thus should be abandoned，that a-
441 广 西 植 物 35卷
gain imply large incongruence between phylogenetic re-
lationship and morphological similarity (Weber et al．，
2011)． The incongruence between phylogenetic rela-
tionship and morphological similarity had often led er-
rors in generic level taxonomic treatments for some spe-
cies and a large proportion of taxa should be trasferred
from their genera originally assigned by traditional tax-
onomy to another according to the molecular phyloge-
netics (Liu et al．，2010;Mller et al．，2011;Weber et
al．，2011)． All these would suggest complicated mor-
phological evolution in Gesneriaceae and necessity to
combine molecular evidence with taxonomic treatment．
Hence we also tested the generic placement and parti-
cular intrageneric relationships of the new species to
other Gensericeae species from the surrounding areas
where the new species was found based on a molecular
framework，a matrix of matK gene modified from our
previous study (Wu et al．，2012)．
1 Materials and Methods
We newly sampled the new species and four other
Primulina taxa within the same region in the present
study，each with one individual as the representative of
the corresponding species． Further，we also newly sam-
pled 14 taxa representing other 14 genera from the same
large region． These newly sampled taxa would be added
to our previously obtained matK sequences from other 41
Primulina taxa mainly from the same large region．
Total genomic DNA was extracted from silica dried
leaves using the CTAB protocol (Doyle et al．，1987)．
DNA sequences were amplified using a PCＲ procedure
of initial denaturation at 94 ℃ for 3 min，followed by 34
cycles of 30 sec at 94 ℃，40 sec at 52 ℃，and 90 sec at
72 ℃，with a final extension at 72 ℃ for 10 min． PCＲ
products were purified using TIANgel Midi Purification
Kit (Tiangen Biotech)according to the manufacturer’s
instructions and then sequenced on an ABI 3730 DNA
Sequencer (Applied Biosystems International，Foster
City，CA，USA)，using Big Dye Terminator (Applied Bi-
osystems，Shanghai，China)． The primers used for am-
plification and sequencing are the same as in Wu et al．
(2011)．
DNA sequences were manually edited and spliced
using the software Bioedit 5． 0． 9 (Hall 1999)according
to the bidirectional chromatograms for each of the se-
quences． All sequences after edition were aligned in
Clustal X 1． 81 (Thompson et al．，1997)and adjusted
manually in Bioedit 5． 0． 9． The flank regions at both
ends with many missing sites were excluded and the in-
dels within sequences were filled by gaps． Then，the
matrix of these newly obtained sequences was added to
our previous larger matrix and aligned visually in Bi-
oedit 5． 0． 9． The genbank accession numbers and
sources of the plant materials are as listed in the supple-
mentary material (Table 1)and the final matrix is a-
vailable upon request from the corresponding author．
Phylogenetic analyses were conducted using neigh-
bor joining (NJ) in Mega 4． 0． 2 (Tamura et al．，
2007)，Maximum Parsimony (MP)in PAUP version 4．
0b10 (Swofford 2002)and Bayesian Inference (BI)in
MrBayes v3． 1． 2 (Huelsenbeck et al．，2001)，respec-
tively． For the NJ method，we chose and calculated
Kimura 2-parameter distance with pairwise deletion of
the gaps and reconstruct the phylogeny based on the dis-
tance matrix． Branch Support (BSnj)was calculated
with 100 bootstraps，also using NJ method and the same
parameter settings． The MP analysis was conducted u-
sing the following settings:100 replicates of random
stepwise addition with tree bisection-reconnection
(TBＲ)branch swapping performed using heuristic sear-
ches，with all most-parsimonious trees saved at each rep-
licate (MulTree on)． Branch Support (BSmp)was as-
sessed using bootstrap analysis with 100 bootstrap repli-
cates，each with 10 stepwise additions． As the inadequa-
cy of the parsimony informative sites of the matK se-
quences would have yielded excessive equally most par-
simonious trees in the tentative MP heuristic search，
therefore，500 000 trees were set as the maximum limit
for the first round of heuristic searches based on the o-
riginal matrix and 50 000 maxi
5412期 黎乾坤等:广西报春苣苔属一新种及其系统位置
Table 1 Genbank accession numbers and the plant material sources used in the present study
Taxa MatK Collector (Voucher)/Location
Aeschynanthus austroyunnanensi KJ137885 Ｒ． C． Hu ＆ Q． Zhang /Huanjiang County，Guangxi，China
Beccarinda tonkinensis KJ137886 W． B． Xu /Shangsi County，Guangxi，China
Chiritopsis glandulosa KJ137887 W． B． Xu /Pingle County，Guangxi，China
Chiritopsis sp． KJ137888 W． B． Xu /Zhongshan County，Guangxi，China
C． lingchuanensis KJ137889 L． Wu ＆ B． Pan /Lingcuan County，Guangxi，China
C． glandulosa var． yangshuoensis KJ137890 W． B． Xu /Yangshuo County，Guangxi，China
Dayaoshania cotinifolia KJ137891 N． Z． Huang (Cultivated in Botanical Garden of Guangxi Institute of Botany (GXIB))
Didymocarpus glandulosus KJ137892 S． S Mo ＆ Q． Zhang /Huanjiang County，Guangxi，China
Gyrocheilos retrotrichum KJ137893 S． S Mo ＆ Q． Zhang /Huanjiang County，Guangxi，China
Hemiboea cavaleriei KJ137894 Y． Liu /Lingcuan County，Guangxi，China
Lagarosolen coriaceiforlium KJ137895 L． Wu ＆ B． Pan /Yangshuo County，Guangxi，China
Lysionotus fengshanensis KJ137896 S． S Mo ＆ Q． Zhang /Huanjiang County，Guangxi，China
Ophithandra sinohenryi KJ137897 W． B． Xu /Fangcheng City，Guangxi，China
Oreocharis xiangguiensis KJ137898 W． B． Xu /Lingcuan County，Guangxi，China
Paraboea swinhoii KJ137899 S． S Mo ＆ Q． Zhang /Huanjiang County，Guangxi，China
Paraisometrum mileense KJ137900 Q． Gao /Xingyi County，Guizhou，China
Petrocodon sp． KJ137901 B． Pan /Jiangyong County，Hunan，China
Priminlina swinglei KJ137902 B． Pan /cult． In GXIB
Henckelia anachoreta KJ137903 W． B． Xu /Ningming County，Guangxi，China
Primulina baishouensis KJ137904 B． Pan /Yongfu County，Guangxi，China
P． bobaiensis KJ137905 B． Pan /Bobai County，Guangxi，China
P． cordifolia KJ137906 Q． Gao ＆ B． Pan /Yangshuo County，Guangxi，China
P． dryas JX195976 A． Chautems ＆ M． Perret 01 －019 (G)
P． eburnea KJ137908 Q． Gao ＆ B． Pan /Guilin City，Guangxi，China
P． fimbrisepala KJ137909 Q． Zhang /Shaoguan City，Guangdong，China
P． gongchengensis KJ137910 Y． S． Huang ＆ W． B． Xu (10496)/Gongcheng County，Guangxi，China
P． gueilinensis KJ137911 Q． Gao ＆ B． Pan /Guilin City，Guangxi，China
P． guihaiensis KJ137912 Q． Gao ＆ B． Pan /Guilin City，Guangxi，China
P． hedyotidea KJ137913 W． B． Xu /cult． In GXIB
P． hochiensis KJ137914 W． B． Xu /cult． in GXIB
P． langshanica KJ137915 W． B． Xu ＆ B． Pan (10021)/ Quanzhou County，Guangxi，China
P． laxiflora KJ137916 Y． S． Huang /Longzhou County，Guangxi，China
P． liboensis KJ137917 W． B． Xu /Guangxi
P． liguliformis KJ137918 B． Pan /Hechi City，Guangxi，China
P． lijiangensis KJ137919 W． B． Xu /Yangshuo County，Guangxi，China
P． linearifolia KJ137920 Y． Liu /Guangxi
P． longangensis KJ137921 Y． S． Huang /Longzhou County，Guangxi，China
P． longicalyx KJ137922 L． Wu ＆ B． Pan /Guilin City，Guangxi，China
P． longii KJ137923 Y． S． Huang＆ B． Pan /Yongfu County，Guangxi，China
P． lunglinensis var． amblyosepala KJ137924 W． B． Xu /Huanjiang County，Guangxi，China
P． lutea KJ137925 W． B． Xu /cangwu County，Guangxi，China
P． lutea KJ137926 S． S． Mo /Hezhou City，Guangxi，China
P． macrodonta KJ137927 L． Wu ＆ B． Pan /Lingcuan，Guangxi，China
P． medica KJ137928 B． Pan /Pingle County，Guangxi，China
P． minutimaculata KJ137929 W． B． Xu /Longzhou County，Guangxi，China
P． debaoensis KJ137930 W． B． Xu /cult． in GXIB，Guangxi，China
P． ningmingensis KJ137931 W． B． Xu /ningming County，Guangxi，China
P． pseudoheterotricha KJ137932 W． B． Xu /Zhongshan County，Guangxi，China
P． pungentisepala KJ137933 W． B． Xu /Guangxi，China
P． ronganensis KJ137934 Y． S． Huang ＆ B． Pan /Ｒongan County，Guangxi，China
P． rongshuiensis KJ137935 Y． S． Huang ＆ B． Pan /Ｒongshui County，Guangxi，China
P． shouchengensis KJ137936 W． B． Xu /Yongfu County，Guangxi，China
P． subrhomboidea KJ137937 B． Pan /Yangshuo County，Guangxi，China
P． tabacum KJ137938 W． B． Xu /Hezhou City，Guangxi，China
P． tribracteata var． zhuana KJ137939 W． B． Xu /cult． in GXIB，Guangxi，China
P． wangiana KJ137940 Y． S． Huang ＆ B． Pan /Ｒongan County，Guangxi，China
P． wentsaii KJ137941 W． B． Xu /Ningming County，Guangxi，China
P． yangshuoensis KJ137942 Q． Gao ＆ B． Pan /Yangshuo County，Guangxi，China
P． yungfuensis KJ137943 J． Liu et al． /Yongfu County，Guangxi，China
P． sinovietnamica KJ137944 W． B． Xu /Longzhou County，Guangxi，China
641 广 西 植 物 35卷
Table 2 Difference of the nucleotides of the new species to its close congeners
Species 32 63 113 149 173 174 298 511 525 567 702 752 834 1005
P． bobaiensis C T C G T C T C A C A C G A
P． swinglei ? ? C A T A C C A C G T G A
P． laxiflora T C T G G C C T T T G C C T
Note:The first row represents the site numbers in the matrix． The mark (?)represents ambiguous state for the site． The different bases of P． swinglei and P． laxiflora to
the new species are highlighted in bold．
Fig． 1 Phylogenetic placement of the new species P． bobaiensis(highlighted in bold)to its close relatives． Nodal supports bigger than 50%
from NJ and MP are sequentially (before or behind the left slash)listed above the branches，while PP values bigger than 0． 50 from BI method are listed
below the branches in italic．
mum trees were set for each of the bootstrap pseudorep-
lications． Prior the Bayesian analysis，we used Modeltest
(Posada et al．，1998;Posada et al．，2004)to select the
fittest DNA substitution model based on the Akaike In-
formation Criterion (AIC)． Two runs of Metropolis-cou-
pled Markov Chain Monte Carlo (MCMCMC)analyses
were performed simultaneously，with a random starting
tree and four chains for each run (one cold and three
heated)． The MCMCMC length was 10 000 000 genera-
tions and the chain was sampled every 1000th generation
7412期 黎乾坤等:广西报春苣苔属一新种及其系统位置
from the cold chain． Bayesian clade posterior probabili-
ties and average branch lengths were calculated based
on the sampled trees combined from the two runs after
the first 15% of the total trees were discarded as burn-
in． The burn-in for each run was validated by plotting
the likelihood values against the generations．
2 Ｒesults and Analysis
2． 1 Phylogenetic placement of the new species
After pruning the flank regions with many missing
characters，the matK region obtained varies from 950 bp
to 1 156 bp for all the sampled 61 accessions with an av-
erage GC content of 32% and the final aligned matrix
has 1 156 bp in length with 115 informative sites as well
as 165 additional variable but uninformative sites． TVM
+G model was selected as the fittest substitution model
for the matK gene by the Modeltest．
Through three methods，we yielded phylogenetic
trees with generally consistent topology and the Bayesian
50% majority consensus tree was more resolved than
those from the other two methods． Maximum constraint
of 500 000 trees was hit in the first replication of MP
heuristic searches，with the shortest tree length of 401
steps，consistency index (CI)of 0． 79 and Ｒetention in-
dex (ＲI)of 0． 84． The results from all three methods
indicated that all the Primulina taxa sampled in present
study formed as a monophyletic group with low NJ and
MP bootstrap support，but high Bayesian posterior proba-
bility (BSnj =56%;BSmp =58%;PP =1． 00)． The new
species was embedded within Primulina，being sister
(BSnj = 100%;BSmp = 99%;PP = 1． 00)to a lineage
(BSnj =84%;BSmp =82%;PP = 0． 98)composed of P．
swinglei and P． laxiflora (Fig． 1)． Further examination
of the matK genes indicated the new species differed
from P． swinglei and P． laxiflora by 5 and 11 nucleo-
tide bases，respectively，implying potential long-term of
isolation between the species or higher substitution rate
of the matK gene in these taxa (Table 2)．
2． 2 Taxonomic treatment
P． bobaiensis Q． K． Li，Q． Zhang ＆ W． L． Li，
sp． nov． Fig． 2 ＆ Fig． 3．
Herbs perennial． Ｒhizome subterete，2 － 7 cm
long，1 －3 cm across． Leaves 5 －9，crowded at the apex
of rhizome，petiolate;petiole applanate，3 －12 cm long，5
－15 mm across;leaf blade herbaceous，elliptic or oval，
4 －12 ×2 － 6 cm，apex acute to obtuse，base cuneate to
broadly cuneate，margin shallowly serrate，pubescent on
both surfaces，lateral veins 3 －6 on each side，impressed
adaxially and prominent abaxially． Cymes 2 － 4，1 － 3-
branched，5 －12-flowered;peduncle 5 －18 cm long，1 －
3 mm across，pubescent;pedicel 1． 2 － 3． 1 cm long，pu-
bescent;bracts opposite，oblanceolate，8 － 22 × 4 － 9
mm，margin repand，apex obtuse，pubescent on both
sides． Calyx 5-parted nearly to the base，lobes narrowly
lanceolate，3 － 5 × 0． 8 － 1． 1 mm，apex acuminate，out-
side pubescent，inside puberulent，margins entire． Corol-
la deep purple，16 － 18 mm long，outside pubescent，in-
side sparsely puberulent;tube ca． 7 － 9 mm long，ca． 4
mm in diam． at the base，ca． 6 －8 mm in diam． at the
mouth;limb distinctly 2-lipped，adaxial lip 2-parted to
over the middle，lobes broadly ovate，3 － 4 × 3 － 3． 5
mm;abaxial lip 3-parted to over the middle，lobes ob-
long，6 －8 ×3 － 4 mm． Stamens 2，adnate to ca． 5 mm
above the corolla tube base，filaments linear，5 － 8 mm
long，purple，twisted at over middle，glabrous;anthers
reniform，dorsifixed，ca． 2 mm long，glabrous;staminodes
3，adnate to ca． 6 mm above the corolla tube base，later-
al ones 2 mm long，apex capitate，middle one 0． 5 mm
long． Disk annular，ca． 1． 1 mm in height，margin repand．
Pistil 12 －15 mm long，ovary 4 －6 mm long，ca． 1． 3 mm
across，pubescent;style pubescent，6 － 8 cm long;stigma
obovate，1 －1． 5 mm long． Capsule not seen． The species
is only known growing on Danxia rock from one locality in
Wangmao Town，Bobai County，Guangxi，China． It is in
flowering from September to October． The specific epithet
is derived from the type locality，Bobai County，Guangxi．
P． bobaiensis is special in floral morphology which can be
easily distinguished from all other Gesneriaceae． It is e-
ven very different from its sampled rela
tives in floral appearance． The details of diagnostic
841 广 西 植 物 35卷
Fig． 2 A． Habitat of the new species (Primulina bobaiensis)in Danxia landscape;B． P． bobaiensis;
C． Flower of P． swinglei;D． Flower of P． laxiflora．
Table 3 Distinguishing morphological characters among P． bobaiensis，P． swinglei and P． laxiflora
P． bobaiensis P． swinglei P． laxiflora
Corolla White at mouth，upper lobes pointed at termi-
nal;round spotted upper lobes oblong，upper la-
bia ca． 1 /2 in length to the lower．
White at mouth，upper lobes pointed at ter-
minal;striped upper lobes ovate，upper la-
bia slightly shorter than the lower．
Yellow at mouth，upper lobes round at ter-
minal，lower lobes near round，upper labia
slightly shorter than the lower．
morphological characters of P． bobaiensis to its phyloge-
netic close relatives of P． swinglei and P． laxiflora are
listed in Table 3．
China，Guangxi，Yulin City，Bobai County，Wang-
mao Town，Yanshi temple，on rock face of Danxia land-
scape，rare，elevation 123 m，9 October 2010，Yu-Song
HUANG and Bo Pan100839 (holotype IBK;isotype
PE)． Guilin Botanical Garden，introduced from Bobai
County，Wangmao Town，31 August 2009，Wei-Bin XU
et Bo Pan 091775 (IBK)．
3 Conclusion and Discussion
The new species embedded within Primulina seems
robust，strongly supported by both morphological obser-
vation and molecular characters． The status as an inde-
pendent species is also easy to be confirmed by its very
special appearance． According to our knowledge the ex-
tremely unsymmetrical labia with the lower one extended
to two folds or more in length than the upper one is rare
in other recognized Gesneriaceae taxa in this area;the
two spotted honey guides on the upper two lobes but not
any marks on the lower three lobes are also rare． All the
floral characters seem stable as several cultivated indi-
viduals on yellow soils in greenhouse show no discern-
able morphological variations compared with those ob-
served for the wild population on Danxia rocks，unlike
many other taxa which show observable or striking mor-
phological variations between and /or within populations
(Wei et al．，2010)． Although the species status is easy
to be confirmed，whereas，the exact relationships and
whether there is more closely-related known species for
9412期 黎乾坤等:广西报春苣苔属一新种及其系统位置
Fig． 3 P． bobaiensis Q． K． Li，＆ W． L． Li A． Habit;B．
Corolla opened showing stamens and staminodes;C． Stamens;D．
Pistil and calyx;E． Stigma． (Drawn by ZHU Yun-Xi from the holo-
type)．
the new species still needs further molecular analysis
based on denser sampling．
Acknowledgements We are grateful to Mr． Zhu
Yun-Xi (IBK)for the handsome drawing． We thank Xu
Wei-Bin，Gao Qi，Hu Ｒen-Chuan，Huang Nin-Zhen and
Liu Jing from IBK for collecting the plant materials．
Ｒeferences:
Chung KF，Huang HY，Peng CI，et al． 2013． Primulina mabaensis
(Gesneriaceae)，a new species from a limestone cave of northern
Guangdong，China［J］． Phytotaxa，92:40 －48
DoyleJJ，Doyle JL． 1987． A rapid DNA isolation procedure for small
quantities of fresh leaf tissue［J］． Phytochem Bull，19:11 －15
Hall TA． 1999． BioEdit:a user-friendly biological sequence align-
ment editor and analysis program for Windows 95 /98 /NT［J］．
Nucleic Acids Symp Ser，41:95 －98
Hance HF． 1883． Primulina tabacum Hance［J］． J Bot． 21:169
Hong X，Zhou SB，Wen F． 2012． Primulina chizhouensis sp． nov．
(Gesneriaceae)，a new species from a limestone cave in Anhui，
China［J］． Phytotaxa，50:13 －18
Huang YS，Xu WB，Liu Y． 2011． Chirita rongshuiensis，a new spe-
cies of Gesneriaceae from northern Guangxi，China［J］． Taiwan-
ia，56:54 －57
Huang YS，Xu WB，Wu L，et al． 2012． Primulina gongchengensis
(Gesneriaceae)，a new species from Guangxi，China［J］． Ann
Bot Fenn，40:107 －110
Huelsenbeck JP． Ｒonquist F． 2001． MrBayes:Bayesian inference
of phylogenetic trees［J］． Bioinformatics，17:754 －755
Li JM，． Mller M． 2009． Chirita grandibracteata (Gesneriaceae)，a
new species from Yunnan，China［J］． Ann Bot Fenn，46:125 －129
Li JM，Zhu SX． 2010． Chirita auriculata (Gesneriaceae)，a new
species from China，placed in sect． Chirita based on molecular
data［J］． Ann Bot Fenn，47:221 －225
Li ZY，Wang YZ． 2004． Plants of Gesneriaceae in China［M］． Zheng-
zhou:Henan Science and Technology Publishing House:721
Liu Y，Xu WB，Pan B． 2010． Wentsaiboea tiandengensis sp． nov．
and W． luochengensis sp． nov． (Gesneriaceae)from Karst caves
in Guangxi，southern China［J］． Nord J Bot，28:739 －745
Mller M，Forrest A，Wei YG，et al． 2011． A molecular phylogenet-
ic assessment of the advanced Asiatic and Malesian didymo-
carpoid Gesneriaceae with focus on non-monophyletic and mono-
typic genera［J］． Plant Syst Evol，292:223 －248
Posada D，Buckley TＲ． 2004． Model selection and model averaging
in phylogenetics:advantages of Akaike information criterion and
Bayesian approaches over likelihood ratio tests［J］． Syst Biol，
53:793 －808
Posada D，Crandall KA． 1998． Modeltest:testing the model of DNA
substitution［J］． Bioinformatics，14:817 －818
Swofford DL． 2002． PAUP* ． Phylogenetic analysis using parsimo-
ny (* and other methods)． Version 4［M］． Sinauer Associates，
Sunderland，Massachusetts
Tamura K，Dudley J，Nei M，et al． 2007． MEGA4:Molecular evolu-
tionary genetics analysis (MEGA)software version 4． 0［J］． Mol
Biol Evol，24:1 596 －1 599
Thompson JD，Gibson TJ，Plewniak F，et al． 1997． The ClustalX
windows interface:flexible strategies for multiple sequence align-
ment aided by quality analysis tools［J］． Nucleic Acids Ｒes，25:
4 876 －4 882
Wang WT，Pan KY，Li ZY． 1990． Gesneriaceae［M］/ /Wang WT
(eds)． Flora Ｒeipublicae Popularis Sinicae． Beijing:Science
Press:125 －581
Wang WT，Pan KY，Li ZY，et al． 1998． Gesneriaceae［M］/ / Wu
ZY，Ｒaven PH(eds)． Flora of China． Beijing:Science Press;
St． Louis:Missouri Botanical Garden Press:244 －401
Wang YZ，Mao ＲB，Liu Y，et al． 2011． Phylogenetic reconstruction
of Chirita and allies (Gesneriaceae)with taxonomic treatments
［J］． J Syst Evol，49:50 －64
Weber A，Middleton DJ，Forrest A，et al． 2011． Molecular systemat-
ics and remodelling of Chirita and associated genera (Gesneri-
aceae)［J］． Taxon，60:767 －790
Wei YG，Wen F，Mller M，et al． 2010． Gesneriaceae of South Chi-
na［M］． Nanning:Guangxi Science and Technology Publishing
House:777
Wen F，Zhao B，Liang GY，et al． 2013． Primulina lutvittata
(Gesneriaceae)，a new species from a limestone cave in Guang-
dong，China［J］． Ann Bot Fenn，50:87 －90
Wu L，Zhang Q，Xu WB，et al． 2012． Primulina guigangensis
(Gesneriaceae):a new species from limestone area in Guangxi，
China［J］． Phytotaxa，38:19 －23
Wu WH，Xu WB，Nong DX，et al． 2011． Chirita ninminensis
(下转第 177页 Continue on page 177 )
051 广 西 植 物 35卷
不同物种在胚乳细胞的长、宽、长宽比以及形状上都
呈现差异，表明胚乳细胞特征在一定程度上具有共
属分种的意义，但不能很好地反映属以及基因组间
的差异。按照染色体组分类方法，含相同染色体组
的物种亲缘关系较近。E． Canadensis (2)和 E．
sibiricus (6)都具有 StH 染色体组，亲缘关系很近，
但是 E． Canadensis 和 E． sibiricus 在细胞形状和长
宽比上的差异都很大。然而具有同样 P 染色体组
的 A． cristatum (2)和 A． mongolicum (2)和同样具
有 St 染色体组的 P． libanotica (10)和 P． spicata
(11)在细胞形状和长宽比上的差异又很小，甚至
P． libanotica (10)和 P． spicata (11)的长宽比为相
同的 1． 57。就考察的胚乳细胞的各项指标来看，在
一些种间胚乳细胞的的大小和宽厚差异不及形状差
异明显。例如 E． nutans (5)和 E． sibiricus (6)的大
小和长宽比差异不大，但是前者主要是椭球形，后者
以圆形和长体形为主，而且前者较密后者稀疏。还
有些物种各项指标仅有少许差异，如 P． libanotica
(10)和 P． spicata (11)，利用胚乳细胞特征分析它
们关系时应该慎重。比较本实验 4 个属 11 个物种
的胚乳细胞，发现不同属、同一属内不同物种的胚乳
细胞在形状和长宽比上都差异较大，且无明显规律，
表明胚乳细胞特征不能能较好地反映属以及基因组
间的亲缘关系。所以对披碱草属植物的分类地位、
起源以及系统进化需要综合分析形态学、细胞学、细
胞遗传学以及分子生物学的资料才能作出正确的
分析。
参考文献:
Cai LB(蔡联炳)． 2000． Characters of endosperm cells in the
seeds of some species of Ｒoegneria and their taxonomics ignifi-
cance (鹅观草属一些种种子胚乳细胞的特征及其分类学意
义的探讨)［J］． Acta Bot Boreal-Occident Sin (西北植物学
报)，20(6):1 070 －1 075
Dong YC(董玉琛)． 1992． Study and Utilization of Wild Ｒelatives
of Wheat in China(小麦野生近缘植物的研究和利用)［M］．
Beijing(北京):China Science and Technology Press(中国科学
技术出版社):39 －44
Guo BZ(郭本兆)，Lu LS(卢莲生)，Sun YH(孙永华)，et al．
1987． Flora of China． 9 (3) (中国植物志第 9 卷 3 分册)
［M］． Beijing(北京):Science Press(科学出版社):52 －61
Jensen KB． 1989． Cytology and origin of Elymus abolinii，and its
F1 hybrids with Pseudoroegneria spicata，Elymus lanceotalus，E．
dentatus ssp． ugamicus，and E． drobovii (Poaceae:Triticeae)
［J］． Genome，32(5):468 － 474
Liu J(刘静)，Yang ＲW(杨瑞武)，Zhang HQ(张海琴)，et al．
2007． Study on diversity of endosperm cells in seeds of Hystrix
and related genera in Triticaae(猬草属及其近缘属种子胚乳细
胞多样性研究)［J］． J Sichuan Agric Univ (四川农业大学学
报)，25(1):14 －18
Lu BＲ． 1994． Meiotic analysis of the intergeneric hybrids between
Pseudoroegneria and tetraploid Elymus［J］． Cathya，6(2):1 －14
Lu BＲ，Bothmer ＲV． 1990． Intergeneric hybridization between
Hordeum and asiatic Elymus［J］． Hereditas，112:109 －116
Lu BＲ． 1993． Genomic relationships within the Elymus parviglumis
group (Triticeae:Poaceae)［J］． Plant Syst Evol． 187(1-4):
191 －211
Wang XL(王晓丽)，Tang JY(汤加勇)，Liu J(刘静)，et al． 2008．
Diversity of endosperm cells in Triticeae species with StH genomes
(小麦族 StH 基因组物种的胚乳细胞多样性)［J］． J Sichuan
Agric Univ (四川农业大学学报)，26(1):11 －15
Yang ＲW(杨瑞武)，Zhou YH(周永红)，Zheng YL(郑有良)，et
al． 2004． Study on diversity of endosperm cells in seeds of Ley-
mus (赖草属种子胚乳细胞多样性研究)［J］． Acta Bot Boreal-
Occident Sin (西北植物学报)，24(1):
櫠櫠櫠櫠櫠櫠櫠櫠櫠櫠櫠櫠櫠櫠櫠櫠櫠櫠櫠櫠櫠櫠櫠櫠櫠櫠櫠櫠櫠櫠櫠櫠櫠櫠櫠櫠櫠櫠櫠櫠櫠櫠櫠櫠櫠櫠
38 －42
(上接第 150页 Continue from page 150 )
(Gesneriaceae)，a new species from Guangxi，China［J］． Ann Bot
Fenn，48:422 －424
Xu H，Li ZY，Jiang H． 2008． A new species of Chirita(Gesneriace-
ae)from Yunnan，China［J］． Bot J Linn Soc，158:269 －273
Xu WB，Huang YS，Wu L，et al． 2011a． Chirita luochengensis
(Gesneriaceae)，a new species from limestone areas in northern
Guangxi，China［J］． Brittonia，63:314 －317
Xu WB，Pan B，Huang YS，et al． 2010． Chirita leprosa sp． nov．
(Gesneriaceae) from limestone areas in Guangxi，China［J］．
Nord J Bot，28:705 －708
Xu WB，Pan B，Huang YS，et al． 2011b． Chirita lijiangensis
(Gesneriaceae)，a new species from limestone area in Guangxi，
China［J］． Ann Bot Fenn，48:188 －190
Xu WB，Pan B，Liu Y，et al． 2012． Two new species，Primulina
multifida and P． pseudomollifolia (Gesneriaceae)，from karst ca-
ves in Guangxi，China［J］． Bot Study，53:165 －175
Zhao B，Pan B，Zhang Y，et al． 2013． Primulina guizhongensis
(Gesneriaceae)，a new species from Guangxi，China［J］． Phyto-
taxa，109:27 －35
7712 期 高刚等:披碱草属及其近源属植物种子胚乳细胞多样性研究