全 文 :浙江大学学报(农业与生命科学版) 39(4):375~380,2013
Journal of Zhejiang University(Agric.&Life Sci.)
http://www.journals.zju.edu.cn/agr
E-mail:zdxbnsb@zju.edu.cn DOI:10.3785/j.issn.1008-9209.2012.08.271
Foundation item:Project supported by Science and Technology Department of Zhejiang Province(No.2011C22101).
*Corresponding author:JIN Xiaofeng,Tel:+86-571-28866880;E-mail:docxfjin@163.com
Biography:YE Lixin,E-mail:lqyelx@163.com
Received:2012-08-27;Accepted:2013-03-19;Published online:2013-07-16
URL:http://www.cnki.net/kcms/detail/33.1247.S.20130716.1447.003.html
Pseudopyxis heterophylla subsp.monilirhizoma (Rubiaceae,tribe
Paederieae),a new combination from Zhejiang of China,based on
morphological trait analysis
YE Lixin1,ZHOU Yingying2,LIU Lingjuan1,CHEN Weijie2,LIU Shenglong1,ZHENG Chaozong3,JIN Xiaofeng2*
(1.Administration of Zhejiang Fengyangshan-Baishanzu National Natural Reserve,Longquan,Zhejiang 323700,China;
2.College of Life and Environment Sciences,Hangzhou Normal University,Hangzhou 310036,China;3.College of
Life Sciences,Zhejiang University,Hangzhou 310058,China)
Summary Pseudopyxis Miq.,agenus distributed in eastern China and Japan,comprises three species,in
which Pseudopyxis monilirhizoma Tao Chen is endemic to China.The variation analysis of the diagnostic
characters among these three species of Pseudopyxis was carried out.Based on principal coordinate analysis
(PCoA)of eighteen morphological characters,the individuals of P.depressa were separated distinctly from P.
heterophylla+P.monilirhizoma,whereas the separation between P.heterophyllaand P.monilirhizoma was
not clear.The observations of stem indumentum and seed micromorphology showed the difference between P.
depressaand P.heterophylla+P.monilirhizoma,while that of P.heterophylla and P.monilirhizoma was
consistent.In case of the observations of stem indumentum and seed micromorphology and PCoA of
morphological characters,P.monilirhizomais consequently combined as a subspecies of P.heterophylla(Miq.)
Maxim.,and the combination of P.heterophyllasubsp.monilirhizoma(Tao Chen)L.X.Ye,C.Z.Zheng &
X.F.Jin is proposed.
Key words Pseudopyxis heterophylla subsp.monilirhizoma;new combination;morphological characters;
scanning electron microscope(SEM);taxonomy
CLC number Q 949 Document code A
基于形态性状分析的浙江茜草科胀节假盖果草亚种新组合 (英文).浙江大学学报 (农业与生命科学
版),2013,39 (4):375-380
叶立新1,周莹莹2,刘玲娟1,陈伟杰2,刘胜龙1,郑朝宗3,金孝锋2* (1.浙江凤阳山-百山祖国家级自然保护区
凤阳山管理处,浙江 龙泉323700;2.杭州师范大学生命与环境科学学院,杭州310036;3.浙江大学生命科
学学院,杭州310058)
摘要 茜草科(Rubiaceae)假盖果草属(Pseudopyxis)共3种,分布于中国和日本,其中胀节假盖果草(P.
monilirhizoma)为近年发表的新种,特产浙江凤阳山。在文献查阅、野外采集和标本整理鉴定的基础上,对产于日
浙江大学学报(农业与生命科学版)
本的P.depressa、P.heterophylla和特产于中国的胀节假盖果草形态性状进行主坐标分析(PCoA),并对茎的毛
被和种子微观形态进行扫描电镜观察。结果表明:在PCoA散点图中,P.heterophylla和胀节假盖果草不能明
显区分,而P.depressa与这二者区别明显;P.heterophylla和胀节假盖果草在茎的毛被方面也无区别,但胀节
假盖果草种子表面除有纵肋外,还有瘤块状突起,与P.heterophylla有所区别。通过形态性状和种子微观形态
分析,认为假盖果属的已有3个种应归并为2个种,将胀节假盖果草作为P.heterophylla的亚种处理,组合为
P.heterophyllasubsp.monilirhizoma(Tao Chen)L.X.Ye,C.Z.Zheng &X.F.Jin.
关键词 胀节假盖果草;新组合;形态性状;扫描电镜;分类
Pseudopyxis Miq.(Rubiaceae,tribe Paederieae)
was described in 1867,with the type species P.
depressa Miq.as its only species.However,in the
protologue,Miquel also described a large number of
species including Oldenlandia heterophylla[1].Later,
Maximowicz[2] transfered O. heterophylla to
Pseudopyxis,as P.heterophylla (Miq.)Maxim.,
which was the second species in Pseudopyxis at that
time.Franchet et al.[3] described P.longituba
Franch.&Sav.from Japan,and it was subsequently
synonymized to P.depressa. Based upon the
basionym of Lysimachia quadriflora Petitm.,
Handel-Mazzetti[4]published Koidzumi’s combination
P.quadriflora (Petitm.)Koidz.,and it was also
reduced to the synonymy of P.heterophylla[5].Until
that time,Pseudopyxis comprised only the two
mentioned species, and was considered to be
distributed from central to southern Honshu and
Shikoku of Japan[6-7].Based on the materials from
China,Zheng[8] recognized P.heterophylla from
Zhejiang Province firstly,and it was accepted by Lo[9]
in a later time.Because Chinese materials exhibited
diference from Japanese ones,Chen[5]recognized three
species in the genus,describing the Chinese plants as a
new species,P.monilirhizoma Tao Chen.
During recent visits to the herbaria of KYO
(Herbarium of University Museum,Kyoto Univeristy),
TI(Herbarium of University Museum,University
of Tokyo)and OSA(Herbarium of Osaka Museum
of Natural History)in April 2005and August
2009,the senior author examined and compared
the colections of Pseudopyxis.The folowing
morphological characters were used to circumscribe
Pseudopyxis for purposes of this inquiry:stem
covered with pilose pubescence; calyx
campanulate,5-ridged,5-lobed;corola tubular-
funnelform or thinly tubular-funnelform
(salverform),5-lobed;stamens 5;style with 4-5
stigmatic lobes;capsule opening by an apical lid;
and seeds minutely longitudinal ridged and
striped[1,7,10-11].Based on statistical analysis of
morphological characters and on direct observation
of specimens,we recognize the Chinese materials
of Pseudopyxis as distinct from the Japanese P.
heterophyllaat the subspecific rank,as described
below.
1 Materials and methods
Al specimens of P.monilirhizoma that we
studied from China were housed in the herbaria of
HTC(Herbarium of Hangzhou Normal University,
formerly Hangzhou Teacher’s Colege)and HZU
(Herbarium of Zhejiang University).The specimens
of P.depressaand P.heterophyllafrom Japan were
housed in KYO,TI and OSA.Eighteen sheets(each a
separate individual)of P.heterophylla,23sheets of
P.depressa and 14sheets of Chinese materials,al
with flowers,were measured and analyzed(Table 1).
Ordination using principal coordinate analysis
(PCoA) was conducted on 18 morphological
characters,including three ordinal characters,12
binary characters, and three quantitative
characters (corola length,corola width,and
flower number;Table 1).PCoA was implemented
in MVSP version 3.13bon data gathered from al
55specimens.
Stem indumentum and seed morphology of P.
depressa,P.heterophylla and P.monilirhizoma
were observed under scanning electron microscope
(SEM).Stems and seeds of these three species
were al gathered from herbarium specimens.The
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stems and seeds were cleaned in 70%ethanol for
30min.After air-dried,the stems and seeds were
mounted on studs using double-sided adhesive
tape,and coated with a layer of gold.The coated
materials were observed and photographed under a
XL-30ESEM.
Table 1 Variation of the diagnostic characters of P.depressa,P.heterophyllaand P.monilirhizoma
Taxon P.depressa P.heterophylla P.monilirhizoma
Height(cm) <15(0)[23] >15(1)[18] <15(0)[14]
Rhizome nodes Inconspicuous(0)[23] Inconspicuous(0)[18] Tuberous(1)[14]
Indumentum of stem Pilosulous throughout(1)[23] Pilosulous in two rows(0)[18] Pilosulous in two rows(0)[14]
Stem branch Absent(0)[23] Present(1)[18] Absent(0)[14]
Dimorphic leaves Absent(0)[23] Present(1)[18] Present(1)[14]
Leaf shape on upper stem Broadly ovate(2)[23] Oblong-ovate(0)[12],ovate(1)[6] Ovate(1)[1],broadly ovate(2)[13]
Indumentum adaxialy Densely pubescent(1)[23] Sparsely pubescent(0)[18] Sparsely pubescent(0)[14]
Corola shape Tubular-funnelform(0)[23] Salverform(1)[18] Salverform(1)[14]
Corola color White(0)[1],pink(1)[22] White(0)[18] White(0)[3],pink(1)[11]
Corola length(mm) 17.26±0.45[23] 7.44±0.16[18] 10.20±0.26[14]
Corola diameter(mm) 10.48±0.42[23] 7.09±0.21[18] 11.48±0.31[14]
Indumentum on calyx Densely pubescent(1)[23] Sparsely pubescent(0)[18] Sparsely pubescent(0)[14]
Stamen vs.corola Concealed(0)[23] Exserted(1)[18] Exserted(1)[14]
Filament Separate from corola(0)[23] Adnate to base of corola lobes(1)[18] Adnate to base of corola lobes(1)[14]
Indumentum of corola lobes Glabrous(0)[23] Glabrous(0)[18] Ciliate(1)[14]
Indumentum of inside corola Pubescent(1)[23] Glabrous(0)[18] Glabrous(0)[14]
Indumentum of outside corola Pubescent(1)[23] Glabrous(0)[18] Pubescent(1)[14]
Flower number 1.17±0.08[23] 3.44±0.29[18] 2.86±0.23[14]
The values of quantitative characters measured from specimens/individuals are shown as mean± SD (standard deviation).The other
values are morphological characters,which are divided into gradual status.The values in brackets correspond to the assigned value,and those
in square brackets correspond to the number of measured individuals.
2 Results
2.1 PCoA of morphological characters
The first two PCoA axes explained 81.8%and
13.4% of the total morphological variations
respectively(Fig.1).Individuals of P.depressa
(black dots in Fig.1)separated distinctly from P.
heterophylla (triangles)and P.monilirhizoma
(diamonds). The separation between P.
heterophyllaand P.monilirhizoma was not clear.
2.2 SEM micromorphology of stem indumentum
The stem indumentums of P.depressa,P.
heterophylla and P. monilirhizoma were al
pilosulous.Stem pubescences in P.heterophylla
and P.monilirhizoma were aligned in two rows,
whereas P.depressa had even stem pubescent
(Fig.2).
2.3 SEM micromorphology of seed
Both P.heterophylla and P.monilirhizoma
had globose seeds,but with diferent surface
sculpturing.The surface sculpture of P.heterophylla
was minutely longitudinal ridged and densely
verrucose,and the ridges were undulate(Fig.3D-F).
The surface sculpture of P.monilirhizoma was also
minutely longitudinal ridged,however,it was not
verrucose but nubbed(Fig.3A-C).The seeds of P.
depressa were long-ovoid with longitudinal ridges and
deep sulcules,and there were sparse verrucae in the
sulcules(Fig.3G-I).
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Euclidean distance was used in axis 1and 2to
represent the PCoA case scores. Axis 1
(horizontal)explains 81.8%of the total variation,
and axis 2 (lengthways)explains 13.4% of the
total variation.
Fig.1 Scatter plots of principal coordinate analysis
(PCoA)of P.depressa (black dots),P.
heterophylla (triangles ) and P.
monilirhizoma(diamonds)
(A-B):P.monilirhizoma,from X.F.Jin 0830,HTC;(C-D):P.heterophylla,from M.Togashi s.n.,KYO;(E-F):P.
depressa,fromT.Tagagi 44967,KYO.
Fig.2 Stem indumentum of P.depressa,P.heterophyllaand P.monilirhizomaunder SEM
3 Taxonomic treatments
According to above reasons,we recognize P.
monilirhizomaas a subspecies of P.heterophylla,
and propose a combination as folow.
3.1 Description
Pseudopyxis heterophylla (Miq.)Maxim.,
Bul.Acad.Imp.Sci.Saint-Pétersbourg 29:175.
1883.Basionym:Oldenlandia heterophylla Miq.,
Ann.Mus.Bot.Lugduno-Batavi 3:109.1867.
TYPE:Japan,without precise locality,Keiske s.
n.(holotype,L).
1a)Pseudopyxis heterophylla (Miq.)Maxim.
subsp.heterophylla.
1b)Pseudopyxis heterophylla (Miq.)Maxim.
subsp.monilirhizoma(Tao Chen)L.X.Ye,C.Z.
Zheng & X.F.Jin,stat. & comb.nov.
Basionym:Pseudopyxis monilirhizoma Tao Chen,
Edinburgh J.Bot.64:304,Fig.1.2007.TYPE:
China,Zhejiang,Longquan County,Mt.Fengyang,
Shuikou,altitude 1 600m,21July 1972,Zhejiang
Med.Fl.Exped.2634 (holotype,A!;isotype,
HZU!).Fig.4.
Herbs perennial,rhizomes slender,ligneous,
creeping, with conspicuously tuberous nodes;
nodes 2-4;internodes 1-11cm long,stems 6-
15cm,quadrate,pilosulous in two interpetiolar
lows.Leaves 2-4pairs,membranous;lowest 1-
2pairs scale-like,blades obovate-spatulate,(1.5
-4)mm × (1.5-3)mm,upper pairs rhombic-
ovate to broadly ovate,(1-4)cm × (0.9-3)
cm,acute or obtuse at apex,cuneate at base,
entire and ciliate at margin,both adaxialy and
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(A-C):P.monilirhizoma,from X.F.Jin 0830,HTC;(D-F):P.heterophylla,fromG.Koidzumi s.n.,KYO;(G-I):
P.depressa,fromG.Murata 44812,KYO.(A,D,G):Seed shape;(B-C,E-F,H-I):Surface sculpture.
Fig.3 Seed micromorphology of P.depressa,P.heterophyllaand P.monilirhizomaunder SEM
A:Natural population in Mt.Fengyang,Zhejiang,China;
B:Natural habitat in Mt.Fengyang,Zhejiang,China;C:Flower,
showing corola lobes,stamens and style;D:Flower,showing corola
shape and calyx.Scale bar=5cm in(A),1cm in(C)and(D).
Fig.4 Photographs of P.monilirhizoma
abaxialy sparsely pilosulous,densely pilosulous
on midribs;petioles 0.7-2.5cm long,sparsely
pilosulous. Stipules triangular, 3-lobed,
persistent. Inflorescences 2-6-flowered;
peduncles ca.5mm long.Calyx campaniform,5-
lobed;lobes triangular-ovate or ovate,acute at
apex,sparsely pubescent,(2-2.5)mm ×ca.1
mm,ca.5 mm long in fruiting,patent;tubes
ridged,ridges sparsely pubescent.Corola pink,
rarely white, thinly tubular-funnelform
(salverform),9-12mm long,9-13.5mm in
diameter,5-lobed;tubes 6-8 mm long;lobes
ovate-lanceolate,(6-7)mm × (2-2.5)mm,
acuminate at apex,externaly glabrous,sparsely
pubescent internaly and at margin.Stamens 5,
exserted,adnate to base of corola lobes;filaments
6-7mm long,glabrous;anthers yelow,linear.
Style glabrous,14-16mm long,exserted;stigma
4-5-lobed.Capsules subglobose,ridged,ca.5mm
in diameter;seeds globose or broadly elipsoid,grayish
black, minutely longitudinal ridged and striped.
Flowering and fruiting from June to August.
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3.2 Distribution and habitat
P.heterophyllasubsp.monilirhizomais known
from Shuikou and Fengyan Temple of Mt.Fengyang,
Longquan County of Zhejiang in eastern China.It is
found on rocks or along stream banks in forest
understories at 1 450mto 1 600mabove sea level.
The known distribution is somewhat limited.
3.3 IUCN Red List Category and Criteria
According to IUCN Red List Category and
Criteria (IUCN 2001),P.heterophylla subsp.
monilirhizoma should be considered criticaly
endangered(CE),and there is need for the future
conservation.Based on botanical trips,two closed
populations have been found in Mt.Fengyang and
the habitats are wel preserved.However,its area
of distribution is less than 2 km2 and in two
populations no more than 3 000individuals occur.
3.4 Additional specimens examined
Zhejiang in China:Longquan County, Mt.
Fengyang,Shuikou,altitude 1 600m,along stream
banks in forest understories,10July 2009,S.L.Liu
s.n.(fl.,HTC),the same locality,21June 1976,
C.Z.Zheng 8307(fl.,HZU),30June 2011,X.F.
Jin 2758(fl.,HTC),2759(fl.,HTC),2760(fl.,
HTC),25 August 2004,X.F.Jin 0830 (fr.,
HTC);Fengyang Temple,on rocks along stream,
altitude 1 450m,1July 2009,X.F.Jin & H.W.
Zhang s.n.(fl.,HTC),the same locality,2July
2011,X.F.Jin 2761 (fl.,HTC),2762 (fl.,
HTC),15Aug.2011,X.F.Jin 2763(fr.,HTC).
3.5 Key to taxa of Pseudopyxis
1a.Stems pilosulous throughout;corola pink,rarely white,tubular-funnelform;filaments short,concealed within corola;seeds
elipsoid,with longitudinal ridges and sulcules. 1.P.depressa Miq.…………………………………………………………………
1b.Stems pilosulous in two rows;corola pink or white,salverform;filaments exserted from corola;seeds globose or broadly
elipsoid,with longitudinal ridges and stripes.
2a.Rhizomes slender,nodes inconspicuous;stems branched;corola 6-9mm long,white,glabrous on both surfaces.
2a.P.heterophylla(Miq.)Maxim.subsp.heterophylla.………………………………………………………………
2b.Rhizomes slender,with tuberous nodes;stems not branched;corola 9-12mm long,pink or white,internaly pubescent.
2b.P.heterophyllasubsp.monilirhizoma(Tao Chen)L.X.Ye,C.Z.Zheng &X.F.Jin.……………………
References:
[1] Miquel F A G.Prolusio florae Japonicae.Annales Musei Botanici
Lugduno-Batavi,1867,3:91-209.
[2] Maximowicz C J.Diagnoses des nouveles plantes asiatiques.V.
Bulletin de L’Academie Impériale des Sciences de St.-
Pétersbourg,ser.3,1884,29:51-228.
[3] Franchet A,Savatier L.Enumeratio Plantarum in Japonia
Sponte Crescentium,Vol.2.Paris:Imp.Simox Racox et Comp,
1878:391-392.
[4] Handel-Mazzetti H. A revision of the Chinese species of
Lysimachia,with a new system of the genus.Notes from the
Royal Botanic Garden,Edinburgh,1928,16:51-121.
[5] ChenT.Observations and a new species in the genus Pseudopyxis
(Rubiaceae).Edinburgh Journal of Botany,2007,64(3):
303-309.
[6] Ohwi J.Flora of Japan.Tokyo:Shibundo,1956:1088.
[7] Iwatsuki K.Flora of Japan,IIIa.Tokyo:Kodansha,1993:
209-240.
[8] 郑朝宗.假盖果草属Pseudopyxis Miq:中国茜草科的一个新记
录属.杭州大学学报:自然科学版,1981,28(1):116-118.
Zheng C Z.Pseudopyxis Miq.,a newly recorded genus of
Rubiaceae from China.Journal of Hangzhou University:
Natural Science Edition,1981,28(1):116-118.(in Chinese with
English abstract)
[9] 罗献瑞.中国植物志:第71(2)卷.北京:科学出版社,1999:
153-155.
Lo H S.Flora Republicae Popularis Sinicae:Vol.71(2).
Beijing:Science Press,1999:153-155.(in Chinese)
[10] 郑朝宗.浙江植物志:第6卷.杭州:浙江科学技术出版社,
1993:93-134.
Zheng C Z.Flora of Zhejiang:Vol.6.Hangzhou:Zhejiang
Science and Technology Publishing House,1993:93-134.(in
Chinese)
[11] Wu Z Y,Raven P H,Hong D Y.Flora of China,Vol.19.
Beijing:Science Press and St.Louis:Missouri Botanical Garden
Press,2011:57-368.
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