全 文 ：Cytology of ten species in Anemone , one in Anemoclema
and six in Clematis (Trib.Anemoneae ,
　 Ranunculaceae)from China 　
YANG Qin_Er
(Laboratory of Systematic and Evolutionary Botany , Institute of Botany , the Chinese Academy of Sciences , Bei jing 100093 ,China)
Abstract　The somatic chromosome number and detailed chromosome morphology have been studied
in ten species of Anemone , one species of Anemoclema , and six species of Clematis , all from
China , namely Anemone davidii Franch.(2n=4x=32), A.stolonifera Maxim.(2n=2x=16),
A.flaccida Fr.Schmidt(2n=2x=14), A.rivularis Buch._Ham.(2n=2x=16), A.begoni-
ifolia Lévl.et Vant.(2n=2x=16), A.hupehensis Lem.f.alba W.T.Wang(2n=2x =16),
A.tomentosa (Maxim.)Péi(2n=2x=16), A.rupestris Hook.f.et Thoms.(2n=2x=14),
A.trullifolia var.colestina (Franch.)Finet et Gagnep.(2n =2x =14), A.trullifolia var.
holophylla Diels(2n=2x=14), A.demissa Hook.f.et Thoms.(2n=2x =14), Anemoclema
glaucifolium (Franch.)W.T.Wang (2n=2x =16), Clematis kockiana Schneid.(2n=2x=
16), C.rehderiana Craib.(2n=2x=16), C.ranunculoides Franch.(2n=2x=16), C.pu-
berula var.ganpiniana (Lévl.et Vant.)W.T.Wang(2n=2x=16), C.brevicaudata DC.(2n
=2x=16), C.chrysocoma Franch.(2n=2x=16).The species of x=8 in Anemone , and those
in Anemoclema and Clematis have very similar karyotypes which consist of five pairs of large medi-
an_centromeric(rarely one pair of which are submedian_centromeric)and three pairs of subterminal_
or terminal_centromeric chromosomes.
Key words　Ranunculaceae;Anemone L., Anemoclema (Franch.)W.T.Wang;Clematis L.;
Chromosome number;Karyotype
1　Introduction
Anemone L., with about 150 species in the world , and Clematis L., with about 240 species ,
are the two largest genera in the tribe Anemoneae of the Ranunculaceae(Tamura , 1995).These two
genera are very richly represented in China , where about 60 species in Anemone andmore than 100
species in Clematis L.have been recorded(Wang ,1979).
Cytological studies in Anemone and Clematis have been reported in various countries (for
Anemone:Moffett , 1932;Kurita , 1957a;Heimburger , 1959;Boraiah &Heimburger , 1964;
Baumberger , 1970;Bhattarai , 1989;and for Clematis:Meurman &Therman , 1939;Kurita ,
1957b , 1962;Pringle , 1971;Dennis , 1976;Dawson , 1993).The Chinese species in these two
genera , particularly in Anemone , however , are cytologically very poorly known.Only 18 species in
Clematis and one in Anemone from China have had reports of chromosome counts and chromosome
morphology (Gong et al., 1985;Zhang &He , 1990;Zhang , 1991;Yang et al., 1994).
Molecular data have shown that there are two large clades within Anemone , clade Ⅰ with the
basic chromosome number x=8 , and clade Ⅱ with the basic chromosome number x=7(Hoot et
　Received 4 September 2001;accepted 4 July 2002.
Supported by a Key Project Grant f rom the Chinese Academy of Sciences(Natural Resources and Environment).
植 物 分 类 学 报　40(5):396～ 405(2002)
Acta Phytotaxonomica Sinica
al., 1994;Ehrendorfer &Samuel , 2001).This indicates that the chromosome number is of impor-
tant significance in determining phylogenetic relationships of the species in Anemone.This paper is
to report the chromosome number and morphology of ten species in Anemone , one in Anemoclema
and six in Clematis.
2　Material and Methods
All the plants studied were collected in the field from Yunnan , Sichuan , Shanxi and Qinghai of
China(Table 1).They were cultivated in pots in the experimental garden of Kunming Institute of
Botany , the Chinese Academy of Sciences , before their root tips were harvested for the cytological
study.Actively growing root tips were pretreated in 0.1% colchicine at about 20℃ for 3 hr before
they were fixed in Carnoy I(glacial acetic acid∶absolute ethanol=1∶3)at 4℃ for 30 min , then
they were macerated in 1∶1 mixture of 1 mol/L hydrochloric acid and 45% acetic acid at 60℃ for
2 min , and stained and squashed in 1% aceto_orcein.
Karyotype formulas were based on the measurements of somatic chromosomes.The symbols
used to describe the karyotypes followed Levan et al.(1964):m=median_centromeric chromo-
some , sm=submedian_centromeric chromosome , st=subterminal_centromeric chromosome , and t=
terminal_centromeric chromosome.
The voucher specimens were deposited in the Herbarium of Institute of Botany , the Chinese A-
cademy of Sciences(PE).Taxonomic treatments and systematic arrangements of the species followed
Wang (1979 , 2000).
3　Results
3.1　Anemone L.
The chromosome counts obtained are given for 11 taxa in Table 1.Of these , two are in each of
sections Anemonanthea and Anemone , one in each of sections Stolonifera , Rivularidum , Begoni-
ifolia , Omalocarpus , and three in section Himalayicae.Some of these are illustrated in Figures 1&
2 and Figures 5 ～ 15 and show the chromosome morphology that is typical of the genus , with five
large pairs of median_centromeric(rarely one pair of which are submedian_centromeric), and three
pairs of subterminal_ or terminal_centromeric chromosomes.
Of the eleven taxa studied , one species , A.davidii , is tetraploid with 2n=32 based on x=
8 , five species , A.stolonifera , A.rivularis , A.begoniifolia , A.hupehensis f.alba , A.to-
mentosa , are diploid with 2n=16 based on x=8 , and five taxa , A.flaccida , A.rupestris , A.
trullifolia var.colestina , A.trullifolia var.holophylla and A.demissa are diploid with 2n=14
based on x=7.The chromosome counts in A.davidii , A.begoniifolia , A.rupestris , A.trulli-
folia var.colestina and A.trullifolia var.holophylla represent new records.Those in the remain-
ing species confirm the reports by previous workers(Kurita , 1957a;Heimburger , 1959;Baumberg-
er , 1970;Bhattarai , 1989).
The chromosomes of A.davidii can be arranged into 8 groups of 4 homologues(Fig.5).It is
noteworthy that the chromosomes of the fifth group are submedian_centromeric with the arm ratio up
to 2.6.The chromosome number of this species from Xinning , Hunan , was counted to be 2n=16
(Luo Yan , personal communication), indicating ploidy variation in this species.
The five species with 2n=16(Fig.6 , Figs.8 ～ 11)have very similar karyotypes , which con-
sist of five pairs of large median_centromeric(in Anemone stolonifera , the fifth pair being submedi-
YANG Qin_Er:Cytology of ten species in Anemone , one in Anemoclema and six in Clematis
No.5 (Trib.Anemoneae , Ranunculaceae)from China 397　
Table 1　Chromosome counts and source of material studied
Taxon Provenance Voucher 2n
Anemone L.
　Sect.Anemonanthea DC.
　　A.davidii Franch. Weixi , Yunnan Q.E.Yang &H.Z.Kong 98_450 32
　　A.stolonifera Maxim. Zhongdian , Yunnan Q.E.Yang &H.Z.Kong 98_110 16
　Sect.Stolonifera(Ulbr.)Juz.
　　A.flaccida Fr.Schmidt Dali , Yunnan
Weixi , Yunnan
Q.E.Yang 94_13
Q.E.Yang 94_73
14
14
　Sect.Rivularidium Jancz.
　　A.rivularis Buch._Ham. Lijiang , Yunnan
Dali , Yunnan
Zhongdian , Yunnan
Dawu , Sichuan
X.Gong 92_14
Q.E.Yang 94_23
X.Gong 92-57
Y.B.Luo 88
16
16
16
16
　Sect.Begoniifolia(Ulbr.)Tamura
　　A.begoniifolia Lévl.et Vant. Nanchuan , Sichuan Q.E.Yang &M.Feng 95_12 16
　Sect.Anemone
　　A.hupehensis Lem.
　　　 f.alba W.T.Wang Zhongdian , Yunnan
Dali , Yunnan
X.Gong 92_42
Q.E.Yang 94_12
16
16
　　A.tomentosa(Maxim.)Pé i Xunhua , Qinghai
Jishan , Shanxi
Q.E.Yang 95_05
Y.B.Luo 20
16
16
　Sect.Himalayicae(Ulbr.)Juz.
　　A.rupestris Hook.f.et Thoms. Zhongdian , Yunnan Q.E.Yang 94_105 14
　　A.trullifolia Hook.f.et Thoms.
var.colestina(Franch.)
　Finet et Gagnep.
var.holophyl la Diels
Dali , Yunnan
Weixi , Yunnan
Zhongdian , Yunnan
Q.E.Yang 94_09
Q.E.Yang 94_68
Q.E.Yang &H.Z.Kong 98_528
14
14
14
　Sect.Omalocarpus DC.
　　A.demissa Hook.f.et Thoms. Zhongdian , Yunnan
Weixi , Yunnan
X.Gong 92_41
Q.E.Yang &H.Z.Kong 98_185
14
14
Anemoclema(Franch.)W.T.Wang
　　A.glaucifolium (Franch.)
　　　W.T.Wang Lijiang , Yunnan Q.E.Yang 93_201 16
Clematis L.
　Sect.Viorna(Reichb.)Prantl
　　Subsect.Connatae Koehne
　　C.kockiana Schneid. Yunlong , Yunnan Q.E.Yang 93_02 16
　　C.rehderiana Craib. Zhongdian , Yunnan Q.E.Yang &H.Z.Kong 98_175 16
Düqün , Yunnan Q.E.Yang &H.Z.Kong 98_405 16
　　C.ranunculoides Franch. Yanyuan, Sichuan
Zhongdian , Yunnan
Q.E.Yang 93_09
Q.E.Yang &H.Z.Kong 98_297
16
16
　　Subsect.Vitalbae Prantl
　　C.puberula Hook.f.et Thoms.var.gan-
piniana(Lévl.et Vant.)W.T.Wang Lijiang , Yunnan Q.E.Yang 94_143 16
　　C.brevicaudata DC. Düqün , Yunnan Q.E.Yang 98_384 16
　Sect.Cheiropsis DC.
　　C.chrysocoma Franch. Lijiang , Yunnan Q.E.Yang 94_150 16
398　 植 物 分 类 学 报　Acta Phytotaxonomica Sinica Vol.40
Figs.1～ 4　Photomicrographs of chromosomes of four species in the Ranunculaceae(all ×1940)　1.Anemone stolonifera Maxim.;
2.A.demissa Hook.f.et Thoms.(Q.E.Yang &H.Z.Kong 98-185);3.Anemoclema glaucifolium (Franch.)W.T.Wang;
4.Clematis puberula var.ganpiniana (Lé vl.et Vant.)W.T.Wang
YANG Qin_Er:Cytology of ten species in Anemone , one in Anemoclema and six in Clematis
No.5 (Trib.Anemoneae , Ranunculaceae)from China 399　
Figs.5～ 9　Karyotypes in five species of Anemone L.(all ×1940)　5.A.davidii Franch.;6.A.stolonifera Maxim.;
7.A.flaccida Fr.Schmidt(Q.E.Yang 94-13);8.A.rivularis Buch._Ham.(X.Gong 92-57);
9.A.begoniifolia Lévl.et Vant.
400　 植 物 分 类 学 报　Acta Phytotaxonomica Sinica Vol.40
Figs.10～ 14　Karyotypes in f ive taxa of Anemone L.(all ×1940)　10.A.hupehensis Lem.f.alba W.T.Wang(X.Gong
92-42);11.A.tomentosa(Maxim.)Pé i(Q.E.Yang 95-05);12.A.rupestris Hook.f.et Thoms.;13.A.trullifolia
var.colestina (Franch.)Finet et Gagnep.(Q.E.Yang 94-09);14.A.trullifolia var.holophylla Diels.
YANG Qin_Er:Cytology of ten species in Anemone , one in Anemoclema and six in Clematis
No.5 (Trib.Anemoneae , Ranunculaceae)from China 401　
an_centromeric), and three pairs of subterminal_or terminal_centromeric chromosomes.On the short
arms of one or two terminal_centromeric chromosomes small satellites are attached.
Anemone stolonifera has been previously recorded to occur in China s Heilongjiang and Tai-
wan , and in Korea as well as in Japan.Its occurrence in Yunnan is a new record of its distribution ,
indicating that it is of a disjunctive distribution pattern.Its karyotype is different from those of the
other four species studied with 2n=16 by having the fifth chromosome pair being submedian_cen-
tromeric.
3.2　Anemoclema (Franch.)W.T.Wang
Anemoclema is a monotypic genus endemic to NW Yunnan and SWSichuan , including the only
species A.glaucifolium (Franch.)W.T.Wang.This species has a chromosome number of 2n=
16(Figs.3 , 16).Its karyotype is very similar to those of Anemone species with 2n=16 , consist-
ing of five pairs of large median_centromeric , one pair of subterminal_centromeric and two pairs of
terminal_centromeric chromosomes.Two small satellites are attached on the short arms of one termi-
nal_centromeric chromosome pair.The chromosome number is reported for the first time.
Figs.15～ 16　Karyotypes in Anemone demissa Hook.f.et Thoms.(Q.E.Yang &H.Z.Kong 98-185)(Fig.15)and
Anemoclema glaucifolium (Franch.)W.T.Wang (Fig.16)(both ×1940)
3.3　Clematis L.
The chromosome counts obtained are given for six species in Table 1.Of these , five are in sec-
tion Viorna , and one in section Cheiropsis.Some of these are illustrated in Figure 4 and Figures 17
～ 22 and show the chromosome morphology that is typical of the genus , with five pairs of large me-
dian_centromeric , one pair of subterminal_centromeric and two pairs of terminal_centromeric chromo-
somes.All the species have the same chromosome number of 2n=16 based on x=8.The chromo-
some counts in C.kockiana , C.rehderiana , C.ranunculoides and C.chrysocoma represent new
records.
402　 植 物 分 类 学 报　Acta Phytotaxonomica Sinica Vol.40
Figs.17～ 22　Karyotypes in six species of Clematis L.(all ×1940).17.C.kockiana Schneid.;18.C.rehderiana Craib.
(Q.E.Yang &H.Z.Kong 98-175);19.C.ranunculoides Franch.(Q.E.Yang 93-09);20.C.puberula var.
ganpiniana(Lévl.et Vant.)W.T.Wang;21.C.brevicaudata DC.;22.C.chrysocoma Franch.
4　Discussion
4.1　Basic chromosome numbers in Anemone L.
The chromosome counts obtained confirm previous reports that there are two basic chromosome
YANG Qin_Er:Cytology of ten species in Anemone , one in Anemoclema and six in Clematis
No.5 (Trib.Anemoneae , Ranunculaceae)from China 403　
numbers , x=7 and 8 , in the genus Anemone(Langlet , 1932;Moffett , 1932;Gregory , 1941;Ku-
rita , 1957a;Heimburger , 1959;Baumberger , 1970).Of the ten species studied , six species ,
which represent four sections , have x=8 , and four species , which represent three sections , have x
=7.The most conspicuous difference of karyotypic constitution between the x=8 and x=7 species
lies in that there is one more pair of median_centromeric chromosomes but two less pairs of subtermi-
nal_centromeric or terminal_centromeric chromosomes in the x=7 species.
4.2　High similarity of karyotypes within Clematis
The uniformity of chromosome number in the six species studied , with karyotypes so similar
that no discernible differences are evident , agrees with the findings of previous workers studying
Chinese and non_Chinese species of Clematis (Kurita , 1957b , 1962;Pringle , 1971;Dennis ,
1976;Gong et al., 1985;Zhang &He , 1990;Zhang , 1991;Dawson , 1993).AlthoughGong et
al.(1985)reported the presence of a heteromorphic submedian_centromeric chromosome pair in
some of the Chinese species they karyotyped , the generalized karyotype of the genus remains the
same , consisting of five pairs of median_centromeric , one pair of subterminal_centromeric and two
pairs of terminal_centromeric chromosomes.It is noteworthy that Clematis is a morphologically very
diverse and geographically wide_ranging genus but its chromosome complement is remarkably stable.
4.3　High similarity of karyotypes among Anemone species of x =8 , Anemoclema and
Clematis
The chromosomes of Anemone , Anemoclema and Clematis are large and of the Ranunculus_
type described by Langlet(1927 , 1932).Chromosome morphology is very similar among the Ane-
mone species of x=8 , and those of Anemoclema and Clematis;their karyotypes all consists of five
pairs of metacentric(median_ and submedian_centromeric)and three pairs of acrocentric(subtermi-
nal_ and terminal_centromeric)chromosomes.This indicates that the three genera are closely relat-
ed.
Acknowledgments　I would like to express my sincere thanks to Mr.Fang Zhen_dong of Gesang Flower Co.Ltd.,
Zhongdian County , Yunnan , and Drs.Luo Yi_bo and Kong Hong_zhi of Laboratory of Systematic and Evolutionary
Botany , Institute of Botany , the Chinese Academy of Sciences , for their help in collecting living materials.I am
grateful to Professor Gu Zhi_jian of Kunming Institute of Botany , the Chinese Academy of Sciences , for kindly leaving
his research facilities at my disposal.
References
Baumberger H , 1970.Chromosomenzahlbestimmungen und Karyotypanalysen bei den Gattungen Anemone , Hepatica
und Pulsatilla.Ber Schweiz Bot Ges , 80:17～ 96
Bhattarai S , 1989.Karyomorphological studies in four species of Anemone.Cytologia , 54:709～ 713
Boraiah G , Heimburger M , 1964.Cytotaxonomic studies on new world Anemone(section Eriocephalus)with woody
rootstocks.Can J Bot , 42:891～ 922
Dawson M I , 1993.Contributions to a chromosome atlas of the New Zealand flora — 31.Clematis(Ranunculaceae).
New Zealand J Bot , 31:91 ～ 96
Dennis W M , 1976.Chromosome morphology of Clematis , subsection Viornae (Ranunculaceae).Can J Bot , 54:
1135～ 1139
Ehrendorfer F , Samuel R, 2001.Contributions to a molecular phylogeny and systematics of Anemone and related gen-
era(Ranunculaceae_Anemoninae).Acta Phytotax Sin , 39:293 ～ 307
Gong W Z , Long Y I , Li M X , 1985.Karyotype studies on Clematis from Beijing , China.JWuhan Bot Res , 3:371～ 379
Gregory W C , 1941.Phylogenetic and cytological studies in the Ranunculaceae Juss.Trans Am Phil Soc , 31:443 ～
521
Heimburger M , 1959.Cytotaxonomic studies in the genus Anemone.Can J Bot , 37:587 ～ 612
Hoot S B , Reznicek A A , Palmer J D , 1994.Phylogenetic relationships in Anemone(Ranunculaceae)based on mor-
phology and chloroplast DNA.Syst Bot , 19:169～ 200
404　 植 物 分 类 学 报　Acta Phytotaxonomica Sinica Vol.40
Kurita M , 1957a.Chromosome studies in Ranunculaceae Ⅰ .Karyotypes of the subtribe Anemoninae.Rep Biol Inst
Ehime Univ , 1:1 ～ 10
Kurita M , 1957b.Chromosome studies in Ranunculaceae Ⅳ.Karyotypes of the three genera , Paeonia , Nigella and
Clematis.Rep Biol Inst Ehime Univ , 2:9～ 16
Kurita M , 1962.Chromosome studies in Ranunculaceae.ⅩⅩ .Karyotypes of ten species.Mem Ehime Univ , Sect
Ⅱ(Sci), Ser B(Biol), 4:395 ～ 401
Langlet O , 1927.Beiträge zur Zytologie der Ranunculazeen.Svensk Bot Tidskr , 21:1 ～ 17
Langlet O , 1932. ber Chromosomenverhä ltnisse und Systematik der Ranunculaceae.Svensk Bot Tidskr , 26:381 ～
400
Levan A , Fredga K , Sandberg A A , 1964.Nomenclature for centromeric position on chromosomes.Hereditas , 52:
201～ 220
Meurman O , Therman E , 1939.Studies on the chromosome morphology and structural hybridity in the genus Clema-
tis.Cytologia, 10:1 ～ 14
Moffett A A , 1932.Chromosome studies in Anemone Ⅰ .A new type of chiasma behaviour.Cytologia , 4:26～ 37
Pringle J S , 1971.Taxonomy and distribution of Clematis , section Atragene(Ranunculaceae), in North America.
Brittonia , 23:361～ 393
Tamura M , 1995.Ranunculaceae.In:Hiepko P ed.Die Natǜrlichen Pflanzenfamilien , zwei Aufl , 17a(4).Berlin:
Duncker and Humblot
Wang W T , 1979.Trib.Anemoneae DC.In:Flora Reipublicae Popularis Sinicae.28:1 ～ 241
Wang W T , 2000.Notes on the genus Clematis(Ranunculaceae)(Ⅰ).Acta Phytotax Sin , 38:305～ 336
Yang Q E , Luo Y B , Hong D Y , 1994.A karyotypic study of six species in the Ranunculaceae from Hunan , central
China.Guihaia , 14:27～ 36
Zhang Y L , 1991.Chromosome studies on 7 species of Clematis in China.J Wuhan Bot Res , 9:107 ～ 111
Zhang Y L , He S Y , 1990.Chromosome studies on 6 species of Clematis in China.JWuhan Bot Res , 8:115 ～ 121
国产毛茛科银莲花族十七种植物的细胞学研究
杨 亲 二
(中国科学院植物研究所系统与进化植物学重点实验室　北京 100093)
摘要　研究了国产毛茛科银莲花族 Trib.Anemoneae 17 种植物的染色体数目和核型。 10 种银莲花属
Anemone L.植物中 , 1 种(西南银莲花 A.davidii)为 x=8 的四倍体(2n=4x=32), 5种 (匍枝银莲花 A.
stolonifera 、草玉梅 A.rivularis 、卵叶银莲花 A.begoniifolia 、水棉花 A.hupehensis f.alba 、大火草 A.tomen-
tosa)为 x=8 的二倍体(2n=2x=16), 4 种(鹅掌草 A.flaccida 、湿地银莲花 A.rupestris 、蓝匙叶银莲花
A.trullifolia var.colestina 、拟条叶银莲花 A.trullifolia var.holophylla 、展毛银莲花 A.demissa)为 x=7 的
二倍体(2n=2x=14)。罂粟莲花 Anemoclema glaucifolium 为 x=8 的二倍体。6 种铁线莲属 Clematis L.植
物(滇川铁线莲 C.kockiana 、长花铁线莲 C.rehderiana 、毛茛铁线莲 C.ranunculoides 、扬子铁线莲 C.
puberula var.ganpiniana 、短尾铁线莲 C.brevicaudata 、金毛铁线莲 A.chrysocoma)均为 x=8 的二倍体。银
莲花属中 x=7的种类的核型彼此十分相似 , 均由 6对大型具中部着丝点的染色体和 1对具端部着丝点
的染色体组成;x=8的二倍体种类的核型与罂粟莲花属和铁线莲属植物的核型十分相似 , 均由 5对大型
具中部着丝点和 3对具端部或近端部着丝点的染色体组成。
关键词　毛茛科;银莲花属;罂粟莲花属;铁线莲属;染色体数目;核型
YANG Qin_Er:Cytology of ten species in Anemone , one in Anemoclema and six in Clematis
No.5 (Trib.Anemoneae , Ranunculaceae)from China 405