全 文 ：毛茛科金莲花族和升麻族细胞学的比较研究

杨 亲 二
(中国科学院植物研究所系统与进化植物学开放研究实验室　北京 100093)
Cytology of the tribe Trollieae and of the tribe Cimicifugeae
in the Ranunculaceae:a comparative study
YANG Qin_Er
(Laboratory of Systematic and Evolutionary Botany , Inst itute of Botany , the Chinese Academy of Sciences , Beijing 100093)
Abstract　Cytology of the tribe Trollieae and of the tribe Cimicifugeae in the Ranunculaceae was
comparatively investigated.Caltha palustris L.is found to represent a polyploid series with 2n =32
(4x), 48 (6x)and 64 (8x)in NW Yunnan , with the tetraploid cytotype being commoner and
showing obvious karyotypic variation among the populations.The chromosomes of Caltha , Calath-
odes , Megaleranthis and Trollius are basically similar in size among the genera , all medium_sized
and belonging to the R_type.The close affinity of Calathodes , Megaleranthis and Trollius is sup-
ported by both cytological and palynological evidence.The karyotypes of Beesia , Anemonopsis ,
Souliea , Cimicifuga and Actaea are very similar among the genera but remarkably different from
those of Caltha , Calathodes , Megaleranthis and Trollius with respect to the chromosome size and
chromosome morphology.From cytological data it is clear that Beesia has closer affinity with
Anemonopsis , Souliea , Cimicifuga and Actaea than with Caltha , Calathodes , Megaleranthis and
Trollius , and therefore may find its better position in the tribe Cimicifugeae.
Key words　Calathodes Hook .f.et Thoms.;Beesia Balf .f et W .W.Smith ;Trollieae Langlet;
Cimicifugeae Torr.et Gray;Ranunculaceae;Chromosome;Systematic position
1　Introduction
The tribe Trollieae Langlet of the subfamily Helleboroideae in the Ranunculaceae includes four
genera:Caltha , Calathodes , Trollius and Megaleranthis , and the tribe Cimicifugeae Torr.et Gray
in the same subfamily includes five genera:Beesia , Souliea , Anemonopsis , Cimicifuga and Actaea
(Wang , 1979).In a very recent classification of the Ranunculaceae , Tamura(1990 , 1995)placed
Caltha , Calathodes , Trollius and Megaleranthis in the subtribe Calthinae Benth .of the tribe
Helleboreae , Helleboroideae , while Beesia in the subtribe Beesiinae Tamura , and Anemonopsis ,
Souliea , Cimicifuga and Actaea in the tribe Cimicifugeae.It is clear that Wang s viewpoint on the
systematic position of the genus Beesia is different from Tamura s.As emphasized by Tamura
(1995), karyological characters are of the utmost importance in the phylogenetic classification of the
Ranunculaceae.Although the chromosome number and karyotypes of all the above genera have been
reported (e.g.Kurita , 1957 , 1965;Kawano et al ., 1966;Hasegawa , 1969;Emura , 1970a ,
1970b;Hasegawa &Feng , 1991;Yang , 1995 , 1998 , 1999a , 1999b), the data are quite scat-
　2001-05-15收稿 , 2001-08-08收修改稿。
基金项目:中国科学院资源与环境重点项目资助课题。
植 物 分 类 学 报　40(1):52 ～ 65(2002)
Acta Phytotaxonomica Sinica
tered in literature.In order to get an exact and whole picture of the karyological characters of the
above genera and thereby have a better understanding of their relationships , it is important to make
a cytological observation on these genera using same methods.In this paper , all the above genera
but Megaleranthis , a monotypic genus endemic to southern Korea , and Anemonopsis , a monotypic
genus endemic to Japan , are cytologically investigated.
2　Material and Methods
All the plants studied were collected in the field from Yunnan , Gansu 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.Ac-
tively growing root tips were pretreated in 0.1% colchicine at about 20℃ for three 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 two
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).
The voucher specimens were deposited in the Herbarium of Institute of Botany , the Chinese A-
cademy of Sciences (PE).
3　Results
3.1 　Caltha L.
Caltha is a small genus of about 12 species in the world (Tamura , 1995).Four species have
been recorded in this genus from China(Wang , 1979).One species , C.palustris L.was here ex-
amined.
Caltha palustris with several varieties is very widely distributed in the northern hemisphere and
shows great morphological variation (Smit , 1973;Tamura , 1995).Its chromosome number has
been reported in various countries (Leoncini , 1951 , 1952;Maugini , 1953 , 1957;Reese , 1954;
Wcislo , 1967;Malik &Mary , 1970;Kootin_Sanwu &Woodell , 1969 , 1970;Nishikawa , 1983 ,
1987;Chrtkov &Jarolimov , 1999).Polyploid series(2n=32 , 48 , 56 , 64 , 88), aneuploidy as
well as the occurrence of B chromosomes were reported for various populations.The Chinese materi-
al , however , has never been cytologically investigated.
Twelve populations were examined (Table 1).Eight were found to be tetraploids with 2n=4x
=32 (Figs.1 ～ 10 , all the results are not shown), one hexaploid with 2n=6x=48 (Figs.11 ～
12), and three octoploids with 2n=8x=64 (Figs.13 ～ 14 , only the result of the Wutoudi popula-
tion from Lijiang , Q.E.Yang &H.Z.Kong 98_471 , is shown).Neither aneuploidy nor B chro-
mosomes have been observed in all these populations.
The karyotypes of the tetraploid cytotype showed obvious variation among populations.The 32
chromosomes in the Tieciliang population from Tewo , Gansu could be quite well arranged into 8
groups of 4 homologues (Fig.6).The karyotype consisted of 20 median_centromeric(m), 8 sub-
median_centromeric (sm)and 4 subterminal_centromeric (st)chromosomes and was formulated as
2n =20m+8sm+4st.Those in the two Dali populations (Fig.7 , only the result of the Huadianba
1 期 杨亲二:毛茛科金莲花族和升麻族细胞学的比较研究 53　
Table 1　Chromosome counts and source of material studied
Taxon Provenance Voucher 2n
Caltha palustris
Tieciliang , Tewo , Gansu
Huadianba , Cangshan Mt., Dali , Yunnan
Xiaohuadianba , Cangshan Mt., Dali , Yunnan
Pantiange , Weixi , Yunnan
Kangpu , Weixi , Yunnan
Kena , Weixi , Yunnan
Napahai , Zhongdian , Yunnan
Tuguancun , Zhongdian , Yunnan
Wenhai , Yulongshan Mt., Lijiang , Yunnan
Wutoudi , YulongshanMt., Li jiang , Yunnan
Yuhu , Yulongshan Mt., Lijiang , Yunnan
Baimangxueshan Mt., Deqen , Yunnan
H.Z.Kong 98_79
Q.E.Yang 94_01
Q.E.Yang 94_18
Q.T.Zhang 94_05
Q.E.Yang &H.Z.Kong 98_438
Q.E.Yang &H.Z.Kong 98_186
X.Gong 92_26
Q.E.Yang &H.Z.Kong 98_540
Q.E.Yang &H.Z.Kong 98_503
Q.E.Yang &H.Z.Kong 98_471
Q.E.Yang &H.Z.Kong 98_464
Q.E.Yang 94_81
32
32
32
32
32
32
32
48
32
64
64
64
Calathodes oxycarpa Huadianba , Cangshan Mt., Dali , Yunnan Q.E.Yang 025 16
Trollius yunnanensis Yulongshan Mt., Lijiang , Yunnan X.Gong 92_13 16
T.farreri Yakou , Datong , Qinghai Q.E.Yang 95_32 16
T.ranunculoides Qiajin, Xunhua , Qinghai Q.E.Yang 95_14 16
Beesia calthifolia
Tieciliang , Tewo , Gansu
Kangpu , Weixi , Yunnan
Bisanggu , Zhongdian, Yunnan
Tianci , Zhongdian , Yunnan
H.Z.Kong 98_76
Q.E.Yang &H.Z.Kong 98_451
Q.E.Yang &H.Z.Kong 98_220
Q.E.Yang &H.Z.Kong 98_132
16
16
16
16
Souliea vaginata
Tieciliang , Tewo , Gansu
Xianrendong , Zhongdian , Yunnan
Hongshan , Zhongdian , Yunnan
Baimangxueshan Mt., Deqen , Yunnan
Wutoudi , YulongshanMt., Li jiang , Yunnan
H.Z.Kong 98_75
Q.E.Yang &H.Z.Kong 98_111
Q.E.Yang &H.Z.Kong 98_332
Q.E.Yang &H.Z.Kong 98_386
Q.E.Yang &H.Z.Kong 98_476
16
16
16
16
16
Cimicifuga yunnanensis Baimangxueshan Mt., Deqen , Yunnan Q.E.Yang &H.Z.Kong 98_408 16
Actaea asiatica Xianrendong , Zhongdian , Yunnan Q.E.Yang &H.Z.Kong 98_230 16
population , Q.E.Yang 94_01 , is shown)and the three Weixi populations (Fig.8 , only the result
of the Pantiange population , Q .T.Zhang 94_05 , is shown)could be roughly arranged into 8
groups of 4 homologues but there existed obvious heteromorphy between the two pairs of homologues
in the fifth and seventh groups.The karyotypes of the two Dali populations were formulated as 2n=
18m+6sm+8st and those of the three Weixi populations as 2n =16m+8sm+8st respectively.The
karyotype of the Napahai population from Zhongdian (Fig.9)and that of the Wenhai population
from Lijiang(Fig.10)were very similar to each other and both characterized by having more st or
t chromosomes and quite obvious heteromorphy between the two pairs of homologues in the eighth
group.The karyotype of the former was formulated as 2n=8m+6sm+14st+4t and that of the lat-
ter as 2n=6m+4sm+22st , respectively.
Most of the chromosomes of the hexaploid population could be arranged into groups of 6 homo-
logues except for the last two groups (Fig.12).The karyotype was formulated as 2n=31m+11sm
+6st.
The chromosomes of the octoploid population could be arranged into groups of 8 homologues ex-
cept for the last three groups (Fig.14).The karyotype was formulated as 2n =40m+22st+2t.
54　 植　物　分　类　学　报 40 卷
Figs.1～ 5　Photomicrographs of metaphase chromosomes in five populations of Caltha palustris 　1.Tieciliang population f rom Tewo ,
S Gansu (H.Z.Kong 98_79);2.Huadianba population from Dali , NW Yunnan (Q.E.Yang 94_01);3.Pantiange population
f romWeixi , NW Yunnan(Q.T.Zhang 94_05);4.Napahai population f rom Zhongdian , NW Yunnan(X.Gong 92_26);5.Wenhai
population from Li jiang ,NW Yunnan(Q.E.Yang &H.Z.Kong 98_503)(all × 1940).
1 期 杨亲二:毛茛科金莲花族和升麻族细胞学的比较研究 55　
Figs.6～ 10　Karyotypes of five populations of Caltha palustris　1.Tieci liang population from Tewo , S Gansu(H.Z.Kong 98_79);
2.Huadianba population f rom Dali , NW Yunnan(Q.E.Yang 94_01);3.Pantiange population from Weixi , NW Yunnan(Q.T.
Zhang 94_05);4.Napahai population from Zhongdian , NW Yunnan (X.Gong 92_26);5.Wenhai population f rom Lijiang , NW
Yunnan (Q.E.Yang &H.Z.Kong 98_503)(all × 1940).
56　 植　物　分　类　学　报 40 卷
Figs.11～ 12　Photomicrograph of metaphase chromosomes(Fig.11)and karyotype(Fig.12)of the Tuguancun population of Caltha
palustris from Zhongdian , NW Yunnan (Q.E.Yang &H.Z.Kong 98_540)(all × 1940).
It should be noted that although these populations show so great variation in the chromosome
number and karyotype , gross_morphologically they can not be distinctly differentiated from each oth-
er.
3.2 　CalathodesHook.f.et Thoms.
Calathodes is a small genus of four species (Wang , 1979;Wang , 2000).One species , C .
oxycarpa Sprague , was examined.The chromosomes were counted to be 2n=16 , consisting of 8 sm
and 8 st ones(Figs.15 , 19).A pair of small satellites was observed on the short arms of the sixth
chromosome pair.The karyotype was formulated as 2n =8sm+8st(2sat).The chromosomes were
medium_sized , very similar in size to those of Caltha , Megaleranthis (Lee &Yeau , 1985)and
Trollius (see below)and should be categorized as the Ranunculus_type (R_type), rather than the
Thalictrum_type(T_type)as suggested by Zhang (1982).
3.3 　Trollius L.
Trollius is a genus of 31 species in the world (Tamura , 1995).Sixteen species have been
recorded in China (Wang , 1979).Three species , T.yunnanensis (Franch.)Ulbr., T .farreri
Stapf and T.ranunculoides Hemsl., were examined here.Their chromosomes were all counted to
be 2n=2x=16 (Figs.16 ～ 18).Their karyotypes are also basically similar to each other , only
1 期 杨亲二:毛茛科金莲花族和升麻族细胞学的比较研究 57　
Figs.13～ 14　Photomicrograph of metaphase chromosomes (Fig.13)and karyotype(Fig.14)of the Wutoudi population of
Caltha palustris from Lijiang , NW Yunnan(Q.E.Yang &H.Z.Kong 98_471)(all × 1940).
with some minor differences among the species(Figs.20 ～ 22).On the short arms of the sixth chro-
mosome pair in T.farreri , a pair of medium_sized satellites was observed (Fig.21).The kary-
otype of both T .yunnanensis and T .ranunuculoides was formulated as 2n =4m+4sm+8st , and
that of T.farreri as 2n=4m+4sm+8st(2sat), respectively.
3.4 　Beesia Balf.f .et W.W.Smith
Beesia is a small genus of two species (Hsiao , 1979).Both of them , B .calthifolia (Max-
im.)Ulbr.and B .deltophylla C .Y.Wu ex Hsiao , are distributed in China.The latter is a
diploid with 2n =16 (Yang , 1995)and the former is of both diploid and tetraploid cytotypes
(Yang , 1999a).The four populations of B .calthifolia examined here (Table 1 , only the result of
the Tianci population from Zhongdian , Q.E.Yang &H.Z.Kong 98_132 , is shown)were all
found to be diploids with 2n=16 , consisting of 10 m , 2 sm , 2 st and 2 t chromosomes (Figs 23 ,
27).A pair of large satellites was observed on the short arms of the second and the third chromo-
some pairs , and a small satellite on the short arm of the 15th chromosome.The karyotype was for-
mulated as 2n=10m(4sat)+2sm+2st+2t(1sat).
58　 植　物　分　类　学　报 40 卷
Figs.15～ 22　Photomicrographs of metaphase chromosomes (Figs.15～ 18)and karyotypes(Figs.19～ 22)in one species of
Calathodes and three species of Trollius　15, 19.C .oxycarpa ;16 , 20.T.yunnanensis;17 , 21.T.farreri ;18 , 22.
T.ranunculoides (all × 1940).
3.5 　Souliea Franch.
Souliea is a monotypic genus with the only species S .vaginata (Maxim.)Franch.(Hsiao ,
1979).
The chromosomes of the five populations examined (Table 1 , only the result of the Wutoudi
population from Lijiang , Q.E.Yang &H.Z.Kong 98_476 , is shown)were all counted to be 2n
1 期 杨亲二:毛茛科金莲花族和升麻族细胞学的比较研究 59　
Figs.23～ 26　Photomicrographs of metaphase chromosomes in Beesia calthifolia (Q.E.Yang &H.Z.Kong 98_132)
(Fig.23), Souliea vaginata (Q.E.Yang &H.Z.Kong 98_476)(Fig.24), Cimicifuga yunnanensis(Fig.25)and
Actaea asiatica (Fig.26)(all × 1940).
60　 植　物　分　类　学　报 40 卷
Figs.27～ 30　Karyotypes in Beesia calthifolia (Q.E.Yang &H.Z.Kong 98_132 , Fig.27), Souliea vaginata (Q.E.Yang &
H.Z.Kong 98_476 , Fig.28), Cimicifuga yunnanensis (Fig.29)and Actaea asiatica (Fig.30)(al l × 1940).
=2x=16 (Fig.24), confirming the previous report on material from W Sichuan by Hasegawa and
Feng (1991).The karyotype consisted of 10m , 2sm , 2st and 2t chromosomes (Fig.28).A pair of
large satellites was observed on the short arms of the third chromosome pair , and a small satellite on
the short arm of the 15th chromosome.On the long arms of the seventh chromosome pair two sec-
ondary constrictions were observed.The karyotype was formulated as 2n =10m(2sat)+2sm+2st
(2sec)+2t(1sat).The present result of karyotype analysis is very similar to that reported by
Hasegawa and Feng (1991)but they did not find secondary constrictions on the long arms of the
seventh chromosome pair.
3.6 　Cimicifuga L.
Cimicifuga is a genus of 18 species in the world (Tamura , 1995).Eight species have been
recorded in China(Hsiao , 1979)and mostly examined cytologically (Wang et al ., 1994;Yang ,
1999b).Just like C.yunnanensis (Fig.25)here examined , they are diploids with 2n =16 with
1 期 杨亲二:毛茛科金莲花族和升麻族细胞学的比较研究 61　
Fig.31　A map showing the dist ribution of the polyploid series of Caltha palustris in NW Yunnan.
the exception of C .foetida , in which both diploid and tetraploid cytotypes have been reported
(Yang , 1999b).If putting satellites and secondary constrictions out of consideration , the karyotypes
are very similar among the species , usually consisting of 10m , 2sm , 2st and 2t chromosomes , as is
shown in Fig.29.
62　 植　物　分　类　学　报 40 卷
3.7 　Actaea L.
Actaea is a small genus of about eight species in the world(Tamura , 1995).Two species , A .
asiatica Hara and A.erythrocarpa Fisch., have been recorded in China(Hsiao , 1979)and both
cytologically examined (Wang et al., 1994;Yang , 1998).The karyotype of this genus is different
from those of Beesia , Anemonopsis , Souliea and Cimicifuga mainly by the absence of short arms of
the eighth chromosome pair , as is shown in Figs.26 and 30.The karyotype of A .asiatica reported
by Kurita(1957), and Wang et al .(1994), and that of the North American species A .pachypo-
da Ell.reported by Utech (1988), represent a mistake of observation.All these authors mistook the
eighth chromosome pair as being median_centromeric.The karyotype of A.asiatica examined here
was formulated as 2n =10m(6sat)+2sm+2st+2T.
4　Discussion
4.1　Caltha palustris is found to represent a polyploid series of 2n=32(4x),48(6x)and 64(8x)
in NW Yunnan (Fig.31).The tetraploid cytotype seems to be commoner than the hexaploid and oc-
toploid ones.Neither aneuploidy nor B chromosomes have been observed in the populations exam-
ined.The karyotype of the tetraploid cytotype shows obvious variations among the populations and
falls roughly into three types:the Tewo type represented by the Tieciliang population from Tewo , S
Gansu , the Weixi type represented by the Weixi and Dali populations from NW Yunnan , and the
Zhongdian type represented by the Napahai population of Zhongdian and the Wenhai population of
Lijiang.In the Tewo type , the chromosomes can be quite well arranged into 8 groups of 4 homo-
logues , and in the Weixi type the chromosomes can be roughly arranged into 8 groups of 4 homo-
logues , though with obvious heteromorphy between the two pairs of homologues in the fifth and sev-
enth groups , while in the Zhongdian type there are more st or t chromosomes in the karyotype , with
obvious heteromorphy between the two pairs of homologues in the eighth chromosome group.More
populations need be studied for a better understanding of the geographical distribution pattern of the
three types.
4.2　The chromosomes of Caltha , Calathodes , Megaleranthis and Trollius are basically similar in
size among the genera , all medium_sized and belonging to the R_type.It is unacceptable that Zhang
(1982)separated Calathodes from the tribe Trollieae to establish an independent tribe Calathodeae ,
regarding chromosomes of the genus as the Thalictrum_type (T_type)because of their small size ,
and that Fu (1990)transferred it from the subfamily Hellebroideae to the subfamily Thalictroideae.
In fact , the present results have shown that the chromosomes of Calathodes are somewhat larger than
those of Trollius .Obviously , when examining the chromosomes of Calathodes oxycarpa Zhang
(1982)may have ignored the work of Kurita (1965)on the karyotype of C.polycarpa Ohwi from
Taiwan.Based on his karyotype analysis on C .polycarpa and six species of Trollius Kurita(1965)
pointed out that the karyotypes of these two genera mainly consist of st chromosomes and are so simi-
lar in asymmetry that from a cytotaxonomical viewpoint it is unreasonable to treat Calathodes as a
separate genus from Trollius.Their close affinity has also been strongly supported by palynological
data (Lee , 1989 , 1992;Lee &Blackmore , 1992;Xi et al ., 1993).The pollen grains of both
Calathodes and Trollius have striate ornamentation on the exine surface.In the Ranunculaceae ,
such pollen grains have been found only in these two genera and their close ally , Megaleranthis , a
monotypic genus endemic to South Korea.Nevertheless , the present results have shown that the
karyotype of Calathodes is different from that of Trollius by the absence of m chromosomes in the
1 期 杨亲二:毛茛科金莲花族和升麻族细胞学的比较研究 63　
chromosome complement , so these two genera is cytologically distinguishable in spite of the high
similarity in their karyotypic asymmetry.Calathodes should be maintained as an independent genus
in the tribe Trollieae.
4.3 　The karyotypes of Beesia , Souliea , Cimicifuga and Actaea are very similar to each other if
putting satellites and secondary constrictions out of consideration , all consisting of five pairs of large
m , one pair of sm , one pair of st , one pair of t or T chromosomes.The monotypic genus
Anemonopis endemic to Japan also has the same chromosome number and karyotype as the above four
genera (Kurita , 1957).Tamura (1990 , 1995)did not place Beesia together with Anemonopis ,
Souliea , Cimicifuga and Actaea into the tribe Cimicifugeae but established for it a monotypic sub-
tribe , Beesiinae , in the tribe Helleboreae , and considered that Beesia may occupy a transitional po-
sition between Caltha and the tribe Cimicifugeae.The present results have shown that cytologically
Beesia is very distinctly distinguishable from Caltha , Calathodes , Megaleranthis and Trollius with
respect to the chromosome size and chromosome morphology but very similar to Anemonopis ,
Souliea , Cimicifuga and Actaea , and therefore may find its better position in the tribe Cimi-
cifugeae.
Acknowledgments　I would like to express my sincere thanks to Profs.Liu Shang_Wu and He Ting_Long , Institute of
Plateau Biology , the Chinese Academy of Sciences , Mr.Gong Xun and Prof.Zhang Qi_Tai , Kunming Institute of
Botany , the Chinese Academy of Sciences , and Dr.Kong Hong_Zhi , Laboratory of Systematic and Evolutionary
Botany , Institute of Botany , the Chinese Academy of Sciences , for their help in collecting living material.I thank
Prof.Gu Zhi_Jian , Kunming Institute of Botany , the Chinese Academy of Sciences , for kindly leaving his research fa-
cility at my disposal.
References
Chrtkov A , Jarolimov V , 1999.Cytotaxonomical study of Caltha palustris.Preslia , 71:349 ～ 360
Emura (Hasegawa)K M , 1970a.A cytotaxonomical study on the Eurasiatic species of the genus Cimicifuga (1).J
Jpn Bot , 45:297 ～ 308
Emura (Hasegawa)K M , 1970b.A cytotaxonomical study on the Eurasiatic species of the genus Cimicifuga (2).J
Jpn Bot , 45:362 ～ 377
Fu D Z , 1990.Phylogenetic consideration on the subfamily Thalictroideae (Ranunculaceae).Cathaya , 2:181～ 190
Hasegawa I , Feng P , 1991.Cytological study of Chinese monotypic genus Souliea (Ranunculaceae).J Phytogeogr
Taxon , 39:96
Hasegawa K M , 1969.Cytotaxonomic notes on Cimicifuga foetida from Bhutan.J Jpn Bot , 44:76～ 84
Hsiao P K , 1979.Cimicifugeae.In:Flora Reipublicae Popularis Sinicae.Vol 27.Beijing:Science Press.88 ～ 106
Kawano S , Kane J M , Iltis H H , 1966.Chromosome morphology of the North American species of Actaea(Ranuncu-
laceae).Canad J Bot , 44:1231 ～ 1234
Kootin_SanwuM , Woodell S R J , 1969.Supernumerary chromosomes in Caltha palustris.Chromosomes today , 2:
192～ 196
Kootin_Sanwu M , Woodell S R J , 1970.The cytology of Caltha palustris(sensu lato):distribution and behaviour of
the chromosome races.Caryologia , 23:225 ～ 239
Kurita M , 1957.Chromosome studies in Ranunculaceae Ⅱ .Karyotype of the subtribe Cimicifuginae.Rep Biol Inst
Ehime Univ , 1:11 ～ 17
Kurita M , Chromosome studies in Ranunculaceae ⅩⅩⅢ .Mem Ehime Univ , sect Ⅱ , ser B(Biol), 5:89～ 95
Lee S T , 1989.Palynological evidence for the relationships between Megaleranthis saniculifolia and Trollius species.
Pollen et Spores , 29:173～ 185
Lee S T , 1992.Palynological relationships among Calathodes and its relative genera.Kor J Plant Tax , 22:23 ～ 31
Lee S T , Blackmore S , 1992.A palynotaxonomical study of the genus Trollius (Ranunculaceae).Grana , 31:81 ～
100
Lee Y N , Yeau S H , 1985.Taxonomic characters of Megaleranthis saniculifolia Ohwi(Ranunculaceae).Kor J Plant
64　 植　物　分　类　学　报 40 卷
Tax , 15:127～ 131
Leoncini M L , 1951.Biotipi cariologici e sistematici di Caltha in Italia.Caryologia , 3:336 ～ 350
Leoncini M L , 1952.Nuove osservazioni cariologiche sul gen.Caltha.Caryologia , 4:367～ 371
Levan A , Fredga K , Sandberg A A , 1964.Nomenclature for centromeric position on chromosomes.Hereditas , 52:
201～ 220
Malik C P , Mary T N , 1970.Cytological studies in Caltha palustris Linn.var.normalis Hook.Beitr Biol Pflanzen ,
47:63～ 70
Maugini E , 1953.Contributo alla conoscenza cariologica e sistematica del genere Caltha in Italia.Caryologia , 5:297
～ 305
Maugini E , 1957.Ricerche cito_sistematiche sul genere Caltha in Italia.Caryologia , 9:408 ～ 435
Nishikawa T , 1983.A new chromosome number of Caltha palustris var.enkoso Hara.J Jpn Bot , 58:319～ 320
Nishikawa T , 1987.Chromosome numbers and geographical distribution of Caltha (Ranunculaceae) in northern
Japan.Mem Nat Sci Mus(Tokyo), 20:91 ～ 101
Reese G , 1954.Euploidie , Aneuploidie und B_Chromosomen bei Caltha palustris L.Planta , 44:203～ 268
Smit P G , 1973.A revision of Caltha(Ranunculaceae).Blumea , 21:119～ 150
Tamura M , 1990.A new classification of the family Ranunculaceae 1.Acta Phytotax Geobot , 41:93 ～ 101
Tamura M , 1995.Ranunculaceae.In:Hiepko P ed.Die Natǖrlichen Pflanzenfamilien.Zwei Aufl , 17a(4).Berlin:
Duncker und Humblot
Utech F H , 1988.Correction:karyotype of Caulophyllum thalictroides.Ann Carnegie Mus , 57:255
Wang W T , 1979.Ranunculaceae.In:Flora Reipublicae Popularis Sinicae.Vol 27.Beijing:Science Press
Wang Z , 2000.Calathodes.In:Fu L G , Chen T Q , Lang K Y et al.eds.Higher Plants of China.Qingdao:Qing-
dao Publishing House.3:392～ 393
Wang X Q , Hong D Y , Li Z Y, 1994.A karyomorphological study of nine species in four genera of Ranunculaceae.
Cathaya , 6:43～ 56
Wcislo H , 1967.Karyological studies in Caltha palustris L.s.l.Acta Biologica Cracov , ser Bot , 10:1～ 23
Xi Y Z , Ning J C , Fu X P , 1993.Pollen morphology of the tribe Trollieae and its taxonomic significance.Cathaya ,
5:115～ 130
Yang Q E , 1995.On the chromosomes of Calathodes (Ranunculaceae) and its systematic position.Acta Phytotax
Sin , 33:453 ～ 460
Yang Q E , 1998.Does Actaea asiatica have the most symmetric and primitive karyotype in the Ranunculaceae?Acta
Phytotax Sin , 36:490～ 495
Yang Q E , 1999a.Correction of karyotype of diploid Beesia calthifolia and discovery of a tetraploid cytotype.Acta
Phytotax Sin , 37:1～ 9
Yang Q E , 1999b.Karyomorphology of four species in Cimicifuga(Ranunculaceae)from China , with some cytogeo-
graphical notes on C.foetida .Acta Phytotax Sin , 37:433～ 444
Yang Q E , Gu Z J , Sun H , 1995.The karyotype of Beesia deltophylla and its systematic significance.Acta Phytotax
Sin , 33:225 ～ 229
Zhang Z Y , 1982.Chromosome observation of three ranunculaceous genera in relation to their systematic positions.
Acta Phytotax Sin , 20:402 ～ 409
摘要　对毛茛科金莲花族 Trollieae和升麻族 Cimicifugeae的细胞学进行了比较研究。发现驴蹄草 Caltha
palustris L.在云南西北部形成一个多倍体系列(2n=32 , 48 , 64),四倍体细胞型(2n=4x=32)较为常见 ,
其核型有明显的居群间变异。驴蹄草属 Caltha 、鸡爪草属 Calathodes 、Megaleranthis以及金莲花属Trollius
的染色体在大小上基本相似 ,都属于中等大小的 R_型染色体。细胞学和花粉学证据都支持鸡爪草属与
Megaleranthis和金莲花属有较近的亲缘关系。铁破锣属 Beesia 、Anemonopsis 、黄三七属 Souliea 、升麻属
Cimicifuga 以及类叶升麻属Actaea 的核型彼此基本相似 , 在染色体大小和形态上都与驴蹄草属 、鸡爪草
属 、Megaleranthis以及金莲花属的核型明显有别。细胞学证据表明铁破锣属应是升麻族中的成员。
关键词　鸡爪草属;铁破锣属;金莲花族;升麻族;毛茛科;染色体;系统位置
(责任编辑　白羽红)
1 期 杨亲二:毛茛科金莲花族和升麻族细胞学的比较研究 65