全 文 ：西北植物学报 2003, 23( 7): 1091— 1097
Acta Bot . Boreal .-Occident. Sin.
文章编号: 1000-4025( 2003) 07-1091-07
基于形态学资料星叶草属系统位置的探讨
任　毅1 ,李智军 2 ,胡正海 3
( 1陕西师范大学 生命科学学院 , 西安 710062; 2太白山国家级自然保护区管理局 , 陕西眉县 722300;
3西北大学生命科学学院 ,西安 710069)
摘　要: 基于以往的形态学研究资料 ,对星叶草属与毛茛目其它成员进行了比较 . 结果表明 ,该属在一些基本的或
原始的性状上与其它毛茛目成员相似 ,特别是它与毛茛科一些植物幼苗间的相似性表明 ,这个属应该属于毛茛目
的成员 ,而且有可能通过幼态成熟的方式起源于前毛茛目植物 . 星叶草属与其它毛茛目成员间的区别 ,尤其是在维
管系统和胚胎发育方面的区别表明 ,星叶草属在毛茛目内发生以后 ,可能是沿着一条特化和减化的途径发展的 .
关键词: 星叶草属 ;形态学 ;起源 ;系统位置
中图分类号: Q949　　　文献标识码: A
Approaches to the systematic position of Circaeaster based
on the morphological data
REN Yi1 , L I Zhi-jun2 , HU Zheng-hai3
( 1 Col leg e of Life Science, Sh aanxi No rm al University, Xi an 710062, China; 2 Taibaishan National Nature Reserve Managem ent
Bureau, Meixian, Sh aanxi 722300, China; 3 Ins ti tute of Botany, Northw es t Universi ty, Xi an 710069, China)
Abstract: Circaeaster was compared with other Ranunculales ( sensu Takhtajan, 1980) based on the mo r-
pho logical data in the present paper. The resul ts w ere show ed as fo llow s. Circaeaster is simi lar to other
Ranunculales in some basic o r prima ry characteristics. In pa rticular, the similarities betw een Circaeaster
and some seedlings of Ranunculaceae show ed that Circaeaster should be a member of Ranunculales and
might o riginate neo teinicly f rom pro-Ranunculales. The di fferences, especially in vascular system and em-
bryogeny , betw een Circaeaster and o ther Ranunculales show tha t Circaeaster probably developed in a spe-
ciali zed and reduced clade af ter i t s o rigination in Ranunculales.
Key words: Circaeaster ; morpho logy; o rigin; systema tic po sitio n
　　
1　 Int roduction
Circaeaster was established by M ax imowicz
( 1881) based on the only species C. agrest is Max-
im. which is an annual herb dist ributed in Yun-
nan, Sichuan, Tibet , Shaanxi, Gansu, Qinghai,
Xinjiang o f China and Nepal , Sikkim and Bhutan.
Many bo tanists paid g reat at tention to this g enus
because it ha s some special characteristics such as
the open dichotomous leaf v enation. A g roup of re-
searchers in Bio logy Depa rtment of No rthw est U-
收稿日期: 2003-05-06;修改稿收到日期: 2003-05-15
基金项目:国家自然科学基金 (No. 30130030和 39870068)资助 . Th e sub ject w as supported by th e National Nature Science Foundation
of China(No. 30130030and 39870068) .
作者简介:任　毅 ( 1959- ) ,教授 ,博士生导师 . 西北大学生物系植物学专业 1996年毕业的博士 .
niv ersity of China have been studying this g enus
since 1987 by using the wi ld material ( Ren et al .
1998; Ren and Hu, 1998, 1995; Hu et al . 1990;
Hu and Yang, 1987) . Based on those results and
also the resul ts of the predecesso rs, the taxonomic
posi tio n and the phy log eny of this mono typic g enus
w ere approached in the present paper.
2　 Histo ry of s tudy
2. 1　 Studies on the characteristics of Circaeaster
Except fo r many mo rphological observ ations
and descriptions ( Ren and Hu, 1995; Wu and Ku-
bi tski , 1993; W ang , 1980; Handel-Mazzetti ,
1931; Oliv er, 1895; Hooker, 1890; Bentham and
Hooker, 1883; Maximow icz, 1881) , the studies of
the characteristics of Circaeaster were mainly fo-
cused on the ana tomy and embryo logy. The fi rst
anatomic description was made by Sco tt ( see Oliv-
er, 1895) . He described that the st ructure of the
stem th roughout is tha t of a dia rch hypoco ty l,
there is sca rcely any dif ference betw een stem and
main roo t, a small amount of seconda ry wood and
phloem is fo rmed on ei ther side of the diarch
xylem-plate, a sing le bundle enters each co tyle-
don. Metcalf and Chalk ( 1950 ) described the
st ructure o f this g enus briefly and the ranuncula-
ceous stomata of the leaf w ere repo rted. Cheadle
( 1953) observ ed that the vessel perfo ration of this
g enus is simple, but Chen and Li ( 1990) observ ed
the vessels wi th simple, scala riformed and of
Gnetalean type perfo ra tion. Junell ( 1931) found
that the anther has tw o pol len sacs, the pol len
tube is mesogamous, there are tw o ovules but only
one is ferti le and the ovule is uni tegmic and tenuin-
ucellate, the embryo sac is Po lygonum-type, the
endosperm is cellular, of Adoxa-type and di fferen-
tiated, the seeds have no testa , and the basic num-
ber of chromosome is 15. These characteristics are
all special in Ranunculales.
Foster ( 1971, 1970, 1968, 1966, 1963) paid
g rea t at tention to this g enus and he found tha t the
node is unilacunar, the co ty ledon trace div erg es
f rom a co rresponding pro tox ylem pole, the co tyle-
don has two closely approxima ted st rands of tra-
cheids, w hich may sepa ra te as tw o very sho rt end-
ing s below the blunt apes, there are fiv e types of
anastomosis in the dicho tomous venation, and the
anastomosis is formed by the approx imate veins.
There is secondary xy lem and phlo em in the upper
pa rt o f the hypocotyl. The tepals, the stamens and
the pistals are 1-traced. Li and Dong ( 1987) re-
po rted tha t there are 2～ 3 layers of cambium like
cells which do no t have the characters o f th e cam-
bium cells betw een the primary xy lem and phloem.
There is no vascula r t ransfo rm betw een the roo t
and the stem, like general dico ty ledons, and the
fi rst and the second leaf t races a re opposi te. The
fur ther embryological studies of Hu and Yang
( 1987) show that the developing process of the
embryo of this genus is simila r to that of
Caryophyllad-type but the second or the thi rd cell
under the terminal cell divides longi tudinally first
w hen the proembryo is 8-celled.
The anatomic studies o f Hu et al . ( 1990) on
the f low er and the inf lorescent show that the inflo-
rescent is a cyme, the vascular cy linder o f the pedi-
cel is a protostele wi th mesarch xylem , like that of
roo t and stem of pteridophy tes, and the ovule is
amphi tropous. Ren and Hu ( 1995) studied the
vegetative org ans o f this genus again. They co r-
rected some mistakes of predecesso rs and found
tha t there is a small amount of second xylem and
phlo em in the root and the hypoco ty l. The phloem
of the cotyledonary t race is from the branches of
two vascular bundles, the xy lem of the co tyledon
is simple and V-shaped, the stomata of the co tyle-
don is on the upper surface and of ranunculous-
type. The phyl lotax is t ransfo rms from opposi te to
alternate. Ren et al . ( 1998, 1997) studied the ve-
na tion of this g enus and found many degenerating
commissural v eins of the anastomosis.
Erdtman ( 1952) and Hu et al . ( 1990) repo rt-
ed that the po llen of this t ricolpa te under the light
microscope. Nowicke and Skvarla ( 1982) repo rted
tha t the tectum of this g enus, like that of Kingdo-
nia and some genera of Ranunculaceae and Berberi-
daceae, is a compound layer of st riae under the
scanning electron micro scope. Kong and Yang
1092 西　北　植　物　学　报 23卷
( 1997) reported that the ka ryomo rpholo gical char-
acteristics of interphase nuclei and prophase chro-
mosomes o f Circaeaster are simila r to that of King-
donia. Oxelman and Liden ( 1995) and Hoo t and
Crane ( 1995) studied the nuclear and chlo roplast
DN A. Their result show ed the similarity betw een
Circaeaster and Kingdonia. Ba rthlo t t and Theisen
( 1995) observ ed that th e epicuticular w ax ul tra-
st ructure o f Circaeaster is Berberis-type tha t is
common in Ranunculi flo rae. Ren and Hu ( 1998)
observ ed tha t there are mo re than two roo t hai r
zones in Circaeaster .
2. 2　 Viewpoint of taxonomic position and rela-
tionship of Circaeaster
Maximowicz ( 1881 ) included Circaeaster in
Chlo ranthaceae w hen he established this genus.
Bentham and Hooker ( 1883) and Hooker ( 1890)
follow ed M ax imowicz ( 1881) and considered that
Circaeaster is a v ery distinct genus in Chlo ran-
thaceae but wi th essential characteristics relating it
to Chlo ranthus, especially in i ts habi ta t, to some
specimens of C. japonicus Sieb. Post and Kuntze
( 1904) also include Circaeaster in Chlo ranthaceae
but t rea ted i t as a distinct subfamily , Circaeast-
ereae.
Oliv er ( 1895) considered tha t Circaeaster al-
lied perhaps to Anemoneae of Ranunculaceae be-
cause the character o f a pendulous orthot ropous
ovule is one of the mo re notewo rthy characteris-
tics. Hal lier ( 1903) included Circaeaster in Ranun-
culaceae. Diels ( 1932) st rong ly insisted that Cir-
caeaster should be regarded as the most ex t remely
reduced member o f Ranunculaceae and should be
closely related to Kingdonia because these tw o
genera have a so li tary , o rtho t ropous, pendulous
ovule and dicho tomous venation of the leaves.
Janchen ( 1949 ) accepted the opinion o f Diels
( 1932) , and included Circaeaster in Kingdoniinae
o f Clematidae of Ranunculoideae tog ether wi th
Kingdonia. Wang ( 1980) and Thorne ( 1974) ac-
cepted this t rea tment but included these two gener-
a in Kingdonioideae. The point o f view o f Cir-
caeaster being related wi th Kingdonia was suppor t-
ed by the studies of pollen mo rpholog y ( Now icke
and Skvarla, 1982 ) , cladistics ( Loconte and
Estes, 1989) , nuclea r and chlo roplast DN A (Oxel-
man and Liden, 1995; Hoot and Crane, 1995) ,
ka ryomorpho logy ( Kong and Yang , 1997 ) , and
the roo t morpho log y ( Ren and Hu, 1998) .
Hutchinson ( 1926) disag reed wi th the opinion
of Max imox icz ( 1881 ) and Hallier ( 1903) and
trea ted Circaeaster as a distinct fami ly, Circaeast-
eraceae, ev en though the idea of t reating this
genus as a fami ly had been put forw ard since the
genus was established ( Hallier, 1912; Po st and
Kuntze, 1904; Maximowic z, 1881) . The opinion
of Hutchinson ( 1926 ) w as approved by many
bo tanists ( Takh tajan, 1997, 1980, 1969; Tamura
1995, 1963; Wu and Kubi tzki, 1993; Tho rne, 1992,
1983; Cronquist, 1981, 1968; Dahlg ren, 1977;
Hutchinson, 1959; Handel-Mazzet ti , 1931 ) , and
w as suppor ted by many results of studies ( Kong
and Yang , 1997; Oxelman and Liden, 1995; Ren
and Hu, 1995; Hu and Yang , 1987; Now icke and
Skvarla, 1982; Foster, 1963; Junell, 1931) .
Circaea steraceae w as rega rded generally as a
mono typic family ( Takh tajan, 1997, 1980, 1969;
Tamura 1995, 1963; Wu and Kubi tzki, 1993;
Dahlg ren, 1977; Hutchinson, 1959, 1926; Han-
del-Mazzet ti, 1931) . Some authors included King-
donia in this fami ly ( Tho rne, 1983; Cronquist,
1981, 1968 ) . Circaeasteraceae w as generally in-
cluded in Berberidales ( Tho rne 1974; Hutchinson,
1959, 1926) o r Ranunculales ( Thorne 1992; Cron-
quist 1981, 1968; Takhta jan 1980, 1969; Dahlg ren,
1977) , and was considered to be related wi th Ra-
nunculaceae ( Takhta jan, 1980) . But recent ly Hoo t
and Crane ( 1995) considered tha t Circaeaster i s in
the same clade wi th Lardizabalaceae and Kingdonia
based on the nuclear and chlo roplast DN A data.
Takhta jan ( 1997) changed his opinion o f negating
the relationship betw een Circaeaster and Kingdonia
( Takh tajan, 1980, 1969 ) , included Circaea ster-
aceae to gether wi th Kingdoniaceae in a new o rder,
Circaeasterales, and considered that i t i s related
wi th Hydrastidaceae and Tribe Epimedieae of
Berberidaceae.
Gundersen ( 1950) considered that Circaeaster
10937期 任　毅 ,等: 基于形态学资料星叶草属系统位置的探讨 (英 )
should be included in Berberidaceae during this pe-
riod. But this opinion was no t accepted by any
bo tanist.
3　 Approach es of the relationship be-
tween Circaeaster and oth er mem-
bers of Ranunculales (sensu Tak-
h tajan 1980)
　　Circaeaster i s similar to the members o f Ra-
nunculales in the fo llowing aspects: 1) annual cy-
cle is similar to some Ranunculaceae ( excluding
Kingdonia , the same bellow ) ; 2) the vessel perfo-
ra tion is simi lar to some of Ranunculaceae,
Berberidaceae and La rdizabalaceae ( Ren and Hu,
1995; Chen and Li, 1990; Cheadle, 1953) ; 3) root
top wi th 3 g roups of ini tials, diarch primary xylem
and a small amount o f second g row th , a re simi lar
to some species of Ranunculaceae ( Ren and Hu,
1995; Tamura , 1964; Metcalfe and Chalk, 1950) ;
4) anomocy tic stomata of leaves ( Ren and Hu,
1995; M etcalfe and Chalk, 1950) ; 5) the change
o f the phy llotaxis is simi lar to some seedlings o r
y oung shoo ts of Ranunculaceae ( Ren and Hu 1995;
Tamura, 1963) ; 6) t rimerous f low er; 7) bisexual
f low er is similar to Ranunculaceae and Berberi-
daceae; 8) the distinct f low er parts ( pistil of
Berberidaceae) is pseudomonomerous; 9) 3-colpa te
and 2-celled po llen g rains are common in Ranuncu-
lales and the tectum is simi lar to some of Ranuncu-
laceae and Berberidaceae ( Nowicke and Skvarla,
1982, 1981; Erdtman, 1952) ; 10) 1-ovuled pisti l
is similar to some of Ranunculaceae and Sarg ento-
doxaceae; 11) the Polyg onum-type o f embryo sac
( Davis, 1966; Junell, 1931) ; 12) the persistent
antipodal cells ( Hu and Yang , 1987; Sast ri , 1969;
Bhanda ri 1968; Davis, 1966) ; 13) the achene is
simi lar to some o f Ranunculaceae; 14) V-shaped
xylem of the co tyledon and the petiole ( Ren and
Hu, 1995 ) is simila r to the xy lem o f Ranuncu-
laceae ( Tamura, 1964) ; 15) the base ch romosome
number and the cellular endosperm is simila r to
some Lardizabalaceae ( Davis, 1966; Bhatnagar,
1965; Swamy, 1953) , but the prima ry endo sperm
cell div ides longi tudinally in Circaeaster ( Hu and
Yang , 1987; Junel l, 1931 ) and transversely in
Lardizabalaceae ( Swamy, 1953) .
Except fo r the above-mentioned cha racteris-
tics, Circaeaster i s similar in the following aspects
to a few genera o r species o f Ranunculales: 1) no
mechanical ti ssue is simi lar to the aqua ticBatrachi-
um of Ranunculaceae ( Ren and Hu, 1995; Tamu-
ra, 1963 ) ; 2 ) the ex t remely elongated and X-
shaped xylem o f hypoco tyl i s similar to the
seedlings of a few species o f Ranunculaceae ( Ren
and Hu, 1995; Gu et al . , 1994, 1990) ; 3) no vas-
cular change betw een roo t and stem is simila r to
Ranunculus japonicus of Ranunculaceae ( Ren and
Hu, 1995; Gu et al . , 1990; Li and Dong , 1987) ;
4) the homogeneous mesophyll is similar to Hyd-
tastis of Ranunculaceae ( Ren and Hu, 1995; Tobe
and Keating , 1985) ; 5) w ith more than one ovule
in the early developing stag e but only one ferti le
ovule, i s simila r to Anemone obtusiloba and Adonis
aest ival is of Ranunculaceae ( Hu et al . , 1990;
Bhandari, 1968; Bhandari, 1963; Junell , 1931) ;
6) the open ca rpel in the early developing stag e in
Circaeaster ( Hu et al . , 1990; Junell, 1931) is sim-
i la r to Copt is of Ranunculaceae ( Tamura, 1965;
Kumazaw a, 1930) and Akebia quinata o f Lardiza-
balaceae ( Payne and Seago, 1968) but inCopt is the
open si tuation can retain the f rui t; 7) unitegmic
ovule is simi lar to a few Ranunculaceae ( Hu and
Yang , 1987; Bhandari , 1968; Junell, 1931; Ku-
mazaw a, 1930) ; 8) the embryo wi th tw o devel-
oped co ty ledons is similar to a few Ranunculaceae
( Engell , 1995; Hu and Yang , 1987; Jalan, 1963) ;
9) the unilacunar node is similar to some Ranuncu-
laceae ( Ren and Hu, 1995; Dé camps, 1979; Fos-
ter , 1963) but in these Ranunculaceae, the node is
2-traced.
Contrasting wi th above-mentioned facts, Cir-
caeaster di ffers from Ranunculaceae in the follow-
ing cha racteristics: 1) wi th more than tw o roo t
hai r zones ( Ren and Hu, 1998) ; 2) the ex t remely
sho rtened stem and inf lo rescent ( Ren and Hu,
1995; Hu et al. 1990; Foster, 1963) ; 3) the per-
sistent and 1-veined co tyledons ( Ren and Hu,
1995) ; 4) the st ructure of the node ( Ren and Hu,
1094 西　北　植　物　学　报 23卷
1995; Foster, 1963) ; 5) a sing le v ascular bundle
in the petiole ( Ren and Hu, 1995) ; 6) the open di-
cho tomous venation ( Ren et al . , 1997; Fo ster,
1963) ; 7) the persistent and un-veined tepals ( Hu
et al . , 1990) ; 8) a sing le v ascula r bundle in the
receptacle ( Hu et al . , 1990) ; 9) the anther wi th
tw o pollen sacs ( Junell , 1931 ) ; 10 ) the un-
branched ventral vascular bundle of the ca rpel
( Foster, 1963) ; 11) the hemitropous ovule and the
integument covers half of the nucella ( Hu et al . ,
1990; Hu and Yang , 1987 ) ; 12 ) mesogamous
pollen tube ( Junell , 1931) ; 13) the embryogeny
type ( Hu and Yang , 1987) ; 14) the Adoxa-type,
cellula r, thick-cell-w alled and dif ferentiated en-
dosperm ( Hu and Yang , 1987; Junell, 1931) ; 15)
the integument disappears and the seeds have no
testa ( Hu and Yang , 1987; Junell, 1931) .
We consider that there is no point in dis-
cussing the relationship betw een Circaeaster and
Kingdonia before the embryo log y of Kingdonia i s
fully revealed. How ever , mo re and more da ta
show that these monotypic g enera are clo sely simi-
lar to one ano ther ( Ren and Hu, 1998; Kong and
Yang, 1997; Oxelman and Liden, 1995; Hoo t and
Crane, 1995; Loconte and Estes, 1989; Now icke
and Skva rla, 1982) .
The above-mentioned analyses show tha t the
di fferences, especially the di fferences in the vascu-
lar system and the embryogeny , a re more obvious
than the simila ri ties betw een Circaeaster and o ther
Ranunculales, and that there are mo re speciali zed
and reduced characters in Circaeaster than in o ther
Ranunculales. The dif ferences o f characters, espe-
cially in the vascula r system and the embryo logy ,
betw eenCircaeaster and oth er Ranuncula les a re ob-
vious and show that they might be in di fferent ev o-
lutiona ry clade. Therefo re, w e ag reed wi th
Hutchinson ( 1926) to t rea t this g enus as a distinct
family. On the o ther hand, the similarities in some
basic o r primary cha racters, especia lly the distinct
f low er pa rts that a re ra ther ranunculous, betw een
Circaeaster and Ranunculaceae and o ther families
o f Ranunculales show tha t they a re connected to
one another.
4　 Approaches of the po ssible o rigin
of Circaeaster
　　 There are di fferent opinions about the possible
origin of Circaeaster . Oliv er ( 1895) considered i t
as a deg raded fo rm and allied perhaps to
Anemoneae of Ranunculaceae. Diels ( 1932) con-
sidered it as an ex t remely reduced fo rm of Ranun-
culaceae. Janchen ( 1949) also considered i t as a
secondari ly simplified g roup, deriv ed f rom an
Anemoninae-like ancesto r. But Hutchinson ( 1959)
regarded it as a very reduced relativ e of the
Podophyllaceae o r Berberidaceae. Fo ster ( 1963 )
disag reed w ith tho se opinions and he regardedCir-
caeaster as a g roup wi thout an obvious al ly.
We considered that the above-mentioned opin-
ions a re baseless. It is dif ficult to imagine that Cir-
caeaster , w hich has characteristics of mesogamy,
the embryogeny like Caryophyllad-type, the Adox-
a-type and cellular endosperm, can be deriv ed f rom
Anemoninae o f Ranunculaceae, Podophy llaceae
and Berberidaceae, which have the characteristics
of porogamy , Onagrad-type embryogeny, nuclea r
endosperm. Furthermo re, the pisti ls of Podophy l-
laceae and Berberidaceae are pseudomonomerous.
Compared wi th o ther members of Ranuncu-
lales, Circaeaster has four g roups of characters: 1)
primary or basic characteristics, such as the habi t,
the vessel perfo ra tion, the change o f the phy l-
lotaxis, the mo rpholog y of the flow ers, the open
ca rpel in i ts early ontog eny, wi th mo re than one
ovule in the early development stag e, the pollen
mo rpholog y , the type o f th e embryo sac, the per-
sistent antipodal cel ls, and the persistent antipodal
cells; 2) speciali zed cha racteristics, such as the at-
tachment and the st ructure of the ovule,
mesogamy , the tectum of the po llen, the embryo-
geny and the development and the st ructure of the
endosperm; 3) reduced cha racteristics, such as no
mechanical ti ssue, the node, the venation, the
number o f the f lo ral pa rts, the vascular tissue in
the flo ral parts, the behavio r o f the integument in
the process of embryo development , the seed w ith-
out testa. Among these characteristics, the theo-
10957期 任　毅 ,等: 基于形态学资料星叶草属系统位置的探讨 (英 )
retic considera tion of the open dicho tomous vena-
tion to be a primitiv e o r reduced one had been the
source of confusing the possible origin until Ren et
al. ( 1998, 1997) revealed the reduced morpho logi-
cal na ture after a series o f carefully observ ations;
4) the characteristics of adul t plant retained from
the seedling o r the characteristics of g row th being
checked, such as the ex tremely elongated hypoco t-
y l, the persistent cotyledons, the ex tremely shor t-
ened stem and the inf lo rescent, and the limited
secondary g row th in root and hypocotyl. Therefore
w e conjectured that Circaeaster should be a tax on
w ith a few primary, but more specialized and re-
duced characters, and being fix ed a t the seedling
stag e. The similarities in some prima ry character-
istics show tha t Circaeaster may have the same o ri-
gin as o ther members of Ranunculales. The spe-
ciali zed characteristics show that Circaeaster di ffer-
entia tes st rong ly f rom other members of Ranuncu-
lales at the early stag e o f i t s phylogeny and this
dif ferentiation was fix ed in the early pro cess of em-
bryogeny. On the o ther hand, this mono typic
genus is dist ributed in east Himalaya and g row s in
moist and shaded habitat of alt. 2 800～ 4 000 m in
the mountains. This habi ta t may be the fo rce to
make this g enus speciali zed and reduced. There-
fo re w e conjecture that Circaeaster originated
neo teinicly f rom pro-Ranunculales and developed
in the clade of specialization and reduction under
the adversi ty.
Acknowledgement: I would like to expr ess my since re thanks to Professor Lu An-min, Labo rato r y o f Sy stematic& Evo lution-
ar y Bo tany , Institute o f Bo tany , Chinese Academy o f Sciences, fo r his help in going over the present manuscript.
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