全 文 ：植 物 分 类 学 报 43(5): 431–488(2005) doi:10.1360/aps040130
Acta Phytotaxonomica Sinica http://www.plantsystematics.com
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2004-10-29收稿, 2005-04-27收修改稿。
基金项目: 国家自然科学基金资助项目(30470126) (Supported by the National Natural Science Foundation of China (Grant
No. 30470126))。
铁线莲属一新分类系统
王文采 李良千
(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)
A new system of classification of the genus Clematis
(Ranunculaceae)
WANG Wen-Tsai LI Liang-Qian
(Laboratory of Systematic and Evolutionary Botany, Institute of Botany, the Chinese Academy of Sciences, Beijing 100093, China)
Abstract A new system of classification of the genus Clematis is proposed. The taxonomic
history of the genus is briefly reviewed. On the basis of the analyses of various morphological
and palynological characters, evolutionary trends of some characters are revealed as follows:

2. Seed germination: epigeal→hypogeal.
3. Leaf: (a) phyllotaxy: alternate→opposite→verticillate.

(c) texture: herbaceous→papery→subcoriaceous or coriaceous.
4. Inflorescence: (a) position: terminal→terminal and axillary→axillary.
(b) type: pedunculate, bibracteate, several-flowered cyme.
(i) 3–4 times branched, 20–35-flowered cyme→many-branched ca.
100-flowered panicle.
(ii) 1-flowered cyme→a single flower, pedicellate, but with peduncle
and bracts wanting.
5. Flower: (a) Sexuality: bisexual→unisexual.
(b) Posture: erect, with straight pedicel→pendulous, with recurved pedicel.
6. Sepal: (a) number per flower: 4 or 5→6–8.
(b) aestivation: valvate→imbricate.
(c) colour: white or yellow→blue, purple or red.
(d) spreading direction: spreading→ascending→erect.
(e) texture: herbaceous or papery→subcoriaceous or coriaceous.
(f) shape: obovate→obovate-oblong, oblanceolate, lanceolate or oblong→
植 物 分 类 学 报 43卷 432
narrowly oblong→linear.
(g) apex: rounded or obtuse→acute→acuminate→attenuate
(h) margin indumentum: glabrous or puberulous→with a narrow velutinous strip.
(i) margin dilation: not dilated→dilated.
(j) indumentum of adaxial surface: glabrous→puberulous→velutinous.
7. Stamen: (a) indumentum: glabrous→filament hairy yet anther glabrous→both filament and
anther hairy.
(b) filament: (i) linear→proximally or distally widened, or entirely so.
(ii) flat→rugose.
(c) anther: oblong, 0.8–3.5 mm long→narrowly oblong, 2–6 mm long→linear, up
to 7–10 mm long.
(d) connective: not projected→apex minutely apiculate, with an apiculum ca. 0.1
mm long→apex with a conic or subglobose projection 0.1–0.2 mm long→apex
with a subulate projection up to 4 mm long or with a narrowly lamellate
projection 1–10 mm long.

9. Staminode: (a) absent in bisexual flower→present in bisexual flower.
(b) number per pistillate flower: numerous→few→zero.
10. Style: not elongate→elongate and plumose→slightly elongate and appressed-puberulous,
then tail-like or subulate in shape→very short, slightly or not elongate, appressed-
puberulous or glabrous.
11. Fruit: not complanate, nor rimmed, nor marginate→complanate, narrowly to tumidly
rimmed→complanate, broadly marginate or winged.
According to the evolutionary trends mentioned above and mainly according to the floral
structure, the relationships among the 15 sections of Clematis are discussed, and four
evolutionary stocks are recognized. Of them, the C. montana stock, in which C. brevipes, C.
potaninii, C. montana etc., all with spreading, white, obovate sepals and glabrous stamens, are
included, is regarded as the primitive group of the genus, whereas the other three, C. vitalba
stock, C. connata stock, and C. alpina stock, are regarded as more advanced. All these four
stocks might be derived from a putative extinct primitive group of the genus, which might
possess the most primitive vegetative and reproductive characters. The four stocks are all
treated as subgenera in the present system. Finally, a systematic enumeration of all the
infrageneric taxa is presented and their brief diagnoses are given.
Key words Clematis, Ranunculaceae, evolutionary trend, relationship, classification.
摘要 提出毛茛科Ranunculaceae铁线莲属Clematis一新分类系统。首先,简要回顾了此属的分类学研究
历史,继对此属的营养器官和生殖器官的重要形态特征和花粉特征进行了分析,揭示出一系列演化趋势。
这些趋势,尤其是萼片和雄蕊的演化趋势说明在现存的铁线莲属植物中,绣球藤Clematis montana与其少
数近缘种的花构造最接近比铁线莲属原始的银莲花属Anemone,因此,包括绣球藤等植物的绣球藤组sect.
Cheirpsis被认为是铁线莲属的原始群。主要根据花构造对铁线莲属现存的15组的亲缘关系进行了分析,
发现它们是在4条演化干(绣球藤干、欧洲铁线莲干、尾叶铁线莲干、长瓣铁线莲干)的演化过程中先后形
成的。本文将此4条演化干处理为亚属。最后做出属下各级分类群的系统排列,并给出简要的形态特征。
关键词 铁线莲属; 毛茛科; 演化趋势; 亲缘关系; 分类
5期 王文采等: 铁线莲属一新分类系统 433
1 分类简史*
Linnaeus(1753)建立了铁线莲属Clematis L.,描述了9种,将这些种分为两类,第一类为
攀援类(Scandentes),包括7种:Clematis viticella L.、C. viorna L. 、C. crispa L. 、C. orientalis L. 、
C. vitalba L. 、C. cirrhosa L.和C. flammula L.; 第二类为直立类(Erectae),包括2种:C. recta L.
和C. integrifolia L.。此外 ,还建立了长瓣铁线莲属Atragene L.,并描述4种: Atragene
zeylanica L. 、A. alpina L. 、A. sibirica L.和A. capensis L.。后来,de Candolle (1818,见下)将此
属降级作为铁线莲属的一个组sect. Atragene (L.) DC.,这种处理得到以后多数铁线莲属研
究者的赞同,只是在俄国,长瓣铁线莲属一直得到承认。其第一种A. zeylanica的花有花瓣,
而与铁线莲属有重要区别,de Candolle(1818)根据此种建立锡兰莲属Naravelia DC.,此种遂
即被转移到新属中,即N. zeylanica (L.) DC.。其第四种A. capensis为直立草本,花序具总苞,
应是属于银莲花属Anemone L.的植物,de Candolle(1818)发现此问题,遂将此种移至银莲花
属中,做出新组合Anemone capensis (L.) DC.。至于其他具花瓣状退化雄蕊的2种Atragene
alpina和A. sibirica已由Miller(1768)转移到铁线莲属中并做出新组合Clematis alpina (L.)
Mill.和C. sibirica (L.) Mill.。
Moench(1794)承认Linnaeus之前的2属Clematitis Tourn.和Viticella Dill.,并描述了共5
种:Clematitis crispa Moench (=Clematis crispa L.)、Clematitis flava Moench (=Clematis
orientalis L.) 、Clematitis integrifolia Moench (=Clematis integrifolia L.) 、Clematitis vitalba
Moench (=Clematis vitalba L.)和Viticella deltoidea Moench (=Clematis viticella L.)。
de Candolle(1818)对铁线莲属进行了第一次修订,收载此属当时已知的85种,利用总
苞、退化雄蕊、花柱等特征将这些种类划分为4组: sect. Flammula DC.、 sect. Viticella (Dill.
ex Moench) DC. (包括C. viticella和C. florida Thunb.等4种) 、sect. Cheiropsis DC. (包括C.
cirrhosa L.、C. napaulensis DC.和C. montana Buch.-Ham. ex DC.等5种)和sect. Atragene (L.)
DC.(包括C. alpina和C. sibirica等4种)。他正确地将Viticella和Atragene从属级下降作为铁
线莲属的两个组处理,这两组以及他新建立的sect. Cheiropsis均是自然群,但是第一组sect.
Flammula包括约70种,内容混杂,如具开展萼片、无毛雄蕊的C. apiifolia DC.和C. vitalba,单
性的C. virginiana L.等种与具斜升或直立萼片和有毛雄蕊的C. brachiata Thunb.、C. villosa
DC.、C. connata DC.以及C. viorna等种混在一起。
Spach(1839)在关于trib. Clematideae的报告中承认过去已建立的Atragene、Viticella
Dill. ex Moench、Viorna Reichb.和Clematis等4属,此外还将sect. Cheiropsis DC.升为属级,做
出新组合Cheiropsis (DC.) Spach,并根据Clematis glauca Willd.和C. orientalis建立新属
Meclatis Spach,但未涉及Naravelia DC.。在他的狭义的Clematis中,除承认过去的sect.
Flammula DC.以外,又建立了2新组: sect. Vitalba Spach(包括两性的C. vitalba和单性的C.
virginiana)和sect. Aspidanthera Spach(包括1种,即C. aristata R. Br.)。
Baillon(1867–1869)研究铁线莲属时承认sect. Atragene (L.) DC.、sect. Cheiropsis DC.、
sect. Flammula DC.和sect. Viticella (Moench) DC.等4组,并将Naravelia DC.、Meclatis Spach
和Atragene subgen. Viorna Pers.降到组的等级。

* 本文 1、2两节由王文采、李良千撰写，3、4两节由王文采撰写。
植 物 分 类 学 报 43卷 434
Kuntze(1885)发表了铁线莲属的专著,对此属当时已发表的种类做出了全面整理,承
认66种。他对不少种的范围处理过宽,如他将亚洲的C. gouriana Roxb. ex DC.、C. grata
Wall.、C. brevicaudata DC.、C. javana DC.、C. subumbellata Kurz和C. taurica Bess.都放在欧
洲的C. vitalba之下作为亚种处理。关于铁线莲属属下类群的划分,他首先用茎攀援或直立
来划分大群,在茎攀援的一类中再用有无芽鳞来划分,这样建立了3个大群:无鳞攀援组
sect. 1. Scandentes eperulatae,有鳞攀援组sect. 2. Scandentes perulatae,非攀援组sect. 3.
Escandentes。在每组之下大致依次用花柱是否强烈伸长,雄蕊花丝是否有毛,萼片开展方
向,花药长度的大小,药隔顶端是否突出,以及花为两性或单性等特征来进行分类。Kuntze
发现并利用了上述用于铁线莲属分类和表现演化水平的多数重要形态特征,因此对铁线
莲属分类学研究做出了重要贡献。由于利用了上述特征,在多数情况下,Kuntze将亲缘相近
的种放在一起,例如将C. pinnata Maxim. 、C. tatarinowii Maxim.、C. heracleifolia DC.等3种
正确地放在一起,实属难能可贵,因为后来一些学者如Handel-Mazzetti (1939)、丁志遵
(1980)、Johnson (1997)、Grey-Wilson (2000)等未能认识到上述植物的相近亲缘关系,均将C.
heracleifolia和C. tatarinowii(实与C. pinnata为同一种,见史京华, 2003)归入sect. Tubulosae
中 ,并与C. connata群(本文系统的sect. Viorna subsect. Connatae)相联系 ,另外则将C.
pinnata放在C. vitalba群(本文系统的sect. Clematis subsect. Clematis)中。Kuntze不承认
Naravelia属,将此属的植物收入其铁线莲属专著中。
Prantl(1888)对铁线莲属做了全面研究,发表了一个分类系统:
Sect. 1. Pseudanemone Prantl 萼片覆瓦状排列;雄蕊花丝有毛,药隔不突出。
1. Spathulifoliae Prantl 叶或小叶宽,不分裂或羽状分裂。包括C. spatulifolia Kuntze、
C. chrysocarpa Welw. ex Oliv.等种。
2. Villosae Prantl 叶三角形,细裂。包括C. villosa、C. anethifolia Hook.等种。
Sect. 2. Viorna (Reichb.) Prantl 萼片镊合状排列,直立或斜上方开展,稀平展,有狭边缘;雄
蕊有毛,药隔不突出。
3. Crispae Prantl 叶或小叶全缘;花单生或组成二歧聚伞花序;萼片厚。包括C.
crispa、C. integrifolia等种。
4. Tubulosae (Decne.) Prantl 叶或小叶边缘有齿;花组成二歧聚伞花序或聚伞总状
花序;萼片厚。包括C. heracleifolia群和C. connata群。
5. Atragene (L.) Prantl 单花自腋芽抽出的短枝顶端生出;萼片薄;花丝有毛。包括
C. alpina、C. sibirica等种。
6. Cirrhosae Prantl 单花自腋芽中生出;萼片薄;花丝无毛。包括C. cirrhosa、C.
napaulensis等种。
Sect. 3. Viticella (Moench) DC. 萼片有褶,有宽边缘,平展或斜上展;雄蕊无毛或上部有少
数毛,药隔无明显突起。
7. Euviticellae Prantl 萼片4,斜上展;花丝比花药短。包括C. viticella、C. campaniflora
Brot.等种。
8. Floridae Prantl 萼片(4–)6–8,平展;花丝多少与花药等长。包括C. florida、C. patens
Morr. & Decne.、C. lanuginosa Lindl.等种。
5期 王文采等: 铁线莲属一新分类系统 435
Sect. 4. Flammula DC. 萼片镊合状排列,平展;雄蕊无毛或基部有毛,药隔不突起。
9. Rectae Prantl 雄蕊无毛;花药长度大于宽度数倍,无突起。包括C. songarica、C.
fruticosa Turcz.、C. flammula、C. meyeniana Walp.、C. crassifolia Benth.、C. chinensis
Osb.、C. apiifolia、C. montana等多数种。
10. Vitalbae Prantl 雄蕊无毛,花药长度至多大于宽度2倍,无突起。
a. Euvitalbae Prantl 花两性;茎不中空。包括C. vitalba、C. gouriana等种。
b. Saxicolae Prantl 花雌雄异株;茎中空。包括C. saxicola Hils. & Bojer ex Baill.
(= C. insidiosa)等种。
c. Dioicae Prantl 花雌雄异株;茎不中空。包括C. virginiana、C. dioica L.、C.
ligusticifolia Nutt.、C. brasiliana DC.、C. campestris St. Hil.、C. lasiantha Nutt.、
C. pauciflora Nutt.等种。
d. Hexapetalae Prantl 花雌雄异株;萼片远长于雄蕊。包括C. microphylla DC.、
C. hexapetala L. f. (=C. fosteri Gmelin) 、C. aphylla Kuntze (=C. afoliata
Buchan.)等种。
11. Aristatae Prantl 雄蕊无毛;花药长度大于宽度数倍 ,顶端有长突起。包括C.
gentianoides DC.、C. aristata、C. clitorioides DC.等种。
12. Orientales Prantl 雄蕊有毛;叶羽状分裂,裂片全缘。包括C. orientalis、C. glauca
等种。
13. Wightianae Prantl 雄蕊有毛;叶为羽状复叶或三出复叶,小叶边缘有齿。包括C.
wightiana、C. mauritiana Lam.、C. brachiata等种。
Sect. 5. Naravelia (DC.) Prantl 萼片镊合状排列;雄蕊药隔顶端突起,或外方的雄蕊变成蜜
叶(花瓣)。包括无蜜叶的C. smilacina Bl. (=C. smilacifolia Wall.)等种和有蜜叶的
C. zeylanica (L.) DC.等种。
Prantl已有生物演化的观点,他推测分布于古热带且与银莲花属最为相似的类型可能
是铁线莲属的原始群,而sect. Pseudanemone(可能由于此组植物的萼片为覆瓦状排列)就
最像这个原始群,于是被放在其系统的开始位置。也正由于这点,雄蕊被毛的C. connata群
和C. viorna群等被放在此原始群之后。Prantl用于分类的形态特征更为广泛,除了上述
Kuntze利用的之外,又增加了萼片卷叠式、厚度、边缘,花药本身长度与宽度的比例,花药长
度与花丝长度的比例等特征,因此划分出更多比较自然的群,所制定的新系统为铁线莲属
分类学研究奠定了良好基础,对以后的有关工作产生了较大影响,如Schneider (1906)、
Rehder和Wilson (1913)、Handel-Mazzetti (1939)、Rehder (1951)、Eichler (1958)、Tamura
(1967)、张美珍等(1980)等学者有关铁线莲属的分类学著作均采用了Prantl系统,但也常做
出或多或少的补充或修改。在他建立的属下分类群中有一些混乱或不自然的情况,例如在
sect. Pseudanemone中混入Sect. Brachiatae的二种C. triloba Thunb.和C. commutata Kuntze;
C. heracleifolia群与C. connata群亲缘关系并不相近,却被同放在sect. Viorna 4. Tubulosae
中;在sect. Viorna 5. Atragene中混入不具花瓣状退化雄蕊的C. japonica Thunb.;不承认雄蕊
无毛的sect. Cheiropsis, 但根据此组的C. cirrhosa等种另立6. Cirrhosae放在雄蕊有毛的
sect. Viorna中,同时将sect. Cheiropsis的另一种C. montana误置于sect. Flammula 9. Rectae
植 物 分 类 学 报 43卷 436
中;将雄蕊无毛的直立小灌木C. ispahanica Boiss.误置于雄蕊有毛的sect. Flammula 12.
Orientales中;C. smilacifolia与Naravelia zeylanica亲缘关系较远,却被同放在sect. Naravelia
中;de Candolle建立的Naravelia属的花有花瓣,与无花瓣的Clematis属区别明显,建立后受
到多数学者承认,只有Kuntze(1885)、Baillon(1867)、Prantl(1888)以及最近Johnson(1997)均
将此属降级作为组处理。本文新系统不赞同这样的处理。Schneider(1906)将Prantl系统中
用阿拉伯数字排列的群作为亚组(subsection)处理,将用英文小写字母排列的群作为系
(series)处理,本文系统采取他的处理方式。
Hutchinson(1920)根据特产非洲的C. villosa群具覆瓦状排列萼片的特征建立了新属
Clematopsis(即Prantl的sect. Pseudanemone),并认为此新属的系统位置介于银莲花属和铁
线莲属之间。
Tamura(田村道夫)(1955, 1956, 1963, 1967, 1970, 1987, 1995)自上世纪50年代初起对
铁线莲属的分类学进行了深入的研究,认为铁线莲属在中国西部和东喜马拉雅的温暖湿
润地区起源于银莲花属,还认为他建立的具互生叶的单型属Archiclematis (Tamura) Tamura
的演化水平比铁线莲属为低,与铁线莲属的sect . Campanella(即C. connata群,幼苗叶互生,
萼片直立 ,雄蕊有毛 )有密切的亲缘关系。他建立了5新组 :sect. Archiclematis、sect.
Campanella、sect. Fruticella、sect. Lasiantha和sect. Pterocarpa。根据sect. Cheiropsis的C.
williamsii Gray的萼片也是覆瓦状排列(Tobe, 1980),Tamura(1987)不承认Hutchinson建立的
Clematopsis新属,恢复了Prantl的sect. Pseudanemone。Tamura 1967年以前的系统,如前所述,
采用了Prantl系统的框架,而在他1987年和1995年最后修改的系统,做了重要变更,强调了
幼苗叶的特征,此外还根据萼片、雄蕊的特征将铁线莲属分为四亚属,认为幼苗叶互生的
subgen. Campanella(萼片直立,雄蕊有毛)和subgen. Clematis(萼片平展,雄蕊无毛)是原始
群 ,从他们分别演化出幼苗叶对生的 subgen. Viorna(萼片直立 ,雄蕊有毛 )和subgen.
Flammula(萼片平展 ,雄蕊无毛 ),在这四亚属中 ,与 Archiclematis相近缘的 subgen.
Campanella是铁线莲属的原始群。此系统全貌如下:
Subgen. 1. Campanella Tamura
Sect. 1. Campanella Tamura 包含C. henryi、C. acuminata DC.、C. connata、C. aethusifolia
Turcz.等40种。
Sect. 2. Tubulosae Decne. 包含C. heracleifolia等8种。
Sect. 3. Bebaenanthera Edgew. 包含C. barbellata Edgew.、C. japonica等4种。
Sect. 4. Atragene (L.) DC. 包含C. alpina等种。
Sect. 5. Meclatis (Spach) Tamura 包含亚洲的C. orientalis等种和非洲的C. hirsuta、C.
brachiata等种。
Sect. 6. Pseudanemone Prantl 包含C. villosa等25种。
Subgen. 2. Viorna (Reichb.) Tamura
Sect. 1. Viorna (Reichb.) Prantl 包含C. integrifolia、C. fusca Turcz.、C. crispa等约20种。
Subgen. 3. Clematis
Sect. 1. Clematis 包含C. vitalba、C. apiifolia、C. javana等25种。
Sect. 2. Cheiropsis DC. 包含C. cirrhosa、C. montana、C. acerifolia Maxim.、C. williamsii
5期 王文采等: 铁线莲属一新分类系统 437
等约13种。
Sect. 3. Naraveliopsis Hand.-Mazz. 包含C. smilacifolia等约13种。
Sect. 4. Aspidanthera Spach 包含北美及南美的C. virginiana、C. dioica、C. brasiliana等
20种,澳大利亚和新西兰的C. microphylla、C. aristata、C. paniculata Gmelin等15
种,以及新几内亚、印度尼西亚等地的C. papuasica Merr. & Perry.等10种。
Sect. 5. Lasiantha Tamura 包含C. lasiantha、C. pauciflora 2种。
Subgen. 4. Flammula (DC.) Peterm.
Sect. 1. Flammula DC.
Subsect. 1. Crassifoliae (Tamura) Tamura 含C. crassifolia 1种。
Subsect. 2. Rectae Prantl ex Rehd. & Wils.
Ser. 1. Rectae Prantl ex Rehd. & Wils. 包含C. recta、 C. flammula、 C. meyeniana、C.
chinensis等约20种。
Ser. 2. Uncinatae Tamura 包含C. uncinata Champ. ex Benth.等3种。
Sect. 2. Angustifoliae (Tamura) Serov 包含C. hexapetala 1种。
Sect. 3. Fruticella Tamura 包含C. fruticosa、C. nannophylla Maxim.、C. songarica、C.
delavayi Franch.等约7种。
Sect. 4. Viticella (Moench) DC.
Subsect. 1. Floridae Prantl 包含C. viticella、C. florida等8种。
Subsect. 2. Patentes Tamura 包含C. patens、C. lanuginosa 2种。
Sect. 5. Pterocarpa Tamura 包含C. brachyura Maxim. 1种。
Ziman(1981)根据铁线莲属植物地理分布的统计,认为中国云南和四川及相邻地区是
此属的起源和分化中心。对铁线莲属的形态特征进行了全面分析之后,与Tamura一
样 ,Ziman认为直立萼片
和被毛雄蕊是原始特征。
她还研究了整个铁线莲
亚 族 subtrib. Clemati-
dinae各群的亲缘关系,除
铁 线 莲 属 外 , 还 承 认
Clematopsis、Archiclema-
tis和Naravelia等3属,并绘
出此亚族的系统发育图
(图1)。
Essig(1991)研究了铁线
莲属 53 种和 Clematopsis
(=Clematis sect. Pseudam-
one) 2种1变种的幼苗形
态,发现幼苗叶可区分为
两种类型(表1)。

图1 Ziman (1981)推测的铁线莲亚族系统发育图
Fig. 1. Diagram showing the phylogeny of subtribe Clematidinae inferred by
Ziman (1981).
植 物 分 类 学 报 43卷 438
表1 铁线莲属植物幼苗类型(Essig, 1991)
Table 1 Two seedling types of Clematis (Essig, 1991)
类型1. 幼苗叶互生,3裂或有齿
Type I Seedling leaves alternate, trifid or serrate
类型2. 幼苗叶对生,全缘,稀分裂
Type II Seedling leaves opposite, entire, rarely lobed
Sect. Cheiropsis
C. chrysocoma Franch.*
C. cirrhosa L.
C. napaulensis DC.
Sect. Clematis subsect. Clematis
C. apiifolia DC.
C. grata Wall.
C. peterae Hand.-Mazz.
C. pierotii Miq.
C. vitalba L.
Sect. Aspidanthera
Subsect. Dioicae
C. catesbyana Pursh
C. denticulata Vell.
C. drummondii Torr. & Gray
C. ligusticifolia Nutt.
Subsect. Lasianthae
C. lasiantha Nutt.
Subsect. Aristatae
C. aristata R. Br.
C. gentianoides DC.
C. papuasica Merr. & Perry.
Sect. Brachiatae
C. brachiata Thunb.
Sect. Pseudanemone
C. anethifolia Hook.
C. villosa DC. ssp. villosa
ssp. kirkii (Oliv.) Brummitt
Sect. Meclatis
C. intricata Bunge
C. ladakhiana Grey-Wilson
C. orientalis L.
C. serratifolia Rehd.
C. tangutica (Maxim.) Korsh.
Sect. Naraveliopsis
C. filamentosa Dunn
(=C. loureiriana DC.)
Sect. Tubulosae
C. heracleifolia DC.
Sect. Viorna
subsect. Connatae
C. buchananiana DC.
C. leschenaultiana DC.
C. ranunculoides Franch.
C. rehderiana Craib
subsect. Bebaenanthera
C. barbellata Edgew.
Sect. Atragene
C. alpina (L.) Mill.
C. macropetala Ledeb.
Sect. Clematis subsect. Rectae
C. chinensis Osb.
C. flammula L.
C. kirilowii Maxim.
C. mandshurica Rupr.
C. recta L.
C. terniflora Thunb.
Sect. Aspidanthera subsect. Microphyllae
C. microphylla DC.
Sect. Viticella
C. campaniflora Brot.
C. patens Morr. & Decne.
C. viticella L.
Sect. Viorna subsect. Crispae
C. addisonii Britton
C. baldwinii Torr. & Gray
C. crispa L.
C. fusca Turcz.
C. glaucophylla Small
C. hirsutissima Pursh.
C. integrifolia L.
C. pitcheri Torr. & Gray
C. reticulata Walter
C. texensis Buckley
C. viorna L.

* 本表中所有种按本文系统排列。The species listed above are arranged according to the present new system of
classification of Clematis.
5期 王文采等: 铁线莲属一新分类系统 439
表1中类型2中的C. microphylla的幼苗叶对生,但分裂,Essig(1991)认为是类型1和类型
2的中间类型。他还认为铁线莲属的祖先居群(ancestral population)具有类型1的幼苗叶,其
萼片有颜色,直立,雄蕊有毛。他说的祖先居群当是sect. Viorna subsect. Connatae的植物。
Essig还观察到具类型1幼苗叶植物的瘦果多少两侧扁,但不扁平,而具类型2幼苗叶植物的
瘦果则强烈两侧扁,扁平,并认为这两种相关特征可以作为在铁线莲属中区分亚属的根据。
对此论断,我们不赞同,因为类型2的各群间的亲缘关系相当疏远,其中的sect. Clematis
subsect. Rectae衍生自类型1的sect. Clematis subsect. Clematis,而sect. Viorna subsect.
Crispae衍生自类型1的sect. Viorna subsect. Connatae (Tamura, 1995),如按Essig的论点将类
型1和类型2的植物各处理为亚属,则类型2亚属必是一个不自然的异质群。
Snoeijer(1992)发表了铁线莲属一新分类系统,其系统与Tamura 1987年的系统非常相
似。在subgen. Campanella中,根据分布非洲的C. brachiata等种建立一新组sect. Brachiatae,
实等同于1888年建立的sect. Flammula subsect. Wightianae Prantl。最近,王文采(2004b)承认
此新组,并做出包括24种的全面修订。
Johnson(1997)发表了铁线莲属分类学研究巨著,收载此属植物314种,划分为19组,是
继de Candolle和Kuntze之后对铁线莲属的第三次全面修订。在其系统中对不少组进行了
亚组的划分,全部19组的排列如下:
Sect. 1. Archiclematis Tamura
Sect. 2. Pseudanemone Prantl
Sect. 3. Novae-Zealandiae M. Johnson
Sect. 4. Atragene (L.) DC.
Subsect. 1. Atragene
Subsect. 2. Brachyblasti M. Johnson
Sect. 5. Naravelia (DC.) Prantl
Sect. 6. Tubulosae Decne.
Sect. 7. Connatae (Koehne) M. Johnson
Sect. 8. Meclatis (Spach) Tamura
Sect. 9. Bebaenanthera Edgew.
Sect. 10. Cheiropsis DC.
Subsect. 1. Cheiropsis
Subsect. 2. Williamsianae M. Johnson
Subsect. 3. Montanae Schneid.
Sect. 11. Clematis
Subsect. 1. Clematis
Subsect. 2. Pierotianae (Tamura) Tamura
Subsect. 3. Potaninianae M. Johnson
Subsect. 4. Africanae M. Johnson
Subsect. 5. Dioicae (Prantl) Tamura
Sect. 12. Lasiantha Tamura
植 物 分 类 学 报 43卷 440
Sect. 13. Aspidanthera Spach
Subsect. 1. Aspidanthera
Subsect. 2. Papuasicae (H. Eichler) M. Johnson
Sect. 14. Naraveliopsis Hand.-Mazz.
Sect. 15. Viorna (Reichb) Prantl
Subsect. 1. Viorna
Subsect. 2. Baldwinianae Ericks.
Subsect. 3. Crispae Prantl
Subsect. 4. Hirsutissimae Ericks.
Subsect. 5. Integrifoliae Ericks. ex M. Johnson
Subsect. 6. Fuscae (Tamura) M. Johnson
Sect. 16. Flammula DC.
Subsect. 1. Flammula
Subsect. 2. Angustifoliae (Tamura) M. Johnson
Subsect. 3. Chinenses (Tamura) M. Johnson
Subsect. 4. Crassifoliae (Tamura) M. Johnson
Subsect. 5. Fasciculiflorae (Tamura) M. Johnson
Subsect. 6. Meyenianae (Tamura) M. Johnson
Subsect. 7. Uncinatae (Tamura) M. Johnson
Sect. 17. Pterocarpa Tamura
Sect. 18. Fruticella Tamura
Subsect. 1. Fruticella
Subsect. 2. Ispahanicae (Serov) M. Johnson
Subsect. 3. Phlebanthae M. Johnson
Subsect. 4. Songaricae (Serov) M. Johnson
Sect. 19. Viticella (Moench) DC.
Subsect. 1. Viticella
Subsect. 2. Floridae (Prantl) Tamura
Subsect. 3. Lanuginosae M. Johnson
Subsect. 4. Patentes Tamura
Johnson不承认Archiclematis属,恢复了其组的等级,并放在其系统的开始位置。他同
Tamura一样,也不承认Clematopsis属,恢复sect. Pseudanemone,并将此组放在其系统第二位
的位置上,从这里可以看到Tamura系统和Prantl系统对其系统的影响。其系统主要与Prantl
和Tamura的系统相同,把雄蕊被毛的群放在雄蕊无毛的诸群之前,但把花单性,雄蕊无毛,
并与sect. Clematis近缘的新组sect. Novae-Zeelandiae放在雄蕊被毛的诸群之间;还把雄蕊
被毛、与sect. Connatae近缘的sect. Viorna放在雄蕊无毛的诸群之间。如前所述,他同Kuntze、
Prantl一样,不承认Naravelia属,将其作为组处理。
王文采(1989)在对铁线莲属的研究中揭示了此属萼片由平展到向上直展和雄蕊从无
5期 王文采等: 铁线莲属一新分类系统 441
毛到有毛的演化趋势,并指出具钟状花萼和被毛雄蕊的C. connata群是相当进化的群。后
来,王文采(1998)进一步揭示了铁线莲属花丝和花药形态的演化趋势,并根据上述认识对
中国铁线莲属诸组做出了新的排列次序: sect. 1. Clematis、sect. 2. Fruticella、sect. 3.
Cheiropsis、sect. 4. Viticella、sect. 5. Naraveliopsis、sect. 6. Meclatis、sect. 7. Tubulosae、sect. 8.
Viorna、sect. 9. Atragene。1999年,王文采在英国邱皇家植物园标本馆(K)和巴黎显花植物
研究所标本馆(P)研究了他们收藏的世界各大洲的铁线莲属植物标本之后,肯定了上述演
化趋势,并又认识到铁线莲属萼片形状、毛被等方面的演化趋势,以及主要分布于中国西南
部的C. brevipes Rehd.、C. potaninii Maxim.、C. montana、C. gracilifolia Rehd. & Wils.、C.
chrysocoma和C. venusta M. C. Chang等种的花构造(萼片平展,白色,倒卵形,雄蕊无毛,花药
长圆形或狭长圆形,药隔顶端不突起)与银莲花属最相似,是现存铁线莲属植物中原始的
种。他还根据对全属各组亲缘关系的分析,发现此属存在4条演化干,其中原始的是绣球藤
干(C. montana stock),其他3条进化的是欧洲铁线莲干(C. vitalba stock) 、尾叶铁线莲干(C.
connata stock)和长瓣铁线莲干(C. alpina stock) (王文采, 2000b, 2002, 2003, 2005)。
Grey-Wilson(2000)继Johnson 3年之后发表了铁线莲属的第四次全面修订,收载297种,
划分为9亚属18组:
Subgen. 1. Clematis
Sect. 1. Clematis
Subsect. 1. Clematis
Subsect. 2. Pierotianae (Tamura) Tamura
Subsect. 3. Potaninianae M. Johnson
Subsect. 4. Dioicae (Prantl) Tamura
Sect. 2. Lasiantha Tamura
Sect. 3. Naraveliopsis Hand.-Mazz.
Sect. 4. Aspidanthera Spach
Subsect. 1. Aspidanthera
Subsect. 2. Papuasicae H. Eichler
Sect. 5. Novae-Zeelandiae M. Johnson
Subgen. 2. Cheiropsis (DC.) Peterm.
Sect. 1. Montanae (Schneid.) Grey-Wilson
Sect. 2. Cheiropsis
Subsect. 1. Cheiropsis
Subsect. 2. Williamsianae M. Johnson
Sect. 3. Fasciculiflorae (M. Johnson) Grey-Wilson
Subgen. 3. Flammula (DC.) Peterm.
Sect. 1. Flammula
Subsect. 1. Flammula
Subsect. 2. Meyenianae (Tamura) M. Johnson
Subsect. 3. Chinenses (Tamura) M. Johnson
植 物 分 类 学 报 43卷 442
Subsect. 4. Uncinatae (Tamura) M. Johnson
Subsect. 5. Angustifoliae Tamura
Subsect. 6. Crassifoliae (Tamura) Tamura
Sect. 2. Viticella (Moench) DC.
Subsect. 1. Viticella
Subsect. 2. Floridae (Prantl) Tamura
Subsect. 3. Patentes Tamura
Subsect. 4. Lanuginosae M. Johnson
Sect. 3. Pterocarpa Tamura
Sect. 4. Fruticella Tamura
Subsect. 1. Fruticella
Subsect. 2. Songaricae Serov
Subsect. 3. Phlebanthae M. Johnson
Subgen. 4. Archiclematis (Tamura) Grey-Wilson
Subgen. 5. Campanella Tamura
Sect. 1. Campanella Tamura
Sect. 2. Bebaenanthera Edgew.
Sect. 3. Meclatis (Spach) Tamura
subsect. 1. Africanae M. Johnson
Subsect. 2. Meclatis
Subgen. 6. Atragene (L.) Grey-Wilson
Subgen. 7. Tubulosae (Decne.) Grey-Wilson
Subgen. 8. Pseudanemone (Prantl) Grey-Wilson
Subgen. 9. Viorna Tamura
Sect. 1. Viorna
Subsect. 1. Viorna
Subsect. 2. Crispae Prantl
Subsect. 3. Fuscae (Tamura) M. Johnson
Sect. 2. Integrifoliae Serov
Subsect. 1. Integrifoliae Ericks. ex M. Johnson
Subsect. 2. Baldwinnae Ericks. ex M. Johnson
Sect. 3. Hirsutissimae (Ericks. ex M. Johnson) Grey-Wilson
Grey-Wilson系统将雄蕊无毛的亚属subgen. Clematis、Cheiropsis、Flammula置于雄蕊
被毛的亚属之前,这与上述王文采1998年的处理相似;与Tamura一样,不将Naravelia降级作
为Clematis的一个组,同时接受了Tamura的subgen. Clematis、Flammula、Campanella、Viorna,
但另外又增加了subgen. Cheiropsis、Archiclematis、Atragene、Tubulosae、Pseudanemone;接受
了Johnson系统中有关组的组下分类,但做出了一些重要改正,如将subsect. Africanae(实等
同于 sect. Flammula 13. Wightianae Prantl及 sect. Brachiatae Snoeijer)正确地从 sect.
5期 王文采等: 铁线莲属一新分类系统 443
Clematis移到sect. Meclatis中,并置于subsect. Meclatis之前,再如将subsect. Fasciculiflorae正
确 地 从 sect. Flammula 移 回 sect. Cheiropsis 中 ; 此 外 , 还 将 Johnson 的 新 组 sect.
Novae-Zeelandiae(实等同于sect. Flammula 10. Vitalbae d. Hexapetalae Prantl)正确地从sect.
Pseudanemone、sect. Atragene附近移至sect. Aspidanthera之后;至于Johnson根据属于sect.
Cheiropsis的C. potaninii建立subsect. Potaninianae并放在sect. Clematis中,Grey-Wilson则加
以接受,未做修正。
Brandenburg(2000)发表了黄花铁线莲组sect. Meclatis研究一书,在此组修订之前,他先
对铁线莲属进行了分支系统学(cladistics)研究,对此属的营养器官和生殖器官的形态特征
的演化趋势进行了全面分析,正确地揭示了铁线莲属不少形态特征的演化趋势,如habit:
perennial 1 subshrub 2 shrub 3; leaf: simple 1 compound 2; inflorescence: flowering on the
young wood 1 flowering on the old wood 2; flower: flat, erect 1 broadly campanulate 2
campanulate 3 tubulose, urceolate 4; filament: glabrous 1 ciliate 2; anther: shorter 1 as long as
2 longer than filament 3; connective: not elongate 1 elongate 2。在萼片形状方面,他认为倒卵
形萼片为进化特征,系由菱形萼片演化而来,可能与实际情况相悖。根据此项研究,他拟定
出铁线莲属的一包括18组的新排列方式,并对其中一组做出组下分类:
Sect. 1. Angustifoliae (Prantl) Brandenb.
Sect. 2. Aristatae (Prantl) Brandenb.
Sect. 3. Atragene (L.) DC.
Sect. 4. Bebaenanthera Edgew.
Sect. 5. Cheiropsis DC.
Sect. 6. Clematis
Sect. 7. Crassifoliae (Tamura) Brandenb.
Sect. 8. Dioicae (Prantl) Brandenb.
Sect. 9. Fruticella (Tamura) Brandenb.
Sect. 10. Lasiantha (Tamura) Brandenb.
Sect. 11. Meclatis (Spach) Baill.
Sect. 12. Papuasicae (H. Eichler) Brandenb.
Sect. 13. Naraveliopsis (Hand.-Mazz.) Brandenb.
Sect. 14. Pierotianae (Tamura) Brandenb.
Sect. 15. Pterocarpa (Tamura) Brandenb.
Sect. 16. Rectae (Prantl) Brandenb.
Sect. 17. Viorna (Reichb.) Prantl
subsect. 1. Viorna
subsect. 2. Connatae Koehne
subsect. 3. Crispae (Prantl) Tamura
Sect. 18. Viticella (Moench) DC.
Brandenburg在有关群亲缘关系的讨论中赞同Clematopsis作为属从铁线莲属分出,但
对Archiclematis和Naravelia二属从铁线莲属分出表示怀疑,认为应将这二群放在铁线莲属
植 物 分 类 学 报 43卷 444
中。其系统未包括sect. Tubulosae,但将Prantl(1888)建立的subsect. Aristatae、Ser. Dioicae和
subsect. Rectae, Eichler(1958)建立的subsect. Papuasicae,以及Tamura(1955)建立的subsect.
Angustifoliae、ser. Crassifoliae和ser. Pierotianae均提升到组的等级。令人不解的是,在此系
统中,不少亲缘关系相近的组被分开不放在一起,如sect. Clematis与sect. Pierotianae极为相
近,但二者却被sect. Crassifoliae、sect. Dioicae等7个组隔开;再如,具单性花的2组sect.
Dioicae和sect. Lasiantha极为相近,中间却被具两性花的sect. Fruticella隔开。再令人奇怪
的是,Handel-Mazzetti建立的sect. Naraveliopsis, Tamura建立的3个组sections Lasiantha、
Pterocarpa、Fruticella,不知为何Brandenburg再做出新组合(combinatio nova)(见上表)。
2 形态和花粉特征分析
2.1 习性
铁线莲属植物多为木质藤本,少数为多年生草本、亚灌木或灌木。由于具藤本的习性
以及受精后花柱强烈伸长等特化特征,在银莲花族trib. Anemoneae中铁线莲属处于进化的
地位,可能由此族原始的草本银莲花属演化而来(Hutchinson, 1920; Tamura, 1995)。因此,
在铁线莲属,多年生直立草本当是原始的,而直立亚灌木、灌木或木质藤本是后来出现的
(Ziman, 1981; Brandenburg, 2000)。但在sect. Meclatis中看到上述习性演化趋势逆转现象,
此组的C. tangutica var. tangutica和C. tibetana var. tibetana均为木质藤本,前者广布于我国
青藏高原、新疆及哈萨克斯坦,在青海南部发现一居群为矮小直立灌木;后者分布于尼泊
尔,我国西藏西部和新疆西南部,其萼片外面无毛,但在帕米尔高原出现直立矮小、萼片外
面被毛灌木的居群。
2.2 发芽
Section Cheiropsis、sect. Clematis subsect. Clematis、sect. Pseudanemone、sect. Viorna
subsect. Connatae、sect. Archiclematis的种子出土发芽,而sect. Clematis subsect. Rectae和
sect. Viorna subsect. Crispae的种子留土发芽。留土发芽可能是适应自温暖地区到寒冷地
区的不同气候的结果,是由出土发芽演变而来(Tamura et al., 1977; Ziman, 1981; Tamura,
1995; Grey-Wilson, 2000)。
2.3 叶
铁线莲属中多数群的幼苗叶互生,少数群的幼苗叶对生(Essig, 1991, 见上),成长的茎
生叶通常为对生,稀轮生(如在 sect. Pseudanemone的 C. teuszii (Kuntze) Engler),而只在单
种组 sect. Archiclematis的 C. alternata Kitam. & Tamura,所有叶均为互生(Tamura, 1995)。
根据毛茛科大多数植物的茎生叶通常互生,铁线莲属的叶序演化趋势当是:互生→对生→
轮生。
Kuntze(1885)在其专著中绘出铁线莲属叶从一回三出到两回三出和1–3回羽状分裂
的示意图,似乎是表示叶分裂的演化趋势。Tamura(1967)认为毛茛科中较普遍的叶为基部
心形、掌状分裂、边缘具缺刻或牙齿的单叶,由此演化到不分裂的单叶,另一方面演化到三
出复叶及1–4回分裂的羽状复叶。这种情况在 sect. Brachiatae(王文采, 2004b)和 sect.
Pseudanemone(王文采, 2004c)可大致看到。但在陕南单叶铁线莲 C. henryi var. ternata M.
Y. Fang (sect. Viorna subsect. Connatae),其枝条的多数叶为不分裂的单叶,枝条顶部的叶
5期 王文采等: 铁线莲属一新分类系统 445
则为三出复叶;在 C. dimorphophylla W. T. Wang 和 C. variifolia W. T. Wang (sect.
Aspidanthera subsect. Dioicae),前者枝条下部叶为不分裂的单叶,上部叶为三出复叶,后者
小枝基部叶为不分裂的单叶,其上一对叶为三出复叶,其余上部叶则均为具5小叶的羽状复
叶,从上述情况可以看到由不分裂的单叶演变为三出复叶和羽状复叶的趋势。另外,在单花
锡金铁线莲 C. siamensis var. monantha W. T. Wang & L. Q. Li (sect. Viorna subsect.
Connatae),枝条多数叶为三出复叶,枝顶部的叶为不分裂的单叶,这里可看到三出复叶由于
二小叶消失而演变成单叶的演化趋势逆转现象。
在铁线莲属中,多数组中只有少数种具单叶,在 sect. Meclatis、sect. Brachyura、sect.
Viticella、sect. Viorna subsect. Bebaenanthera、sect. Atragenopsis和 sect. Atragene,全部种只
有复叶,而无单叶。在具单叶的种中,具近似上述原始形态的单叶的只有4种1变种: C.
acerifolia、C. cirrhosa var. semitriloba (Lag.) Butt. (sect. Cheiropsis), C. trifida Hook. (sect.
Pseudanemone), C. ranunculoides (sect. Viorna subsect. Connatae)和 C. alternata (sect.
Archiclematis); 另外,C. marmoraria Sneddon (sect. Aspidanthera subsect. Hexapetalae)的单
叶掌状细裂,C. nannophylla (sect. Fruticella)的单叶羽状全裂。其他具不分裂单叶的植物有
24种: C. cirrhosa (sect. Cheiropsis), C. lancifolia、C. songarica、C. xinhuiensis (sect. Clematis),
C. dubia (Endl.) P. S. Green (sect. Aspidanthera), C. actinostemmatifolia W. T. Wang (sect.
Brachiatae), C. fruticosa、C. tomentella (Maxim.) W. T. Wang & L. Q. Li、C. viridis (W. T.
Wang & M. C. Chang) W. T. Wang (sect. Fruticella), C. smilacifolia、C. fulvicoma Rehd. &
Wils.、C. metouensis (sect. Naraveliopsis), C. chiupehensis M. Y. Fang、C. henryi、C. yui W. T.
Wang、C. repens、C. kweichowensis Pei、C. pinchuanensis W. T. Wang & M. Y. Fang (sect.
Viorna subsect. Connatae), C. flabellata Nakai、C. integrifolia、C. ochroleuca Aiton、C.
albicoma Wherry、C. viticaulis Steele、C. fremontii S. Watson (sect. Viorna subsect. Crispae)。
另一方面,具3–4回分裂复叶的种与具掌状分裂单叶的种一样也相当少,只有8种,这又可分
为两类:C. ulbrichiana Pilger、C. seemannii Kuntze、C. millefoliolata、C. rutoides W. T. Wang
(sect. Aspidanthera)等4种的叶为2–3回羽状复叶,C. falciformis Viguier & Perrier、C. dissecta
Baker (sect. Brachiatae), C. anethifolia (sect. Pseudanemone), C. aethusifolia (sect. Viorna
subsect. Connatae)等4种的叶3–4回羽状细裂。由上可见,铁线莲属大部分种的叶为1–2回三
出复叶或1–2回羽状复叶,具叶分裂方面的中等演化水平。
在质地方面,像银莲花属植物,铁线莲属多数种的叶为纸质或草质。一些种的叶呈亚
革质,如亚热带低山的 C. meyeniana、C. finetiana Lévl. & Van.、C. uncinata、C. crassifolia
(sect. Clematis),以及干旱地区的C. songarica (sect. Clematis), C. fruticosa (sect. Fruticella)。
少数种的叶呈革质,如亚热带、热带低山区的 C. armandii Franch.,皱叶铁线莲 C. uncinata
var. coriacea Pamp. (sect. Clematis), C. smilacifolia (sect. Naraveliopsis), C. kweichowensis
(sect. Viorna subsect. Connatae),热带高山的 C. seemannii、C. millefoliolata (sect.
Aspidanthera subsect. Dioicae),干旱地区的 C. nannophylla;产热带新几内亚岛的 C.
sclerophylla W. T. Wang和 C. cruttwellii H. Eichler ex W. T. Wang的三出复叶呈厚革质,
是铁线莲属中最厚、最坚硬的叶。
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2.4 花序
银莲花属植物通常具顶生的1至数花的聚伞花序,据此推测铁线莲属花序的原始类型
可能近似这样的花序。这时,花序的花通常中等大,直径2–4 cm。由这种类型的花序发生二
演化方向(王文采, 1998),一是花序分枝稍增,花数目随之增加,达20–35朵;在少数情况下,如
C. gouriana (sect. Clematis)和 C. sclerophylla (sect. Aspidanthera subsect. Aristatae),花序长
达30 cm,多回分枝,约具100朵花,花小,直径仅0.8–1 cm。另一方向是减化(reduction),花序变
成不分枝,只含1朵花,这时在有些种如 C. kockiana Schneid.、C. ranunculoides (sect. Viorna
subsect. Connatae)和花序分枝发生减化同时,花序梗缩短,苞片变小,这种具1花的花序再进
一步减化的结果是花序梗和苞片一同消失,只剩下1朵仅具花梗的花(王文采, 2002),如 C.
montana (sect. Cheiropsis), C. viridis (sect. Fruticella), C. patens (sect. Viticella)等。这时,单
朵且顶生的花通常中等大 ,有时很大 ,直径可达 7 cm,如 C. chrysocarpa (sect.
Pseudanemone), 或达7–15 cm,如 C. patens, C. lanuginosa (sect. Viticella)。
在直立的草本,如 C. hexapetala、C. recta (sect. Clematis)或小灌木,如 C. songarica、C.
delavayi (sect. Clematis), C. fruticosa (sect. Fruticella), 其花序均顶生;但小灌木 C. viridis
(sect. Fruticella)是个例外,其花序减化为1朵只具花梗的花,生于腋生短枝的顶端,顶生花序
则完全消失(王文采, 2002)。在大多数藤本种类,花序或同时在当年生枝上顶生并腋生,如
C. apiifolia、C. grata、C. grandidentata (Rehd. & Wils.) W. T. Wang、C. vitalba、C.
brevicaudata、C. meyeniana、C. flammula、C. terniflora (sect. Clematis), C. dioica、C.
ligusticifolia、C. microphylla、C. pickeringii Gray、C. glycinoides DC. (sect. Aspidanthera), C.
simensis Fresen.、C. hirsuta (sect. Brachiatae), C. tangutica、C. tibetana var. vernayi (C. E. C.
Fisch.) W. T. Wang (sect. Meclatis), C. pinnata (sect. Tubulosae), C. lasiandra Maxim.、C.
aethusifolia、C. ranunculoides (sect. Viorna subsect. Connatae),或在当年生枝上全部腋生,顶
生花序消失,如 C. potaninii、C. tongluensis (Brühl) Tamura (sect. Cheiropsis), C. tenuipes W.
T. Wang、C. vaniotii Lévl. & Port. (sect. Clematis), C. virginiana、C. polygama Jacq.、C.
campestris、C. linearifolia Steud.、C. aristata、C. paniculata、C. foetida Raoul. (sect.
Aspidanthera), C. mauritiana、C. commutata、C. graveolens、C. dissecta (sect. Brachiatae), C.
orientalis、C. akebioides (Maxim.) Veitch、C. intricata (sect. Meclatis), C. smilacifolia、C.
fulvicoma、C. tashiroi (sect. Naraveliopsis), C. courtoisii Hand.-Mazz.、C. florida、C.
hancockiana Maxim.、C. cadmia Buch.-Ham. ex Hook. f. & Thoms. (sect. Viticella), C. henryi、
C. connata、C. leschenaultiana、C. rehderiana (sect. Viorna subsect. Connatae)。最后一种情
况是在一些藤本种类 ,花序或花与叶同自老枝的腋芽中生出 :在 C. pauciflora (sect.
Aspidanthera subsect. Lasianthae),花枝与叶同自一老枝腋芽中生出;在 C. zemuensis W. W.
Smith (sect. Clematis), C. lasiantha (sect. Aspidanthera subsect. Lasianthae)和 C. japonica
(sect. Viorna subsect. Bebaenanthera),花序与叶同自一老枝腋芽中生出;在 C. montana、C.
gracilifolia、C. cirrhosa (sect. Cheiropsis), C. barbellata、C. pseudopogonandra Finet &
Gagnep. (sect. Viorna subsect. Bebaenanthera),只具花梗的单花与叶同自一老枝腋芽中生
出。另外,在 C. fasciculiflora Franch. (sect. Cheiropsis)和 C. armandii (sect. Clematis),只具花
梗的花或花序分别单独由一老枝腋芽中生出。上述花序或花自老枝腋芽生出的现
5期 王文采等: 铁线莲属一新分类系统 447
象,Tamura (1963, 1967, 1995)认为是对寒冷气候的适应结果。
2.5 花
铁线莲属大多数种的花为两性,只有 sect. Aspidanthera 的全部种(王文采, 2004a)和
sect. Naraveliopsis的少数种(王文采, 2000a)以及 sect. Tubulosae的一些种,花的一种性器
官退化和消失而成为单性。
在铁线莲属,多数种的花梗直,花不下垂,在一些花构造较进化的群如 sect. Viorna
subsect. Crispae 和 sect. Archiclematis(萼片直立,雄蕊被毛),其花梗顶部反曲,花变为下垂,
这当是进化的现象。
2.6 萼片
在毛茛科Ranunculaceae的原始属驴蹄草属Caltha L. 、 鸡爪草属Calathodes Hook. f. &
Thoms.、金莲花属 Trollius L.以及银莲花属的较原始种,如西南银莲花 Anemone davidii
Franch.,萼片为5片,平展,呈黄色或白色,膜质或纸质,倒卵形,顶端圆形或钝,外面近边缘处
被贴短柔毛或无毛,内面无毛;其雄蕊无毛,花丝狭条形,花药呈长圆形,顶端不突起。在铁线
莲属中具符合上述萼片形态的植物只有大约6种,即 sect. Cheiropsis 中的 C. brevipes、C.
potaninii、C. montana、C. gracilifolia、C. venusta和 C. chrysocoma (Wang, 2002)。对本属植
物进行全面观察后可以看到以下演化趋势:(a)数目:铁线莲属中大多数种的花具4萼片,少
数为5片或更多,如在 C. paniculata (sect. Aspidanthera subsect. Hexapetalae)为5–8片,在 C.
florida为5–6片,在 C. patens (sect. Viticella)为6–8片。从上述情况可见4或5片当是原始现
象。(b)卷叠式:铁线莲属大多数种的萼片在花芽时期呈镊合状排列,但在 sect. Cheiropsis
subsect. Williamsianae、sect. Aspidanthera subsect. Hexapetalae及 sect. Pseudanemone诸群
植物的萼片部分覆瓦状排列,部分镊合状排列,这种混合的卷叠式可能衍生于镊合状卷叠
式(Kuntze, 1885; Hutchinson, 1920; Godley, 1977; Tobe, 1980; Tamura, 1987, 1995)。(c)颜色:
根据上述毛茛科原始属萼片的颜色,以及银莲花属萼片主要为白色,可见在铁线莲属中,萼
片白色或黄色是原始特征,而蓝色、紫色、红色等为进化特征。(d)根据上述有关原始属的萼
片均为平展,可见铁线莲属萼片开展的演化趋势是平展→斜上展→直立(王文采, 1989,
1998) (图2)。(e)质地:铁线莲属的萼片通常与银莲花属的萼片一样,呈草质或纸质,但在少
数种,如 C. villosa、C. katangensis (sect. Pseudanemone), C. tibetana var. vernayi (sect.
Meclatis), C. viorna (sect. Viorna subsect. Crispae), 萼片质地变厚,呈亚革质或革质。(f)形
状:如前所述,在铁线莲属,只有 C. montana等6种的萼片呈倒卵形,其他种的萼片则多少变
狭呈倒卵状长圆形、倒披针形、披针形或长圆形,在少数种或变种如毛果扬子铁线莲 C.
puberula var. tenuisepala (Maxim.) W. T. Wang (sect. Clematis)和 C. microphylla (sect.
Aspidanthera subsect. Microphyllae), 萼片呈狭长圆形或宽条形,而后一亚组的 C. fawcettii
Muell.和 C. delicata H. Eichler ex W. T. Wang的萼片呈狭条形,宽仅1.1–1.8 mm,是铁线莲
属中最狭长的萼片(Wang, 2004a) (图3)。(g)顶端:从上述铁线莲属萼片形状从宽到狭的演
变同时可看到其顶端的演化趋势:圆形或钝→急尖→渐近→渐狭(图3)。(h)边缘毛被:在
sect. Cheiropsis 的大多数种,以及在具覆瓦状排列萼片的 sect. Aspidanthera subsect.
Hexapetalae 和 sect. Pseudanemone,萼片边缘本身无毛或有柔毛,但不被密短绒毛,在外面
沿边缘有贴伏的短柔毛,这种情况与银莲花属的萼片相似。在 sect. Cheiropsis 的 C.
植 物 分 类 学 报 43卷 448
wenshanensis W. T. Wang, subgen. Clematis的大多数种(上述具覆瓦状排列萼片的群除外),
以及 subgen. Viorna的全部种,在萼片边缘上有一狭条密短绒毛带,萼片具有这种毛被会对
花芽起到更好的保护作用。(i)边缘延展:铁线莲属的多数种的萼片边缘不延展,只在少数种

图2 3种铁线莲属植物的花,显示萼片的3种开展方向 A, 绣球藤的花,具平展萼片; B, 甘川铁线莲的花,具斜上展
萼片; C, 毛木通的花,具直立萼片。
Fig. 2. Flowers each in three species of Clematis, showing three spreading directions of sepals. A, flower of Clematis
montana Buch.-Ham. ex DC., with sepals horizontally spreading; B, C. akebioides (Maxim.) Veitch, with sepals ascending;
C, C. buchananiana DC., with sepals erect.

图3 11种铁线莲属植物的萼片,显示萼片的不同形状 A, 绣球藤(根据俞德浚8282号标本绘); B, 光叶铁线莲(根据
王晓东,孙坤251号标本绘); C, 短毛铁线莲(根据Polunin et al. 3051号标本绘); D, 菝葜叶铁线莲(根据蒋英12935号标
本绘); E, C. commutata Kuntze (根据Winter & Marais 4942号标本绘); F, 裂叶铁线莲(根据236 Exped. 1353号标本绘);
G, C. tongluensis (Brühl) Tamura (根据Anderson 335号标本绘); H, C. crispa L. (根据B. F. Bush无号标本绘); I, 光柱铁
线莲(根据中德考察队56-904号标本绘); J, C. microphylla DC. (根据I. B. Wilson 508号标本绘); K, C. fawcettii Muell
(根据无名氏24313号标本绘).
Fig. 3. Sepals in 11 species of Clematis, showing their different shapes. A, Clematis montana Buch.-Ham. ex DC. (from
T. T. Yu 8282, PE); B, C. glabrifolia K. Sun & M. S. Yan (from X. D. Wang & K. Sun 251, PE); C, C. puberula Hook. f. &
Thoms. (from Polunin et al. 3051, GH); D, C. smilacifolia Wall. (from Y. Tsiang 12935, PE); E, C. commutata Kuntze
(from Winter & Marais 4942, K); F, C. parviloba Gardn. & Champ. (from 236 Exped. 1353, PE); G, C. tongluensis (Brühl)
Tamura (from Anderson 335, K); H, C. crispa L. (from B. F. Bush s.n., US); I, C. longistyla Hand.-Mazz. (from
Sino-German Exped. 56–904, PE); J, C. microphylla DC. (from I. B. Wilson 508, GH); K, C. fawcettii Muell. (from
Anonymous 24313, K).

5期 王文采等: 铁线莲属一新分类系统 449
发生延展现象:在sect. Fruticella诸种,其卵形萼片在花开放后自边缘生出狭长的膜质边缘;
在sect. Viticella, 除C. hancockiana以外的大多数种的狭披针形萼片边缘在花开放后全长
向外强烈延展成宽翅,使整个萼片呈椭圆形或狭倒卵形,从侧生基出脉向外分出多数平行
细脉;在C. crispa和C. baldwinii (sect. Viorna subsect. Crispae),其披针形的萼片在花开放后
边缘中部以上延展成翅(图3)。(j)萼片内面毛被:铁线莲属多数种的萼片内面像银莲花属的
萼片一样无毛,但也有不少种的萼片内面被毛:在sect. Clematis subsect. Clematis、sect.
Aspidanthera subsect. Dioicae、sect. Brachiatae、sect. Pseudanemone和sect. Meclatis的多数
种,sect. Viorna subsect. Connatae和sect. Atragene的部分种的萼片内面被短柔毛;另外,少数
种如C. fulvicoma (sect. Naraveliopsis)、C. villosa、C. uhehensis Engler、C. katangensis (sect.
Pseudanemone)的萼片内面密被短绒毛,此情况可作为与组内种类区别的进化特征。
2.7 雄蕊
根据上述毛茛科原始属以及银莲花属的雄蕊形态,可以看到铁线莲属雄蕊的以下演
化趋势(Wang, 1998): (a)毛被:无毛→花丝有毛,花药无毛→花丝和花药均有毛。(b)花丝:
条形或狭条形→下部或上部变宽→整体变宽,如 C. robertsiana Aitch & Hemsl. (sect.
Atragenopsis); 平→皱缩(C. crassifolia, sect. Clematis)。(c)花药:长圆形,长0.8–3.5 mm→狭
长圆形→条形,长3–10 mm; C. katangensis (sect. Pseudanemone)的条形花药长8–9.5 mm
(Wang, 2004c), C. lanuginosa (sect. Viticella)的条形花药长达10 mm (Fang, 1980),均是铁线
莲属中最长的花药。(d)药隔:不突出→稍突出,突起小尖头长约0.1 mm,如在 C. meyeniana
(sect. Clematis),或突起圆锥状或近球状 ,长 0.1–0.2 mm,如在 C. pickeringii (sect.
Aspidanthera)→明显突出,或突起呈钻形,长达4 mm,如在 C. aristata (sect. Aspidanthera),或
突起呈狭片形,如 sect. Naraveliopsis的种,一般长1–3 mm,在 C. papillosa H. Eichler长达8.5
mm,在 C. macgregorii Merr.长达10 mm (王文采, 2000a, 2004a) (图4)。
2.8 花粉
铁线莲属的花粉包含三沟、散沟、散孔3种类型,大多数种具三沟花粉,后者是本属花粉
的主要类型(Tobe, 1974; Tarasevich & Serov, 1986; Kapoor et al., 1989; 杨宗愈,黄增泉,
1992; Yano, 1992; Nowicke & Skvarla, 1995; 张镱锂, 1998)。张镱锂(1998)对中国铁线莲属
多种植物的花粉进行了深入研究,认为其演化趋势是:由三沟花粉经过散沟花粉演化出散
孔花粉,或由三沟花粉直接形成散沟花粉。Kapoor et al. (1989)研究了32种印度铁线莲属的
花粉,其中多数种具三沟花粉,只1种即 C. cadmia 具散孔花粉,他们在做过分析之后指出:
“The pollen features in Clematis exhibit a predominently negatively reticulate sexine pattern
and are of little phylogenetic significance”。但从上引有关铁线莲属花粉研究工作可以了
解花粉形态对解释 sect. Viticella和 sect. Tubulosae的系统发育以及分类均有重要意义:sect.
Viticella的 subsect. Floridae具散孔花粉,而 subsect. Viticellae则具三孔花粉;sect. Tubulosae
的原始群 subsect. Pinnatae 具三沟花粉,而进化群 subsect. Tubulosae 具散孔花粉。另外,
在 sect. Viorna subsect. Crispae,产亚欧二洲的C. fusca和Ｃ. integrifolia具三沟花粉,而产美
国的 C. viorna等种则具散孔花粉,对了解此亚组的系统发育及分类等方面,花粉也可起重
要的作用。
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2.9 退化雄蕊
铁线莲属的大多数种均具无退化雄蕊的两性花,只有 C. metouensis、C. loureiriana、C.
liboensis Z. R. Xu (sect. Naraveliopsis)的两性花的外方雄蕊的花药消失而成为狭条形的退
化雄蕊, 以及重瓣铁线莲 C. florida var. plena D. Don (sect. Viticella)和 sect. Atragene组诸
种的两性花具花瓣状退化雄蕊。此外,在具单性花的 sect. Aspidanthera的雌花中,可以看到
退化雄蕊在数目上的演化趋势:多数(subsect. Dioicae、subsect. Lasianthae)→少数(subsect.
Microphyllae、subsect. Aristatae、subsect. Hexapetalae→0 (subsect. Insidiosae) (王文采,
2004a)。
2.10 花柱
铁线莲属大多种的花柱丝形,长(3–)6–10(–20) mm,密被直展的长柔毛,花期后强烈伸
长,毛(长达4–5 mm)随之开展,整个宿存花柱呈羽毛状(plumose),通常长达2.5–6 cm,在

图4 14种铁线莲属植物的雄蕊 A, 美花铁线莲(根据四川考察队8231号标本绘); B, 绣球藤(根据冯国楣960号标本
绘); C, 女萎(根据刘昉勋2626号标本绘); D, C. virginiana L. (根据Churchill 3920号标本绘); E, 毛柱铁线莲(根据王战
30632号标本绘); F, 威灵仙(根据李学根200084号标本绘); G, 大花威灵仙(根据安徽调查队59-252号标本绘); H, 毛
叶铁线莲(根据杭州植物园737号标本绘); I, 菝葜叶铁线莲(根据蒋英12935号标本绘); J, 甘青铁线莲(根据郑思绪425
号标本绘); K, 芹叶铁线莲(根据孔宪武921号标本绘); L, 单叶铁线莲(根据李洪钧1586号标本绘); M, 贵州铁线莲(根
据安顺调查队1245号标本绘); N, C. robertsiana Aitch & Hemsl. (根据Maarsukh 14962号标本绘)。
Fig. 4. Stamens in 14 species of Clematis. A, Clematis potaninii Maxim. (from Sichuan Exped. 8231, PE); B, C.
montana Buch.-Ham. ex DC. (from K. M. Feng 960, PE); C, C. apiifolia DC. (from F. X. Liu 2626, PE); D, C. virginiana
L. (from Churchill 3920, MO); E, C. meyeniana Walp. (from C. Wang 30632, PE); F, C. chinensis Osb. (from X. G. Li
200084, PE); G, C. courtoisii Hand.-Mazz. (from Anhui Exped. 59-252, PE); H, C. lanuginosa Lindl. (from Hangzhou Bot.
Gard. 737, PE); I, C. smilacifolia Wall. (from Y. Tsiang 12935, PE); J, C. tangutica (Maxim.) Korsh. (from S. X. Zheng
425, PE); K, C. aethusifolia Turcz. (from H. W. Kung 921, PE); L, C. henryi Oliv. (from H. J. Li 1586, PE); M, C.
kweichowensis Pei (from Anshun Exped. 1245, PE); N, C. robertsiana Aitch. & Hemsl. (from Maarsukh 14962, GH).
5期 王文采等: 铁线莲属一新分类系统 451
C. drummondii (sect. Aspidanthera subsect. Dioicae) (Wang, 2004a)和 C. baldwinii (sect.
Viorna subsect. Crispae) (Pringle, 1997)可达10 cm,在 C. grandiflora DC.和 C. longicauda
Steud. ex A. Rich. (sect. Viorna subsect. Connatae)可达10–12 cm,而与白头翁属 Pulsatilla
Adans.的宿存花柱十分相似,是对风力传播果实的适应。但铁线莲属的一些种,如 C. florida
(Finet & Gagnepain, 1903)、C. huchouensis、C. viticella、C. campaniflora (图5: F) (sect.
Viticella), C. crispa (图5: E)、C. pitcheri (sect. Viorna subsect. Crispae)的花柱长约10 mm,在
花期之后稍伸长,其上的毛较短且贴伏,整个宿存花柱呈尾状或钻状,而不呈羽毛状。另有2
种 C. brachyura (sect. Pterocarpa)和 C. cadmia (sect. Viticella)的花柱很短,长1–2 mm,花期
后不伸长或稍伸长,呈钻状,被贴伏短毛或无毛。比铁线莲属原始的银莲花属的花柱通常短,
长1–2 mm,或不存在,稀较长(Ulbrich, 1905)。因此,Brandenburg (2000)认为铁线莲属的宿存
花柱的演化趋势为: not elongate 1 elongate 2自有其道理。但对上述具较短、非羽毛状花柱
的诸种的果实 ,以及有关近缘群的花柱和果实加以考虑 ,则会得出其他的认识。C.
brachyura的瘦果十分特化,强烈扁压,具宽翅(图5: H),此种可能由 sect. Clematis演化而来,
而在 sect. Clematis,宿存花柱均伸长呈羽毛状,瘦果均无翅。C. cadmia (图5: G) 、C.
huchouensis、C. viticella、C. campaniflora (图5: F)的瘦果均强烈扁压,有粗或肿胀的边缘框;
而在 sect. Viticella 同组中的原始种 C. hancockiana 以及其他近缘种 C. longistyla、 C.
courtoisii、C. patens等植物,宿存花柱均伸长呈羽毛状,瘦果不强烈扁压,无肿胀边缘框。C.
crispa (图5: E)和Ｃ. pitcheri 的瘦果均强烈扁压,具肿胀的边缘框;而在同一亚组 subsect.
Crispae 中的多数种的瘦果均具伸长的羽毛状宿存花柱,均无肿胀的边缘框;sect. Viorna
subsect. Crispae源自 subsect. Connatae (Tamura, 1995),而在 subsect. Connatae,所有种的宿
存花柱均伸长呈羽毛状,瘦果不强烈扁压,无肿胀边缘框。从上述情况可以推测 C. viticella、
C. crispa等种的宿存花柱弱度伸长,不呈羽毛状的现象,以及Ｃ. brachyura和 C. cadmia花
柱缩短现象乃是随着瘦果特化发生的花柱由短变长演化趋势的逆转现象。至于花柱原始
的由短到长的演化,可能存在于已灭绝的铁线莲属原始群(见下)中,在现存的种类中已不
能见到了。
2.11 果实
铁线莲属多数种的瘦果两侧扁压,侧面稍鼓起,通常卵形或椭圆形,长2–5(–7) mm,宽
1–3(–5) mm (图5: A)。再稍进化的瘦果或强烈两侧扁,近扁平,如 C. montana (sect.
Cheiropsis)的瘦果,或边缘出现狭边缘框,如 C. meyeniana (sect. Clematis)和 C. pickeringii
(图5: B) (sect. Aspidanthera subsect. Aristatae)。更进化的形态是强烈两侧扁,或具肿胀的边
缘框,果体常增大,如在 C. terniflora (sect. Clematis),瘦果呈椭圆形,长6–9 mm (图5: C), 在
C. viticella、C. campaniflora (图5: F) (sect. Viticella), 瘦果呈近圆形,直径约7 mm,再如 C.
crispa (sect. Viorna subsect. Crispae)的瘦果呈椭圆形,长约8 mm (图5: E);或瘦果具延展的
宽边缘,如在 C. puberula (sect. Clematis), 瘦果呈圆形,如一小饼,周围有平的宽边缘(Wang,
2000a),另外在 C. microphylla (sect. Aspidanthera subsect. Microphyllae), 瘦果椭圆形,周围
的宽边缘有横皱(图5: D)。上述诸类型的瘦果或具伸长羽毛状宿存花柱,或具尾状或钻状
稍伸长的宿存花柱,而下面的2种的宿存花柱很短,长约2 mm,被贴伏短毛或无毛,其中 C.
cadmia (sect. Viticella)的瘦果强烈扁压,有边缘框,但边缘不延展,而 C. brachyura (sect.
植 物 分 类 学 报 43卷 452

图5 6种铁线莲属植物的瘦果 A, 粗齿铁线莲(根据王作宾13281号标本绘); B, C. pickeringii Gray (根据Nyman 110
号标本绘); C, 圆锥铁线莲(根据杭州植物园58-983号标本绘); D, C. microphylla DC. (根据R. H. Fisher无号标本绘); E,
C. crispa L. (根据Kral无号标本绘); F, C. campaniflora Brot. (根据Rothmaler 13874号标本绘); G, 短柱铁线莲(根据
Master无号标本绘); H, C. brachyura Maxim. (根据Jack无号标本绘)。
Fig. 5. Achenes in six species of Clematis. A, Clematis grandidentata (Rehd. & Wils.) W. T. Wang (from T. P. Wang
13281, PE); B, C. pickeringii Gray (from Nyman 110, UPS); C, C. terniflora Thunb. (from Hangzhou Bot. Gard. 58-983,
PE); D, C. microphylla DC. (from R. H. Fisher s.n., UPS); E, C. crispa L. (from Kral s.n., US); F, C. campaniflora Brot.
(from Rothmaler 13874, S); G, C. cadmia Buch.-Ham. ex Hook. f. & Thoms. (from Masters s.n., K); H, C. brachyura
Maxim. (from Jack s.n., GH).

Pterocarpa)的瘦果也强烈扁压,沿背、腹缝线向外延展成宽翅(图5: H),这是对风力传播果
实的另一种适应,表现出高度特化水平。
3 属下各群间的亲缘关系
根据上述铁线莲属形态特征的演化趋势可以推测这属的原始群当是:多年生直立草
本植物;幼苗叶互生,茎生叶互生或近互生,为掌状分裂的单叶;花序为顶生的聚伞花序,具
不多的花;花萼辐状,萼片平展,倒卵形,边缘不具狭短绒毛带;雄蕊无毛,花药长圆形,药隔不
突出;瘦果稍两侧扁,宿存花柱不强烈伸长。在现存的铁线莲属植物中,只有 C. alternata的
叶互生,为掌状浅裂的单叶,近似上述原始群的植物,但为攀援亚灌木,花下垂,萼片直立,紫
5期 王文采等: 铁线莲属一新分类系统 453
红色,雄蕊被毛,这些都是很特化的特征;sect. Cheiropsis的 C. potaninii、C. montana等6种
如上所述,其花构造与上述原始群的近似,但均为木质藤本,叶均对生,为三出或1–2回羽状
复叶,瘦果强烈扁压,这些都属进化特征。所以,在现存铁线莲属植物中已不能找到上述的
原始群,推测此原始群早已灭绝,现存的群当均由其演化而出。
主要根据花构造可以看到在现存铁线莲属植物中存在4条演化干(王文采, 2003,
2005)。包含 C. montana等6原始种的 sect. Cheiropsis代表第一演化干,即绣球藤干,其特征
为花两性,萼片倒卵形或其他变狭的形状,边缘上无毛或有短柔毛,无狭短绒毛带(但有一
例外,C. wenshanensis的萼片边缘有狭短绒毛带),雄蕊无毛,花药药隔通常无突起。在组内,
由于萼片从平展变到直展,形状从倒卵形演变为长圆形、披针形等形状,花序从在当年生枝
上腋生转变为自老枝腋芽中生出,且常退化为只具花梗的单花,而出现组下类群的分化。由
于包含 C. montana等6原始种,sect. Cheiropsis被视为铁线莲属的现存原始群。
第二演化干为欧洲铁线莲干,其特征为花两性,稀单性;萼片平展,稀斜上展或直展,呈
长圆形、倒披针形、披针形或条形,边缘有一条狭短绒毛带(具覆瓦状排列萼片的 sect.
Aspidanthera subsect. Hexapetalae和 sect. Pseudanemone例外);雄蕊无毛,稀有毛;花药长圆
形至条形,药隔顶端无或有突起。此演化干包括10组,基部群为 sect. Clematis。此组的花两
性,萼片平展,白色,雄蕊无毛,花丝条形,原始的 subsect. Clematis 的花药长圆形,进化的
subsect. Rectae的花药条形。sect. Aspidanthera与 sect. Clematis极为相近,但花为单性。sect.
Brachiatae也与 sect. Clematis极为相近,主要区别为雄蕊花丝有柔毛。sect. Pseudanemone
的雄蕊花丝有毛,与 sect. Brachiatae相近,但茎直立,萼片卷叠式常部分为覆瓦状排列,部分
为镊合状排列。sect. Meclatis的雄蕊花丝也被毛,也与 sect. Brachiatae相近,但萼片常斜上
展,呈黄色,雄蕊花丝下部变宽而不同。sect. Fruticella与 sect. Clematis相近,但萼片斜上展,
呈黄色,花开放后边缘延展成狭翅。sect. Naraveliopsis也与 sect. Clematis相近,此组植物的
药隔顶端具短或长的狭片状突起。单型组 sect. Pterocarpa也与 sect. Clematis相近,但花柱
短,不密被长柔毛,果期不强度伸长,不呈羽毛状,瘦果扁平,具宽翅。sect. Viticella的特征为
萼片狭披针形,花开放后边缘强度延展成宽翅,但其原始种 C. hancockiana 的萼片长圆形,
边缘不延展,其花构造与 sect. Clematis 的相似。sect. Tubulosae 含2群,原始群 subsect.
Pinnatae为藤本,花两性,萼片斜上展,白色或淡紫色,长圆形,雄蕊无毛或花丝顶部有少数毛,
花粉具三沟,上述特征与 sect. Clematis 相近;进化群 subsect. Tubulosae 的茎直立,花杂性,
萼片直立,蓝紫色,花丝顶部常有疏毛,花粉具散孔。sect. Pseudanemone 可能源自 sect.
Brachiatae,其他8组均可能源自 sect. Clematis。
第三演化干为尾叶铁线莲干,其萼片直立,边缘有狭短绒毛带,雄蕊花丝自下部至顶端
密被柔毛,花药无毛或有毛。含2组:sect. Viorna 的茎生叶对生,幼苗叶互生或对生;sect.
Archiclematis的幼苗叶及茎生叶都为互生。尾叶铁线莲干与欧洲铁线莲干亲缘关系相近,
二者可能有共同的起源。
第四演化干为长瓣铁线莲干,其特征为木质藤本,萼片薄草质,黄色、蓝色或紫色,边缘
无狭短绒毛带(在 C. sibirica和 C. macropetala,萼片在花芽时期边缘有密柔毛,在花开放后
毛变稀疏),雄蕊被毛,外轮雄蕊的花丝及药隔均展宽,整个雄蕊呈狭倒披针形、披针状条形
或匙形。含2组,sect. Atragenopsis的雄蕊全部能育,sect. Atragene的外轮雄蕊不育,成为花
植 物 分 类 学 报 43卷 454
瓣状退化雄蕊。此演化干可能与尾叶铁线莲干的 sect. Viorna有亲缘关系。
上述四演化干在本文系统中均作为亚属处理,它们及其所包含的诸组之间的亲缘关
系如图6所示。

图6 铁线莲属各群间可能的系统发育亲缘关系示意图
Fig. 6. Diagram showing the putative phylogenetic relationships among the groups of the genus Clematis.

4 属下分类群系统排列
铁线莲属
Clematis L., Sp. Pl. 1: 543. 1753; et Gen. Pl., ed. 5, 242, no. 616. 1753; DC., Syst. 1: 131.
1818; et Prodr. 1: 2. 1824; Benth. in Benth. & Hook. f., Gen. Pl. 1: 3. 1862; Kuntze in Verh.
Bot. Ver. Brand. 26: 83. 1885, p.p.; Prantl in Bot. Jahrb. 9: 249. 1888, p.p. excl. sect.
Naravelia; et in Engler & Prantl, Nat. Pflanzenfam. 3 (2): 62. 1888, p.p. excl. sect. Naravelia;
Koehne, Deuts. Dendr. 152. 1893; Schneid., Ill. Handb. Laubh. 1: 273. 1906; Rehd., Man. Cult.
Trees & Shrubs, ed. 2, 207. 1951; Tamura in Sci. Rep. Osaka Univ. 4: 44. 1955 et 16 (2): 31.
1967; et in Acta Phytotax. Geobot. 38: 38. 1987; et in Hiepko, Nat. Pflanzenfam., Zwei. Aufl.,
17a (4): 368. 1995; Snoeijer in Clematis 1992: 7. 1992; M. Johnson, Klematis 139. 1997, p.p.
excl. sect. Naravelia; Grey-Wilson, Clematis 24. 2000; Brandenb., Meclatis in Clematis 38.
2000. Lectotype: C. vitalba L. (Britton & Brown, 1913; Tamura, 1995).
Clematitis Moench, Meth. 296. 1794.
木质藤本,稀为直立多年生草本、亚灌木或灌木。叶对生,稀互生,为单叶或复叶,具掌
5期 王文采等: 铁线莲属一新分类系统 455
状脉。花序聚伞状。花两性,稀单性;萼片4(–5–8),花瓣状,镊合状排列,稀覆瓦状排列;花瓣
不存在;雄蕊多数,有时外部雄蕊不育成为退化雄蕊;心皮多数,稀少数,子房具1胚珠,花柱
在果期宿存,通常伸长呈羽毛状。瘦果多少两侧扁。
约355种,广布世界各大洲。
本文系统承认本属包含4亚属15组如下。
亚属1 绣球藤亚属
subgen. 1. Cheiropsis Peterm., Deuts. Fl. 3. 1846; Grey-Wilson, Clematis 75. 2000.
Lectotype: C. cirrhosa L.
含1组,形态特征及地理分布见下。
组1 绣球藤组
sect. 1. Cheiropsis DC., Syst. 1: 162. 1818; et Prodr. 1: 9. 1824; Tamura in Sci. Rep.
Osaka Univ. 16 (2): 34. 1967; et in Hiepko, Nat. Pflanzenfam., ed. 2, 17a (4): 379. 1995; M.
Johnson, Klematis 365. 1997; W. T. Wang in Acta Phytotax. Sin. 40: 196. 2002.——
Cheiropsis (DC.) Bercht. & Presl, Rostl. i. Ranunculac. 11. 1823.——Cheiropsis (DC.) Spach,
Hist. Nat. Veg. Phan. 7: 260. 1839. Lectotype: C. cirrhosa L. (Tamura, 1955).
花两性。萼片镊合状排列,稀覆瓦状排列,平展或有时斜上展或直立,倒卵形、长圆形、
长椭圆形、倒披针形或披针形,除1种(文山铁线莲)之外,边缘不被狭短绒毛带。雄蕊无毛;
花丝条形;花药通常长圆形或狭长圆形,稀条形。宿存花柱伸长,羽毛状。
Flower bisexual. Sepals valvate, rarely imbricate, spreading or sometimes ascending or
erect, obovate, oblong, long elliptic, oblanceolate or lanceolate, on margin without a narrow
velutinous strip (except in one species, C. wenshanensis). Stamens glabrous; filaments linear;
anthers oblong or narrowly oblong, rarely linear. Persistent styles elongate, plumose.
约23种,隶属8亚组,分布于亚洲,欧洲南部,非洲北部。
亚组1 美花铁线莲亚组
Subsect. 1. Potaninianae W. T. Wang in Acta Phytotax. Sin. 36: 162. 1998, sphalm.
Potanianae, et 38: 401, 499. 2000 et 40: 200. 2002.——sect. Clematis subsect. Potaninianae
M. Johnson, Klematis 410, 451. 1997, p.p. excl. C. trichotoma Nakai; Grey-Wilson, Clematis
45. 2000, p.p. excl. C. trichotoma. Type: C. potaninii Maxim.
木质藤本,叶为1–2回羽状复叶。花组成含1–3朵花的聚伞花序,生于当年生枝叶腋。
萼片4–7,平展,倒卵形。花药长圆形或狭长圆形。
Woody vines. Leaves 1–2-pinnate. Flowers in 1–3-flowered cymes arising from leaf axils
of hornotinous branch. Sepals 4–7, spreading, obovate. Anthers oblong or narrowly oblong.
2种: 1. 短梗铁线莲 C. brevipes Rehd., 2. 美花铁线莲 C. potaninii Maxim.。特产中国
西南部。
亚组2 Subsect. 2. Heynianae W. T. Wang in Acta Phytotax. Sin. 38: 402, 499. 2000 et 40:
203. 2002. Type: C. heynei Rau.
木质藤本。叶为三出复叶。聚伞花序顶生或腋生,有3至多数花。萼片4–6,平展,倒披
针形。花药条形。
植 物 分 类 学 报 43卷 456
Woody vines. Leaves ternate. Flowers in 3–many-flowered, terminal or axillary cymes.
Sepals 4–7, spreading, oblanceolate. Anthers linear.
1种: 3. C. heynei Rau。特产印度中部。
亚组3 槭叶铁线莲亚组
Subsect. 3. Acerifoliae W. T. Wang in Acta Phytotax. Sin. 36: 101. 1998 et 38: 499.
2000 et 40: 203. 2002. Type: C. acerifolia Maxim.
直立小灌木。单叶掌状浅裂。萼片5–8,平展,狭倒卵形,无毛。
Small erect shrub. Leaves simple, palmatilobed. Sepals 5–8, spreading, narrowly obovate,
glabrous.
1种: 4. 槭叶铁线莲 C. acerifolia Maxim.。特产中国北京。
亚组4 绣球藤亚组
Subsect. 4. Montanae Schneid., Ill. Handb. Laubh. 290. 1906; Hand.-Mazz. in Acta Hort.
Gotob. 13: 210. 1939, p.p.; M. Johnson, Klematis 386. 1997, p.p.; W. T. Wang in Acta
Phytotax. Sin. 40: 204. 2002.——Clematis sect. Anemoniflora Loudon, Encycl. Trees and
Shrubs 14. 1842. ——subgen. Cheiropsis sect. Montanae (Schneid.) Grey-Wilson, Clematis
75. 2000. Type: C. montana Buch.-Ham. ex DC.
木质藤本。叶为三出复叶,稀羽状复叶。花只具花梗,1或数朵与叶一同自老枝腋芽中
生出,稀单朵或成对生于当年生枝叶腋。萼片4,平展。
Woody vines. Leaves ternate, rarely pinnate. Flowers only pedicellate, 1 or several
together with leaves arising from an axillary bud of old branch, rarely singularly or in pair
arising from leaf axils of hornotinous branch. Sepals 4, spreading.
12种,自中国台湾向西经过中国亚热带地区到喜马拉雅西部。
系1 绣球藤系
Ser. 1. Montanae Rehd. & Wils. in Sarg. Pl. Wils. 1: 331. 1913; Tamura in Sci. Rep.
Osaka Univ. 4: 51. 1955; et in Acta Phytotax. Geobot. 16: 80. 1956; W. T. Wang in Acta
Phytotax. Sin. 38: 500. 2000 et 40: 205. 2002. Lectotype: C. montana Buch.-Ham. ex DC.
萼片倒卵形,顶端圆形或钝。
Sepals obovate, apex rounded or obtuse.
6种: 5. 绣球藤 C. montana Buch.-Ham. ex DC., 6. 薄叶铁线莲 C. gracilifolia Rehd. &
Wils., 7. 丽叶铁线莲 C. venusta M. C. Chang, 8. 金毛铁线莲 C. chrysocoma Franch., 9. 深
裂铁线莲 C. tripartita W. T. Wang, 10. 宁静山铁线莲 C. ningjingshanica W. T. Wang。分
布同绣球藤亚组。
系2 糙毛铁线莲系
Ser. 2. Tongluenses W. T. Wang in Acta Phytotax. Sin. 38: 501. 2000 et 40: 224. 2002.
Type: C. tongluensis (Brühl) Tamura.
萼片长椭圆形、披针形或倒披针形,顶端急尖,渐尖,或渐狭。
Sepals long elliptic, lanceolate, or oblanceolate, apex acute, acuminate, or attenuate.
6种: 11. C. wallichii W. T. Wang, 12. C. manipurensis (Brühl) W. T. Wang, 13. C.
5期 王文采等: 铁线莲属一新分类系统 457
khasiana (Brühl) W. T. Wang, 14. 糙毛铁线莲 C. laxistrigosa (W. T. Wang & M. C. Chang)
W. T. Wang, 15. C. tongluensis (Brühl) Tamura, 16. 文山铁线莲 C. wenshanensis W. T.
Wang。分布于孟加拉,不丹,中国西南部,印度,缅甸北部,尼泊尔东部。
亚组5 戟状铁线莲亚组
Subsect. 5. Hastatae (W. T. Wang) W. T. Wang in Acta Phytotax. Sin. 38: 503. 2000 et
40: 226. 2002.——sect. Clematis subsect. Rectae ser. Hastatae W. T. Wang in Acta Phytotax.
Sin. 36: 159. 1998. Type: C. hastata Finet & Gagnep.
本亚组似绣球藤亚组,但枝条和叶无毛而不同。
This subsection is similar to subsect. Montanae, but differs in its glabrous branches and
leaves.
2种: 17. 戟状铁线莲 C. hastata Finet & Gagnep., 18. 光叶铁线莲 C. glabrifolia K.
Sun & M. S. Yan。特产中国中部。
亚组6 滑叶藤亚组
Subsect. 6. Fasciculiflorae (Tamura) W. T. Wang in Acta Phytotax. Sin. 38: 503. 2000 et
40: 230. 2002.——sect. Flammula subsect. Rectae ser. Fasciculiflorae Tamura in Acta
Phytotax. Geobot. 16 (3): 8. 1956.——sect. Flammula subsect. Fasciculiflorae (Tamura) M.
Johnson, Klematis 604. 1997.——sect. Cheiropsis subsect. Montanae ser. Fasciculiflorae
(Tamura) W. T. Wang in Acta Phytotax. Sin. 36: 162. 1998.——subgen. Cheiropsis sect.
Fasciculiflorae (Tamura) Grey-Wilson, Clematis 94. 2000. Type: C. fasciculiflora Franch.
木质藤本。叶为三出复叶。花2–4朵单独或与叶同自一老枝腋芽中生出,只具花梗。
萼片4,镊合状排列,直立,长圆形。
Woody vines. Leaves ternate. Flowers 2–4 alone or sometimes together with leaves
arising from an axillary bud of old branch, pedicellate only. Sepals 4, valvate, erect, oblong.
1种: 19. 滑叶藤 C. fasciculiflora Franch.。分布中国西南部,缅甸北部,越南北部。
亚组7 Subsect. 7. Williamsianae M. Johnson, Klematis 366. 1997; W. T. Wang in Acta
Phytotax. Sin 38: 504. 2000 et 40: 233. 2002; Grey-Wilson, Clematis 93. 2000. Type: C.
williamsii Gray.
本亚组似绣球藤亚组,但花组成具1花并具花序梗和2苞片的聚伞花序,萼片渐升,覆瓦
状排列而不同。
This subsection is similar to subsect. Montanae, but differs in its flowers in 1-flowered
pedunculate and bibracteate cymes and its ascending imbricate sepals.
1种: 20. C. williamsii Gray。特产日本南部。
亚组8 合苞铁线莲亚组
Subsect. 8. Cirrhosae Prantl in Bot. Jahrb. 9: 259. 1888; M. Johnson, Klematis 365. 1997;
W. T. Wang in Acta Phytotax. Sin. 38: 504. 2000 et 40: 234. 2002. Lectotype: C. cirrhosa L.
Sect. Cheiropsis subsect. Cheiropsis M. Johnson, Klematis 365. 1997; Grey-Wilson,
Clematis 90. 2000. Type: C. cirrhosa L.
本亚组与 subsect. Williamsianae相似,但具1花的聚伞花序通常具总苞,萼片镊合状排
植 物 分 类 学 报 43卷 458
列,直立而不同。
This subsection is similar to subsect. Williamsianae, but differs in its usually involucrate
cymes and valvate, erect sepals.
2种: 21. C. cirrhosa L., 22. 合苞铁线莲 C. napaulensis DC.。分布于中国西南部,喜马
拉雅地区,亚洲西南部,欧洲南部,非洲北部。
亚属2 欧洲铁线莲亚属
Subgen. 2. Clematis. Keener & Dennis in Taxon 31: 38. 1982; Tamura in Acta Phytotax.
Geobot. 38: 40. 1987; et in Hiepko, Nat. Pflanzenfam., Zwei. Aufl., 17a (4): 377. 1995;
Grey-Wilson, Clematis 25. 2000. Type: C. vitalba L.
Subgen. Flammula (DC.) Peterm., Deuts. Fl. 3. 1846, p.p.; Tamura in Acta Phytotax.
Geobot. 38: 42. 1987; et in Hiepko, Nat. Pflanzenfam., Zwei. Aufl., 17a (4): 381. 1995;
Grey-Wilson, Clematis 27. 2000. Lectotype: C. flammula L.
萼片镊合状排列,稀覆瓦状排列,平展,稀斜上展或直立,白色,稀黄色,蓝色或紫色,边缘
有狭短绒毛带(C. sichotealinensis Ulanova、sect. Aspidanthera subsect. Hexapetalae和 sect.
Pseudanemone例外)。雄蕊无毛,稀花丝有毛。
Sepals valvate, rarely imbricate, spreading, rarely ascending or erect, white, rarely yellow,
blue or purple, margin with a narrow velutinous strip (however, in C. sichotealinensis Ulanova,
sect. Aspidanthera subsect. Hexapetalae and sect. Pseudanemone, the velutinous strip is
wanting). Stamens glabrous, rarely with hairy filaments.
约264种,隶属10组,分布世界各大洲。
组1 威灵仙组
Sect. 1. Clematis. Tamura in Sci. Rep. Osaka Univ. 16 (2): 33. 1967, p.p.; M. C. Chang
in Fl. Reip. Pop. Sin. 28: 156. 1980; Tamura in Acta Phytotax. Geobot. 38: 40. 1987; et in
Hiepko, Nat. Pflanzenfam., Zwei. Aufl., 17a (4): 378. 1995; M. Johnson, Klematis 407. 1997,
p.p.; Grey-Wilson, Clematis 34. 2000, p.p.; W. T. Wang in Acta Phytotax. Sin. 41: 6. 2003.
Type: C. vitalba L.
Sect. Flammula DC., Syst. 1: 133. 1818, p.p.
叶为1–2回三出复叶或1–2回羽状复叶,稀为单叶或3回羽状全裂。花两性。萼片4(–5–8),
镊合状排列,平展,白色,稀粉红色,长圆形、倒披针形或披针形,边缘不延展。雄蕊无毛;花丝
条形;花药长圆形或条形,顶端钝或有小尖头。瘦果无翅,具伸长羽毛状宿存花柱。
Leaves 1–2-ternate or 1–2-pinnate, rarely simple or 3-pinnatisect. Flower bisexual. Sepals
4(–5–8), valvate, spreading, white, rarely pink, oblong, oblanceolate or lanceolate, margin not
dilated. Stamens glabrous; filaments linear; anthers oblong or linear, apex obtuse or minutely
apiculate. Achenes not winged; Styles elongate, plumose.
约74种,隶属5亚组,广布于亚洲,欧洲和非洲北部和东部。
亚组1 钝萼铁线莲亚组
Subsect. 1. Clematis. Tamura in Acta Phytotax. Geobot. 38: 41. 1987; M. Johnson,
Klematis 421. 1997; Grey-Wilson, Clematis 34. 2000, p.p.——sect. Vitalba Spach, Hist. Nat.
5期 王文采等: 铁线莲属一新分类系统 459
Veg. Phan. 7: 276. 1839, p.p.——sect. Clematis subsect. Vitalbae Tamura in Sci. Rep. Osaka
Univ. 16 (2): 33. 1967. Type: C. vitalba L.
木质藤本。幼苗叶互生。萼片4(–5),被长达1 mm的短柔毛。花丝不皱缩;花药通常长
圆形或狭长圆形。
Woody vines. Seedling leaves alternate. Sepals 4(–5), with hairs up to 1 mm long.
Filaments not rugose; anthers oblong or narrowly oblong.
29种,隶属2系,广布于亚洲和欧洲。
系1 钝萼铁线莲系
Ser. 1. Clematis.——sect. Vitalbae ser. Euvitalbae Prantl in Bot. Jahrb. 9: 260. 1888;
Schneid., Ill. Handb. Laubh. 1: 290. 1906; Tamura in Sci. Rep. Osaka Univ. 4: 51. 1955.——
Ser. Vitalbae Koehne, Deuts. Dendr. 12. 1893; Rehd., Man. Cult. Trees & Shrubs, ed. 2, 218.
1940. Type: C. vitalba L.
Ser. Pierotianae Tamura in Sci. Rep. Osaka Univ. 4: 52. 1955.——sect. Clematis subsect.
Pierotianae (Tamura) Tamura in Sci. Rep. Osaka Univ. 16 (2): 33. 1967; et in Acta Phytotax.
Geobot. 38: 41. 1987; M. Johnson, Klematis 409. 1997; Grey-Wilson, Clematis 43. 2000, p.p.
Type: C. pierotii Miq.
瘦果不扁平,不具边缘。
Achenes not flattened, nor marginate.
28种: 23. 鼎湖铁线莲 C. tinghuensis C. T. Ting, 24. 女萎 C. apiifolia DC., 25. 马山铁
线莲C. mashanensis W. T. Wang, 26. 细梗铁线莲C. tenuipes W. T. Wang, 27. 台湾铁线莲
C. formosana Kuntze, 28. C. cinnamomoides W. T. Wang, 29. 缅甸铁线莲 C. burmanica
Lace, 30. C. apiculata Hook. f. & Thoms., 31. 云贵铁线莲 C. vaniotii Lévl. & Port., 32. 金
佛铁线莲 C. gratopsis W. T. Wang, 33. 粗齿铁线莲 C. grandidentata (Rehd. & Wils.) W. T.
Wang, 34. C. trichotoma Nakai, 35. 密毛铁线莲 C. pycnocoma W. T. Wang, 36. 两广铁线
莲 C. chingii W. T. Wang, 37. 福贡铁线莲 C. tsaii W. T. Wang, 38. 钝萼铁线莲 C. peterae
Hand.-Mazz., 39. C. vitalba L., 40. 小蓑衣藤 C. gouriana Roxb. ex DC., 41. 秀丽铁线莲 C.
grata Wall., 42. 串鼻龙 C. javana DC., 43. C. mollissima (Hall.) H. Eichler, 44. C.
multistriata H. Eichler, 45. 短尾铁线莲 C. brevicaudata DC., 46. 厚萼铁线莲 C.
wissmanniana Hand.-Mazz., 47. 细木通 C. subumbellata Kurz, 48. 裂叶铁线莲 C. parviloba
Gardn. & Champ., 49. 田村铁线莲 C. tamurae T. Y. Yang & T. C. Huang, 50. C. taeguensis
Y. Lee, 51. C. pierotii Miq.。地理分布同钝萼铁线莲亚组的分布。
系2 短毛铁线莲系
Ser. 2. Puberulae W. T. Wang in Acta Phytotax. Sin. 41: 54. 2003. Type: C. puberula
Hook. f. & Thoms.
瘦果扁平,具宽边缘。
Achenes flattened, broadly marginate.
1种: 52. 短毛铁线莲 C. puberula Hook. f. & Thoms.。分布于中国,缅甸北部,不丹,印度
北部,尼泊尔。
植 物 分 类 学 报 43卷 460
亚组2 厚叶铁线莲亚组
Subsect. 2. Crassifoliae (Tamura) Tamura in Sci. Rep. Osaka Univ. 16 (2): 33. 1967; M.
C. Chang in Fl. Reip. Pop. Sin. 28: 178. 1980; W. T. Wang in Acta Phytotax. Sin. 41: 59. 2003.
——subsect. Rectae ser. Crassifoliae Tamura in Sci. Rep. Osaka Univ. 4: 53. 1955.——sect.
Flammula subsect. Crassifoliae (Tamura) Tamura in Acta Phytotax. Geobot. 38: 43. 1987;
Grey-Wilson, Clematis 28. 2000. ——sect. Flammula subsect. Crassifoliae (Tamura) M.
Johnson, Klematis 604. 1997. Type: C. crassifolia Benth.
本亚组与钝萼铁线莲亚组接近,主要区别在于雄蕊花丝皱缩。
This subsection is related to subsect. Clematis, differing mainly in its rugose stamen
filaments.
1种: 53. 厚叶铁线莲 C. crassifolia Benth.。分布于中国南部,日本南部。
亚组3 保民铁线莲亚组
Subsect. 3. Baominianae (W. T. Wang) W. T. Wang in Acta Phytotax. Sin. 41: 61. 2003.
——subsect. Clematis ser. Baominianae W. T. Wang in l.c. 36: 157. 1998. Type:
C.baominiana W. T. Wang.
本亚组与钝萼铁线莲亚组接近,但花具6枚萼片,萼片的毛较长(1.6–3 mm)而不同。
This subsection is related to subsect. Clematis, differing in its 6-sepalled flower and
sepals with hairs 1.6–3 mm long.
1种: 54. 保民铁线莲 C. baominiana W. T. Wang。分布于中国中部及东部。
亚组4 棉团铁线莲亚组
Subsect. 4. Angustifoliae Tamura in Sci. Rep. Osaka Univ. 4: 55. 1955 et 16 (2): 33.
1967; M. C. Chang in Fl. Reip. Pop. Sin. 28: 1980; W. T. Wang in Acta Phytotax. Sin. 41: 97.
2003.——sect. Flammula subsect. Angustifoliae (Tamura) Tamura in Acta Phytotax. Geobot.
16: 81. 1956 et 38: 43. 1987; M. Johnson, Klematis 603. 1997; Grey-Wilson, Clematis 113.
2000.——sect. Angustifoliae (Tamura) Serov in Bot. Zhurn. 73: 1740. 1988; Tamura in
Hiepko, Nat. Pflanzenfam., Zwei. Aufl., 17a (4): 383. 1995. Type: C. hexapetala Pall.
Sect. Fruticella Tamura in Sci. Rep. Osaka Univ. 16 (2): 34. 1967, p.p.; M. C. Chang in
Fl. Reip. Pop. Sin. 28: 147. 1980, p.p.; M. Johnson, Klematis 651. 1997, p.p.; W. T. Wang in
Acta Phytotax. Sin. 36: 160. 1998, p.p.; Grey-Wilson, Clematis 126. 2000, p.p.
直立小灌木、亚灌木或多年生草本。花序顶生。萼片4–8。
Small erect shrubs, subshrubs, or perennial herbs. Inflorescences terminal. Sepals 4–8.
7种,隶属3系。分布于阿富汗,中国西部达东北部,伊朗,哈萨克斯坦,吉尔吉斯斯坦,蒙
古,尼泊尔,巴基斯坦,俄国西伯利亚,塔吉克斯坦,土库曼斯坦。
系1 准噶尔铁线莲系
Ser. 1. Songaricae Serov in Bot. Zhurn. 73: 1739. 1988; W. T. Wang in Acta Phytotax.
Sin. 41: 98. 2003.——subsect. Songaricae (Serov) M. Johnson, Klematis 653. 1997;
Grey-Wilson, Clematis 129. 2000. Type: C. songarica Bunge.
Ser. Ispahanicae Serov in l.c.——subsect. Ispahanicae (Serov) M. Johnson, l.c. Type: C.
5期 王文采等: 铁线莲属一新分类系统 461
ispahanica Boiss.
直立小灌木或亚灌木。萼片4–7,侧生基出脉具少数细脉。
Small erect shrubs or subshrubs. Sepals 4–7, with lateral basal veins having a few
veinlets.
5种 : 55. 披针叶铁线莲 C. lancifolia Bur. & Franch., 56. 定军山铁线莲 C.
dingjunshanica W. T. Wang, 57. 准噶尔铁线莲 C. songarica Bunge, 58. C. ispahanica
Boiss., 59. 银叶铁线莲 C. delavayi Franch.。分布大致与棉团铁线莲亚组相同,但在中国东
北无分布。
系2 Ser. 2. Phlebanthae (M. Johnson) W. T. Wang in Acta Phytotax. Sin. 41: 107. 2003.
——sect. Fruticella Tamura subsect. Phlebanthae M. Johnson, Klematis 653. 1997;
Grey-Wilson, Clematis 130. 2000. Type: C. phlebantha Williams.
本亚组与准噶尔铁线莲亚组相近,但花单生,萼片侧生基出脉有多数细脉而不同。
This subsection is related to subsect. Songaricae, differing in its solitary flowers and
lateral basal veins of sepal with numerous veinlets.
1种: 60. C. phlebantha Williams。特产尼泊尔。
系3 棉团铁线莲系
Ser. 3. Hexapetalae W. T. Wang in Acta Phytotax. Sin. 41: 108. 2003. Type: C.
hexapetala Pall.
多年生草本。萼片(4–)5–6(–8).
Perennial herbs. Sepals (4–)5–6(–8).
1种: 61. 棉团铁线莲 C. hexapetala Pall.。分布于中国北部及东北部,朝鲜,蒙古,俄国西
伯利亚。
亚组5 威灵仙亚组
Subsect. 5. Rectae Prantl in Bot. Jahrb. 9: 259. 1888, p.p.; Schneid., Ill. Handb. Laubh. 1:
288. 1906; Tamura in Sci. Rep. Osaka Univ. 4: 53. 1955, p.p. et 16 (2): 33. 1967; et in Hiepko,
Nat. Pflanzenfam., Zwei. Aufl., 17a (4): 382. 1995; M. C. Chang in Fl. Reip. Pop. Sin. 28: 158.
1980; W. T. Wang in Acta Phytotax. Sin. 41: 111. 2003. Type: C. recta L.
Sect. Flammula auct. non DC.: M. Johnson, Klematis 599. 1997, p.p.; Grey-Wilson,
Clematis 95. 2000, p.p.
本亚组与钝萼铁线莲亚组很接近,主要区别为幼苗叶对生,花药条形。
This subsection is very closely related to subsect. Clematis, differing mainly in its
opposite seedling leaves and linear anthers.
约35种,隶属6系,广布于亚洲,欧洲,非洲北部及东部。
系1 毛柱铁线莲系
Ser. 1. Meyenianae Tamura in Sci. Rep. Osaka Univ. 4: 54. 1955; et in Acta Phytotax.
Geobot. 16: 81. 1956; W. T. Wang in Acta Phytotax. Sin. 41: 115. 2003.——sect. Flammula
subsect. Meyenianae (Tamura) M. Johnson, Klematis 605. 1997; Grey-Wilson, Clematis 104.
2000. Type: C. meyeniana Walp.
植 物 分 类 学 报 43卷 462
叶为三出复叶或一回羽状复叶,稀为单叶或二回羽状复叶;小叶通常全缘。花序生于
当年生枝叶腋。瘦果不扁平,卵形,椭圆形或披针形,无肿胀边缘框。
Leaves ternate or pinnate, rarely simple or 2-pinnate; leaflets usually entire.
Inflorescences arising from leaf axils of hornotinous branch. Achenes not flattened, ovate,
elliptic or lanceolate, not tumidly rimmed.
15种: 62. C. thaiana Tamura, 63. 那坡铁线莲 C. napoensis W. T. Wang, 64. 迦拉萨铁
线莲 C. jialasaensis W. T. Wang, 65. 城固铁线莲 C. chengguensis W. T. Wang, 66. 辛氏铁
线莲 C. sinii W. T. Wang, 67. C. theobromina Dunn, 68. 新会铁线莲 C. xinhuiensis R. J.
Wang, 69. 屏边铁线莲 C. pingbianensis W. T. Wang, 70. 毛柱铁线莲 C. meyeniana Walp.,
71. 浙江铁线莲 C. chekiangensis Pei, 72. 山木通 C. finetiana Lévl. & Van., 73. 陕西铁线
莲 C. shensiensis W. T. Wang, 74. 五叶铁线莲 C. quinquefoliolata Hutch., 75. 屏东铁线莲
C. akoensis Hayata, 76. 舟柄铁线莲 C. dilatata Pei。多数种分布于亚洲东部,只1种分布于
印度南部。
系2 小木通系
Ser. 2. Armandianae W. T. Wang in Acta Phytotax. Sin. 36: 158. 1998 et 41: 132. 2003.
Type: C. armandii Franch.
本系极近毛柱铁线莲系,其花序单独自老枝腋芽中生出而不同。
This series is most closely related to ser. Meyenianae, differing in its infloresceneces
alone arising from axillary buds of old branch.
3种: 77. 凌云铁线莲 C. lingyunensis W. T. Wang, 78. 小木通 C. armandii Franch., 79.
安顺铁线莲 C. zygophylla Hand.-Mazz.。分布于中国亚热带地区,印度,缅甸,越南。
系3 威灵仙系
Ser. 3. Rectae Prantl ex Rehd. & Wils. in Sarg., Pl. Wils. 1: 325. 1913, p.p.; Rehd., Man.
Cult. Trees & Shrubs, ed. 2, 215. 1951; Tamura in Sci. Rep. Osaka Univ. 4: 54. 1955; et in
Acta Phytotax. Geobot. 16: 81. 1956; W. T. Wang in Acta Phytotax. Sin. 41: 137. 2003.——
sect. Rectae Serov in Bot. Zhurn. 73: 1739. 1988. Type: C. recta L.
Subsect. Rectae ser. Chinenses Tamura in ll. cc.——sect. Flammula subsect. Chinenses
(Tamura) M. Johnson, Klematis 603. 1997; Grey-Wilson, Clematis 110. 2000. Type: C.
chinensis Osb.
Sect. Flammula subsect. Flammula M. Johnson, l.c. 602. 1997; Grey-Wilson, l.c. 95.
2000. Type: C. flammula L.
本系接近毛柱铁线莲系,但瘦果扁平,具肿胀边缘框而不同。
This series is related to ser. Meyenianae, differing in its achenes flattened and tumidly
rimmed.
约13种: 80. 威灵仙 C. chinensis Osb., 81. 巴山铁线莲 C. pashanensis (M. C. Chang)
W. T. Wang, 82. 太行铁线莲 C. kirilowii Maxim., 83.秦岭铁线莲 C. obscura Maxim., 84.
文县铁线莲 C. wenxianensis W. T. Wang, 85. 条形铁线莲 C. linearifoliola W. T. Wang, 86.
圆锥铁线莲 C. terniflora DC., 87. C. papuligera Ohwi, 88. C. flammula L., 89. C. tunisiatica
5期 王文采等: 铁线莲属一新分类系统 463
W. T. Wang, 90. C. elisabethae-carolae Greuter, 91. C. recta L., 92. C. lathyrifolia Bess. ex
Trautv.。广布于亚洲东部及西南部,欧洲,非洲北部。
系4 柱果铁线莲系
Ser. 4. Uncinatae Tamura in Sci. Rep. Osaka Univ. 4: 53. 1955; et in Hiepko, Nat.
Pflanzenfam., Zwei. Aufl., 17a (4): 383. 1995; W. T. Wang in Acta Phytotax. Sin. 36: 159.
1998 et 41: 163. 2003.—— sect. Flammula subsect. Uncinatae (Tamura) M. Johnson, Klematis
606. 1997; Grey-Wilson, Clematis 112. 2000. Type: C. uncinata Champ. ex Benth.
本系接近毛柱铁线莲系,但其瘦果呈钻状圆柱形或狭披针形可以区别。
This series is related to ser. Meyenianae, differing mainly in its subulate-terete or
narrowly lanceolate achenes.
2种: 93. 柱果铁线莲 C. uncinata Champ. ex Benth., 94. C. sigensis Engler。分布于亚洲
东部和非洲东部。
系5 Ser. 5. Zemuenses W. T. Wang in Acta Phytotax. Sin. 38: 505. 2000 et 41: 169. 2003.
Type: C. zemuensis W. W. Smith.
木质藤本。叶为二回羽状复叶;小叶具齿。聚伞花序单条与数叶自老枝一腋芽生出或
单独生当年生枝下部叶腋。
Woody vines. Leaves 2-pinnate; leaflets dentate. Cymes solitarily with several leaves
arising from an axillary bud of old branch or solitarily from lower leaf axils of hornotinous
branch.
1种: 95. C. zemuensis W. W. Smith。分布于不丹,尼泊尔。
系6 Ser. 6. Sichotealinenses W. T. Wang in Acta Phytotax. Sin. 38: 504. 2000 et 41: 171.
2003. Type: C. sichotealinensis Ulanova.
本系稍近似 ser. Zemuenses,但聚伞花序在当年枝上顶生或腋生,萼片无毛而不同。
This series is somewhat similar to ser. Zemuenses, differing in its cymes terminal and
axillary on hornotinous branch and its glabrous sepals.
1种: 96. C. sichotealinensis Ulanova。特产俄国远东地区。
在铁线莲属大多数种,萼片多少有柔毛,只有2种,即这里的 C. sichotealinensis 和 sect.
Cheiropsis的 C. acerifolia的萼片完全无毛。
组2 Sect. 2. Aspidanthera Spach, Hist. Nat. Veg. Phan. 7: 283. 1839; Tamura in Hiepko,
Nat. Pflanzenfam., Zwei. Aufl., 17a (4): 380. 1995, s.l.; Snoeijer in Clematis 1992: 16. 1992;
M. Johnson, Klematis 49. 1997; Grey-Wilson, Clematis 58. 2000; W. T. Wang in Acta
Phytotax. Sin. 38: 505. 2000 et 42: 6. 2004, s.l. Lectotype: C. aristata R. Br. ex Ker Gawler.
木质藤本,稀小亚灌木(C. gentianoides, C. marmoraria)。叶对生,为1–2回三出复叶或
1–2回羽状复叶 ,稀单叶或3回羽状复叶。花单性。萼片4(–5–8),平展 ,稀直立(subtrib.
Insidiosae),白色,稀黄色或黄绿色,长圆形或条形。雄蕊无毛;花丝条形;花药长圆形,稀条形,
药隔不突起或具圆锥状或钻形突起。瘦果无翅,具伸长羽毛状宿存花柱。
Woody vines, rarely low subshrubs. Leaves opposite, 1–2-ternate or 1–2-pinnate, rarely
simple or 3-pinnate. Flower unisexual. Sepals 4(–5–8), spreading, rarely erect, white, rarely
植 物 分 类 学 报 43卷 464
yellow or yellow-green, oblong or linear. Stamens glabrous; filaments linear; anthers oblong,
rarely linear; connectives not projected or with conic or subulate projections. Achenes not
winged, with elongate plumose persistent styles.
约73种,隶属6亚组,广布于北、南美洲,大洋洲,亚洲东南部及马达加斯加。
亚组1 Subsect. 1. Dioicae (Prantl) W. T. Wang in Acta Phytotax. Sin. 38: 506. 2000 et 42: 7.
2003. ——sect. Flammula DC. subsect. Vitalbae Prantl ser. Dioicae Prantl in Bot. Jahrb. 9:
260. 1888; Schneid., Ill. Handb. Laubh. 1: 291. 1906.——sect. Clematis subsect. Dioicae
(Prantl) Tamura in Sci. Rep. Osaka Univ. 16 (2): 33. 1967; et in Acta Phytotax. Geobot. 38: 41.
1987; M. Johnson, Klematis 471. 1997; Grey-Wilson, Clematis 47. 2000.——sect. Dioicae
(Prantl) Brandenb., Meclatis in Clematis 38. 2000. Type: C. dioica L.
花序生于当年生枝叶腋。萼片4(–5–6),镊合状排列,长圆形,倒披针形或卵形,比雄蕊稍
长。雄蕊药隔不突起。退化雄蕊多数。
Inflorescences arising from leaf axils of hornotinous branch. Sepals 4(–5–6), valvate,
oblong, oblanceolate or ovate, slightly longer than stamens. Connectives not projected.
Staminodes numerous.
约35种,隶属2系,广布于北美洲和南美洲。
系1 Ser. 1. Dioicae W. T. Wang in Acta Phytotax. Sin. 38: 507. 2000 et 42: 10. 2004. Type:
C. dioica L.
叶为1–2回三出复叶或羽状复叶,有时为单叶,稀为3回羽状复叶,此时小叶长达5.2 cm。
萼片白色,稀黄色。
Leaves once to twice ternate or pinnate, sometimes simple, rarely thrice pinnate, then
leaflets up to 5.2 cm long. Sepals white, rarely yellow.
约33种: 97. C. dimorphophylla W. T. Wang, 98. C. malacocoma W. T. Wang, 99. C.
virginiana L., 100. C. subtriloba G. Don, 101. C. dioica L., 102. C. rhodocarpa Rose, 103. C.
polygama Jacq., 104. C. populifolia Turcz., 105. C. bonariensis Juss. ex DC., 106. C.
variifolia W. T. Wang, 107. C. uruboensis Lourt., 108. C. coahuilensis Keil, 109. C. plukenetii
DC., 110. C. grossa Benth., 111. C. thalictroides Steud., 112. C. malacoclada W. T. Wang,
113. C. haenkeana Presl., 114. C. guadelopae Pers., 115. C. grahamii Benth., 116. C.
caleoides Standley & Steyerm., 117. C. fulvofurfuracea W. T. Wang, 118. C. drummondii
Torr. & Gray, 119. C. catesbyana Pursh, 120. C. ligusticifolia Nutt., 121. C. alborosea Ulbr.,
122. C. flammulastrum Griseb., 123. C. flammulastroides W. T. Wang, 124. C. acapulcensis
Hook. & Arn., 125. C. brasiliana DC., 126. C. affinis St. Hil., 127. C. campestris St. Hil., 128.
C. peruviana DC., 129. C. ulbrichiana Pilger。分布同本亚组。
系2 Ser. 2. Millefoliolatae W. T. Wang in Acta Phytotax. Sin. 36: 507. 2000 et 42: 58. 2003.
Type: C. millefoliolata Eichler.
叶为3或2回羽状复叶;小叶小,长2–15(–22) mm。萼片常淡黄色。
Leaves thrice or twice pinnate; leaflets small, 2–15(–22) mm long. Sepals often
yellowish.
5期 王文采等: 铁线莲属一新分类系统 465
2种: 130. C. seemannii Kuntze, 131. C. millefoliolata Eichler。分布于玻利维亚和秘鲁
安第斯山高山地区。
亚组2 Subsect. 2. Lasianthae (Tamura) W. T. Wang in Acta Phytotax. Sin. 38: 507. 2000 et
42: 61. 2004.——sect. Lasiantha Tamura in Sci. Rep. Osaka Univ. 16 (2): 34. 1967; et in
Hiepko, Nat. Pflanzenfam., Zwei. Aufl., 17a (4): 381. 1995; M. Johnson, Klematis 489. 1997;
Grey-Wilson, Clematis 53. 2000. Type: C. lasiantha Nutt.
本亚组极近 subsect. Dioicae,区别在于本亚组的当年生花枝或花序与数叶同自老枝一
腋芽中生出。
This subsection is most closely related to subsect. Dioicae, differing in its hornotinous
flowering branch or inflorescence arising together with several leaves from an axillary bud of
old branch.
2种: 132. C. pauciflora Nutt., 133. C. lasiantha Nutt.。分布于美国西南部及墨西哥北
部。
亚组3 Subsect. 3. Microphyllae W. T. Wang in Acta Phytotax. Sin. 38: 507. 2000 et 42: 64.
2004. Type: C. microphylla DC.
本亚组近似 subsect. Dioicae,区别在于萼片通常比雄蕊长2–4倍,狭长圆形或条形,退化
雄蕊较少,2–16。
This subsection is similar to subsect. Dioicae, differing in its sepals usually 2–4 times
longer than stamens, narrowly oblong or linear in outline, and in having fewer (2–16)
staminodes.
7种: 134. C. rhodocarpoides W. T. Wang, 135. C. queenslandica W. T. Wang, 136. C.
fawcettii Muell., 137. C. brachystemon Gunn ex W. T. Wang, 138. C. linearifolia Steud., 139.
C. microphylla DC., 140. C. delicata H. Eichler ex W. T. Wang。特产澳大利亚。
亚组4 Subsect. 4. Aristatae (Prantl) W. T. Wang in Acta Phytotax. Sin. 38: 509. 2000 et 42:
97. 2004. ——sect. Flammula DC. subsect. Aristatae Prantl in Bot. Jahrb. 9: 260. 1888; H.
Eichler in Bibl. Bot. 124: 32. 1958; Tamura in Sci. Rep. Osaka Univ. 16 (2): 33. 1967.——
sect. Aristatae (Prantl) Brandenb., Meclatis in Clematis 38. 2000. Lectotype: C. aristata R. Br.
ex Ker Gawler.
Sect. Clematis subsect. Papuasicae H. Eichler in l.c. 35; Tamura in l.c. ——sect.
Aspidanthera subsect. Papuasicae (H. Eichler) M. Johnson, Klematis 494. 1997; Grey-Wilson,
Clematis 63. 2000.——sect. Papuasicae (H. Eichler) Brandenb., l.c. Type: C. papuasica Merr.
& Perry.
Sect. Aspidanthera Spach subsect. Aspidanthera M. Johnson, l.c. 493; Grey-Wilson, l.c.
58. Type: C. aristata R. Br. ex Ker Gawler.
本亚组相似于 subsect. Dioicae, 区别在于本亚组的雄蕊药隔明显突起,退化雄蕊较
少,2–14(–18)。
This subsection is similar to subsect. Dioicae, differing in its conspicuously projected
connectives and fewer (2–14–18) staminodes.
植 物 分 类 学 报 43卷 466
16种,隶属于3系,分布于澳大利亚,印度尼西亚东部,斐济,新喀里多尼亚,巴布亚新几
内亚,所罗门群岛,东帝汶。
系1 Ser. 1. Pickeringianae W. T. Wang in Acta Phytotax. Sin. 38: 510. 2000 et 42: 99. 2004.
Type: C. pickeringii Gray.
木质藤本。萼片4。药隔突起呈球状圆锥形,长0.1–0.2 mm。
Woody vines. Sepals 4. Connective projections globose-conic, 0.1–0.2 mm long.
7种: 141. C. dubia (Endl.) P. S. Green, 142. C. cruttwellii H. Eichler ex W. T. Wang, 143.
C. tuaensis H. Eichler ex W. T. Wang, 144. C. clemensiae H. Eichler, 145. C. archboldiana
Merr. & Perry, 146. C. novocaledoniensis W. T. Wang, 147. C. pickeringii Gray。分布于澳大
利亚北部,印度尼西亚东部,斐济,新喀里多尼亚,巴布新几内亚,所罗门群岛,东帝汶。
系2 Ser. 2. Aristatae W. T. Wang in Acta Phytotax. Sin. 38: 510. 2000 et 42: 105. 2004.
Type: C. aristata R. Br. ex Ker Gawler.
木质藤本。萼片4。药隔突起呈细柱形或钻形,长0.3–4.5 mm.
Woody vines. Sepals 4. Connective projections thinly columnar or subulate, 0.3–4.5 mm
long.
8种 : 148. C. glycinoides DC., 149. C. phanerophlebia Merr. & Perry, 150. C.
sclerophylla W. T. Wang, 151. C. tenuimarginata H. Eichler, 152. C. stenanthera H. Eichler,
153. C. papuasica Merr. & Perry, 154. C. aristata R. Br. ex Ker Gawler, 155. C. clitorioides
DC.。广布于澳大利亚,印度尼西亚东部,巴布亚新几内亚,所罗门群岛。
系3 Ser. 3. Gentianoides W. T. Wang in Acta Phytotax. Sin. 42: 116. 2004. Type: C.
gentianoides DC.
小亚灌木。叶为单叶或三出复叶。萼片4–6。药隔突起呈钻状柱形,长0.5–0.9 mm.
Low subshrubs. Leaves simple or ternate. Sepals 4–6. Connective projections
subulate-columnar, 0.5–0.9 mm long.
1种: 156. C. gentianoides DC.。特产澳大利亚塔斯马尼亚。
亚组5 Subsect. 5. Hexapetalae (Prantl) W. T. Wang in Acta Phytotax. Sin. 38: 510. 2000 et
42: 117. 2004.——sect. Flammula DC. subsect. Vitalbae Prantl ser. Hexapetalae Prantl in Bot.
Jahrb. 9: 260. 1888, p.p. excl. C. microphylla DC. ——sect. Clematis subsect. Hexapetalae
(Prantl) Snoeijer in Clematis 1992: 15. 1992. Lectotype: C. hexapetala L. f. (= C. fosteri
Gmelin).
Sect. Novae-Zeelandiae M. Johnson, Klematis 159. 1997; Grey-Wilson, Clematis 65.
2000. Type: C. fosteri Gmelin.
萼片覆瓦状排列,或部分镊合状排列。退化雄蕊2–13。
Sepals imbricate, or partly valvate. Staminodes 2–13.
11种,隶属于4系,特产新西兰。
系1 Ser. 1. Afoliatae W. T. Wang in Acta Phytotax. Sin. 38: 512. 2000 et 42: 119. 2004.
Type: C. afoliata Buchan.
木质藤本。叶片通常不存在。花梗无小苞片。萼片4。
5期 王文采等: 铁线莲属一新分类系统 467
Woody vines. Leaf blades usually wanting. Pedicels not bracteolate. Sepals 4.
1种: 157. C. afoliata Buchan。
系2 Ser. 2. Maratae W. T. Wang in Acta Phytotax. Sin. 38: 512. 2000 et 42: 119. 2004.
Type: C. marata Armstr.
木质藤本。叶为三出复叶。花梗有明显对生的小苞片。萼片4。退化雄蕊2–4。
Woody vines. Leaves ternate. Pedicels with conspicuous opposite bracteoles. Sepals 4.
Staminodes 2–4.
2种: 158. C. marata Armstr., 159. C. quadribracteolata Colenso。
系3 Ser. 3. Hexapetalae W. T. Wang in Acta Phytotax. Sin. 38: 512. 2000 et 42: 121. 2004.
Type: C. fosteri Gmelin.
木质藤本。叶为1–2回三出复叶。萼片(4–)5–8,披针形或狭长圆形。退化雄蕊3–5(–12)。
Woody vines. Leaves 1–2-ternate. Sepals (4–)5–8, lanceolate or narrowly oblong.
Staminodes 3–5(–12).
7种: 160. C. paniculata Gmelin, 161. C. cunninghamii Turcz., 162. C. foetida Raoul.,
163. C. fosteri Gmelin, 164. C. hookeriana Allan, 165. C. australis Kirk, 166. C. petriei Allan.
系4 Ser. 4. Marmorariae W. T. Wang in Acta Phytotax. Sin. 38: 512. 2000 et 42: 129. 2004.
Type: C. marmoraria Sneddon.
小亚灌木。叶为单叶,掌状全裂,全裂片细裂。萼片5–8,倒卵状椭圆形。退化雄蕊8–13。
Low subshrubs. Leaves simple, palmatisect; segments dissected. Sepals 5–8,
obovate-elliptic. Staminodes 8–13.
1种: 167. C. marmoraria Sneddon。
亚组6 Subsect. 6. Insidiosae W. T. Wang in Acta Phytotax. Sin. 38: 315, 513. 2000 et 42:
130. 2004. Type: C. insidiosa Baill.
木质藤本。萼片4(–5–6),镊合状排列,雄花萼片平展,雌花萼片直立。雌花无退化雄蕊。
Woody vines. Sepals 4(–5–6), valvate; those of staminate flower spreading, and those of
pistillate flower erect. Staminodes wanting in pistillate flowers.
2种,隶属2系,特产马达加斯加。
系1 Ser. 1. Saxicolae (Prantl) W. T. Wang, comb. nov.——sect. Flammula DC. subsect.
Vitalbae Prantl ser. Saxicolae Prantl in Bot. Jahrb. 9: 260. 1888, p.p. quoad C. saxicola Hils. &
Bojer tantum. Lectotype: C. saxicola Hils. & Bojer ex Baill. (=C. insidiosa Baill.).
Subsect. Insidiosae ser. Insidiosae W. T. Wang in Acta Phytotax. Sin. 38: 513. 2000.
Type: C. insidiosa Baill.
叶为具5小叶的羽状复叶; 小叶长(2.5–)3–6 cm。
Leaves 5-foliolately pinnate; leaflets (2.5–)3–6 cm long.
1种: 168. C. insidiosa Baill.。
系2 Ser. 2. Rutoides W. T. Wang in Acta Phytotax. Sin. 38: 513. 2000 et 42: 132. 2004.
Type: C. rutoides W. T. Wang.
叶为2–3回羽状复叶;小叶多数,小,长0.3–2.2 cm。
植 物 分 类 学 报 43卷 468
Leaves 2–3-pinnate; leaflets numerous, small, 0.3–2.2 cm long.
1种: 169. C. rutoides W. T. Wang。
组3 Sect. 3. Brachiatae Snoeijer in Clematis 1992: 12. 1992, ut “Brachiata”; W. T. Wang in
Acta Phytotax. Sin. 42: 292. 2004. Type: C. brachiata Thunb.
Sect. Flammula DC. subsect. Wightianae Prantl in Bot. Jahrb. 9: 261. 1888.——sect.
Meclatis (Spach) Tamura subsect. Wightianae (Prantl) W. T. Wang in Acta Phytotax. Sin. 38:
321. 2000 et 39: 325. 2001. Lectotype: C. wightiana Wall. ex Wight & Arn.
Sect. Clematis subsect. Africanae M. Johnson, Klematis 416. 1997, p.p. excl. C. ibarensis
Baker ssp. edentata (Baker) Viguier & Perrier et C. sigensis Engler.——sect. Meclatis subsect.
Africanae (M. Johnson) Grey-Wilson, Clematis 155. 2000, p.p. Type: C. hirsuta Perr. & Guill.
本组极近 sect. Clematis subsect. Clematis,主要区别为雄蕊花丝被毛。
This section is most closely related to sect. Clematis subsect. Clematis, differing in its
hairy stamen filaments.
24种,隶属2系,多数种分布于非洲大陆,马达加斯加及邻近岛屿,少数种分布于亚洲南
部。
系1 Ser. 1. Wightianae (W. T. Wang) W. T. Wang in Acta Phytotax. Sin. 42: 295. 2004.
——sect. Meclatis (Spach) Tamura subsect. Wightianae (Prantl) W. T. Wang ser. Wightianae
W. T. Wang in Acta Phytotax. Sin. 38: 421. 2000. Type: C. wightiana Wall. ex Wight & Arn.
叶为三出复叶或1–2回羽状复叶,稀为单叶或3回羽状复叶;小叶通常卵形,长圆形或披
针形。腋生聚伞花序具(1–)少数到多数花。萼片长圆形,卵形或披针形。
Leaves ternate or 1–2-pinnate, rarely simple or 3-pinnate; leaflets usually ovate, oblong,
or lanceolate. Axillary cymes (1–)few–many-flowered. Sepals oblong, ovate, or lanceolate.
22种: 170. C. actinostemmatifolia W. T. Wang, 171. C. strigillosa Baker, 172. C.
mauritiana Lam., 173. C. zaireensis W. T. Wang, 174. C. ibarensis Baker, 175. C. simensis
Fresen., 176. C. kakoulimensis Schnell, 177. C. massoniana DC., 178. C. tibestica Quézel, 179.
C. dolichopoda Brenan, 180. C. wightiana Wall. ex Wight & Arn., 181. C. hirsuta Perr. &
Guill., 182. C. brachiata Thunb., 183. C. viridiflora Bertol., 184. C. commutata Kuntze, 185.
C. comoresensis W. T. Wang, 186. C. oweniae Harvey, 187. C. bowkeri Burtt Davy ex W. T.
Wang, 188. C. triloba Thunb., 189. C. thalictrifolia Engler, 190. C. graveolens Lindl.。分布与
组的分布相同。
系2 Ser. 2. Dissectae (W. T. Wang) W. T. Wang in Acta Phytotax. Sin. 42: 328. 2004.——
sect. Meclatis (Spach) Tamura subsect. Wightianae (Prantl) W. T. Wang ser. Dissectae W. T.
Wang in Acta Phytotax. Sin. 38: 423. 2000. Type: C. dissecta Baker.
叶为3–4回羽状全裂复叶,末回裂片条状披针形、条形或狭三角形。腋生花序具1花。
萼片宽椭圆形。
Leaves 3–4-pinnatisect; ultimate lobes linear-lanceolate, linear, or narrowly triangular.
Axillary cymes 1-flowered. Sepals broadly elliptic.
2种: 191. C. falciformis Viguier & Perrier, 192. C. dissecta Baker。特产马达加斯加。
5期 王文采等: 铁线莲属一新分类系统 469
组4 Sect. 4. Pseudanemone Prantl in Bot. Jahrb. 9: 257. 1888, p.p. excl. C. welwitschii
Hiern., C. commutata Kuntze et C. dissecta Baker; et in Engler & Prantl, Nat. Pflanzenfam. 3
(2): 63. 1888; Tamura in Acta Phytotax. Geobot. 38: 40. 1987; et in Hiepko, Nat. Pflanzenfam.,
Zwei. Aufl., 17a (4): 376. 1995; Snoeijer in Clematis 1992: 12. 1992; M. Johnson, Klematis
140. 1997; Brummitt in Kew Bull. 55: 102. 2000; W. T. Wang in Acta Phytotax. Sin. 42: 385.
2004.——subgen. Pseudanemone (Prantl) Grey-Wilson, Clematis 30: 195. 2000. Lectotype: C.
pimpinellifolia Hook. (Tamura, 1995).
Clematopsis Bojer ex Hutch. in Bull. Misc. Inform. Kew 1920: 12. 1920 et 1923: 84.
1923; Exell & Mendonca in Carrisso, Consp. Fl. Angol. 1: 4. 1937; Staner & Léonard in Bull.
Soc. R. Bot. Belg. 82: 338. 1950; Léonard & Milne-Redhead in Bull. Soc. R. Bot. Belg. 83: 47.
1951; Tamura in Sci. Rep. Osaka Univ. 16 (2): 31. 1967; Brummitt in Kew Bull. 31: 156.
1976; Raynal in Adansonia, ser. 2, 18: 3. 1978.
直立亚灌木、小灌木或多年生草本。叶对生,稀轮生,为单叶,三出复叶,羽状复叶或2–4
回羽状全裂复叶。花两性。萼片4(–5–6),覆瓦状排列,或部分镊合状排列,平展,边缘无短绒
毛带。雄蕊花丝条形,有柔毛;花药条形或狭长圆形,无毛;药隔顶端不突出或有1小尖头。
瘦果不扁平,不具翅,具伸长羽毛状宿存花柱。
Subshrubs erect, low shrubs or perennial herbs. Leaves opposite, rarely verticillate,
simple, ternate, pinnate, or 2–4-pinnatisect. Flower bisexual. Sepals 4(–5–6), imbricate or
partly valvate, spreading, on margin without a narrow velutinous strip. Stamen filaments linear,
pubescent; anthers linear or narrowly oblong, glabrous; connective apex not projected or with
a minute apiculum. Achenes not flattened, nor winged; persistent styles elongate, plumose.
约16种,隶属3系,分布于非洲大陆和马达加斯加。
系1 Ser. 1. Pimpinellifoliae W. T. Wang in Acta Phytotax. Sin. 42: 2004. Type: C.
pimpinellifolia Hook.
萼片纸质或膜质,内面无毛或被短柔毛。花药条形或狭长圆形,长2.2–4 mm。
Sepals papery or membranous, inside glabrous or puberulous. Anthers linear or narrowly
oblong, 2.2–4 mm long.
7种: 193. C. trifida Hook., 194. C. macrophylla (Raynal) W. T. Wang, 195. C. bojeri
Hook., 196. C. pseudoscabiosifolia Perrier, 197. C. oligophylla Hook., 198. C. pimpinellifolia
Hook., 199. C. anethifolia Hook.。特产马达加斯加。
系2 Ser. 2. Villosae (Prantl) W. T. Wang in Acta Phytotax. Sin. 42: 399. 2004. ——sect.
Pseudanemone subsect. Villosae Prantl in Bot. Jahrb. 9: 258. 1888, p.p. Type: C. villosa DC.
萼片纸质或亚革质,内面被短绒毛或密短柔毛。花药条形,稀狭长圆形,长2.5–4.5(–5)
mm。
Sepals papery or subcoriaceous, inside velutinous or densely puberulous. Anthers linear,
rarely narrowly oblong, 2.5–4.5(–5) mm long.
3种: 200. C. villosa DC., 201. C. africolineariloba W. T. Wang, 202. C. stanleyi Hook.。
特产非洲大陆。
植 物 分 类 学 报 43卷 470
系3 Ser. 3. Chrysocarpae W. T. Wang in Acta Phytotax. Sin. 42: 407. 2004. Type: C.
chrysocarpa Welw. ex Oliv.
? Sect. Pseudanemone subsect. Spatulifoliae Prantl in Bot. Jahrb. 9: 258. 1888.
萼片纸质或亚革质,内面被短柔毛或短绒毛,稀无毛。花药条形,长达5.8–9.5 mm。
Sepals papery or subcoriaceous, inside puberulous or velutinous, rarely glabrous. Anthers
linear, up to 5.8–9.5 mm long.
6种: 203. C. intraglabra W. T. Wang, 204. C. uhehensis Engler, 205. C. grandifolia
(Staner & Leonard) M. Johnson, 206. C. teuszii (Kuntze) Engler, 207. C. chrysocarpa Welw.
ex Oliv., 208. C. katangensis (Hutch.) M. Johnson。特产非洲大陆。
组5 黄花铁线莲组
Sect. 5. Meclatis (Spach) Baill., Hist. Pl. 1: 57. 1867–1869; Brandenb., Meclatis in
Clematis 100. 2000.——Meclatis Spach, Hist. Nat. Veg. Phan. 7: 272. 1839. ——sect.
Meclatis (Spach) Tamura in Sci. Rep. Osaka Univ. 16 (2): 32. 1967; Ling in Fl. Reip. Pop. Sin.
28: 139. 1980; Tamura in Acta Geobot. 38: 39. 1987; et in Hiepko, Nat. Pflanzenfam., Zwei.
Aufl., 17a (4): 375. 1995, p.p.; Serov in Bot. Zhurn. 73: 1738. 1988; Grey-Wilson in Kew Bull.
44: 34. 1989, p.p. excl. C. graveolens Lindl.; Snoeijer in Clematis 1992: 11. 1992; M. Johnson,
Klematis 329. 1997, p.p. excl. C. graveolens Lindl.; Grey-Wilson, Clematis 155. 2000, p.p.
quoad subsect. Meclatis, excl. subsect. Africanae. Lectotype: Meclatis orientalis (L.) Spach
(=C. orientalis L.) (Tamura, 1987).
Sect. Flammula DC. subsect. Orientales Prantl in Bot. Jahrb. 9: 260. 1888; Schneid., Ill.
Handb. Laubh. 1: 293. 1906, p.p. excl. C. graveolens Lindl.; Hand.-Mazz. in Acta Hort. Gotob.
13: 217. 1939; Rehd., Man. Cult. Trees & Shrubs, ed. 2, 219. 1951, ut ser. Orientales.——sect.
Viorna (Reichb.) Prantl subsect. Orientales (Prantl) Tamura in Sci. Rep. Osaka Univ. 4: 48.
1955; et in Acta Phytotax. Geobot. 16: 80. 1956.——sect. Meclatis subsect. Orientales (Prantl)
Tamura in Sci. Rep. Osaka Univ. 16 (2): 32. 1967; et in Acta Phytotax. Geobot. 38: 40. 1987;
Grey-Wilson in Kew Bull. 44: 34. 1989. Lectotype: C. orientalis L. (Tamura, 1987).
Sect. Flammula subsect. Tanguticae Schneid., l.c.——sect. Viorna subsect. Tanguticae
(Schneid.) Tamura in Sci. Rep. Osaka Univ. 4: 49. 1955; et in Acta Phytotax. Geobot. 16: 80.
1956.——sect. Meclatis subsect. Tanguticae (Schneid.) Tamura in Acta Phytotax. Geobot. 38:
40. 1987. Lectotype: C. tangutica (Maxim.) Korsh. (Tamura, 1987).
木质藤本,稀为小灌木(C. tangutica, C. tibetana)。叶为1–2回羽状复叶。萼片4,通常斜
上展,黄色。雄蕊花丝狭披针形,被柔毛;花药通常狭长圆形,无毛,药隔顶端不突起或具小尖
头。瘦果不扁平,无翅,具伸长羽毛状宿存花柱。
Woody vines, rarely small shrubs. Leaves 1–2-pinnate. Sepals 4, usually ascending,
yellow. Stamen filaments narrowly lanceolate, pubescent; anthers usually narrowly oblong,
glabrous; connective apex not projected or with a minute apiculum. Achenes not flattened, nor
winged; styles elongate, plumose.
约13种 : 209. 甘青铁线莲 C. tangutica (Maxim.) Korsh., 210. 甘川铁线莲 C.
5期 王文采等: 铁线莲属一新分类系统 471
akebioides (Maxim.) Veitch, 211. 黄花铁线莲 C. intricata Bunge, 212. 中印铁线莲 C.
tibetana Kuntze, 213. C. ladakhiana Grey-Wilson, 214. 粉绿铁线莲 C. glauca Willd., 215.
扎达铁线莲 C. zandaensis W. T. Wang, 216. 角萼铁线莲 C. corniculata W. T. Wang, 217.
尾尖铁线莲 C. caudigera W. T. Wang, 218. 齿叶铁线莲 C. serratifolia Rehd., 219. C.
hilariae Koval., 220. C. sarezica Ikonn., 221. 东方铁线莲 C. orientalis L.。广布亚洲北部及
西部干旱地区或山地,西达欧洲东南部。
组6 灌木铁线莲组
Sect. 6. Fruticella Tamura in Sci. Rep. Osaka Univ. 16 (2): 34. 1967, p.p.; M. C. Chang
in Fl. Reip. Pop. Sin. 28: 147. 1980, p.p. excl. C. songarica Bunge, C. lancifolia Bur. &
Franch. et C. delavayi Franch.; Tamura in Acta Phytotax. Geobot. 38: 44. 1987, p.p.; et in
Hiepko, Nat. Pflanzenfam., Zwei. Aufl. 17a (4): 384. 1995, p.p. excl. C. delavayi etc.; Snoeijer
in Clematis 1992: 18. 1992, p.p.; M. Johnson, Klematis 651. 1997, p.p. excl. C. delavayi, C.
lancifolia, C. ispahanica Boiss., C. phlebantha Williams, C. asplenifolia Schrenk et C.
songarica; Grey-Wilson, Clematis 126. 2000, p.p. excl. C. delavayi, C. lancifolia, C.
songarica, C. asplenifolia, C. ispahanica et C. phlebantha; W. T. Wang & L. Q. Li in Acta
Phytotax. Sin. 43: 196. 2005. Type: C. fruticosa Turcz.
小灌木。叶为单叶,不分裂或分裂。萼片4,镊合状排列,斜上展,黄色,卵形,花开放后边
缘延展成膜质狭翅。雄蕊无毛。瘦果不扁平,无翅,具伸长羽毛状宿存花柱。
Small shrubs. Leaves simple, undivided or divided. Sepals 4, valvate, ascending, yellow,
with margins dilated after anthesis into narrow membranous wings. Stamens glabrous.
Achenes not flattened, nor winged; persistent styles elongate, plumose.
5种,隶属2系,分布于中国西部及北部,蒙古。
系1 灌木铁线莲系
Ser. 1. Fruticosae (Tamura) W. T. Wang & L. Q. Li in Acta Phytotax. Sin. 43: 197. 2005.
——sect. Flammula DC. subsect. Fruticosae Tamura in Sci. Rep. Osaka Univ. 4: 55. 1955; et
in Acta Phytotax. Geobot. 16: 81. 1956. Type: C. fruticosa Turcz.
Sect. Fruticella Tamura subsect. Fruticella M. Johnson, Klematis 651. 1997, p.p. excl. C.
delavayi et C. lancifolia; Grey-Wilson, Clematis 126. 2000, p.p. excl. C. delavayi. Type: C.
fruticosa Turcz.
花组成顶生的具花序梗和2苞片的聚伞花序。
Flowers in terminal, pedunculate, bibracteate cymes.
4种: 222. 灌木铁线莲 C. fruticosa Turcz., 223. 小叶铁线莲 C. nannophylla Maxim.,
224. 毛灌木铁线莲 C. canescens (Turcz.) W. T. Wang & M. C. Chang, 225. 灰叶铁线莲 C.
tomentella (Maxim.) W. T. Wang & L. Q. Li。地理分布同组的分布。
系2 绿叶铁线莲系
Ser. 2. Virides W. T. Wang & L. Q. Li in Acta Phytotax. Sin. 43: 207. 2005. Type: C.
viridis (W. T. Wang & M. C. Chang) W. T. Wang.
花单生于腋生短枝的顶端,只具花梗。
植 物 分 类 学 报 43卷 472
Flowers singularly arising from the apexes of axillary short branches, only pedicellate.
1种: 226. 绿叶铁线莲 C. viridis (W. T. Wang & M. C. Chang) W. T. Wang。特产中国
横断山区北部。
组7 菝葜叶铁线莲组
Sect. 7. Naraveliopsis Hand.-Mazz. in Acta Hort. Gotob. 13: 219. 1939; Tamura in Sci.
Rep. Osaka Univ. 4: 55. 1955; H. Eichler in Bibl. Bot. 124: 43. 1958; Tamura in Sci. Rep.
Osaka Univ. 16 (2): 35. 1967; M. Y. Fang in Fl. Reip. Pop. Sin. 28: 226. 1980; Tamura in
Acta Phytotax. Geobot. 38: 42. 1987; et in Hiepko, Nat. Pflanzenfam., Zwei. Aufl., 17a (4):
380. 1995; Snoeijer in Clematis 1992: 16. 1992; M. Johnson, Klematis 511. 1997;
Grey-Wilson, Clematis 54. 2000; W. T. Wang in Acta Phytotax. Sin. 38: 412. 2000. Lectotype:
C. smilacifolia Wall. (Eichler, 1958).
Sect. Pseudonaravelia Janchen in Denkschr. Akad. Wis. Wien, Math.-Nat. Kl. 108(4): 43.
1949, nom. seminud.
Sect. Naraveliocarpa Tamura in Acta Phytotax. Geobot. 51: 127. 2001. Type: C. eichleri
(Tamura) Tamura.
木质藤本。叶为单叶,三出复叶或1–2回羽状复叶。花两性,稀单性。萼片4或6。雄蕊
无毛,稀被毛,有时最外部雄蕊不育;药隔顶端突出成狭片状突起。瘦果不具翅,有伸长羽毛
状宿存花柱。
Woody vines. Leaves simple, ternate, or 1–2-pinnate. Flowers bisexual, rarely unisexual.
Sepals 4 or 6. Stamens glabrous, rarely pubescent, sometimes outermost stamens sterile and
transformed into linear staminodes; connective apex projecting into narrow lamella. Achenes
not winged; persistent styles elongate, plumose.
约21种,隶属3亚组,广布于亚洲热带。
亚组1 菝葜叶铁线莲亚组
Subsect. 1. Smilacifoliae W. T. Wang in Acta Phytotax. Sin. 36: 363. 1998 et 38: 414.
2000. Type: C. smilacifolia Wall.
Subsect. Liboenses ser. Loureirianae W. T. Wang in l.c. 38: 420. 2000. Type: C.
loureiriana DC.
花两性。雄蕊无毛,全部能育,稀最外部雄蕊不育。
Flowers bisexual. Stamens glabrous, all fertile, rarely those outermost sterile.
约18种: 227. 菝葜叶铁线莲 C. smilacifolia Wall., 228. 滇南铁线莲 C. fulvicoma Rehd.
& Wils., 229. 墨脱铁线莲 C. metouensis M. Y. Fang, 230. 粗柄铁线莲 C. crassipes Chun &
How, 231. C. gialaiensis Serov, 232. C. andersonii (Clarke ex Kuntze) H. Eichler, 233. C.
munroiana Wight, 234. 丝铁线莲 C. loureiriana DC., 235. C. hedysarifolia DC., 236. 勐腊
铁线莲 C. menglaensis M. C. Chang, 237. C. bourdillonii Dunn, 238. 片马铁线莲 C.
pianmaensis W. T. Wang, 239. 国楣铁线莲 C. fengii W. T. Wang, 240. C. herrei H. Eichler,
241. C. korthalsii H. Eichler, 242. 长萼铁线莲 C. tashiroi Maxim., 243. C. papillosa H.
Eichler。分布于亚洲热带地区。
5期 王文采等: 铁线莲属一新分类系统 473
亚组2 荔波铁线莲亚组
Subsect. 2. Liboenses W. T. Wang in Acta Phytotax. Sin. 36: 164. 1998 et 38: 420. 2000.
——subsect. Liboenses ser. Liboenses W. T. Wang in Acta Phytotax. Sin. 38: 420. 2000. Type:
C. liboensis Z. R. Xu.
花两性。雄蕊被柔毛,最外部雄蕊不育。
Flowers bisexual. Stamens pubescent, those outermost sterile.
1种: 244. 荔波铁线莲 C. liboensis Z. R. Xu。特产中国贵州东南部。
亚组3 Subsect. 3. Macgregorianae W. T. Wang in Acta Phytotax. Sin. 38: 421. 2000. Type:
C. macgregorii Merr.
花单性。雄蕊和雌花退化雄蕊均无毛。
Flowers unisexual. Stamens and staminodes of pistillate flower all glabrous.
2种: 245. C. antonii (Elmer) H. Eichler, 246. C. macgregorii Merr.。特产菲律宾。
组8 Sect. 8. Pterocarpa Tamura in Sci. Rep. Osaka Univ. 4: 50. 1955 et 16 (2): 34. 1967; et
in Hiepko, Nat. Pflanzenfam., Zwei. Aufl., 17a (4): 385. 1995; Snoeijer in Clematis 1992: 18.
1992; M. Johnson, Klematis 649. 1997; Grey-Wilson, Clematis 125. 2000. Type: C. brachyura
Maxim.
小亚灌木。叶为三出复叶或羽状复叶。聚伞花序顶生。萼片4–6,平展,白色,倒披针形。
雄蕊无毛;花丝条形;花药狭长圆形。瘦果扁平,有宽翅,具钻形短宿存花柱。
Small subshrubs. Leaves ternate or pinnate. Cymes terminal. Sepals 4–6, spreading, white,
oblanceolate. Stamens glabrous; filaments linear; anthers narrowly oblong. Achenes flattened,
broadly winged, with short subulate persistent styles.
1种: 247. C. brachyura Maxim.。特产朝鲜。
组9 铁线莲组
Sect. 9. Viticella (Dill. ex Moench) DC., Syst. 1: 160. 1818; et Prodr. 1: 8. 1824; Prantl
in Bot. Jahrb. 9: 259. 1888; et in Engler & Prantl, Nat. Pflanzenfam. 3 (2): 63. 1888; Koechne,
Deuts. Dendr. 152. 1893; Schneid., Ill. Handb. Laubh. 1: 285. 1906; Hand.-Mazz. in Acta Hort.
Gotob. 13: 198. 1939; Rehd., Cult. Trees & Shrubs, ed. 2, 212. 1951; Tamura in Sci. Rep.
Osaka Univ. 16 (2): 34. 1967; M. Y. Fang in Fl. Reip. Pop. Sin. 28: 199. 1980; Tamura in
Acta Phytotax. Geobot. 38: 43. 1987; et in Hiepko, Nat. Pflanzenfam., Zwei. Aufl., 17a (4):
384. 1995; M. Johnson, Klematis 665. 1997; Grey-Wilson, Clematis 114. 2000.——Viticella
Dill. ex Moench, Method. 296. 1794; Spach, Hist. Nat. Veg. Phan. 7: 263. 1839; Small, Fl.
Southeast U.S. 437. 1903. ——Clematis subgen. Viticella (Dill. ex Moench) Keener &
Dennis in Taxon 31: 142. 1982; Snoeijer in Clematis 1992: 18. 1992; Pringle in Morin et al.,
Fl. N. Amer. 3: 164. 1997. Type: Viticella deltoidea Moench (= Clematis viticella L.).
Clematis sect. Tessen Tamura in Sci. Rep. Osaka Univ. 4: 49. 1955; et in Acta Phytotax.
Geobot. 16: 80. 1956. Type: C. florida Thunb.
木质或草质藤本。叶为1–2回三出复叶或羽状复叶,稀为单叶。花两性。萼片4–8,平
展或斜上展,边缘在花开放后强烈延展,稀不延展。雄蕊无毛,或花丝近顶部有毛;花药条形,
植 物 分 类 学 报 43卷 474
有时狭长圆形。瘦果不扁平,具伸长羽毛状宿存花柱,或扁平,有肿胀边缘框,具较短尾状或
钻形宿存花柱。
Woody or herbaceous vines. Leaves 1–2-ternate or pinnate, rarely simple. Flowers
bisexual. Sepals 4–8, spreading or ascending, with margins strongly dilated after anthesis,
rarely not dilated. Stamens glabrous, or filaments near apex pilose; anthers linear, sometimes
narrowly oblong. Achenes not flattened, and with elongate plumose persistent styles, or
flattened, tumidly rimmed, and with shorter tail-like or subulate persistent styles.
11种,隶属3亚组,分布于亚洲东部和西南部和欧洲南部。
本组的 C. hancockiana的花具4枚长圆形萼片,边缘不延展,其花构造与 sect. Clematis
的相同,因此是 sect. Viticella的原始种。但是,此种的萼片紫色,花粉具散孔,这些均是进化
特征,这说明 sect. Viticella 的真正原始群(可能花具4枚白色萼片和三沟花粉)早已灭绝。
Erickson (1943)认为本组的 C. viticella应属于 sect. Viorna s. str.。Tamura (1967, 1995)根据
C. crispa (sect. Viorna subsect. Crispae)的小叶坚纸质,全缘,花单生叶腋,较大,萼片边缘延
展等特征与 sect. Viticella相似,也认为 sect. Viticella与 C. viorna群有亲缘关系。但是上面
关于 C. hancockiana系统地位的讨论说明 sect. Viticella系由 sect. Clematis演化而出;另一
方面,C. viorna 群系由 C. connata 群演化而出(Tamura, 1995),所以,sect. Viticella 与 sect.
Viorna的相似实为两条演化路线发展结果的趋同演化(convergence)现象。
亚组1 毛萼铁线莲亚组
Subsect. 1. Hancockianae W. T. Wang, subsect. nov. Type: C. hancockiana Maxim.
Folia 1–2-pinnata vel ternata. Flores in cymas axillares 1-floras manifeste 2-bracteatas
dispositi. Sepala 4, patentia, purpurea, oblonga, margine post anthesin haud dilatata, trinervia,
nervis basalibus lateralibus nervillis carentibus. Stamina glabra. Pollenia pantoporata. Achenia
haud complanata, stylis persistentibus valde elongatis plumosis.
叶为1–2回羽状复叶,或为三出复叶。花组成腋生、具1花和2枚明显苞片的聚伞花序。
萼片4,平展,紫色,花期后边缘不展宽,具3条基出脉,侧生基出脉无细脉。雄蕊无毛。花粉具
散孔。瘦果不扁平;宿存花柱强烈伸长,羽毛状。
1种: 248. 毛萼铁线莲 C. hancockiana Maxim.。特产中国东部。
亚组2 铁线莲亚组
Subsect. 2. Floridae Prantl in Bot. Jahrb. 9: 259. 1888; Schneid., Ill. Handb. Laubh. 1:
286. 1906; Hand.-Mazz. in Acta Hort. Gotob. 13: 198. 1939; Tamura in Sci. Rep. Osaka Univ.
4: 50. 1955; et in Acta Phytotax. Geobot. 38: 43. 1987; et in Hiepko, Nat. Pflanzenfam., Zwei.
Aufl., 17a (4): 385. 1995; M. Johnson, Klematis 675. 1997; Grey-Wilson, Clematis 120. 2000.
Lectotype: C. florida Thunb. (Tamura, 1955).
叶为单叶,1–2回三出复叶或1–2回羽状复叶。花组成腋生、具1花和2苞片的聚伞花序,
或单独顶生。萼片5–8,平展,白色或紫色,狭披针形,具3条脉,边缘在花期之后强烈展宽,侧基
生脉具多数平行、1–3回二叉状分枝的细脉。雄蕊无毛。花粉具散孔。瘦果扁压,但通常不
扁平;宿存花柱通常强烈伸长,羽毛状,稀为尾状或几乎不伸长。
Leaves simple, 1–2-ternate, or 1–2-pinnate. Flowers in axillary, 2-bracteate, 1-flowered
5期 王文采等: 铁线莲属一新分类系统 475
cymes, or solitary and terminal. Sepals 5–8, spreading, white or purple, narrowly lanceolate,
3-veined; margins strongly dilated after anthesis; lateral basal veins with numerous parallel
veinlets 1–3 times dichotomous. Stamens glabrous. Pollen pantoporate. Achenes compressed,
usually not flattened; persistent styles usually strongly elongate, plumose, rarely tail-like, or
nearly not elongate.
6种,隶属2系,分布于亚洲东部。
系1 铁线莲系
Ser. 1. Floridae W. T. Wang, ser. nov. Type: C. florida Thunb.
Subgen. Viticella sect. Floridae Snoeijer in Clematis 1992: 19. 1992, nom. illegit.
Folia 1–2-ternata vel 1–2-pinnata. Flores in cymas axillares 1-floras manifeste
2-bracteatas dispositi. Sepala 5–6. Styli persistentes valde elongati, plumosi, raro caudiformes
vel fere haud elongati.
叶为1–2回三出复叶或1–2回羽状复叶。花组成腋生、具1花和2显著苞片的聚伞花序。
萼片5–6。宿存花柱强烈伸长,羽毛状,稀呈尾状或几乎不伸长。
4种: 249. 光柱铁线莲 C. longistyla Hand.-Mazz., 250. 大花威灵仙 C. courtoisii
Hand.-Mazz., 251. 铁线莲 C. florida Thunb., 252. 短柱铁线莲 C. cadmia Buch.-Ham. ex
Hook. f. & Thoms.。分布于中国亚热带地区,越南北部,缅甸北部,印度东北部,孟加拉。
系2 转子莲系
Ser. 2. Patentes (Tamura) W. T. Wang, st. nov. ——sect. Viticella subsect. Patentes
Tamura in Sci. Rep. Osaka Univ. 4: 50. 1955; in Acta Phytotax. Geobot. 38: 43. 1987; et in
Hiepko, Nat. Pflanzenfam., Zwei. Aufl., 17a (4): 385. 1995; M. Johnson, Klematis 682. 1997;
Grey-Wilson, Clematis 123. 2000.——Clematis sect. Patentes (Tamura) Tamura in Sci. Rep.
Osaka Univ. 16(2): 34. 1967.——subgen. Viticella sect. Patentes Snoeijer in Clematis 1992:
19. 1992. Type: C. patens Morr. & Decne.
Sect. Viticella subsect. Lanuginosae M. Johnson, Klematis 681. 1997; Grey-Wilson,
Clematis 124. 2000. Type: C. lanuginosa Lindl.
叶为单叶或三出复叶,稀为羽状复叶。花单独顶生。萼片5–8。宿存花柱强烈伸长,羽
毛状。
Leaves simple or ternate, rarely pinnate. Flowers solitary, terminal. Sepals 5–8. Persistent
styles strongly elongate, plumose.
2种: 253. 毛叶铁线莲 C. lanuginosa Lindl., 254. 转子莲 C. patens Morr. & Decne.。分
布于中国东部和东北部,朝鲜,日本。
亚组3 湖州铁线莲亚组
Subsect. 3. Viticella Tamura in Sci. Rep. Osaka Univ. 16 (2): 34. 1967; M. Johnson,
Klematis 670. 1997; Grey-Wilson, Clematis 114. 2000.——subgen. Viticella sect. Viticella
Snoeijer in Clematis 1992: 18. 1992, p.p. excl. C. cadmia. Type: C. viticella L.
Subsect. Euviticellae Prantl in Bot. Jahrb. 9: 259. 1888; Schneid., Ill. Handb. Laubh. 1:
285. 1906.
植 物 分 类 学 报 43卷 476
叶为1–2回羽状复叶。花组成腋生、有时顶生、具1–7花的聚伞花序。萼片4,斜上展,
狭披针形,具3条脉,边缘在花开放后多少强度延展;侧基出脉具平行细脉。雄蕊花丝顶部被
短缘毛,稀无毛。花粉具三沟。瘦果扁平,具肿胀边缘框,宿存花柱稍伸长,尾状。
Leaves 1–2-pinnate. Flowers in axillary, sometimes also terminal, 1–7-flowered cymes.
Sepals 4, ascending, narrowly lanceolate, 3-veined, with margins more or less strongly dilated
after anthesis; lateral basal veins with parallel veinlets. Stamen filaments near apex ciliolate,
rarely glabrous. Pollen 3-colpate. Achenes flattened, tumidly rimmed, with slightly elongate
tail-like persistent styles.
4种,隶属2系,分布于中国中部和东部以及亚洲西南部和欧洲南部。
系1 湖州铁线莲系
Ser. 1. Huchouenses W. T. Wang, ser. nov. Type: C. huchouensis Tamura.
Sepala alba, margine post anthesin minus dilatata. Stamina glabra.
萼片白色,边缘在花开放后弱度延展。雄蕊无毛。
1种: 255. 湖州铁线莲 C. huchouensis Tamura。特产中国中部和东部。
系2 Ser. 2. Viticellae W. T. Wang, ser. nov. Type: C. viticella L.
Sepala purpurea, coerulea vel rubra, raro alba, margine post anthesin valde dilatata.
Staminum filamenta apice ciliolata, raro glabra.
萼片紫色、蓝色或红色,稀白色,边缘在花开放后强度延展。雄蕊花丝顶部被短缘毛,
稀无毛。
3种: 256. C. viticella L., 257. C. campaniflora Brot., 258. C. rigoi W. T. Wang。分布于
欧洲南部和亚洲西南部。
组10 大叶铁线莲组
Sect. 10. Tubulosae Decne. in Nouv. Arch. Mus. Hist. Nat. Paris. Sér. 2, 4: 203. 1881; H.
Eichler in Bibl. Bot. 124: 18. 1958, excl. syn.; M. Johnson, Klematis 265. 1997; W. T. Wang
in Acta Phytotax. Sin. 39: 6. 2001.——subgen. Tubulosae (Decne.) Grey-Wilson, Clematis 30,
189. 2000. Lectotype: C. heracleifolia DC. (Eichler, 1958).
木质藤本、多年生直立草本、矮亚灌木或灌木。叶为三出复叶或1–2回羽状复叶;小叶
有齿。花两性或单性。萼片4,镊合状排列,斜上展或直立,白色、蓝色或紫色,倒卵状长圆形。
雄蕊花丝条形,通常近顶部有短柔毛。瘦果不扁平,无翅,宿存花柱伸长,羽毛状。
Woody vines, erect perennial herbs, low subshrubs or shrubs. Leaves ternate or
1–2-pinnate; leaflets dentate. Flowers bisexual or unisexual. Sepals 4, valvate, ascending or
erect, white, blue or purple, obovate-oblong. Stamen filaments linear, usually near apex pilose.
Achenes not flattened; persistent styles elongate, plumose.
9种,隶属2亚组,分布于亚洲东部。
由于 C. heracleifolia群诸种的萼片直立,雄蕊常有毛,Prantl (1888)、Schneider (1906)、
Handel-Mazzetti (1939)、Johnson (1997)、Grey-Wilson (2000)误将此群与 C. connata群或 C.
viorna群相联系,另一方面,后三位作者则将 sect. Tubulosae的原始种C. pinnata误置于 sect.
Clematis subsect. Clematis中,造成混乱。这里,C. heracleifolia群与C. connata群或C. viorna
5期 王文采等: 铁线莲属一新分类系统 477
群的相似,正如上述的 C. viticella群与 C. viorna群的相似,也是一种趋同演化现象。
亚组1 羽叶铁线莲亚组
Subsect. 1. Pinnatae (W. T. Wang) W. T. Wang in Acta Phytotax. Sin. 39: 9. 2001.——
sect. Clematis subsect. Pinnatae W. T. Wang in Acta Phytotax. Sin. 36: 159. 1998. Type: C.
pinnata Maxim.
Subsect. Pinnatae ser. Tatarinowianae W. T. Wang in Acta Phytotax. Sin. 39: 9. 2001.
Type: C. tatarinowii Maxim.
木质藤本。叶为三出复叶或1–2回羽状复叶。花序顶生和腋生,通常具多数花。萼片
斜上展,白色。花粉具三沟。
Woody vines. Leaves ternate or 1–2-pinnate. Inflorescences terminal and axillary, usually
many-flowered. Flowers bisexual. Sepals ascending, white. Pollen 3-colpate.
2种: 259. 羽叶铁线莲 C. pinnata Maxim., 260. C. takedana Makino。分布于中国河北
西部、北京、辽宁、黑龙江南部,日本。
史京华(2003)经过野外观察,发现 C. pinnata的叶在分裂方面变异很大,C. pinnata与
C. tatarinowii Maxim.实为同一种,遂在其未发表的论文中正确地将后者并入前者,澄清了
分类学多年存在的一个混乱。
本亚组的2种,即中国的 C. pinnata(萼片白色)和日本的 C. takedana(萼片淡紫色)在体
态及花构造方面极为相似,二者的亲缘关系极为相近,可能是有共同祖先的一对姊妹群,也
可能后者由前者演化而出。但是,Makino(1907)、Kitagawa (1937)、Ohwi (1965)、Tamura (1982)
均认为 C. takedana是 sect. Clematis的 C. apiifolia和本组的 C. stans的杂交种。2003年,
史京华(2003)经过研究又认为 C. pinnata是 sect. Clematis的 C. brevicaudata和本组的 C.
heracleifolia的杂交种。这样就产生2个问题,需加解决:一是在中国大陆和在日本从不同亲
本形成的杂交种为何如此高度相似。二是关于 C. heracleifolia 群(本文系统的 subsect.
Tubulosae)的起源问题。王文采(2001)曾认为 subsect. Pinnatae 的系统位置介于 sect.
Clematis和 subsect. Tubulosae之间, subsect. Tubulosae可能由 subsect. Pinnatae演化而来,
如果 subsect. Pinnatae是由杂交形成的群,而 subsect. Tubulosae为其亲本之一,则后者源出
于何祖先群的问题就会产生。
亚组2 大叶铁线莲亚组
Subsect. 2. Tubulosae (Decne.) W. T. Wang in Acta Phytotax. Sin. 39: 9. 2001.——sect.
Tubulosae Decne. in l.c.; Kitagawa in J. Jap. Bot. 13: 343. 1937; Tamura in Acta Phytotax.
Geobot. 38: 39. 1987; et in Hiepko, Nat. Pflanzenfam., Zwei. Aufl., 17a (4): 372. 1995;
Snoeijer in Clematis 1992: 10. 1992.——sect. Viorna subsect. Tubulosae (Decne.) Prantl in
Bot. Jahrb. 9: 258. 1888, p.p.; Schneid., Ill. Handb. Laubh. 1: 28. 1906; Hand.-Mazz. in Acta
Hort. Gotob. 13: 191. 1939, p.p. excl. C. tatarinowii Maxim.; Tamura in Sci. Rep. Osaka Univ.
16 (2): 32. 1967; M. Y. Fang in Fl. Reip. Pop. Sin. 28: 93. 1980, p.p. excl. C. tatarinowii
Maxim.——subgen. Tubulosa T. Y. A. Yang in Syst. Geogr. Pl. 68: 301. 1999; et in Chiu &
Peng, Proc. Cross-str. Symp. Fl. Divers. & Cons. 92. 1998. Lectotype: C. heracleifolia DC.
直立多年生草本植物,小灌木或亚灌木。花序通常顶生。花通常杂性。萼片直立,蓝
植 物 分 类 学 报 43卷 478
色或紫色。花药具散孔。
Erect perennial herbs, small shrubs or subshrubs. Inflorescences usually terminal.
Flowers usually polygamous. Sepals erect, blue or purple. Pollen pantoporate.
约7种,隶属2系,分布于中国,朝鲜和日本。
系1 大叶铁线莲系
Ser. 1. Tubulosae Rehd. & Wils. in Sarg. Pl. Wils. 1: 320. 1913; Rehd., Man. Cult. Trees
& Shrubs, ed. 2, 209. 1951, p.p. excl. C. ranunculoides Franch.; W. T. Wang in Acta Phytotax.
Sin. 39: 10. 2001. Lectotype: C. heracleifolia DC.
叶为三出复叶。花多数,组成顶生花序。
Leaves ternate. Flowers in terminal, many-flowered inflorescences.
约6种: 261. C. urticifolia Nakai ex Kitagawa, 262. 大叶铁线莲 C. heracleifolia DC.,
263. 卷萼铁线莲 C. tubulosa Turcz., 264. 台湾草牡丹 C. psilandra Kitagawa, 265. C.
speciosa Makino, 266. C. stans Sieb. & Zucc.。分布与大叶铁线莲亚组相同。
系2 高山铁线莲系
Ser. 2. Uniflorae W. T. Wang in Acta Phytotax. Sin. 39: 10. 2001. Type: C. tsugetorum
Ohwi.
叶为三出复叶或羽状复叶。花单朵顶生。
Leaves ternate or pinnate. Flowers solitary, terminal.
1种: 267. 高山铁线莲 C. tsugetorum Ohwi。特产中国台湾。
亚属3 尾叶铁线莲亚属
Subgen. 3. Viorna Gray, Syn. Fl. N. Amer. 1 (1): 5. 1895; Keener & Dennis in Taxon 31:
37. 1982; Pringle in Morin et al., Fl. N. Amer. 3: 167. 1997. Lectotype: C. viorna L.
Viorna Reichb., Handb. 277. 1837; Spach, Hist. Nat. Veg. Phan. 7: 268. 1839; Small, Fl.
Southeast. U.S. 437. 1903; Rydb., Fl. Rocky Mts. 291. 1917.——subgen. Viorna (Reichb.)
Tamura in Acta Phytotax. Geobot. 38: 40. 1987; et in Hiepko, Nat. Pflanzenfam., Zwei. Aufl.,
17a (4): 376. 1995; Grey-Wilson, Clematis 200. 2000. Lectotype: Viorna urnigera Spach (=
Clematis viorna L.).
Subgen. Campanella Tamura in Acta Phytotax. Geobot. 38: 38. 1987; et in Hiepko, Nat.
Pflanzenfam., Zwei. Aufl., 17a (4): 372. 1995, p.p.; Snoeijer in Clematis 1992: 10. 1992; T. Y.
A. Yang & Moore in Syst. Geogr. Pl. 68: 300. 1999; Grey-Wilson, Clematis 133. 2000. Type:
C. lasiandra Maxim.
木质藤本植物、直立多年生草本植物、小亚灌木或灌木。花两性。萼片4,镊合状排列,
直立,长圆形,倒披针形或披针形,边缘上有一狭短绒毛带。雄蕊被毛;花丝条形;花药长圆形
或狭长圆形。
Woody vines, erect perennial herbs, small subshrubs or shrubs. Flowers bisexual. Sepals
4, valvate, erect, oblong, oblanceolate, or lanceolate, on margin with a narrow velutinous strip.
Stamens pubescent; filaments linear; anthers oblong or narrowly oblong.
约72种,隶属2组,广布于亚洲,非洲,欧洲和北美洲。
5期 王文采等: 铁线莲属一新分类系统 479
组1 尾叶铁线莲组
Sect. 1. Viorna (Reichb.) Prantl in Bot. Jahrb. 9: 258. 1888, p.p.; et in Engler & Prantl,
Nat. Pflanzenfam. 3 (2): 63. 1888; Koehne, Deuts. Dendr. 152. 1893; Schneid., Ill. Handb.
Laubh. 1: 275. 1906, p.p.; Kraschen. in Kom., Fl. USSR 7: 312. 1937, p.p.; Rehd., Man. Cult.
Trees & Shrubs, ed. 2, 207. 1951, p.p.; Tamura in Sci. Rep. Osaka Univ. 4: 45. 1955 et 16 (2):
32. 1967, p.p.; M. Y. Fang in Fl. Reip. Pop. Sin. 28: 88. 1980, p.p. Lectotype: C. viorna L.
幼苗叶互生。茎生叶对生,为单叶或复叶。
Seedling leaves alternate. Mature cauline leaves opposite, simple or compound.
约71种,隶属3亚组,分布同尾叶铁线莲亚属。
亚组1 尾叶铁线莲亚组
Subsect. 1. Connatae Koehne, Deuts. Dendr. 158. 1893; Schneid., Ill. Handb. Laubh. 1:
283. 1906; Hand.-Mazz. in Acta Hort. Gotob. 13: 191. 1939; H. Eichler in Bibl. Bot. 124: 18.
1958; Tamura in Sci. Rep. Osaka Univ. 4: 46. 1955 et 16 (2): 79. 1967; M. Y. Fang in Fl. Reip.
Pop. Sin. 28: 96. 1980.——sect. Connatae (Koehne) M. Johnson, Klematis 287. 1997. Type:
C. connata DC.
Trigula Noronha, Verh. Batav. Gen. 5, Art. 4: 4. 1790. Type: T. trifoliata Noronha (=
Clematis leschenaultiana DC.).
Sect. Campanella Tamura in Acta Phytotax. Geobot. 38: 38. 1987; et in Hiepko, Nat.
Pflanzenfam., Zwei. Aufl. 17a (4): 372. 1995; Snoeijer in Clematis 1992: 10. 1992; T. Y. A.
Yang & Moore in Syst. Geogr. Pl. 68: 301. 1999, p.p. excl. C. pseudopogonandra Finet &
Gagnep. et C. tatarinowii Maxim.; Grey-Wilson, Clematis 133. 2000, p.p. excl. C.
pseudopogonandra Finet & Gagnep. Type: C. lasiandra Maxim.
幼苗叶互生,稀对生(芹叶铁线莲)。花通常不下垂,组成在当年枝上顶生或腋生的聚伞
花序。萼片白色,稀黄色或紫色。
Seedling leaves alternate, rarely opposite (C. aethusifolia). Flowers usually not pendulous,
in cymes arising from apex or leaf axils of hornotinous branch. Sepals white, rarely yellow or
purple.
43种,隶属5系,广布于亚洲东部和南部及非洲。
系1 尾叶铁线莲系
Ser. 1. Connatae Rehd. & Wils. in Sarg., Pl. Wils. 1: 322. 1913; Rehd., Man. Cult. Trees
& Shrubs, ed. 2, 210. 1951. Type: C. connata DC.
Subsect. Connatae ser. Henryianae Tamura in Sci. Rep. Osaka Univ. 4: 47. 1955; et in
Acta Phytotax. Geobot. 16: 79. 1956.——sect. Connatae subsect. Henryianae (Tamura) M.
Johnson, Klematis 325. 1997. Type: C. henryi Oliv.
叶为单叶,三出复叶或1–2回羽状复叶。聚伞花序顶生或腋生,具花序梗和2枚苞片,有1
至多数花。萼片外面不具纵肋或纵翅。宿存花柱长2–5.5 cm。
Leaves simple, ternate or 1–2-pinnate. Cymes terminal or axillary, pedunculate,
2-bracteate, 1–many-flowered. Sepals outside not longitudinally costate or winged. Persistent
植 物 分 类 学 报 43卷 480
styles 2–5.5 cm long.
33种: 268. 邱北铁线莲 C. chiupehensis M. Y. Fang, 269. 单叶铁线莲 C. henryi Oliv.,
270. 多花铁线莲 C. jingdungensis W. T. Wang, 271. 锡金铁线莲 C. siamensis Drumm. &
Lace, 272. 尾叶铁线莲C. urophylla Franch., 273. 滇川铁线莲C. kockiana Schneid., 274. C.
nagaensis W. T. Wang, 275. 柱梗铁线莲C. teretipes W. T. Wang, 276. C. nainitalensis W. T.
Wang, 277. 平坝铁线莲 C. clarkeana Lévl. & Van., 278. 青城山铁线莲 C.
qingchengshanica W. T. Wang, 279. C. acuminata DC., 280. 云南铁线莲 C. yunnanensis
Franch., 281. 怒江铁线莲 C. nukiangensis M. Y. Fang, 282. 森氏铁线莲 C. morii Hayata,
283. 海南铁线莲C. hainanensis W. T. Wang, 284. 绣毛铁线莲C. leschenaultiana DC., 285.
莓叶铁线莲 C. rubifolia C. H. Wright, 286. 毛木通 C. buchananiana DC., 287. 黄毛铁线莲
C. grewiifolia DC., 288. 俞氏铁线莲C. yui W. T. Wang, 289. 贵州铁线莲C. kweichowensis
Pei, 290. 曲柄铁线莲 C. repens Finet & Gagnep., 291. 华中铁线莲 C. pseudootophora M. Y.
Fang, 292. 宽柄铁线莲 C. otophora Franch. ex Finet & Gagnep., 293. C. staintonii W. T.
Wang, 294. 湖北铁线莲C. hupehensis Hemsl., 295. 毛花铁线莲C. dasyandra Maxim., 296.
长花铁线莲 C. rehderiana Craib, 297. C. roylei Rehd., 298. C. jeypurensis Beddome ex W. T.
Wang, 299. 合柄铁线莲 C. connata DC., 300. 毛蕊铁线莲 C. lasiandra Maxim.。广布于亚
洲东部和南部。
系2 须蕊铁线莲系
Ser. 2. Pogonandrae W. T. Wang in Acta Phytotax. Sin. 36: 170. 1998. Type: C.
pogonandra Maxim.
叶为三出复叶。花单生当年生枝叶腋,只具花梗,无花序梗和苞片。萼片外面无纵肋
和纵翅,稀在中脉上具1翅。宿存花柱长约2.5 mm。
Leaves ternate. Flowers singularly arising from leaf axils of hornotinous branch, only
pedicellate, not pedunculate, nor bracteate. Sepals outside not longitudinally costate, nor
winged, rarely on midrib 1-winged. Persistent styles ca. 2.5 cm long.
2种: 301.须蕊铁线莲 C. pogonandra Maxim., 302. 神农架铁线莲 C. shenlungchiaensis
M. Y. Fang。特产中国中部和西南部。
系3 Ser. 3. Grandiflorae W. T. Wang in Acta Phytotax. Sin. 38: 334. 2000. Type: C.
grandiflora DC.
叶为三出复叶。聚伞花序腋生。萼片厚纸质,外面具3–5条纵肋,无翅。宿存花柱长达
10–12 cm。
Leaves ternate. Cymes axillary. Sepals thickly papery, outside longitudinally 3–5-costate,
not winged. Persistent styles up to 10–12 cm long.
2种: 303. C. longicauda Steud. ex A. Rich., 304. C. grandiflora DC.。分布于非洲。
系4 毛茛铁线莲系
Ser. 4. Acutangulae W. T. Wang in Acta Phytotax. Sin. 31: 220. 1993 et 36: 170. 1998.
Type: C. acutangula Hook. f. & Thoms.
叶为单叶,1–2回三出复叶或羽状复叶;基生叶常存在。聚伞花序顶生并腋生。萼片外
5期 王文采等: 铁线莲属一新分类系统 481
面具2–4条狭纵翅。宿存花柱长1–2.5 cm。
Leaves simple, 1–2-ternate or pinnate; basal leaves often present. Cymes terminal and
axillary. Sepals outside longitudinally narrowly 2–4-winged. Persistent styles 1–2.5 cm long.
5种: 305. 宾川铁线莲 C. pinchuanensis W. T. Wang & M. Y. Fang, 306. 毛茛铁线莲
C. ranunculoides Franch., 307. C. acutangula Hook. f. & Thoms., 308. 思茅铁线莲 C.
pterantha Dunn, 309. 元江铁线莲 C. yuanjiangensis W. T. Wang。分布于中国西南部,印度,
不丹和尼泊尔。
系5 芹叶铁线莲系
Ser. 5. Aethusifoliae Tamura in Sci. Rep. Osaka Univ. 4: 47. 1955; et in Acta Phytotax.
Geobot. 16: 79. 1956.——sect. Aethusifoliae (Tamura) Serov in Bot. Zhurn. 73: 1739. 1988.
——sect. Connatae subsect. Aethusifoliae (Tamura) M. Johnson, Klematis 327. 1997. Type: C.
aethusifolia Turcz.
幼苗叶对生(Zhang et al., 2000)。叶为3–4回羽状细裂复叶。聚伞花序顶生并腋生。萼
片外面无纵肋和纵翅。宿存花柱长1.6–2.7 cm。
Seedling leaves opposite (Zhang et al., 2000). Leaves 3–4-pinnatisect. Cymes terminal
and axillary. Sepals outside not longitudinally costate, nor winged. Persistent styles 1.6–2.7
cm long.
1种: 310. 芹叶铁线莲 C. aethusifolia Turcz.。广布于中国北部和西北部,蒙古和俄国西
伯利亚。
亚组2 西南铁线莲亚组
Subsect. 2. Bebaenanthera (Edgew.) W. T. Wang in Acta Phytotax. Sin. 39: 16. 2001.
——sect. Bebaenanthera Edgew. in Trans. Linn. Soc. London 20: 25. 1851; Tamura in Sci.
Rep. Osaka Univ. 16 (2): 32. 1967; et in Acta Phytotax. Geobot. 38: 39. 1987; et in Hiepko,
Nat. Pflanzenfam., Zwei. Aufl., 17a (4): 373. 1995; M. Johnson, Klematis 359. 1997;
Grey-Wilson, Clematis 153. 2000. Type: C. barbellata Edgew.
Sieboldia Hoffmgg., Preisv. Nachtr. 28. 1842. Type: S. japonica (Thunb.) Hoffmgg. (=
Clematis japonica Thunb.).
Sect. Paratragene Tamura in Sci. Rep. Osaka Univ. 4: 49. 1955; et in Acta Phytotax.
Geobot. 16: 80. 1956. Type: C. japonica Thunb.
幼苗叶互生。具1花的聚伞花序或花与叶同自一老枝腋芽生出。萼片外面无纵肋和
翅。
Seedling leaves alternate. One-flowered cyme(s) or flower(s) arising together with leaves
from an axillary bud of old branch. Sepals outside not longitudinally costate, nor winged.
6种: 311. C. japonica Thunb., 312. C. obvallata (Ohwi) Tamura, 313. C. tosaensis
Makino, 314. C. bracteolata Tamura, 315. C. barbellata Edgew., 316. 西南铁线莲 C.
pseudopogonandra Finet & Gagnep.。分布于日本,中国西南部,印度北部,尼泊尔和巴基斯坦
北部。

植 物 分 类 学 报 43卷 482
亚组3 褐毛铁线莲亚组
Subsect. 3. Crispae Prantl in Bot. Jahrb. 9: 258. 1888, p.p.; Schneid., Ill. Handb. Laubh.
1: 275. 1906; Hand.-Mazz. in Acta Hort. Gotob. 13: 190. 1939; Tamura in Sci. Rep. Osaka
Univ. 4: 47. 1955 et 16 (2): 32. 1967; et in Acta Phytotax. Geobot. 16: 79. 1956; M. Y. Fang
in Fl. Reip. Pop. Sin. 28: 89. 1980. Lectotype: C. crispa L. (Tamura, 1955).
Viorna Reichb., l.c.; Spach, l.c.——Clematis subgen. Viorna (Reichb.) Tamura in l.c.
1987; Grey-Wilson, Clematis 200. 2000.
Viorna sect. Euviorna Spach, Hist. Nat. Veg. Phan. 7: 268. 1839.
Viorna sect. Viornium Spach, l.c. 270.
Sect. Viorna (Reichb.) Prantl in l.c., p.p.; Erickson in Ann. Missouri Bot. Gard. 30: 1.
1943.
Subgen. Viorna Gray, l.c.; Keener & Dennis in l.c.; Pringle in Morin et al., l.c.; T. Y. A.
Yang & Moore in Syst. Geogr. Pl. 68: 301. 1999.
幼苗叶对生。花多下垂。萼片通常蓝色,紫色或红色,质地有时较厚。瘦果有时扁平,
具肿胀边缘框,宿存花柱有时弱度伸展,呈尾状,不呈羽毛状。
Seedling leaves opposite. Flowers often pendulous. Sepals usually blue, purple or red,
sometimes subcoriaceous in texture. Achenes sometimes flattened and tumidly rimmed;
persistent styles sometimes weakly elongate, tail-like, not plumose.
约22种,隶属6系,分布于亚洲东北部和西部,欧洲,美国和墨西哥北部。
系1 褐毛铁线莲系
Ser. 1. Fuscae Tamura in Acta Phytotax. Geobot. 16: 79. 1956. ——sect. Viorna subsect.
Fuscae (Tamura) M. Johnson, Klematis 538. 1997; Grey-Wilson, Clematis 31. 2000. Type: C.
fusca Turcz.
攀援或直立草本或亚灌木。叶为羽状复叶,有时为单叶。萼片厚,外面被褐色毛,边缘
不延展。花粉具三沟。瘦果不扁,无肿胀边缘框;宿存花柱伸长,羽毛状。
Scandent or erect herbs, or subshrubs. Leaves pinnate, sometimes simple. Sepals thick,
outside covered with brown hairs, margin not dilated. Pollen 3-colpate. Achenes not flattened,
nor tumidly rimmed; persistent styles elongate, plumose.
约3种: 317. 褐毛铁线莲 C. fusca Turcz., 318. C. ajanensis (Regel & Tiling) Kuntze,
319. C. flabellata Nakai。分布于中国东部和东北部,俄国西伯利亚东部,朝鲜和日本。
系2 全缘铁线莲系
Ser. 2. Integrifoliae (Erickson ex M. Johnson) W. T. Wang, st. nov. ——sect. Viorna
subsect. Integrifoliae Erickson [in Ann. Missouri Bot. Gard. 30: 35. 1943, sine diagn. Latin.]
ex M. Johnson, Klematis 531. 1997, cum diagn. Latin.; Grey-Wilson, Clematis 31. 2000.
——sect. Integrifoliae Serov in Bot. Zhurn. 12: 1740. 1988; Grey-Wilson, l.c., excl. subsect.
Baldwinianae.——subgen. Integrifolia (Erickson) T. Y. A. Yang in Syst. Geogr. Pl. 68: 302.
1999, p.p.——sect. Integrifolia (Erickson) T. Y. A. Yang in l.c. Type: C. integrifolia L.
直立亚灌木。叶通常为单叶,全缘。萼片纸质,边缘不延展。花粉具3沟(C. integrifolia)
5期 王文采等: 铁线莲属一新分类系统 483
或散孔。瘦果不扁平 ,边缘无或稍有框状隆起 ;宿存花柱伸长 ,羽毛状 ,稀为尾状 (C.
fremontii)。
Erect subshrubs. Leaves usually simple, entire. Sepals papery, margin not dilated. Pollen
3-colpate (C. integrifolia) or pantoporate. Achenes not flattened, margin not or slightly
rimmed; persistent styles elongate, plumose, rarely tail-like (C. fremontii).
6种: 320. 全缘铁线莲 C. integrifolia L., 321. C. ochroleuca Aiton, 322. C. coactilis
(Fernald) Keener, 323. C. albicoma Wherry, 324. C. viticaulis Steele, 325. C. fremontii S.
Watson。分布于亚洲西部和中部,欧洲,美国中部及东部,墨西哥北部。
系3 Ser. 3. Hirsutissimae (Erickson ex M. Johnson) W. T. Wang, st. nov.——sect. Viorna
subsect. Hirsutissimae Erickson [in Ann. Missouri Bot. Gard. 30: 42. 1943, sine diagn. Latin.]
ex M. Johnson, Klematis 531. 1997, cum diagn. Latin.——sect. Hirsutissima (Erickson) T. Y.
A. Yang in Syst. Geogr. Pl. 68: 302. 1999.——sect. Hirsutissimae (Erickson ex M. Johnson)
Grey-Wilson, Clematis 31. 2000. Type: C. hirsutissima Pursh.
Subgen. Integrifolia (Erickson) T. Y. A. Yang in l.c., p.p.
直立多年生草本。叶为羽状复叶,裂片狭。花单朵顶生。萼片薄,边缘不延展。瘦果
小,稍扁压;宿存花柱伸长,羽毛状。
Erect perennial herbs. Leaves pinnate, with narrow segments. Flowers solitary, terminal.
Sepals thin, margin not dilated. Achenes small, slightly compressed; persistent styles elongate,
plumose.
1种: 326. C. hirsutissima Pursh。分布于美国西南部。
系4 Ser. 4. Euviornae (Erickson) W. T. Wang, st. nov. ——sect. Viorna subsect.
Euviornae Erickson in Ann. Missouri Bot. Gard. 30: 14. 1943, sine diagn. Latin.——sect.
Viorna subsect. Viorna M. Johnson, Klematis 528. 1997, cum diagn. Latin.; Grey-Wilson,
Clematis 200. 2000. Type: C. viorna L.
藤本植物,稀灌木状(C. addisonii)或茎直立(C. bigelovii)。叶为羽状或三出复叶。聚伞
花序具1或少数花,腋生,稀花单朵顶生。萼片厚,边缘通常不延展。花粉具散孔。瘦果扁平,
通常有框状边缘;宿存花柱伸长,羽毛状,稀尾状(C. pitcheri, C. bigelovii)。
Vines, rarely bushy (C. addisonii) or stems erect (C. bigelovii). Leaves pinnate or ternate.
Cymes 1–few-flowered, axillary, rarely flowers solitary, terminal. Sepals thick, margin usually
not dilated. Pollen pantoporate. Achenes flattened, usually rimmed; persistent styles elongate,
plumose, rarely tail-like (C. pitcheri, C. bigelovii).
9种: 327. C. viorna L., 328. C. morefieldii Kral, 329. C. addisonii Britton, 330. C.
glaucophylla Small, 331. C. versicolor Small ex Rydb., 332. C. texensis Buckley, 333. C.
reticulata Walter, 334. C. pitcheri Torr. & Gray, 335. C. bigelovii Torr.。分布于美国东部和
南部。
系5 Ser. 5. Viticellae (Erickson) W. T. Wang, st. nov.——sect. Viorna subsect. Viticellae
Erickson in Ann. Missouri Bot. Gard. 30: 31. 1943, sine diagn. Latin. Type: C. crispa L.
Sect. Viorna subsect. Crispae M. Johnson, Klematis 530. 1997, cum diagn. Latin.;
植 物 分 类 学 报 43卷 484
Grey-Wilson, Clematis 30. 2000. Type: C. crispa L.
藤本。叶为羽状复叶。花单朵顶生或组成腋生、具1花的聚伞花序。萼片薄,上部延展。
花粉具散孔。瘦果扁平,具边缘框;宿存花柱尾状,不呈羽毛状。
Vines. Leaves pinnate. Flowers solitary and terminal, or in axillary, 1-flowered cymes.
Sepals thin, margin dilated distally. Pollen pantoporate. Achenes flattened, rimmed; persistent
styles tail-like, not plumose.
1种: 336. C. crispa L.。特产美国东南部。
系6 Ser. 6. Baldwinianae (Erickson ex M. Johnson) W. T. Wang, st. nov. ——sect. Viorna
subsect. Baldwinianae Erickson [in Ann. Missouri Bot. Gard. 30: 41.1943, sine diagn. Latin.]
ex M. Johnson, Klematis 530. 1997, cum diagn. Latin.; Grey-Wilson, Clematis 31. 2000.
——sect. Baldwiniana (Erickson) T. Y. A. Yang in Syst. Geogr. Pl. 68: 302. 1999. Type: C.
baldwinii Torr. & Gray.
Subgen. Integrifolia (Erickson) T. Y. A. Yang in l.c., p.p.
直立多年生草本。叶为单叶,有时为羽状复叶。花1–3朵顶生。萼片狭长圆形,边缘上
部延展,瘦果扁平,具边缘框;宿存花柱羽毛状或尾状。
Erect perennial herbs. Leaves simple, sometimes pinnate. Flowers 1–3, terminal. Sepals
narrowly oblong, margin dilated distally. Achenes flattened, rimmed; persistent styles plumose
or tail-like.
2种: 337. C. baldwinii Torr. & Gray, 338. C. socialis Kral。分布于美国东南部。
组2 互叶铁线莲组
Sect. 2. Archiclematis Tamura in Sci. Rep. Osaka Univ. 4: 45. 1955; et in Acta Phytotax.
Geobot. 16: 79. 1956; M. Johnson, Klematis 139. 1997.——Archiclematis (Tamura) Tamura
in Sci. Rep. Osaka Univ. 16 (2): 31. 1967; et in Acta Phytotax. Geobot. 24: 151. 1970; et in
Hiepko, Nat. Pflanzenfam., Zwei. Aufl., 17a (4): 366. 1995; M. C. Chang in Fl. Reip. Pop. Sin.
28: 74. 1980.——Clematis subgen. Archiclematis (Tamura) Grey-Wilson, Clematis 29, 132.
2000. Type: C. alternata Kitam. & Tamura.
亚灌木状藤本植物。所有叶互生,为单叶,掌状浅裂。花组成具1–3朵花的腋生聚伞花
序,下垂。萼片紫红色。宿存花柱伸长,羽毛状。
Suffrutescent vines. Leaves all alternate, simple, palmatilobate. Flowers in axillary,
1–3-flowered cymes, pendulous. Sepals purple-red. Persistent styles elongate, plumose.
1种: 339. 互叶铁线莲 C. alternata Kitam. & Tamura。分布于中国西藏南部和尼泊尔。
亚属4 长瓣铁线莲亚属
Subgen. 4. Atragene Torr. & Gray, Fl. N. Amer. 1: 10. 1838; Keener & Dennis in Taxon
31: 39. 1982; Pringle in Morin et al., Fl. N. Amer. 3: 165. 1997; Grey-Wilson, Clematis 176.
2000. Type: C. alpina (L.) Mill.
木质藤本植物。叶为1–2回三出复叶或羽状复叶。花两性,通常1朵顶生于与1–2对叶
同自一老枝腋芽生出的短枝上,稀组成生于当年生枝叶腋的具少数花的聚伞花序。萼片4,
镊合状排列,斜上展,黄色、蓝色或紫色,卵形或长圆形,边缘上无一条短绒毛带。雄蕊被柔
5期 王文采等: 铁线莲属一新分类系统 485
毛,外部雄蕊的花丝和药隔变宽,常不育成花瓣状退化雄蕊。瘦果扁压,具伸长、羽毛状宿存
花柱。
Woody vines. Leaves 1–2-ternate or pinnate. Flowers bisexual, usually solitary and
terminal on new short shoots, arising together with 1–2 pairs of leaves from an axillary bud of
old branch, rarely in axillary few-flowered cymes arising from leaf axils of hornotinous branch.
Sepals 4, valvate, ascending, yellow, blue or purple, ovate or oblong, on margins without
narrow velutinous strips. Stamens pubescent, those outer ones with dilated filaments and
connectives often sterile and transformed into petal-like staminodes. Achenes compressed,
with elongate, plumose persistent styles.
约15种,隶属2组,广布于北温带。
组1 Sect. 1. Atragenopsis Boiss., Fl. Orient. Suppl. 2. 1888. Type: C. robertsiana Aitch. &
Hemsl.
叶为二回三出复叶。花单朵顶生。萼片淡黄色,长圆状披针形。雄蕊全部能育。
Leaves 2-ternate. Flowers solitary, terminal. Sepals yellowish, oblong-lanceolate.
Stamens all fertile.
1种: 340. C. robertsiana Aitch. & Hemsl.。产巴基斯坦北部,在阿富汗东部可能也产。
组2 长瓣铁线莲组
Sect. 2. Atragene (L.) DC., Syst. 1: 165. 1818; et Prodr. 1: 9. 1824; Rehd., Man. Cult.
Trees & Shrubs, ed. 2, 211. 1951; Tamura in Sci. Rep. Osaka Univ. 4: 49. 1955 et 16 (2): 34.
1967; Pringle in Baileya 19: 51. 1973; M. Y. Fang in Fl. Reip. Pop. Sin. 28: 138. 1980;
Tamura in Acta Phytotax. Geobot. 38: 39. 1987; et in Hiepko, Nat. Pflanzenfam., Zwei. Aufl.,
17a (4): 373. 1995; M. Johnson, Klematis 171. 1997. ——Atragene L., Sp. Pl. 1: 542. 1753,
p.p.; Spach, Hist. Nat. Veg. Phan. 7: 257. 1839; Maxim. in Bull. Acad. Sci. St.-Petersb. 22:
224. 1876; Small, Fl. Southeast. U.S. 439. 1903; Krasch. in Kom., Fl. URSS 7: 310. 1937.
——sect. Viorna subsect. Atragene (L.) Prantl in Bot. Jahrb. 9: 258. 1888, ut “Atragenae”, p.p.
excl. C. robertsiana Aitch. & Hemsl.; Schneid., Ill. Handb. Laubh. 1: 283. 1906; Hand.-Mazz.
in Acta Hort. Gotob. 13: 196. 1936. Lectotype: Atragene alpina L. (=Clematis alpina (L.)
Mill.) (Britton & Brown, 1913; Tamura, 1995).
Sect. Atragene subsect. Atragene M. Johnson, Klematis 171. 1997. Type: C. alpina (L.)
Mill.
Sect. Atragene subsect. Brachyblasti M. Johnson, l.c. 172. Type: Atragene moissenkoi
Serov.
叶为1–2回三出复叶或羽状复叶。花单朵顶生,稀组成1–3朵花的聚伞花序。萼片黄色、
蓝色或紫色。外部雄蕊不育成退化雄蕊,椭圆状条形,倒披针形或匙形,有时具残留的花药。
Leaves 1–2-ternate or pinnate. Flowers solitary, terminal, rarely in 1–3-flowered cymes.
Sepals yellow, blue or purple. Outer stamens sterile, transformed into staminodes,
elliptic-linear, oblanceolate or spathulate, sometimes with rudimentary anthers.
约14种: 341. C. occidentalis (Hornem.) DC., 342. C. columbiana (Nutt.) Torr. & Gray,
植 物 分 类 学 报 43卷 486
343. 朝鲜铁线莲 C. koreana Kom., 344. C. chiisanensis Nakai, 345. C. fauriei (Boiss.) M.
Johnson, 346. 伊犁铁线莲 C. iliensis Y. S. Hou & W. H. Hou, 347. C. kogenensis W. T.
Wang, 348. 绒萼铁线莲 C. moissenkoi (Serov) W. T. Wang, 349. 长瓣铁线莲 C.
macropetala Ledeb., 350. 长白铁线莲 C. nobilis Nakai, 351. C. crassisepala Ohwi, 352. 西
伯利亚铁线莲 C. sibirica (L.) Mill., 353. C. turkestanica M. Johnson, 354. C. alpina (L.)
Mill.。广布于北温带地区。
致谢 本文第一作者深切感谢BM, E, G, GH, K, KYO, LE, MO, P, S, TI, UPS, US诸研究所
所长和标本馆主任的邀请,到他们标本馆访问,或惠借标本。
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