全 文 ：植 物 分 类 学 报 43(3): 193–209(2005) doi:10.1360/aps040048
Acta Phytotaxonomica Sinica http://www.plantsystematics.com
———————————
Received: 27 April 2004 Accepted: 27 May 2004
Supported by the National Natural Science Foundation of China (Grant Nos. 30070059, 39570051) and the Special Project for
Taxonomic and Floristic Researches from the Chinese Academy of Sciences.
A revision of Clematis sect. Fruticella (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 In this paper Clematis sect. Fruticella is revised. Five species, two varieties, and
three forms are recognized. They are classified into two series, and keyed, described, and
illustrated. Brief taxonomic history and geographical distribution of the section are given, and
the relationships of sect. Fruticella with other sections of the Clematis and of the five species
within the section are discussed. Two new combinations, ser. Fruticosa (Tamura) W. T. Wang
& L. Q. Li, and C. fruticosa f. pinnatisecta (W. T. Wang & L. Q. Li) W. T. Wang & L. Q. Li,
are proposed, and a new series, ser. Virides W. T. Wang & L. Q. Li, is described.
Key words Clematis sect. Fruticella, Ranunculaceae, taxonomic revision.
1 Brief taxonomic history
In 1832, Turczaninow described the first species of the section Fruticella Tamura,
Clematis fruticosa Turcz., and two varieties under it, i.e. var. viridis Turcz. and var. canescens
Turcz., based on specimens collected by Kusnetsov from Nei Mongol, China. In 1877,
Maximowicz described the second species of the section, C. nannophylla Maxim., and
afterwards in 1889, he added two varieties (C. fruticosa var. lobata Maxim. (1889b) and var.
tomentella Maxim. (1889a)) to C. fruticosa and one variety (C. nannophylla var. foliosa
Maxim. (1889c)) to C. nannophylla, based on the specimens collected by Potanin and
Przewalski from the arid regions of North and West China respectively.
In the account of the tribe Clematideae DC. by Spach (1839), C. fruticosa was not
included.
In the monograph of the Clematis published by Kuntze (1885), C. fruticosa and C.
nannophylla were treated as two subspecies of C. recta L., which was placed in sect. 1.
Scandentes aperulatae.
In the classification of the Clematis proposed by Prantl (1888), C. fruticosa and C.
nannophylla together with C. songarica Bunge, C. asplenifolia Schrenk and other allies were
placed in sect. Flammula subsect. Rectae. This treatment was adopted by Rehder & Wilson
(1913) and Handel-Mazzetti (1939).
In 1933, Kozlov described three forms of C. fruticosa (C. fruticosa f. lancifolia Kozlov,
f. atriplexifolia Kozlov and f. chenopodiofolia Kozlov) on the basis of the specimens
collected by Licent and Serre from North and West China. In the present revision, C. fruticosa
f. atriplexifolia Kozlov is accepted to represent the form of C. fruticosa with 3-lobed to 3-sect
leaves.
In 1955, on the basis of C. fruticosa Tamura founded a new subsection, subsect.
Fruticosae under sect. Flammula. Twelve years later in 1967, he elevated this group to
sectional rank, naming it as sect. Fruticella, and giving the diagnosis “sepals are rather thick
and dilated marginate”. However, with the species of the C. fruticosa group, in which the
sepals are ascending, yellow, and narrowly dilated along margin after anthesis, Tamura
Acta Phytotaxonomica Sinica Vol. 43

194
associated C. delavayi Franch., C. songarica and their allies, in which the sepals are
spreading, white, and not dilated after anthesis. Tamura’s sect. Fruticella sensu lato was
adopted by Chang (1980), Snoeijer (1992), Johnson (1997), and Grey-Wilson (2000).
In the account of the Chinese Clematis by Chang (1980), C. fruticosa var. canescens
Turcz. was elevated to specific rank, and a new subspecies of it (C. canescens ssp. viridis W.
T. Wang & M. C. Chang) was described. Unfortunately, the new specific name C. canescens
(Turcz.) W. T. Wang & M. C. Chang was a misidentification, being misapplied to the plants
of C. fruticosa var. tomentella Maxim. In 2001, in the revised account of the Chinese
Clematis published by Wang & Bartholomew, the misidentification just mentioned was
corrected, and C. fruticosa var. tomentella and C. canescens ssp. viridis were all raised to
specific rank, and a new variety, C. nannophylla var. pinnatisecta W. T. Wang & L. Q. Li,
was described.
In the revision of Clematis sect. Clematis published by Wang (2003), C. songarica, C.
delavayi and their allies were transferred to the subsection Angustifoliae of that section from
sect. Fruticella sensu Tamura. After the removal of C. songarica, C. delavayi and their allies,
the section Fruticella defined in the present revision appears to become a monophyletic
group.
2 Relationships with other sections of the Clematis
In having ascending, yellow sepals and usually lanceolate-linear stamen filaments, the
floral structure of sect. Fruticella shows remarkable resemblance to that of sect. Meclatis, and
by the floral characters just mentioned these two sections may be distinguished from all other
sections of the genus Clematis. Sect. Fruticella differs from sect. Meclatis only in its dilated
sepals and its glabrous stamens. In sect. Meclatis, the sepals are not dilated, and the stamen
filaments are hairy. Compared with sect. Meclatis, therefore, sect. Fruticella is more primitive
in having glabrous stamens, and more advanced in having dilated sepals. However, in any
case the striking resemblance of their floral structure appears to indicate that the two sections
are closely related between each other, and might be together derived from a common
ancestry, i.e. sect. Clematis, in which the sepals are spreading, white, and not dilated after
anthesis, the stamens are glabrous, and the filaments are linear in outline (Wang, 2003).
3 Relationships of the five species in sect. Fruticella
Within sect. Fruticella, the five species are strikingly uniform in floral structure,
suggesting their close affinity. They can be classified into two groups according to the
inflorescence position and structure. In the C. fruticosa group consisting of four species, the
flowers are arranged in terminal, pedunculate, bibracteate, usually several-flowered cymes on
the hornotinous branch. In the fifth species, C. viridis (W. T. Wang & M. C. Chang) W. T.
Wang, however, the cymes shift from terminal to lateral position on the hornotinous branch,
their peduncles and bracts disappear, and the flower number is reduced from usually several
to only one. This phenomenon of the solitary flowers terminal to lateral branchlets has been
considered secondary by Wang (2002).
In the C. fruticosa group are observed three evolutionary trends of leaf morphology:
from undivided to pinnatisect, from green to grey-green, and from narrowly ovate to linear.
Of the four species, C. fruticosa with three forms and C. nannophylla with two varieties have
green, narrowly ovate or lanceolate, often subglabrous leaves, and are both strongly variable
in leaf division, and in leaf division there are a series of intermediate forms between C.
fruticosa f. fruticosa with undivided leaves and C. nannophylla with pinnatisect leaves. In
No. 3 WANG Wen-Tsai et al.: A revision of Clematis sect. Fruticella (Ranunculaceae)

195
another two species, C. canescens with undivided, narrowly rhombic-ovate or lanceolate
leaves and C. tomentella (Maxim.) W. T. Wang & L. Q. Li with undivided, linear leaves, the
leaves are grey-green in colour, and more or less densely hairy. Considering that the
pinnatisect, or grey-green, or linear leaves might all be products resulted from adaptation to
the deteriorating climate and strong aridity occurring in Northwest China by the Late Tertiary
(Liou, 1982; Zhou, 1982; Hsu, 1983; Axelrod et al., 1998), we would put forward the
following speculation: C. nannophylla, C. canescens and C. tomentella might all be
differentiated out from C. fruticosa f. fruticosa, the primitive taxon of the sect. Fruticella, and
come into existence later than the latter at that period.
4 Geographical distribution
Sect. Fruticella here defined consists of five species, which fall into two groups as
mentioned above, and are confined to the arid regions and the alpine areas of the Hengduan
Mountains of the eastern Asiatic continent (Fig. 1).

Fig. 1. Map showing distribution of the two series and five species in sect. Fruticella.

The first group, ser. Fruticosae, consists of four species, two varieties, and three forms,
widespread in the Loess Plateau and its adjacent arid regions of China and the Gobi desert of
Mongolia. In the northern Loess Plateau and its adjacent arid regions of Nei Mongol, the
Acta Phytotaxonomica Sinica Vol. 43

196
distribution center of sect. Fruticella, are concentrated the total four species of the series, and
from there C. canescens extends northward to southern Mongolia, C. tomentella westward to
northwestern Gansu and southern Mongolia, with one population disjunctively occurring in
the Mt. Baihuashan of Beijing, and C. nannophylla southwestward to eastern Qinghai, with its
variety foliosa occurring in southern Gansu. The primitive species of the series, C. fruticosa,
consists of three forms: the typical form fruticosa is widely distributed on the eastern Loess
Plateau, southwestern Nei Mongol and the Gobi desert of Mongolia, with a few populations
extending southwestward to southern Gansu; f. atriplexifolia is widespread on the Loess
Plateau and adjacent southwestern Nei Mongol, also with a few populations extending
southwestward to southern Gansu; and f. pinnatisecta is restricted in geographical distribution
to the central hinterland of the Loess Plateau.
The second group, ser. Virides, is monotypic, and its only species, C. viridis, occurs in
alpine areas of the northern Hengduan Mountains of Southwest China. In bearing green, linear
leaves and lateral, solitary flowers on hornotinous branch, this species is more advanced than
C. fruticosa, and might be derived from C. fruticosa f. fruticosa, and migrate to the northern
Hengduan Mountains from the Loess Plateau.
5 Taxonomic treatment
Clematis L. sect. Fruticella Tamura in Sci. Rep. Osaka Univ. 16: 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, Engler’s Nat. Pflanzenfam., ed. 2, 17a (4): 384. 1995, p.p. excl. C. delavayi Franch.
etc.; Snoeijer in Clematis 1992: 18. 1992, p.p.; M. Johnson, Klematis 651. 1997, p.p. excl. C.
delavayi Franch., C. lancifolia Bur. & Franch., C. ispahanica Boiss., C. phlebantha Williams,
C. asplenifolia Schrenk et C. songarica Bunge; Grey-Wilson, Clematis 126. 2000, p.p. Type:
C. fruticosa Turcz.
Small shrubs. Seedling leaves unknown (Essig, 1991). Leaves opposite, simple,
undivided or pinnatilobed to pinnatisect. Flowers bisexual, medium-sized or small, in terminal
cymes, rarely solitarily arising from the apexes of axillary, short or abbreviated branchlets.
Sepals 4, ascending, yellow, valvate, on margin velutinous, after anthesis along margin dilated
into narrow membranous wings. Stamens numerous, glabrous; filaments lanceolate-linear;
anthers narrowly oblong, apex obtuse. Achenes numerous, compressed, bearing elongate
plumose persistent styles.
Five species, occurring on the Loess Plateau and adjacent regions and in the Hengduan
Mountains of China, and in the Gobi desert of Mongolia.
Key to infrasectional taxa
1. Flowers arranged in terminal pedunculate, bibracteate 1-10(-25)-flowered cymes on hornotinous branch
(Ser. 1. Fruticosae).
2. Leaves undivided, sometimes some ones with 1 or 2 lobes near base.
3. Leaves green, on both surfaces sparsely puberulous or subglabrous……....1a. C. fruticosa f. fruticosa
3. Leaves grey-green, on both surfaces more or less densely appressed-puberulous.
4. Leaves narrowly rhombic-ovate to lanceolate, margin usually 1-3-dentate per side, sometimes
entire ..……………………………………………………………… ……..………. 3. C. canescens
4. Leaves linear or lanceolate-linear, margin usually entire…………………...……….4. C. tomentella
2. Leaves near base lobed to sect, or pinnatisect.
5. Leaves near base 3-lobed or 3-parted, sometimes 3-sect……..……….1b. C. fruticosa f. atriplexifolia
5. Leaves pinnatisect.
6. Leaf blade oblong-lanceolate in outline, primary lateral lobes subrhombic, 3-lobulate, with ovate or
No. 3 WANG Wen-Tsai et al.: A revision of Clematis sect. Fruticella (Ranunculaceae)

197
triangular lobules……………………………………………...…….1c. C. fruticosa f. pinnatisecta
6. Leaf blade narrowly ovate or ovate in outline, primary lateral lobes narrowly oblong, linear, or
triangular, undivided or unequally lobed…………………………………………2. C. nannophylla
7. Leaf blade 0.5-1.4(-2)×0.3-0.9(-1.4) cm, primary lobes triangular, narrowly oblong, or
linear….…………………………………..………………………………..2a. var. nannophylla
7. Leaf blade larger, 2.5-4.5×1.6-3 cm, primary lobes linear……..……………2b. var. foliosa
1. Flowers solitary, arising from apexes of short or abbreviated axillary branchlets of hornotinous branch;
leaves green, linear, entire, subglabrous (Ser. 2. Virides)…………………………………..…..5. C. viridis
Ser. 1. Fruticosae (Tamura) W. T. Wang & L. Q. Li, st. et comb. nov. ——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 Franch. et C. lancifolia Bur. & Franch; Grey-Wilson, Clematis 126, p.p. excl. C.
delavayi Franch. Type: C. fruticosa Turcz.
Flowers arranged in terminal pedunculate, bibracteate 1-10(-25)-flowered cymes on
hornotinous branch.
Four species occurring on the Loess Plateau and adjacent regions of China and the Gobi
desert of Mongolia.
1. Clematis fruticosa Turcz. in Bull. Soc. Nat. Mosc. 5: 180. 1832; Franch., Pl. David. 1: 12.
1884; Maxim., Enum. Pl. Mong. 2. 1889; Finet & Gagnep. in Bull. Soc. Bot. France 50: 537.
1903; Hand.-Mazz. in Acta Hort. Gotob. 13: 202. 1939, p.p.; Anonymous in Iconogr. Corm.
Sin. 1: 743, fig. 1485. 1972; M. C. Chang in Fl. Reip. Pop. Sin. 28: 148, pl. 43, figs. 1-6.
1980, p.p.; Grubov, Key Vasc. Pl. Mong. 112. 1982; He, Fl. Beijing, rev. ed., 1: 247. 1984;
Liou f., Fl. Desert. Sin. 1: 481. 1985; J. W. Wang in Fl. Hebei. 1: 470, fig. 474. 1986, p.p.; Y.
Z. Zhao in Fl. Intramong., ed. 2, 2: 517. 1990, p.p.; Y. Z. Ling et al. in Fl. Shanxi. 1: 633.
1992, p.p; M. Johnson, Klematis 656. 1997, p.p.; J. Q. Fu in Fl. Loe. Plat. Sin. 1: 444. 2000,
p.p.; Grey-Wilson, Clematis 127. 2000; Grabovsk. in Grubov, Pl. As. Centr. 12: 78, tab. 6,
fig. 2. 2001, p.p. excl. syn. C. fruticosa var. canescens Turcz. et C. canescens ssp. viridis W.
T. Wang & M. C. Chang; W. T. Wang & Barth. in Fl. China 6: 362. 2001. ——C. fruticosa
var. viridis Turcz. in l.c.; Grabovsk. in Grubov, Cat. Type Specim. Centr. As. Vasc. Pl. 194.
2000. ——C. recta L. ssp. fruticosa (Turcz.) Kuntze in Verh. Bot. Ver. Brand. 26: 112. 1885.
Type: “Mongolia Chinensis: in collibus subarenosis prope Mogoitu”, 1831, Kusnetzov s.n.
(holotype, KW; isotype, LE!).
灌木铁线莲
This species consists of three forms widespread on the Loess Plateau and adjacent
regions of China with the typical form extending to the Gobi desert of Mongolia.
Because C. fruticosa is highly variable in leaf shape and division, and between it and C.
nannophylla Maxim. there are intermediate forms as mentioned above, further study is needed
for elucidation of their relationships.
1a. f. fruticosa Fig. 2: A, B
Small shrub. Stem up to 1 m tall; branches shallowly 4-5-sulcate, sparsely puberulous,
glabrescent. Leaves simple; leaf blades green, thinly coriaceous or coriaceous, lanceolate,
broadly lanceolate, or narrowly triangular, 1.5-4×0.5-1.6 cm, apex acute, base cuneate or
broadly cuneate, margin dentate, denticulate, or entire, on both surfaces sparsely
appressed-puberulous or subglabrous, midrib abaxially slightly prominent; petioles 0.3-1.2
cm long. Cymes terminal and axillary, 1-5-flowered; peduncle 0.7-2 cm long; bracts
foliaceous. Flower 1.8-3.5 cm in diam.; pedicel 0.4-1.3 cm long, puberulous. Sepals 4,
Acta Phytotaxonomica Sinica Vol. 43

198
yellow, elliptic-ovate, 1-1.8(-2)× 0.6-1.2 cm, apex acute, inside glabrous, outside
subglabrous or sparsely puberulous, margin velutinous, marginal wings 1-2 mm broad.
Stamens 8-13 mm long, glabrous; anthers narrowly oblong, 2.5-4 mm long, apex obtuse.
Ovaries pubescent; styles 7-11 mm long, densely villous. Achenes ovate or elliptic, 4-6×
2.2-3.5 mm, villous; persistent styles ca. 2.5 cm long, yellowish-plumose. Fl. Jul.-Aug.
China (SW Gansu, NW Hebei, SW Nei Mongol, Ningxia, N Shaanxi, Shanxi) and
Mongolia. In bushes or on dry slopes; alt. 800-1900 m.
Additional specimens examined:
China. Gansu (甘肃): Têwo (迭部), J. F. Rock 15007 (K). Hebei (河北): Xiaowutai
Shan (小五台山), T. P. Wang (王作宾) 797, Y. Liu (刘瑛) 11119, X. L. Huang (黄秀兰)
5677 (PE); Yu Xian (蔚县), Licent 2253 (PE); Zhuolu (涿鹿), C. G. Yang (杨朝广) 1711
(PE). Nei Mongol (内蒙古): Bailingmiao (百灵庙), Roerich Exped. 696 (US); Baotou (包
头), C. W. Yao (姚仲吾) 925 (PE); Chaggan, J. Eriksson s.n. (S); Liangcheng (凉城), Loess
Plat. Exped. (黄土高原队) 84-4052 (WUK); Zhuozi Shan (卓资山), Y. W. Tsui (崔友文)
1109 (PE). Ningxia (宁夏): Tongxin (同心), Y. P. Hsu (徐养鹏) 1859 (WUK). Shaanxi (陕
西): Jingbian (靖边), K. T. Fu (傅坤俊) 7461 (WUK); Suide (绥德), K. T. Fu (傅坤俊) 6606
(WUK); Zichang (子长), K. T. Fu (傅坤俊) 7643 (WUK). Shanxi (山西): Fanshi (繁峙),
Loess Plat. Exped. (黄土高原队) 84-3602 (WUK); Lishi (离石), Huanghe Exped. (黄河队)
55-1610 (WUK); Ningwu (宁武), Loess Plat. Exped. (黄土高原队) 84-3466 (WUK); Pinglu
(平鲁), Shaanxi Exped. (陕西队) 53-542 (PE); Wuzhai (五寨), W. Y. Hsia (夏纬瑛) 1663
(PE, WUK); Yonghe (永和), R. Z. Zhang (张荣祖) 360 (WUK); Youyu (右玉), Loess Plat.
Exped. (黄土高原队) 84-3826 (WUK).
Mongolia. Bulgan Somon, Norlindh 10385 (S); Gobi, Potanin s.n. (LE, P), Lisovsky
305, Pahomov s.n., Tugarinov 3 (LE); Godu-Altai, Bolkova & Parkovskaya 7016 (LE);
Hedgo, Grubov 6807 (LE); Ikhe Bogdo, R. W. Chaney 516 (LE); South Todusk, Grubov
5944, 5945 (LE); Tuinigora, Klemenz s.n. (PE).
1b. f. atriplexifolia Kozlov in Publ. Mus. Hoangho Paiho Tien Tsin 22: 11, pl. 2. 1933. Type: China.
Shanxi (山西): Kaochan, 1925-08-22, Licent 7618 (syntype, TIE; isosyntype, K!). Beijing (北京):
Yanqing (延庆), Tongtchoang Pou, 1930-08-24, Serre 9967 (syntype, TIE; isosyntype, PE!).
C. fruticosa Turcz. var. lobata Maxim., Enum. Pl. Mong. 3. 1889; W. T. Wang & Barth.
in Fl. China 6: 363. 2001. Type: China. Nei Mongol (内蒙古): Ordos (鄂尔多斯), vallis
Hoangho, 1871-08-18, Przewalski s.n. (lectotype, here designated, LE!; isolectotype, K!);
Ordos, Gunjatagai, 1884-09-06, Potanin s.n. (syntype, LE!; isosyntype, K!).
C. fruticosa f. chenopodiofolia Kozlov in l.c., pl. 3. Type: China. Hebei (河北): Pankiao,
1917-08-07, Licent 3265 (syntype, TIE; isosyntypes, K!, P!, UPS!). Nei Mongol (内蒙古):
Baotou (包头 ), Plaine de Toumet, Palakai, 1919-07-30, Licent 5557 (syntype, TIE;
isosyntypes, K!, P!, PE!).
C. fruticosa auct. non Turcz.: Rehd. in J. Arn. Arb. 9: 41. 1928; Hand.-Mazz. in l.c.
1939, p.p; M. C. Chang in l.c. 1980, p.p.; J. W. Wang in l.c. 1986, p.p.; Y. Z. Zhao in l.c.
1990, p.p.; Y. J. Ling et al. in l.c. 1992, p.p.; J. Q. Fu in l.c. 2000, p.p.
裂叶灌木铁线莲 Fig. 2: C, D
This form differs from f. fruticosa in its leaves 3-lobed, 3-parted, or 3-sect near base.
Leaf blade broadly lanceolate or lanceolate, 2-5.5×0.6-2.2 cm, margin coarsely dentate
or lobulate. Fl. Jul.-Sept.
No. 3 WANG Wen-Tsai et al.: A revision of Clematis sect. Fruticella (Ranunculaceae)

199


Fig. 2. A, B, Clematis fruticosa Turcz. f. fruticosa. A, flowering branch; B, stamen (from Shanxi Exped. 53-542). C, D,
C. fruticosa f. atriplexifolia Kozlov. C, flowering branch; D, stamen (from Kuan & Chen 2303).





Acta Phytotaxonomica Sinica Vol. 43

200


Fig. 3. A, Clematis fruticosa Turcz. f. pinnatisecta (W. T. Wang & L. Q. Li) W. T. Wang & L. Q. Li. flowering branch
(from K. T. Fu 3122). B, C, C. nannophylla Maxim. var. foliosa Maxim. B, flowering branch; C, stamen (from
Potanin s.n.). D, E, C. nannophylla Maxim. var. nannophylla. D, flowering branch; E, stamen (from Y. Q. He 4928).




No. 3 WANG Wen-Tsai et al.: A revision of Clematis sect. Fruticella (Ranunculaceae)

201
China (Beijing, S & C Gansu, NW Hebei, SW Nei Mongol, Ningxia, N Shaanxi,
Shanxi). On dry slopes or cliffs; alt. 700-2300 m.
Additional specimens examined:
China. Beijing (北京): Baihua Shan (百花山), D. F. Jin 504 (PE). Gansu (甘肃):
Dangchang (宕昌), Q. E. Yang (杨亲二) 92007 (PE); Têwo (迭部), J. F. Rock 14566 (G, GH,
P, S); Heshui (合水), T. P. Wang (王作宾) 17381 (PE), H. Y. Zou (邹厚远) 3251 (WUK);
Min Xian (岷县), K. S. Hao (郝景盛) 555 (PE, WUK); Wudu (武都), T. P. Wang (王作宾)
14932 (PE), J. X. Yang (杨金祥) 3358 (WUK); Yan-pu-ko, F. N. Meyer 1786 (LE, P);
Zhenyuan (镇原), S. Q. Zhong (仲世奇) 190 (PE). Hebei (河北): Xiaowutai Shan (小五台
山), C. F. Li (李建藩) 10662, Y. Liu (刘瑛) 11230 (PE); Zhuolu (涿鹿), C. G. Yang (杨朝广)
441, 1473 (PE). Nei Mongol (内蒙古): Baotou (包头), T. P. Wang (王作宾) 2392 (PE);
Daqing Shan (大青山), W. Y. Hsia (夏纬瑛) 2731 (PE); Sartchy, David 2701 (LE, P); Ordos
(鄂尔多斯), Licent 13701 (GH, PE, TIE); Ul Shan (乌拉山), X. Z. Lang (郎学忠) 159 (PE).
Ningxia (宁夏): Guyuan (固原), Gansu Exped. (甘肃队) 56-2294 (PE); Yanchi (盐池),
Yellow River Exped. (黄河队) 56-7859 (PE); Pingluo (平罗), Z. Y. Yu (于兆英) 754
(WUK); Helan Shan (贺兰山), Y. P. Xu (徐养鹏) 2490 (WUK). Shaanxi (陕西): Ansai (安
塞), J. X. Yang (杨金祥) 4771 (WUK); Hengshan (横山), K. T. Fu (傅坤俊) 7159, Yellow
River Exped. (黄河队) 56-7432 (PE); Jingbian (靖边), K. T. Fu (傅坤俊) 7461 (PE); Mizhi
(米脂), K. T. Fu (傅坤俊) 6877 (PE, WUK); Suide (绥德), K. T. Fu (傅坤俊) 6606 (PE);
Wuqi (吴旗), Yellow River Exped. (黄河队) 56-8202 (PE); Yan’an (延安), Yellow River
Exped. (黄河队) 54-1055 (PE); Zhidan (志丹), J. X. Yang (杨金祥) 4903 (WUK); Zichang
(子长), K. T. Fu (傅坤俊) 7643 (PE). Shanxi (山西): Guandishan (关帝山), S. Ma (马塞)
15516 (WUK); Hunyuan (浑源), Loess Plat. Exped. (黄土高原队) 1834 (WUK); Jiexiu (介
休), Loess Plat. Exped. (黄土高原队) 84-2785 (WUK); Kushehshan, T. P. Wang (王作宾)
3706 (K, PE, WUK); Lan Xian (岚县), Loess Plat. Exped. (黄土高原队) 84-3145 (WUK);
Lin Xian (临县), Yellow River Exped. (黄河队) 54-1394 (WUK); Lishi (离石), P. Y. Li (李
培元) 10159 (WUK); Ningwu (宁武), T. Tang (唐进) 1234 (GH, PE); Qinyuan (沁源), K. M.
Liou (刘继孟) 1443 (PE); Without precise locality, 1884-07-05, Potanin s.n. (LE, P); Wutai
Shan (五台山), K. M. Liou (刘继孟) 3696, K. C. Kuan & Y. L. Chen (关克俭, 陈艺林) 2303
(PE); Xi Xian (隰县), T. P. Wang (王作宾) 3467 (PE, WUK); Xiaoyi (孝义), T. P. Wang (王
作宾) 2794 (PE, WUK); Zhongyang (中阳), P. Y. Li (李培元) 10300 (WUK).
1c. f. pinnatisecta (W. T. Wang & L. Q. Li) W. T. Wang & L. Q. Li, st. et comb. nov. ——
C. nannophylla Maxim. var. pinnatisecta W. T. Wang & L. Q. Li in Fl. China 6: 363. 2001.
Type: China. Shaanxi (陕西): Huanglong (黄龙), Huanglong Shan (黄龙山), 1939-08-10, K.
T. Fu (傅坤俊) 3122 (holotype, PE!; isotype, WUK!).
全裂灌木铁线莲 Fig. 3: A
This form differs from f. fruticosa in its pinnatisect leaves and larger oblong-lanceolate
leaf blades up to 5.5 cm long, 3 cm broad. Fl. Jul. –Sept.
China (N Shaanxi and W Shanxi). On dry slopes; alt. 1050-1500 m.
Additional specimens examined:
China. Shaanxi (陕西): Ansai (安塞), J. X. Yang (杨金祥) 4762 (WUK); Jingbian (靖
边), Yellow River Exped. (黄河队) 56-7730 (PE); Suide (绥德), Petrov s.n. (LE); Zhidan (志
丹), J. X. Yang (杨金祥) 4903, 4924 (WUK). Shanxi (山西): Baode (保德), Yellow River
Acta Phytotaxonomica Sinica Vol. 43

202
Exped. (黄河队) 55-3054 (WUK); Jiexiu (介休), Loess Plat. Exped. (黄土高原队) 84-7785
(WUK); Lishi (离石), Yellow River Exped. (黄河队) 55-1740 (WUK); Zhongyang (中阳), P.
Y. Li (李培元) 10381 (WUK).
2. Clematis nannophylla Maxim. in Bull. Acad. Sci. St.-Petersb. 23: 305. 1877; et in Mél.
Mél. Biol. 9: 707. 1877; et Fl. Tangut. 2, tab. 1, figs. 1-9. 1889; Forbes & Hemsl. in J. Linn.
Soc. Bot. 23: 6. 1886; Finet & Gagnep. in Bull. Soc. Bot. France 50: 537. 1903; Rehd. in J.
Arn. Arb. 9: 41. 1928; Kozlov in Publ. Mus. Hoangho Paiho Tien Tsin 22: 13. 1933;
Hand.-Mazz. in Acta Hort. Gotob. 13: 203. 1939; W. W. Smith in Curtis, Bot. Mag. 164: tab.
9641. 1940-1942; Anonymous in Iconogr. Corm. Sin. 1: 744, fig. 7487. 1972; M. C. Chang in
Fl. Reip. Pop. Sin. 28: 147, fig. 18. 1980; Liou f., Fl. Desert. Sin. 1: 481. 1985; Y. Z. Zhao in
Fl. Intramong., ed. 2, 2: 519, pl. 209, figs. 6-8. 1990; Y. P. Hsu in Fl. Sin. Area Tan-yang 2:
160. 1993; L. H. Zhou in Fl. Qinghai. 1: 348. 1997; M. Johnson, Klematis 659. 1997;
Grey-Wilson, Clematis 128. 2000; J. Q. Fu in Fl. Loe. Plat. Sin. 1: 443. 2000; Grabovsk. in
Grubov, Cat. Type Specim. Centr. As. Vasc. Pl. 194. 2000; et Pl. As. Centr. 12: 82, tab. 6, fig.
4. 2001; W. T. Wang & Barth. in Fl. China 6: 363. 2001. ——C. recta L. ssp. nannophylla
(Maxim.) Kuntze in Verh. Bot. Ver. Brand. 26: 112. 1885. Type: China. Qinghai (青海):
Datong (大通), 1872-06-25, Przewalski 161 (lectotype, designated by Grabovskaja, 2000,
LE!); Gansu (甘肃), without precise locality, 1875, Piasezki s.n. (syntype, not seen).
C. nannophylla Maxim. var. bifoliata Y. P. Hsu in Fl. Sin. Area Tan-yang 2: 160. 1993.
Type: China. Ningxia (宁夏): Zhongwei (中卫), alt. 2000 m, 1981-08-01, Z. Y. Yu et al. (于
兆英等) 1434 (holotype, WUK!).
小叶铁线莲
This species consists of two varieties occurring on western Loess Plateau and adjacent
regions of China.
2a. var. nannophylla Fig. 3: D, E
Small shrub. Stem 30-100 cm tall; branches shallowly 4-5-sulcate, densely
appressed-puberulous. Leaves simple; leaf blades green, coriaceous, ovate or narrowly ovate,
0.5-1.4(-2)×0.3-0.9(-1.4) cm, apex acute, base cuneate or subtruncate, pinnatilobed to
pinnatisect, with 1-3(-4) pairs of primary lateral lobes; lobes triangular, narrowly oblong, or
linear, undivided or unequally 2-lobed, on both surfaces sparsely puberulous, glabrescent,
midrib obscure; petioles 1.5-4 mm long. Cymes terminal, 1-3(-7)-flowered; peduncles 1-2
cm long; bracts foliaceous. Flower 1.2-2.2 cm in diam.; pedicel 0.5-1.3 cm long, densely
puberulous. Sepals 4, yellow, oblong or long elliptic, 0.8-1.6×0.4-0.8 cm, apex obtuse or
subtruncate, inside glabrous, outside puberulous or glabrous, margin velutinous, marginal
wings 0.8-1.8 mm broad. Stamens 6-8 mm long, glabrous; anthers narrowly oblong, 2.2-3
mm long, apex obtuse. Ovaries pubescent; styles 5-7 mm long, densely villous. Achenes
elliptic, 3-4×2.2-2.5 mm, densely pubescent; persistent styles ca. 2 cm long, yellowish-
plumose. Fl. Jul.–Aug.
China (C Gansu, SW Nei Mongol, Ningxia, and E Qinghai). On dry slopes; alt.
1200-3200 m.
Additional specimens examined:
China. Gansu (甘肃): Baiyin (白银), Y. Q. He 6207 (PE); Jingtai (景泰), Z. Y. Yu (于
兆英) 3309 (WUK); Jingyuan (靖远), Yellow River Exped. (黄河队) 56-5495 (PE); Lanzhou
(兰州), J. F. Rock 13231 (S), Licent 4488 (PE, TIE), Y. S. Lian (廉永善) 96912 (PE);
No. 3 WANG Wen-Tsai et al.: A revision of Clematis sect. Fruticella (Ranunculaceae)

203
Lichen, 1923-07-06, R. C. Ching (秦仁昌) 241 (US); Liancheng (莲城), Qinggan Exped. (青
甘队) 60-3369 (PE); Lintao (临洮), Gansu Exped. (甘肃队) 56-1635 (PE); Lotani, on road to
Liku, Purdom 1027 (K, US); Xiahe (夏河), T. P. Wang (王作宾) 5708 (PE); Yongdeng (永
登), Loess Plat. Exped. (黄土高原队) 85-4858 (WUK). Nei Mongol (内蒙古): Alxa Zuoqi
(阿拉善左旗), Y. M. Yang (杨永明) 496 (HIMC). Ningxia (宁夏): Tongxin (同心), Loess
Plat. Exped. (黄土高原队) 85-4156 (WUK). Qinghai (青海): Datong (大通), 1880-08-06,
Przewalski s.n. (LE); fl. Hoangho, 1880-07, Przewalski 433 (LE); Guide (贵德), S. W. Liu
(刘尚武) 3121 (PE); Ledu (乐都), K. M. Liou (刘继孟) 5866, K. S. Hao (郝景盛) 745 (PE);
Minhe (民和), S. W. Liu (刘尚武) 2697 (PE); Xunhua (循化), Loess Plat. Exped. (黄土高原
队) 85-5873 (WUK).
2b. var. foliosa Maxim. in Acta Hort. Petrop. 11: 5. 1889; Hand.-Mazz. in Acta Hort. Gotob.
13: 203. 1939; Anonymous in Fl. Tsinling. 1 (2): 291, fig. 250. 1974; M. C. Chang in Fl.
Reip. Pop. Sin. 28: 148. 1980; M. Johnson, Klematis 659. 1997; Grey-Wilson, Clematis 128.
2000; W. T. Wang & Barth. in Fl. China 6: 363. 2001. Type: China. Gansu (甘肃): “valle fl.
Lipagu” (near Wudu 武都), 1885-09-18, Potanin s.n. (holotype, LE!; isotypes, K!, PE!).
多叶铁线莲 Fig. 3: B, C
This variety differs from var. foliosa in its larger leaves 2.5-4.5 cm long, 1.6-3 cm
broad, and in their lobes and lobules all linear. Fl. Jun.-Oct.
China (S Gansu). On dry or gravelly slopes; alt. 1200-1700 m.
Additional specimens examined. Gansu (甘肃): Min Xian (岷县), Hummel 3777, 5376
(S); Chieh Chou, F. N. Meyer 1786, 1794 (K).
3. Clematis canescens (Turcz.) W. T. Wang & M. C. Chang in Fl. Reip. Pop. Sin. 28: 150.
1980, p.p. quoad nomen tantum, excl. descr. et fig. ——C. fruticosa var. canescens Turcz. in
Bull. Soc. Nat. Mosc. 5: 180. 1832; Grabovsk. in Grubov, Cat. Type Specim. Centr. As. Vasc.
Pl. 193. 2000; W. T. Wang & Barth. in Fl. China 6: 363. 2001. ——C. recta L. ssp. fruticosa
var. canescens (Turcz.) Kuntze in Verh. Bot. Ver. Brand. 26: 112. 1885. Type: “Mongolia
chinensis: in salsis ad lacum Kobur”, 1831, Kusnetzov s.n. (holotype, KW; isotype, LE!).
C. salsuginea Bunge ex W. T. Wang in Acta Phytotax. Sin. 38: 411, fig. 2: 1-3. 2000;
Finet & Gagnep. in Bull. Soc. Bot. France 50: 537, pro syn. Type: China. Nei Mongol (内蒙
古): Without precise locality, Bunge s.n. (holotype, K!; isotypes, LE!, P!); Pan chiang, Y. G.
Anderson 772 (paratype, UPS!).
灰灌木铁线莲 Fig. 4
Small shrub. Stem shallowly 6-sulcate, glabrescent, near apex on nodes densely
appressed-puberulous; hornotinous branches velutinous. Leaves simple, shortly petiolate; leaf
blades grey-green, coriaceous, narrowly rhombic-ovate or lanceolate, 1.9-3(-5)×0.25-
0.8(-1.6) cm, apex pungent, base cuneate, margin above base 1-3-dentate per side, sometimes
entire, undivided, seldom 3-lobed, on both surfaces more or less densely appressed-
puberulous, trinerved, the 2 lateral veins obscure; petioles 1.5-3 mm long. Cymes terminal,
3-5-flowered; peduncles 1-1.2 cm long, velutinous; bracts triangular, ca. 2 mm long,
velutinous. Flower ca. 2 cm in diam.; pedicel 3-4 mm long, velutinous. Sepals 4, yellow,
lanceolate or long elliptic, 1.2-1.9×0.4-0.7 cm, apex acute or obtuse, inside glabrous,
outside puberulous, margin velutinous, marginal wings 1-1.8 mm broad. Stamens 8-12 mm
long, glabrous; anthers linear, 2.8-3.2 mm long, apex obtuse. Ovaries densely pubescent;
styles 8-12 mm long, densely villous. Fl. Jul.-Aug.
Acta Phytotaxonomica Sinica Vol. 43

204

Fig. 4. Clematis canescens (Turcz.) W. T. Wang & M. C. Chang. A, flowering branch; B, sepal, outside; C, stamen
(from Bunge s.n.).
No. 3 WANG Wen-Tsai et al.: A revision of Clematis sect. Fruticella (Ranunculaceae)

205
China (SW Nei Mongol) and S Mongolia. By sandy stream banks.
Additional specimens examined:
China. Nei Mongol (内蒙古): Siziwang Qi (四子王旗), Naomugen (脑木根), Y. Z.
Zhao (赵一之) 38 (PE); the same locality, Weijingsumu (卫境苏木), Y. Z. Zhao (赵一之)
1263 (PE).
Mongolia. Sain-shand, Isachenko & Rochhovskaja 1817 (LE).
4. Clematis tomentella (Maxim.) W. T. Wang & L. Q. Li in Fl. China 6: 364. 2001. ——C.
fruticosa Turcz. var. tomentella Maxim., Fl. Tangut. 2. 1889. Type: China. Gansu: “valle fl.
Bajan-gol”, near Liyuan (梨园), west of Zhangye (张掖), 1886-06-01, Potanin s.n. (holotype,
LE!).
C. fruticosa f. lancifolia Kozlov in Publ. Mus. Hoangho Paiho Tien Tsin 22: 10, pl. 1.
1933. Type: China. Nei Mongol (内蒙古): S Ordos (鄂尔多斯), Sjara Osson gol, 1922-08-10,
Licent 6869 (syntype, TIE; isosyntypes, K!, PE!); the same locality, 1923-08-11, Licent 7069
(syntype, TIE!; isosyntypes, G!, K!); the same locality, 1923-08-20, Licent 7087 (TIE!;
isosyntypes, K!, PE!).
C. fruticosa var. canescens auct. non Turcz.: Rehd. in J. Arn. Arb. 9: 41. 1928; Hand.
-Mazz. in Acta Hort. Gotob. 13: 203. 1939.
C. canescens sensu W. T. Wang & M. C. Chang in Fl. Reip. Pop. Sin. 28: 150, pl. 43,
fig. 7. 1980, non C. fruticosa Turcz. var. canescens Turcz.; Anonymous in Iconogr. Corm.
Sin. 1: 743. 1972; Y. Z. Zhao in Fl. Intramong., ed. 2, 2: 517, pl. 209, figs. 4, 5. 1990; M.
Johnson, Klematis 654. 1997; Grey-Wilson, Clematis 126. 2000; J. Q. Fu in Fl. Loe. Plat. 1:
444. 2000.
灰叶铁线莲 Fig. 5: D-G
Small shrub. Stem 60-100 cm tall; branches shallowly 4-6-sulcate, densely puberulous.
Leaves simple, shortly petiolate; leaf blades grey-green, coriaceous, linear, linear-lanceolate,
or broadly linear, 1-4.8×0.2-0.8 cm, apex acute, base cuneate, margin entire, sometimes
below 1-2-dentate, on both surfaces more or less densely appressed-puberulous, midrib
obscure; petioles 2-5 mm long or nearly absent. Cymes terminal, (1-)2-10(-25)-flowered,
sometimes narrowly panicle-like; peduncles 0.7-3 cm long; bracts linear or lanceolate-linear,
0.4-1.5 cm long. Flower 1.3-2.2 cm in diam.; pedicel 0.5-1.8 cm long, densely puberulous.
Sepals 4, yellow, broadly lanceolate, lanceolate, long elliptic, or oblong, 0.8-1.6×0.4-0.9
cm, apex shortly cuspidate, inside glabrous or sparsely puberulous, outside puberulous,
margin velutinous, marginal wings 0.6-2 mm broad. Stamens 7-10 mm long, glabrous;
anthers narrowly oblong, 2-3 mm long, apex obtuse. Ovaries densely pubescent; styles 6-11
mm long, densely villous. Achenes long elliptic or narrowly ovate, 3-5×1.4-2.5 mm, villous;
persistent styles ca. 2 cm long, white-plumose. Fl. Jul.–Aug.
China (Beijing, N Gansu, SW Nei Mongol, and Ningxia) and S Mongolia. On dry sandy
slopes; alt. 1700-2150 m.
Additional specimens examined:
China. Beijing (北京): Baihua Shan (百花山), 1850-1858, Bazilevski s.n. (LE). Gansu
(甘肃): Anxi (安西), Anonymous 312 (PE); Hexi Zoulang (河西走廊), Mazong Shan (马宗
山), Hexi Exped. (河西队) 1394, 1413, 1451 (PE); Richthofen range, Rock 13322 (K);
Shandan (山丹), Hexi Exped. (河西队) 64-877, 64-991 (PE); Sunan (肃南), Hexi Exped. (河
西队) 67-528, 67-547 (PE); Yongchang (永昌), Anonymous 131 (PE); Yumen (玉门), T. N.
Liou (刘慎谔) 2359 (PE, WUK), Anonymous 392 (PE); Zhangye (张掖), P. C. Tsoong (钟补
Acta Phytotaxonomica Sinica Vol. 43

206

Fig. 5. A-C, Clematis viridis (W. T. Wang & M. C. Chang) W. T. Wang. A, flowering branch; B, stamen (from Sichuan
Exped. 74-6565); C, achene (from Qinghai-Xizang Exped. 76-9842). D-G, C. tomentella (Maxim.) W. T. Wang & L. Q. Li.
D, flowering branch; E, stamen (from Licent 7087); F, aggregate fruit; G, achene (from Hexi Exped. 1451).
No. 3 WANG Wen-Tsai et al.: A revision of Clematis sect. Fruticella (Ranunculaceae)

207
求) 8761 (PE, WUK). Nei Mongol (内蒙古): Mt. Alaschen, 1880, Przewalski s.n. (LE); Alxa
Qi (阿拉善旗), Beiyiamu (北依阿母), Anonymous 109 (WUK); Alxa Youqi (阿拉善右旗),
Longshou Shan (龙首山), X. T. Lei et al. (雷喜亭等) 252 (HIMC); Ejin Qi (额济纳旗), X. T.
Lai (雷喜亭) 84684 (HIMC); Siziwang Qi (四子王旗), Mongolia-Ningxia Exped. (蒙宁队)
169 (HIMC); Tengger Shamo (腾格里沙漠), Z. Y. Guo (郭中央) 1105 (HIMC); Uxin Qi (乌
审旗), Y. T. Liu (刘亚田) 152 (HIMC); Ordos (鄂尔多斯), 1871, Przewalski s.n. (K); the
same locality, 1859, Tatarinow s.n. (K, S). Ningxia (宁夏): Taole (陶乐), Z. W. Zhang (张振
万) 512, 577 (PE, WUK).
Mongolia. Gobi, 1886, Potanin s.n. (K).
Ser. 2. Virides W. T. Wang & L. Q. Li, ser. nov. Type: C. viridis (W. T. Wang & M. C.
Chang) W. T. Wang.
Flores ad apices ramulorum rami hornotini axillarium brevium vel abbreviatorum
solitarii, tantum pedicellati; pedunculi absentes.
Flowers solitary, terminal to the axillary short or abbreviated branchlets of hornotinous
branch, pedicellate only; peduncles wanting.
One species endemic to the northern Hengduan Mountains of Southwest China.
5. Clematis viridis (W. T. Wang & M. C. Chang) W. T. Wang in Fl. China 6: 364. 2001.
——C. canescens (Turcz.) W. T. Wang & M. C. Chang ssp. viridis W. T. Wang & M. C.
Chang in Fl. Reip. Pop. Sin. 28: 150, 356. 1980; M. Y. Fang in Fl. Xizang. 2: 90, fig. 25: 4, 5.
1985; M. Johnson, Klematis 654. 1997, p.p. excl. syn.; Grey-Wilson, Clematis 127. 2000.
——C. canescens var. viridis (W. T. Wang & M. C. Chang) W. T. Wang in Acta Phytotax.
Sin. 36: 161. 1998. Type: China. Sichuan (四川): Barkam (马尔康), alt. 2750 m, 1957-06-11,
X. Li (李馨) 71434 (holotype, PE!; isotypes, IBSC!, WUK!); Xiaojin (小金), 1958-07-16, S.
S. Chang et al. (张秀实等) 5927 (paratype, PE!). Xizang (西藏): Jomda (江达), Gamtog (岗
托), alt. 3150 m, 1959-07-29, S. Jiang & T. L. Chin (姜恕, 金存礼) 2605 (paratype, PE!).
C. fruticosa auct. non Turcz.: Rehd. & Wils. in Sarg., Pl. Wils. 1: 326. 1913.
绿叶铁线莲 Fig. 5: A-C
Small shrub. Stem 60-120 cm tall; branches 4-6-angulate, shallowly 4-6-sulcate,
glabrous, rarely puberulous; axillary branchlets 0.2-4 cm long, puberulous. Leaves simple,
shortly petiolate; leaf blades green, chartaceous, lanceolate-linear or linear, rarely lanceolate,
1.4-3.5(-4.5)×0.2-0.6(-0.8) cm, apex acute or pungent, base cuneate or attenuate, margin
entire, on both surfaces sparsely puberulous or subglabrous, midrib abaxially slightly
prominent; petioles 0.5-2.5 mm long or nearly wanting. Flowers solitary, terminal to axillary
short or abbreviated branchlets, only pedicellate, 2-3 cm in diam.; pedicel slender, 1.4-4 cm
long, puberulous. Sepals 4, yellow, ovate or subelliptic, 1.4-2.7×0.7-1.3 cm, apex apiculate,
inside glabrous, outside sparsely puberulous and on margin velutinous, marginal wings
1-3(-4) mm broad. Stamens 6-13 mm long, glabrous; anthers narrowly oblong or oblong,
2.5-4 mm long, apex obtuse. Ovaries densely pubescent; styles 6-9 mm long, densely villous.
Achenes elliptic or narrowly obovate, 4-6×2.5-3.5 mm, densely pubescent, near apex
villous, rimmed; persistent styles (2-)3-3.5 cm long, white-plumose. Fl. Jun.-Jul.
China (S Qinghai, W Sichuan, and E Xizang). On dry slopes or in bushes; alt.
3000-3600 m.
Additional specimens examined:
China. Qinghai (青海): Chindu (称多), T. N. Ho et al. (何廷农等) 1907 (PE); Yushu
Acta Phytotaxonomica Sinica Vol. 43

208
(玉树), T. N. Ho et al. (何廷农等) 3070 (PE). Sichuan (四川): Baiyü (白玉), W. L. Chen et
al. (陈伟烈等) 6860, Z. X. Tang (汤宗孝) 706 (PE); Dege (德哥), Sichuan Exped. (四川队)
74-7262, 74-7452 (PE); Sêrtar (色达), Sichuan Exped. (四川队) 74-6565 (PE); Xiaojin (小
金), E. H. Wilson 2483 (GH, K, LE), F. T. Wang (汪发缵) 21279 (PE). Xizang (西藏):
Jomda (江达), Y. W. Tsui (崔友文) 5530, Qinghai-Xizang Exped. (青藏队) 76-12466, Z. J.
Ren (任再金) s.n. (PE); Gamtog (岗托), P. C. Tsoong (钟补求) 5117, Sichuan Exped. (四川
队) 74-7013, Qinghai-Xizang Exped. (青藏队) 76-9842 (PE).
Acknowledgements We are most grateful to the directors and curators of G, GH, HIMC,
IBSC, K, LE, P, S, TIE, UPS, US, and WUK for kindly inviting us or the senior author to visit
their herbaria or/and sending herbarium material on loan; to HIMC for the gift of two
specimens of Clematis canescens; to ZHU Zong-Yuan for providing valuable information on
distribution and ecology of the species of the C. fruticosa group occurring in Nei Mongol; to
WANG Xue-Ming (TIE) for providing the paper on North China Ranunculaceae written by
Kozlov; to QIN Hai-Ning, YANG Qin-Er, ZHANG Zhi-Yun, WANG Zhong-Tao, and BAN
Qin for various kinds of help during the preparation of the present revision; and to SUN
Ying-Bao for making the drawings.
References
Axelrod D I, Al-Shehbaz I, Raven P H. 1998. History of the modern flora of China. In: Zhang A-L, Wu S-K
eds. Floristic Characteristics and Diversity of East Asian Plants. Beijing: China Higher Education Press.
43-55.
Chang M-C (张美珍). 1980. Clematis sect. Fruticella. In: Flora Reipublicae Popularis Sinicae (中国植物志).
Beijing: Science Press. 28: 147-156.
Essig F B. 1991. Seedling morphology in Clematis (Ranunculaceae) and its taxonomic implications. Sida 14:
377-390.
Grey-Wilson C. 2000. Clematis the genus. Portland, Oregon: Timber Press.
Handel-Mazzetti H. 1939. Plantae Sinenses a Dre. H. Smith annis 1921-1922, 1924 et 1934 lectae. XXXIII.
Ranunculaceae. Acta Horti Gotoburgensis 13: 37-219.
Hsu J (徐仁). 1983. Late Cretaceous and Cenozoic vegetation in China, emphasizing their connections with
North America. Annals of the Missouri Botanical Garden 70: 490-508.
Johnson M. 1997. Släktet Klematis. Sodertalje: M Johnsons Plantskula AB.
Kozlov I. 1933. Herbier du Musée Hoangho Paiho. Renonculacées. Publications du Musée Hoangho Paiho
Tien Tsin 22: 1-43.
Kuntze O. 1885. Monographie der Gattung Clematis. Verhandlungen des Botanischen Vereins der Prov.
Brandenburg 26: 83-202.
Liou Y-X (刘媖心). 1982. Observations on the formation of Chinese desert floras. Acta Phytotaxonomica
Sinica (植物分类学报) 20: 131-141.
Maximowicz C J. 1877. Diagnoses plantarum novarum Asiaticarum. I. Bulletin de I’Académie impériale des
Sciences de St. Pétersbourg 23: 305-391.
Maximowicz C J. 1889a. Flora Tangutica 1. St. Petersburg.
Maximowicz C J. 1889b. Enumeratio plantarum hucusque in Mongolia nec non adjacente parte Turkestaniae
sinensis lectarum 1. St. Petersburg.
Maximowicz C J. 1889c. Plantae Chinenses Potaninianae nec non Piasezkianae. Acta Horti Petropolitani 11:
1-112.
Prantl K. 1888. Beträge zur Morphologie und Systematik der Ranunculaceen. Botanische Jahrbücher 9:
225-373.
Rehder A, Wilson E H. 1913. Clematis. In: Sargent C S ed. Plantae Wilsonianae. Cambridge: The University
Press. 1: 319-343.
No. 3 WANG Wen-Tsai et al.: A revision of Clematis sect. Fruticella (Ranunculaceae)

209
Snoeijer W. 1992. A suggested classification for the genus Clematis. Clematis 1992: 7-20.
Spach E. 1839. Trib. Clematideae. Histoire des Naturelle Végétaux. Phanérogames. Paris 7: 257-284.
Tamura M. 1955. Systema Clematidis Asiae Orientalis. Science Reports Osaka University 4: 43-55.
Tamura M. 1967. Morphology, ecology and phylogeny of the Ranunculaceae 7. Science Reports Osaka
University 16: 21-43.
Tamura M. 1995. Clematis. In: Heipko P ed. Engler’s Die Natürlichen Pflanzenfamilien. 2nd ed. Berlin:
Duncker & Humblot. 17a (4): 368-387.
Turczaninow N. 1832. Decades tres plantarum novarum Chinae boreali et Mongoliae chinensi incolarum.
Bulletin de la Société des naturalistes de Moscou 5: 180-206.
Wang W-T (王文采). 2002. A revision of Clematis sect. Cheiropsis (Ranunculaceae). Acta Phytotaxonomica
Sinica (植物分类学报) 40: 193-241.
Wang W-T (王文采). 2003. A revision of Clematis sect. Clematis (Ranunculaceae). Acta Phytotaxonomica
Sinica (植物分类学报) 41: 1-62, 97-172.
Wang W-T (王文采), Bartholomew B. 2001. Clematis. In: Wu Z-Y, Raven PH eds. Flora of China. Beijing:
Science Press; St. Louis: Missouri Botanical Garden Press. 6: 333-386.
Zhou T-R (周廷儒). 1982. Paleogeography. Beijing: Beijing Normal University Press.
铁线莲属灌木铁线莲组修订
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)
摘要 对毛茛科铁线莲属Clematis的灌木铁线莲组sect. Fruticella进行了全面修订,确定此组共含5种、2
变种和3变型;对此组的分类学简史和地理分布做了介绍;写出本组2系的形态特征和地理分布,组下分类
群检索表,以及各种植物的形态描述、地理分布、生长环境等,并附有各分类群的3幅插图。根据花的构造
认为在铁线莲属中,灌木铁线莲组与黄花铁线莲组sect. Meclatis在亲缘关系上最为接近。二组拥有的共
同特征为: 花的4枚萼片呈黄色,斜上方开展;雄蕊花丝下部变宽,呈披针状条形。二组的区别为: 灌木铁
线莲组的萼片边缘在花开放后展宽成膜质狭翅,雄蕊无毛;而在黄花铁线莲组,萼片边缘不展宽,雄蕊花丝
被柔毛。本组5种的花构造一致;在其他营养器官形态特征方面观察到以下演化趋势: (1)叶由不分裂到羽
状全裂,由绿色到灰绿色,由狭菱状卵形到条形;(2)花序含数朵花,具花序梗和2苞片,生于当年生枝顶端,
到花数目减少到1朵,花序梗和苞片消失,花生于当年生枝的腋生短枝顶端。根据上述演化趋势认为灌木
铁线莲的模式变型C. fruticosa f. fruticosa (叶绿色,狭卵形或披针形;花序通常含数朵花,顶生于当年生枝
上)是本组的原始类型,其他种可能均源出于此类型。根据花序特征,本组被划分为2系。第1系,灌木铁线
莲系ser. Fruticosae,含4种,分布于我国黄土高原及相邻干旱地区,向北达蒙古戈壁荒漠,向南在甘肃越过
秦岭到达甘肃南部。在黄土高原北部及相邻的内蒙古西南部集中分布本系全部4种,这里是灌木铁线莲
组的分布中心。第2系,绿叶铁线莲系ser. Virides,含绿叶铁线莲C. viridis 1种,分布于我国横断山区北部的
高山灌丛草地。
关键词 铁线莲属; 灌木铁线莲组; 毛茛科; 分类学修订