全 文 ：植 物 分 类 学 报 44 (4): 401–436(2006) doi:10.1360/aps050049
Acta Phytotaxonomica Sinica http://www.plantsystematics.com
———————————
Received: 28 March 2005 Accepted: 31 March 2005
Supported by the National Natural Science Foundation of China, Grant No. 30470126.
A revision of Clematis sect. Meclatis (Ranunculaceae)
WANG Wen-Tsai
(State Key Laboratory of Systematic and Evolutionary Botany, Institute of Botany, the Chinese Academy of Sciences,
Beijing 100093, China)
Abstract Clematis sect. Meclatis is revised in this paper. Brief taxonomic history and
geographical distribution of the section are given, its systematic position and the relationships
among the species are discussed, and the evolutionary trends of some characters in the section
are evaluated. Clematis akebioides (Maxim.) Veitch and C. tangutica (Maxim.) Korsh. are
considered the primitive species in the section, whereas C. caudigera W. T. Wang and C.
corniculata W. T. Wang are considered the advanced ones. The western edge of the
Qinghai-Xizang (Tibet) plateau with the Pamirs and the adjacent mountains, the highest land
mass in the world, where 10 species of the section are concentrated, is regarded as the
distribution center, and the eastern edge of the Qinghai-Xizang (Tibet) plateau, where the two
primitive species, C. akebioides and C. tangutica, sympatrically occur, may be the center of
origin of the section. The inclusion of C. ispahanica Boiss. and C. graveolens Lindl. in sect.
Meclatis by some authors is not accepted, with the former being a member of sect. Clematis,
and the latter a member of sect. Brachiatae Snoeijer. A new variety, C. intricata Bunge var.
intrapuberula W. T. Wang, is described, and two new combinations, C. tangutica var.
mongolica (Grey-Wilson) W. T. Wang and C. tibetana Kuntze var. pamiralaica (Grey-Wilson)
W. T. Wang, is proposed. As a result, 13 species and 13 varieties are recognized in sect.
Meclatis. They are keyed, described, and illustrated.
Key words Clematis, Clematis sect. Meclatis, taxonomic revision.
1 Brief taxonomic history
Linnaeus (1753) described C. orientalis L., the first species of Clematis sect. Meclatis.
In the first revision of the genus Clematis, de Candolle (1818) placed the two species of
sect. Meclatis, C. orientalis L. and C. glauca Willd., in the first group of his sect. Flammula,
which is characterized by “pedunculis ramoso-paniculatis, foliis pinnatim sectis”.
In his account of the tribe Clematideae of the Ranunculaceae, Spach (1839) established
the new genus Meclatis on the basis of C. orientalis and C. glauca. In his classification of the
Clematis, Baillon (1867) correctly sunk Spach’s new genus to sectional rank. However, this
treatment had long been overlooked by many authors working on the taxonomy of Clematis
until Brandenburg (2000) recognized this section.
In the monograph on the genus Clematis published by Kuntze (1885), one species of the
C. orientalis group, C. orientalis, was included in his sect. Scandentes aperulatae, and one
other species of that group, C. tibetana Kuntze, in his sect. Escandentes. Here, C. orientalis
was treated in so broad a sense that two species (C. intricata Bunge and C. daurica Pers.)
belonging to the C. orientalis group and five species (C. massoniana DC., C. graveolens
Lindl., C. brachiata Thunb., C. wightiana Wall., and C. simensis Fresen.) belonging to the C.
brachiata group were all treated as infraspecific taxa of C. orientalis.
In his classification of the genus Clematis, Prantl (1888) established a new subsection,
subsect. Orientales, under sect. Flammula. This subsection, consisting of C. orientalis, C.
glauca and C. ispahanica Boiss, was afterwards accepted by Schneider (1906), and by Rehder
Acta Phytotaxonomica Sinica Vol. 44 402
and Wilson (1913), and by Rehder (1951) who relegated it to the series rank. The inclusion of
C. ispahanica, which has white, spreading sepals, glabrous stamens, and linear filaments, and
thus should belong to sect. Clematis, in the C. orientalis group was adopted by Brandenburg
(2000, see below).
Tamura (1955) recognized Prantl’s subsect. Orientales, but transferred it from sect.
Flammula DC. to sect. Viorna (Reichb.) Prantl. Possibly having ignored or overlooked
Baillon’s work on Clematis, Tamura (1967) also reduced Spach’s genus Meclatis as a section
of Clematis, and classified the species of the section into subsect. Orientales, which is
characterized by having flowers in axillary and terminal inflorescences, and into subsect.
Tanguticae, which is characterized by having solitary and terminal flowers. In his revised
classification of the genus Clematis, Tamura (1987) placed sect. Meclatis in his new subgenus
Campanella between sect. Atragene (L.) DC. and sect. Pseudanemone Prantl, and still maintained
his previous subdivision of sect. Meclatis (Tamura, 1967). However, Grey-Wilson (1989)
considereded that there are variations in the characters for distinguishing the two subsections, and
thus that the subdivision proposed by Tamura should not be recognized. Tamura (1995) accepted
Grey-Wilson’s opinion, and abandoned his previous subdivision of sect. Meclatis, and at the same
time added the African species of the C. brachiata group into sect. Meclatis.
Grey-Wilson (1989) published a revision of the C. orientalis group, and recognized 10
species, in which he followed Schneider (1906) to include the western Himalayan C.
graveolens Lindl., a member of sect. Brachiatae (Wang, 2004). In the analytical key, the
absence or presence of hairs on the adaxial surface of sepal is used as a character for the
primary subdivision of the species. In his book entitled Clematis the Genus, Grey-Wilson
(2000) followed Tamura (1987) to place sect. Meclatis in subgen. Campanella but behind sect.
Campanella Tamura (i.e. the C. connata group) and sect. Bebaeanthera Edgew., and
transferred subsect. Africanae, a subsection established by Johnson (1997) on the basis of the
C. brachiata group, from sect. Clematis to sect. Meclatis and placed it before subsect. Meclatis
(i.e. the C. orientalis group). In subsect. Meclatis 14 species are recognized, and the leaf
colour is used as an important character for distinguishing them.
Johnson (1997), in his monograph of the genus Clematis, placed sect. Meclatis just
behind sect. Connatae (Koehne) M. Johnson (i.e. the C. connata group), and recognized 13
species in sect. Meclatis, with the species, like those in other sections, being arranged
alphabetically. He misplaced C. graveolens, a member of sect. Brachiatae as mentioned
above, in sect. Meclatis, as Schneider (1906) and Grey-Wilson (2000) did.
Brandenburg (2000) carried out an intensive study on the taxonomy of sect. Meclatis, in
which the evolutionary trends of most of the characters in this section were analysed, and on
the basis of the results of analysis, a cladogram was reconstructed as follows:

The arrangement of the six species he recognized is as follows: Clematis orientalis L., C.
graveolens Lindl., C. intricata Bunge, C. ispahanica Boiss., C. serratifolia Rehd., and C.
tibetana Kuntze with three subspecies, ssp. tibetana, ssp. tangutica (Maxim.) Brandenb., and
ssp. vernayi (C. E. C. Fisch.) Grey-Wilson. In his treatment of sect. Meclatis, C. hilariae
No. 4 WANG Wen-Tsai: A revision of Clematis sect. Meclatis (Ranunculaceae) 403
Kaval. and C. sarezica Ikonn. were reduced to synonymy under C. intricata; C. akebioides
(Maxim.) Veitch to those under C. tibetana ssp. tibetana and ssp. tangutica; C. ladakhiana
Grey-Wilson to that under C. tibetana ssp. tibetana; C. pamiralaica Grey-Wilson to that under
C. tibetana ssp. tangutica; none of the varieties of C. orientalis were recognized. Besides, C.
ispahanica, a member of sect. Clematis subsect. Angustifoliae (Wang, 2003), and C.
graveolens, a member of sect. Brachiatae (Wang, 2004), were both misplaced in sect.
Meclatis. So, sect. Meclatis defined by Brandenburg (2000) is a heterogeneous group.
2 Systematic position
Sect. Meclatis is closely related to sect. Brachiatae in the similar floral structure (Wang,
2004), but differs by having usually ascending and yellow sepals, and stamen filaments
widened in the lower part. In fact, as mentioned above, Kuntze (1885), Tamura (1995), and
Grey-Wilson (2000) have all associated the C. orientalis group with the C. brachiata group, in
which the sepals are always spreading and white in colour, and the stamen filaments are
narrowly linear in outline, not widened in the lower part. Section Brachiatae with floral
structure showing striking resemblance to that of sect. Clematis is closely related to and may
be derived from sect. Clematis. Being a close ally of sect. Brachiatae, sect. Meclatis may also
be derived from sect. Clematis, and should be a member of subgen. Clematis (Wang, 2003;
Wang & Li, 2006).
3 Relationships among the species
The 13 species of sect. Meclatis are so closely related to each other that no subdivision of
the section can be made (Grey-Wilson, 1989, 2000; Tamura, 1995; Johnson, 1997;
Brandenburg, 2000). After analysing the variational patterns of the various morphological
characters in the section, the evolutionary trends of some characters may be revealed as
follows: (1) the leaf colour may have changed from green to grey-green due to adaptation to
the arid climate; (2) the ovate or broadly ovate, palmately lobed and dentate leaflets may
represent the primitive condition, and the lanceolate to linear, undivided and entire leaflets
may represent the derived condition; (3) the solitary, terminal flower without a peduncle and
two opposite bracts may be derived from the pedunculate, 2-bracteate, few- to many-flowered
cyme (Wang, 2002); (4) the evolutionary trend of the sepal shape is somewhat like that of the
leaflet shape, i.e. from ovate to lanceolate and linear; (5) the sepals glabrous inside may
represent the primitive condition, and the sepals hairy inside may represent the derived
condition; (6) the sepals without any apical projections may represent the primitive condition,
and those with apical projections may represent the derived condition; (7) the anthers may
have changed from oblong, narrowly oblong to linear in outline. According to the evolutionary
trends of characters mentioned above, the two species, C. akebioides and C. tangutica, in
which the leaflets are green, ovate or oblong, dentate or crenate, the cymes are
several-flowered (in C. tangutica the flowers are often singular and terminal), the sepals are
ovate and glabrous inside, and the anthers are narrowly oblong or oblong, seem to be the
primitive species of sect. Meclatis, and the two species, C. caudigera W. T. Wang and C.
corniculata W. T. Wang, in which the leaflets are grey-green, and deeply divided or lanceolate,
the flowers are singular and terminal, and the sepals are oblong-lanceolate, and caudate or
corniculate, are considered the advanced ones. Besides, the species C. orientalis, in which the
leaflets or leaflet lobes are grey-green, often lanceolate, or oblong-lanceolate, the sepals are
puberulous inside, and the anthers are usually linear, may also be an advanced one.
4 Geographical distribution
In the present revision, 13 species and 13 varieties of sect. Meclatis are recognized. They
Acta Phytotaxonomica Sinica Vol. 44 404
are widespread in southeastern Europe and western, central, and northeastern Asia (Fig. 1).
Ten species (C. tangutica, C. tibetana, C. ladakhiana, C. glauca, C. zandaensis W. T. Wang, C.
hilariae, C. sarezica, C. orientalis, C. caudigera, and C. corniculata) and 10 varieties are
concentrated in the western edge of the Qinghai-Xizang (Tibet) plateau with the Pamirs and
the adjacent montane regions, the highest land mass in the world. Of them, the typical variety
of C. orientalis extends from this high land mass westward via the arid regions of western Asia
to the Aegian Islands of Greece and eastward to the arid regions of Xinjiang and north-western
Gansu of China and Mongolia, C. tangutica extends from this land mass eastward to eastern
edge of the Qinghai-Tibet Plateau and the arid regions of the western Loess plateau and
Mongolia (Fig. 2), C. glauca extends from this land mass northeastward to southern Siberia,
and the remaining seven species are endemic to this land mass proper. On the eastern
Qinghai-Xizang plateau and in the arid regions of northern Gansu, Ningxia, and Nei Mongol
of China and Mongolia occur four species (C. akebioides, C. tangutica, C. intricata, and C.
glauca), with C. akebioides and C. intricata extending eastward to the eastern Loess plateau,
northern Hebei and western Liaoning (Fig. 2). In northern Korea, northeastern China, Far East
Region of Russia, and northern Japan is distributed one species (C. serratifolia). The western
edge of the Qinghai-Xizang (Tibet) plateau with the Pamirs and the adjacent montane regions,
where 10 species of sect. Meclatis are concentrated, therefore, may be regarded as the
distribution center of the section, and the eastern edge of the Qinghai-Xizang plateau, where
the two primitive species in the section, C. akebioides and C. tangutica, sympatrically occur
(Fig. 2), may be regarded as the center of origin of the section.





Fig. 1. Map showing the distribution of the section Meclatis.




No. 4 WANG Wen-Tsai: A revision of Clematis sect. Meclatis (Ranunculaceae) 405

Fig. 2. Map showing the distribution of Clematis akebioides, C. tangutica, C. intricata, C. caudigera, and C. corniculata.

5 Taxonomic treatment
Clematis L. sect. Meclatis (Spach) Baillon, Hist. Pl. 1: 57. 1867; Brandenb., Meclatis in
Clematis 100. 2000, p.p. excl. C. ispahanicam Boiss. et. C. graveolentem Lindl.——Meclatis
Spach, Hist. Nat. Veg. Phan. 7: 272. 1839.——Sect. Meclatis (Spach) Tamura in Sci. Rep.
Osaka Univ. 16: 32. 1967; P. P. Ling in Fl. Reip. Pop. Sin. 28: 139. 1980; Tamura in Acta
Phytotax. Geobot. 38: 39. 1987; et in Heipko, Engler’s Nat. Pflanzenfam., Zwei. Auf., 17a (4):
375. 1995. p.p. excl. pl. Afric.; Grey-Wilson in Kew Bull. 44: 33. 1989, p.p. excl. C.
graveolentem; Snoeijer in Clematis 1992: 11. 1992; M. Johnson, Klematis 329. 1997, p.p.
excl. C. graveolentem; Grey-Wilson, Klematis 155. 2000, p.p. excl. subsect. Africanas et C.
graveolentem. Lectotype: Meclatis orientalis (L.) Spach (= C. orientalis L.).
Sect. Flammula DC. subsect. Orientales Prantl in Bot. Jahrb. 9: 261. 1888, p.p., excl. C.
ispahanicam; Schneid., Ill. Handb. Laubh. 1: 293. 1906. p.p., excl. C. graveolentem.——Sect.
Flammula ser. Orientales (Prantl) Rehd. & Wils. in Sarg., Pl. Wils. 1: 342. 1913; Rehd., Man.
Cult. Trees & Shrubs, ed. 2, 219. 1940.——Sect. Viorna (Reichb.) Prantl subsect. Orientales
(Prantl) Tamura in Sci. Rep. Osaka Univ. 4: 48. 1955; et in Acta Phytotax. Geobot. 16: 80.
1956; et in Sci. Rep. Osaka Univ. 16: 32. 1967; et in Acta Phytotax. Geobot. 38: 40. 1987;
Grey-Wilson in Kew Bull. 44: 34. 1989. Lectotype: C. orientalis L.
Acta Phytotaxonomica Sinica Vol. 44 406
Sect. Flammula subsect. Tanguticae Schneid., Ill. Handb. Laubh. 1: 294. 1906.——
Sect.Viorna subsect. Tanguticae (Schneid.) Tamura in Sci. Rep. Osaka Univ. 4: 48. 1955; et in
Acta Phytotax. Geobot. 16: 80. 1956.——Sect. Meclatis subsect. Tanguticae (Schneid.)
Tamura in Sci. Rep. Osaka Univ. 16: 33. 1967; et in Acta Phytotax. Geobot. 38: 40. 1987.
Lectotype: C. tangutica (Maxim.) Korsh.
Sect. Meclatis subsect. Meclatis Grey-Wilson, Clematis 29: 162. 2000, p.p., excl. C.
graveolentem.
Woody vines, rarely dwarf erect shrubs or subshrubs. Seedling leaves alternate (Essig,
1991). Cauline leaves opposite, 1–2-pinnate or 2-ternate. Flowers bisexual, in pedunculate,
2-bracteate, few- to many-flowered cymes, sometimes solitary, terminal, and only pedicellate.
Sepals 4, yellow, sometimes purple or red, ascending, rarely spreading, ovate, lanceolate, or
narrowly oblong, glabrous or puberulous on both surfaces, velutinous on margin. Stamens
numerous; filaments narrowly lanceolate or lanceolate-linear, pubescent; anthers oblong,
narrowly oblong, or linear, apex obtuse or minutely apiculate. Carpels numerous, with long
villous styles. Achenes compressed, ovate or elliptic, with elongate, plumose persistent styles.
Thirteen species with thirteen varieties mostly occurring in central and northern Asia,
only one species extending from central Asia westward via western Asia to the Aegean Islands
of Greece.
Key to species and varieties
1. Sepals glabrous inside.
2. Sepals not corniculate; flowers usually in axillary cymes, sometimes solitary and terminal (C. tangutica).
3. Leaflet margin crenate or entire; flowers usually in axillary cymes.
4. Sepals ovate or narrowly ovate.
5. Leaflets blue-green, broadly oblong, elliptic or ovate, margin crenate or entire⋯1. C. akebioides
5. Leaflets grey-green, usually lanceolate to lanceolate-linear, margin usually entire⋯⋯⋯⋯⋯⋯
⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯3. C. intricata
6. Leaflets all lanceolate or lanceolate-linear; sepals yellow⋯⋯⋯⋯⋯⋯⋯⋯3a. var. intricata
6. Terminal leaflet lanceolate or elliptic-lanceolate, lateral leaflets elliptic or long elliptic; sepals
purple⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯3b. var. purpurea
4. Sepals narrowly oblong⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯6. C. glauca
3. Leaflet margin usually regularly dentate or denticulate; flowers solitary, terminal, sometimes in
1–3-flowered cymes⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯2. C. tangutica
7. Leaflets ovate to lanceolate, with up to 7 teeth per side; usually woody vines, occasionally dwarf
erect shrubs⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯2a. var. tangutica
7. Leaflets narrowly lanceolate, with 8–13 teeth per side; woody vines⋯⋯⋯⋯⋯2b. var. mongolica
2. Sepals outside below apex corniculate; flowers solitary, terminal; leaflets grey-green, linear-lanceolate,
margin entire or 1-denticulate⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯13. C. corniculata
1. Sepals inside puberulous.
8. Dwarf erect subshrubs⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯4d. C. tibetana var. pamiralaica
8. Woody vines.
9. Flowers solitary, terminal, or also in lateral, axillary cymes.
10. Sepal without tail-like projection at apex; flowers solitary, terminal, or also in lateral, axillary,
1–3-flowered cymes.
11. Leaflet margin regularly dentate⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯2c. C. tangutica var. pubescens
11. Leaflet margin usually entire, sometimes few-dentate⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯4. C. tibetana
12. Leaflets narrowly ovate, elliptic, lanceolate, or linear, margin usually entire.
13. Sepals lanceolate, apex attenuate⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯4a. var. tibetana
13. Sepals ovate, apex acute⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯4b. var. vernayi
12. Leaflets rhombic-ovate, 2–3-lobed and few-dentate⋯⋯⋯⋯⋯⋯⋯⋯4c. var. laciniifolia
No. 4 WANG Wen-Tsai: A revision of Clematis sect. Meclatis (Ranunculaceae) 407
10. Sepal with tail-like projection 3–6 mm long at apex; flowers only solitary and terminal, never in
axillary cymes⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯12. C. caudigera
9. Flowers usually in lateral, rarely also in terminal cymes, never solitary and terminal.
14. Sepals brown-purple, outside below apex shortly corniculate; leaflets lanceolate, margin with 1–2
teeth per side⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯10. C. sarezica
14. Sepals yellow (in C. ladakhiana sometimes tinged with purple-brown), not corniculate.
15. Leaflet margin regularly dentate or serrate.
16. Sepals glabrous outside.
17. Leaflets usually narrowly ovate or lanceolate, undivided, apex attenuate, margin
serrate⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯8. C. serratifolia
17. Leaflets broadly ovate or ovate, 2–3-lobed, apex acute, margin irregularly dentate.
18. Sepals ovate or narrowly ovate; anthers narrowly oblong, 1.4–3.2 mm long⋯⋯⋯
⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯7. C. zandaensis
18. Sepals oblong-lanceolate; anthers linear, ca. 4 mm long⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯11b. C. orientalis var. latifolia
16. Sepals puberulous outside.
19. Leaflets green, narrowly ovate or lanceolate, undivided or 3-lobed⋯⋯⋯9. C. hilariae
19. Leaflets grey-green, ovate or broadly ovate, 3-lobed to 3-sect⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯11b. C. orientalis var. albida
15. Leaflet margin entire or with 1–2 teeth per side.
20. Leaflets ovate, narrowly ovate, or elliptic⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯6. C. glauca
20. Leaflets lanceolate to linear.
21. Peduncles short, robust, up to 2 mm in diam., 2–5 mm long, 1-flowered; pedicels 4–8
cm long⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯11g. C. orientalis var. robusta
21. Peduncles usually slender, 0.6–1.8 mm rarely 2 mm (C. hilariae) in diam., up to 1–12
cm long, usually 3–many-flowered.
22. Sepals glabrous outside.
23. Leaflets green, lanceolate-linear, apex attenuate; sepals narrowly ovate or
broadly lanceolate⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯5. C. ladakhiana
23. Leaflets grey-green, linear-lanceolate, apex acute; sepals narrowly oblong-
lanceolate⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯11e. C. orientalis var. tenuifolia
22. Sepals puberulous outside.
24. Sepals narrowly ovate or broadly lanceolate.
25. Leaflets usually entire; peduncles 2–4 mm long, 1–1.2 mm in diam.;
pedicels 2.2–3 cm long; sepals narrowly ovate, outside glabrescent⋯⋯⋯
⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯3b. C. intricata var. intrapuberula
25. Leaflets below sparsely dentate or serrate; peduncles 0.4–6 cm long, 1.8–2
mm in diam.; pedicels 3–10 cm long; sepals narrowly ovate or lanceolate,
puberulous outside⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯9. C. hilariae
24. Sepals oblong-lanceolate.
26. Leaflets (0.4–)0.6–4.5 cm broad.
27. Peduncles 1.4–6.5 cm long; sepals 6.5–15 mm long; persistent styles
2.5–5.5 cm long⋯⋯⋯⋯⋯⋯⋯⋯11a. C. orientalis var. orientalis
27. Peduncles 6–12 cm long; sepals 16–21 mm long; persistent styles up to
8 cm long⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯11c. C. orientalis var. robusta
26. Leaflets linear, 1–4(–5) mm broad⋯11f. C. orientalis var. baluchistanica

1. Clematis akebioides (Maxim.) Veitch, New Hardy Pl. West China 19. 1912; P. P. Ling in
Fl. Reip. Pop. Sin. 28: 145, pl. 42. 1980; M. Y. Fang in Fl. Xizang. 2: 89. 1985; Grey-Wilson
in Kew Bull. 44: 43. 1989; Y. Z. Zhao in Fl. Intramongol., ed. 2, 2: 533, pl. 215, figs. 4–6.
Acta Phytotaxonomica Sinica Vol. 44 408
1990; L. Q. Li in Vasc. Pl. Hengduan Mount. 1: 523. 1993; L. H. Zhou in Fl. Qinghai. 1: 348.
1997; M. Johnson, Klematis 333. 1997; Grey-Wilson, Clematis 166, fig. 120. 2000; J. Q. Fu in
Fl. Loess Plat. 1: 441. 2000; W. T. Wang in Fl. Yunnan. 11: 238. 2000; W. T. Wang & Barth. in
Fl. China 6: 336. 2001.——C. orientalis L. var. akebioides Maxim. in Acta Hort. Petrop. 11: 6.
1890.——C. glauca Willd. var. akebioides (Maxim.) Rehd. & Wils. in Sarg., Pl. Wils. 1: 342.
1913; Rehd. in J. Arn. Arb. 4: 191. 1923; et Man. Cult. Trees & Shrubs 229. 1927;
Hand.-Mazz. in Acta Hort. Gotob. 13: 218. 1939; C. Y. Wu, Ind. Fl. Yunnan. 109. 1984. Type:
China. Sichuan: Heishui, 1885-07-22, Potanin s.n. (holotype, LE!).
C. tangutica auct. non (Maxim.) Korsh.: Kozlov in Publ. Mus. Hoangho Paiho Tien Tsin
22: 14. 1933.
C. glauca auct. non Willd.: P. P. Ling in Fl. Reip. Pop. Sin. 28: 143. 1980, p.p., quoad pl.
Qinghai., Gansu., Shaanxi. et Shanxi.; Y. J. Ling et al. in Fl. Shanxi. 1: 633. 1992; L. H. Zhou
in Fl. Qinghai. 1: 347. 1997; W. T. Wang & Barth. in Fl. China 6: 366. 2001, p.p., quoad pl.
Qinghai., Gansu., Shaanxi., et Shanxi.
C. tibetana auct. non Kuntze: Brandenb., Meclatis in Clematis 165. 2000, p.p., quoad syn.
C. akebioides (Maxim.) Veitch.
C. tibetana Kuntze ssp. tangutica (Maxim.) Brandenb., l.c., p.p., quoad syn. C.
akebioides (Maxim.) Veitch.
甘川铁线莲 Fig. 3: A–C
Woody vine. Branches shallowly 6–10-sulcate, sparsely puberulous or subglabrous.
Leaves 1–2-pinnate; leaflets green, thinly papery or herbaceous, oblong, elliptic, or ovate,
occasionally lanceolate, 1.2–4×0.6–3 cm, apex obtuse, rounded, or slightly acute, base
broadly cuneate or rounded, margin crenate or entire, undivided or 2–3-lobed, adaxially
glabrous, abaxially glaucous, on veins sparsely puberulous or glabrous, basal veins flat;
petioles 3–7.8 cm long. Cymes axillary, 1–3(–5)-flowered; peduncles 0.2–3.5(–6) cm long;
bracts leaflet-like, 1–1.5 cm long. Flower 2–3.4 cm in diam.; pedicel 2.5–7 cm long, sparsely
puberulous or subglabrous. Sepals 4, yellow, greenish-yellow, or sometimes tinged with
purple, ascending, thinly to thickly papery, narrowly ovate or ovate-oblong, 1.6–2.7×0.6–1.1
cm, apex acute or apiculate, inside glabrous, on margin velutinous, outside glabrous or above
sparsely puberulous. Stamens 7–12 mm long; anthers narrowly oblong or oblong, 2–3 mm
long, glabrous, apex obtuse or minutely apiculate. Ovaries densely puberulous; styles 7–12
mm long, densely villous. Achenes obovate or elliptic, 2–3×1–1.8 mm, pubescent, margin
rimmed; persistent styles 2.5–3 cm long, plumose. Fl. Jul. –Sept.
China (SW Gansu, SW Nei Mongol, S & E Qinghai, Shaanxi, Shanxi, W Sichuan, E
Xizang, NW Yunnan). On grassy slopes, in bushes, or by streams; alt. 1200–3600 m.
Additional specimens examined:
China. Gansu (甘肃): Huating (华亭), S. Q. Zhong (仲世奇) 110 (PE); Huining (会宁), Huanghe River
Exped. (黄河队) 56-5100 (PE); Jonê (卓尼), W. Y. Hsia (夏纬瑛) 8499 (NAS, PE); Ka-tian-ko, Hummel
4137, 5050 (S); Lanzhou (兰州), Y. Q. He (何业祺) 4284, 5759 (PE); Liancheng (莲城); Y. Q. He (何业祺)
4961, 5310 (PE); Lianhua Shan (莲花山), Rock 13228 (NAS); Lichen, R. C. Ching (秦仁昌) 286 (US); Low
Rwen, Purdom 1022 (GH, US); Min Xian (岷县), Hummel 5382a (S); Têwo (迭部), P. C. Kuo (郭本兆) 5656
(WUK); Xiahe (夏河), R. C. Ching (秦仁昌) 775 (GH, US), T. P. Wang (王作宾) 7089 (NAS, PE); Yuzhong
(榆中), Huanghe River Exped. (黄河队) 56-3125 (PE); Zhangye (张掖), P. C. Tsoong (钟补求) 8770 (PE).
Hebei (河北): Zhuolu (涿鹿), C. G. Yang (杨朝广) 1410 (PE). Nei Mongol (内蒙古): Helan Shan (贺兰山),
X. T. Lei (雷喜亭) 840766 (NIMC). Qinghai (青海): Datong (大通), K. C. Kuan (关克俭) 77-303 (PE);
Huzhu (互助), P. C. Kuo (郭本兆) 9279, 9501 (PE); Menyuan (门源), K. M. Liou (刘继孟) 6861, 6933 (PE);
Minhe (民和), T. N. Ho (何廷农) 809 (PE); Qilian (祁连), P. C. Tsoong (钟补求) 8532 (PE); Qilian Shan (祁
连山), P. C. Kuo (郭本兆) 12565 (PE); Xining (西宁), K. S. Hao (郝景盛) 853, P. C. Tsoong (钟补求) 8187
(PE). Shaanxi (陕西): Without precise locality, K. T. Fu (傅坤俊) 930 (NAS). Shanxi (山西): Guandi Shan
No. 4 WANG Wen-Tsai: A revision of Clematis sect. Meclatis (Ranunculaceae) 409


Fig. 3. A–C, Clematis akebioides (Maxim.) Veitch. A, flowering branch; B, stamen (from X. Li 75478); C, leaf (from T.
P. Wang 7089). D, E, C. intricata Bunge var. intricata. D, flowering branch; E, stamen (from K. K. Tsoong s.n.).



Acta Phytotaxonomica Sinica Vol. 44 410
(关帝山), W. Y. Hsia (夏纬瑛) 1226 (PE); Hongdong (洪洞), Y. L. Chen (陈艺林) 825 (PE); Huo Xian (霍
县), Shanxi Exped. (山西考察队) 54-469 (PE); Jiaocheng (交城), Licent 2163 (TIE), H. Smith 7214 (GH, S,
TIE, UPS), 6893 (PE, UPS, US), Huanghe River Exped. (黄河队) 57-927 (NAS, PE); Luya Shan (芦芽山), H.
Smith 8122 (S, UPS); Ningwu (宁武), Shanxi Exped. (山西考察队) 57-1810 (PE); Wuzhai (五寨), Guanqin
Shan (管涔山), W. Y. Hsia (夏纬瑛) 1486 (PE); Xiaoyi (孝义), K. M. Liou (刘继孟) 2801 (IBSC, PE); Xi
Xian (隰县), Licent 2163 (PE). Sichuan (四川): Baoxing (宝兴), K. C. Kuan et al. (关克俭等) 2964 (K, PE);
Barkam (马尔康), X. Li (李馨) 71729, 72594, 72767 (NAS, PE), 71886, 72265 (IBSC, NAS); Batang (巴塘),
G. Yao (姚淦) 549 (NAS); Dawu (道孚), S. Jiang (姜恕) 2312 (PE); Daocheng (稻城), Sichuan Exped. (四川
队) 73-2130 (PE); Garzê (甘孜), Y. W. Tsui (崔友文) 4308, S. Jiang (姜恕) 9126, 9349 (PE); Heishui (黑
水), X. Li (李馨) 73482, 73853, 73963 (IBSC, NAS, PE); Hongyuan (红原), Q. S. Zhao (赵清盛) 234 (K, PE,
S, UPS); Jinchuan (金川), X. Li (李馨) 75130, 75277, 75845, 78036, 78321 (IBSC, PE); Kangding (康定),
Soulié 436 (G, K), Hosie s.n. (K), Mussot 6 (P), E. H . Wilson 3131, 3132 (GH, K), H. Smith 10872 (GH, PE,
S, UPS), T. P. Wang (王作宾) 9841, W. K. Hu & C. Ho (胡文光, 何景) 10650, 11347, X. L. Jiang (蒋兴麟)
36960, 37160 (PE); Li Xian (理县), C. L. Wu (吴中伦) 33138 (PE), X. Li (李馨) 46545 (IBSC); Luhuo (炉
霍), T. S. Ying (应俊生) 9347 (PE); Mao Xian (茂县), W. P. Fang (方文培) 1521 (K, GH, IBSC, NAS, P,
PE); Nanping (南坪), Q. X. Li (李全喜) 2694 (PE); Songpan (松潘), E. H. Wilson 4548 (GH, US), W. P.
Fang (方文培) 4267 (GH, NAS), K. T. Fu (傅坤俊) 2201 (PE); Tongolo (东俄洛), Soulié 921 (K, P);
Xiangcheng (乡城), T. T. Yu (俞德浚) 13385, Sichuan Exped. (四川队) 73-2980 (PE); Xiaojin (小金), S. S.
Chang (张秀实) 6506 (PE), 6945 (NAS), Sichuan Exped. (四川队) 75-9811 (PE); Yanyuan (盐源),
Handel-Mazzetti 4601 (P), 5418 (GH, IBSC, US). Xinjiang (新疆): Korla (库尔勒), A. J. Li & J. N. Zhu (李
安仁, 朱家柟) 8642 (PE). Xizang (西藏): Markam (芒康), Qinghai-Xizang Exped. (青藏队) 76-11732 (PE);
Zhag’yab (察雅), Qinghai-Xizang Exped. (青藏队) 76-12354 (PE). Yunnan (云南): Dêqên (德钦), C. W.
Wang (王启无) 70285 (GH), T. T. Yu (俞德浚) 10253, 10444 (PE); Lijiang (丽江), C. W. Wang (王启无)
71589 (NAS), K. M. Feng (冯国楣) 2665 (PE); Without precise locality, Forrest 12861 (K, NAS), 14856,
15636, 16921 (K); Tsekou, Soulié s.n. (P); Zhongdian (中甸), Handel-Mazzetti 4601 (P), T. T. Yu (俞德浚)
12474, 13940, K. M. Feng (冯国楣) 1790 (PE).
In C. akebioides, the populations occurring in northwestern Yunnan, western Sichuan,
eastern Xizang, and the Mt. Helanshan in Nei Mongol have leaflets which are usually broadly
oblong in outline, with obtuse apexes and crenate margins, and those occurring in eastern
Qinghai, Gansu, Shaanxi, and Shanxi have leaflets which are usually long elliptic or ovate in
outline, with acute apexes and entire or subentire margins (Fig. 1: A, C). However, in western
Sichuan, among the populations with oblong and crenate leaflets are scattered some plants
with ovate or elliptic and entire leaflets, such as W. K. Hu & C. Ho 10650 collected from
Kangding, S. Jiang 9349 from Garzê, Sichuan Exped. 75-9811 from Xiaojin, and X. Li 78036
from Jinchuan. Unfortunately, the plants occurring in eastern Qinghai, Gansu, and Shanxi have
been misidentified as C. glauca Willd. by Ling (1980), and by Wang & Bartholomew (2001).
Such plants really show remarkable resemblance to C. glauca, and differ in their narrowly
ovate sepals. In C. glauca, the sepals are narrowly oblong in outline and often puberulous
inside.
2. Clematis tangutica (Maxim.) Korsh. in Bull. Acad. Sci. St.-Petersb. 9: 399. 1898, p.p.,
quoad nomen tantum; Andre in Rev. Hort. 1902: 528. 1902; Schneid., Ill. Handb. Laubh. 1:
294, fig. 185: W–Z. 1906; Rehd. & Wils. in Sarg., Pl. Wils. 1: 343. 1913; Rehd. in J. Arn. Arb.
4: 191. 1923; Krasch. in Kom., Fl. URSS 7: 250. 1937; Hand.-Mazz. in Acta Hort. Gotob. 13:
218. 1939; Gamayou. in Fl. Kazakhst. 4: 74. 1961; Anonymous in Iconogr. Corm. Sin. 1: 741,
fig. 1481. 1972; Anonymous in Fl. Tsinling. 1 (2): 289, fig. 248. 1974; P. P. Ling in Fl. Reip.
Pop. Sin. 28: 145, fig. 17. 1980; Grubov, Key Vasc. Pl. Mongol. 112, t. 52, fig. 238. 1982; M.
Y. Fang in Fl. Xizang. 2: 89. 1985; Liou f., Fl. Desert. Sin. 1: 483. 1985; Grey-Wilson in Kew
Bull. 44: 53, fig. 4: A–C. 1989; Y. Z. Zhao in Fl. Intramongol., ed. 2, 2: 533, pl. 215, figs. 1–3.
1990; L. Q. Li in Vasc. Pl. Hengduan Mount. 1: 523. 1993; J. G. Liou in Fl. Xinjiang. 2: 290.
1994; L. H. Zhou in Fl. Qinghai. 1: 347. 1997; M. Johnson, Klematis 352. 1997; Grubov et al.,
No. 4 WANG Wen-Tsai: A revision of Clematis sect. Meclatis (Ranunculaceae) 411
Cat. Type Specim. Centr. As. Vasc. Pl. Herb. Kom. Bot. Inst. 194. 2000; Grey-Wilson,
Clematis 171, figs.128–130. 2000; J. Q. Fu in Fl. Loess Plat. 1: 441. 2000; Grabovsk. in
Grubov, Pl. As. Centr. 12: 85. 2001; W. T. Wang & Barth. in Fl. China 6: 364. 2001.——C.
orientalis L. var. tangutica Maxim., Fl. Tangut. 3. 1889; et Enum. Pl. Mongol. 4. 1889.——C.
tibetana Kuntze ssp. tangutica (Maxim.) Brandenb., Meclatis in Clematis 169. 2000, p.p.,
excl. syn. C. akebioides (Maxim.) Veitch et C. pamiralaica Grey-Wilson. Type: China.
Qinghai (青海): Datong (大通), 1872-07, Przewalski 105 (lectotype, LE!—Grabovskaya,
2000; isolectotypes, K!, LE!, P!). Gansu (甘肃): Without precise locality, 1875-07, Piasezky
s.n. (syntype, LE!).
C. atragenoides Batalin in Acta Hort. Petrop. 11: 481. 1891. Type: China. Gansu (甘肃):
fl. Gui-dui-scha, 1890-06-21, Grum-Grshimailo 201 (syntype, LE!). Qinghai (青海): North of
Qinghai Lake, 1890-08-01, Grum-Grshimailo 228 (syntype, LE!).
C. tangutica var. obtusiuscula Rehd. & Wils. in Sarg., Pl. Wils. 1: 343. 1913; Rehd., Man.
Cult. Trees & Shrubs 229. 1927; et in J. Arn. Arb. 9: 42. 1928; Hand.-Mazz. in Acta Hort.
Gotob. 13: 218. 1939; W. T. Wang in Acta Phytotax. Sin. 31: 225. 1993; M. Johnson, Klematis
335. 1997; W. T. Wang & Barth. in Fl. China 6: 346. 2001.——C. tangutica ssp. obtusiuscula
(Rehd. & Wils.) Grey-Wilson in Kew Bull. 44: 54. 1989; et Clematis 172. 2000. Type: China.
Sichuan (四川): Kangding (康定), Tapao-shan (大炮山), alt. 2600–3300 m, 1908-07, E. H.
Wilson 2487 (holotype, GH!; isotype, LE!).
? C. chrysantha Ulbr. in Repert. Sp. Nov. Beih. 12: 374. 1922. Type: China. Sichuan (甘
肃): Kanse, Limpricht 2086 (syntype); Dêgê (德格), Limpricht 2146 (syntype).
C. orientalis auct. non L.: Kozlov in Publ. Mus. Hoangho Paiho Tien Tsin 22: 14. 1933.
甘青铁线莲
This species consists of three varieties widespread in west China and adjacent countries.
2a. var. tangutica
Woody vine, occasionally dwarf erect shrub. Branches shallowly 6–8-sulcate, puberulous,
glabrescent. Leaves 1–2-pinnate; leaflets green, papery, rhombic-ovate, narrowly ovate, or
lanceolate, 1–6×0.5–2.8 cm, apex acute or slightly obtuse, base broadly cuneate, rounded, or
subcordate, margin usually dentate or denticulate, with up to 7 teeth per side, near base
2–3-lobed or undivided, sparsely puberulous on veins on both surfaces, basal veins abaxially
nearly flat; petioles 2–6 cm long. Flower solitary, terminal, or sometimes also in axillary
1–3-flowered cymes; peduncle 0.3–3 cm long; bracts shortly petiolate, leaflet-like; pedicels
3.5–16.5 cm long, puberulous or subglabrous. Sepals 4, yellow, sometimes tinged with purple,
ascending, papery or submembranous, ovate, oblong, or lanceolate, 1.5–4×0.6–1.4 cm, apex
acuminate, long acuminate, or acute, inside glabrous, on margin velutinous, outside sparsely
puberulous. Stamens 5–11 mm long; anthers narrowly oblong, 2–3 mm long, glabrous, apex
obscurely apiculate or subobtuse. Ovaries puberulous; styles 9–15 mm long, densely villous.
Achenes rhombic-ovate, ca. 4.5×2.2 mm, puberulous; persistent styles up to 5 cm long,
plumose. Fl. Jun.–Sept.
E Afghanistan, China (Gansu, SW Nei Mongol, Ningxia, Qinghai, S Shaanxi, W Sichuan,
Xinjiang, Xizang), Kashmir Region, E Kazakhstan, Mongolia, and E Tadzhikistan. On grassy
slopes, in bushes, or on gravelly river banks; alt. 1370–4900 m.
甘青铁线莲 Fig. 4: A, B; Fig. 5: C, D
Additional specimens examined:
Afghanistan. Badakhshan, alt. 3650–4100 m, Anders 7381, 7615, 7718 (G).
China. Gansu (甘肃): Dangchang (宕昌), Hummel 5237 (S), Q. E. Yang (杨亲二) 92006 (PE); Gannan
(甘南), Qinghai-Gansu Exped. (青甘调查队) 62-3201, Q. E. Yang (杨亲二) 92033 (PE); Huining (会宁), Z.
Acta Phytotaxonomica Sinica Vol. 44 412
Y. Yu & Y. P. Xu (于兆英, 徐养鹏) 3799 (WUK); Jingtai (景泰), Z. Y. Yu & Y. P. Xu (于兆英, 徐养鹏)
3228 (WUK); Jingyuan (靖远), Z. Y. Yu & Y. P. Xu (于兆英, 徐养鹏) 3047 (WUK); Jiuquan (酒泉),
Qinghai-Gansu Exped. (青甘调查队) 60-3057 (PE); Jonê (卓尼), Rock 12917 (GH, NAS, PE), W. Y. Hsia
(夏纬瑛) 8503 (PE), Purdom 1023 (GH, US); Lanzhou (兰州), T. N. Liou (刘慎谔) 2306, Y. Q. He (何业
祺) 5956 (PE), Licent 4265 (P, TIE); Lianhua Shan (莲花山), Rock 13615 (P); Lichen, R. C. Ching (秦仁昌)
318 (GH, P, US); Lintan (临潭), C. W. Yao (姚仲吾) 1217 (NAS); Linxia (临夏), K. T. Fu (傅坤俊) 843
(PE); Min Xian (岷县), K. S. Hao (郝景盛) 544, T. P. Wang (王作宾) 5035 (PE), Hummel 3791, 5382 (PE);
Pingliang (平凉), Huanghe Exped. (黄河队) 56-1893 (NAS, PE); Shandan (山丹), Y. Q. He (何业祺) 4129
(PE); Subei (肃北), P. C. Kuo (郭本兆) 3355 (PE); Sunan (肃南), Hexi Exped. (河西队) 121, 405 (PE);
Tchapou, Licent 4098 (TIE); Tchen-tsiang, Licent 4493 (K, P, TIE); Tianzhu (天祝), Y. Q. He (何业祺) 4638
(PE); Xiahe (夏河), C. W. Yao (姚仲吾) 471, T. P. Wang (王作宾) 5758 (PE); Yongdeng (永登), Loess
Plateau Exped. (黄土高原调查队) 4767 (WUK); Yuzhong (榆中), P. G. Wu (吴培根) 651053 (NAS);
Zhangye (张掖), Anonymous 191 (PE). Ningxia (宁夏): Guyuan (固原), Huanghe River Exped. (黄河队)
56-2372 (PE); Helan Shan (贺兰山), Ningxia Inst. Drug Contr. Exped. (宁夏药检所队) 2372 (PE). Qinghai
(青海): Burhan Budai Shan (布尔汗布达山), Ladeijin 12 (LE); Da Qaidam (大柴旦), Qinghai-Gansu Exped.
(青甘调查队) 59-58 (PE); Datong (大通), Farrer & Purdom 520 (MO), K. M. Liou (刘继孟) 5930, Geobot.
Group Exped. (地植物组) 1393 (PE); Delingha (德令哈), P. C. Kuo (郭本兆) 11615 (NAS, PE); Gangca (刚
察), P. C. Kuo (郭本兆) 11235 (NAS, PE); Golmud (格尔木), Qinghai-Gansu Exped. (青甘调查队) 59-503
(PE); Gonghe (共和), Z. W. Chang (张振万) 2118 (PE), C. Y. Yao (姚仲吾) 879 (NAS); Guide (贵德), S. W.
Liu (刘尚武) 3324 (PE); Guinan (贵南), S. J. Ma (马世骏) 33 (PE); Haiyan (海晏), P. C. Tsoong (钟补求)
8311 (PE); Huangyuan (湟源), S. X. Zheng (郑斯绪) 425 (PE); Huzhu (互助), P. C. Kuo (郭本兆) 9231 (PE);
Ledu (乐都), W. W. Hou (侯无危) s.n. (PE); Le Hargue gol, Licent 4829 (TIE); Menyuan (门源),
Qinghai-Gansu Exped. (青甘调查队) 60-2718 (PE); Minhe (民和), T. N. Ho (何廷农) 581 (PE); Nangqên (囊
谦), Y. C. Yang (杨永昌) 1145, 1554 (PE); Nomhon (诺木洪), Y. X. Liou (刘媖心) 78, C. Y. Wu et al. (吴征
镒等) 75-224 (PE); E Qaidam basin, Ladeijin 159 (LE); Qinghai Lake (青海湖), Ridley 1 (K), Rock 13277
(LE, NAS, S), K. F. Luo (罗开富) 10 (PE); Tianjun (天俊), Qinghai-Gansu Exped. (青甘调查队) 59-1080
(PE); Tongde (同德), T. N. Ho (何廷农) Bartholomev 72, 134 (GH); Ulan (乌兰), P. C. Kuo (郭本兆) 11484
(PE); Xiangride (香日德), Qinghai-Gansu Exped. (青甘调查队) 59-1296 (PE); Xinghai (兴海), T. P. Wang
(王作宾) 19593 (WUK), T. N. Ho (何廷农) 64, 465 (NAS); Xining (西宁), S. H. Zheng (郑斯绪) 368
(IBSC), Y. R. Ling (林有润) 74-1451 (PE); Yushu (玉树), C. W. Yao (姚仲吾) 754 (PE), 755 (NAS).
Shaanxi (陕西): Taibai Shan (太白山), K. T. Fu (傅坤俊) 1518 (NAS). Sichuan (四川): Aba (阿坝), S. Jiang
(姜恕) 1196 (PE); Baiyu (白玉), W. L. Chen et al. (陈伟烈等) 6827 (PE); Barkam (马尔康), X. Li (李馨)
74740 (IBSC, NAS, PE); Daocheng (稻城), S. Jiang (姜恕) 5354 (PE); Dêgê (德格), Sichuan Exped. (四川
队) 74-7031 (PE); Garzê (甘孜), P. C. Kuo (郭本兆) 21094 (NAS); Gongga Shan (贡嘎山), Rock 17616,
17699 (PE); Hongyuan (红原), Q. S. Zhao (赵清盛) 83 (K, GH, PE, S, UPS); Kangding (康定), Pratt 237,
622, Soulié 2771 (K), T. T. Yu (俞德浚) 992, K. C. Kuan et al. (关克俭等) 36231 (PE); Li Xian (理县), Q. S.
Zhao (赵清盛) 36 (K, GH, PE, S, UPS); Sêrxü (石渠), S. Jiang (姜恕) 9275 (PE); Shuajingsi (刷经寺), X. Li
(李馨) 74654, 74876, 74929 (IBSC, PE); Songpan (松潘), H. Smith 3663 (S, UPS), Beresowsky s.n. (LE), K.
T. Fu (傅坤俊) 1858 (PE); Zoigê (若尔盖), Sichuan Exped. (四川队) 75-10290 (IBSC), 75-10377 (IBSC,
PE). Xinjiang (新疆): Akto (阿克陶), Inst. Northwest Bot. Exped. (西北植物所队) 746 (PE); Barkol (巴里
坤), Q. R. Wang (王庆瑞) 4481 (PE); Hejing (和静), Q. R. Wang (王庆瑞) 3796 (PE); Hoxud (和硕), A. J. Li
& J. N. Zhu (李安仁, 朱家柟) 6986 (PE); Kunlun Shan (昆仑山), Serpuchov 43, 563, Younatov & Y. F.
Yang 230 (LE); Tian Shan (天山), T. Y. Chou et al. (周太炎等) 651317 (IBSC, PE), Merzbacher 1022 (LE),
Brocherel 162 (G); Toksun (托克逊), A. J. Li & J. N. Zhu (李安仁, 朱家柟) 7383 (PE); Turpan (吐鲁番), A.
J. Li & J. N. Zhu (李安仁, 朱家柟) 5734 (PE). Xizang (西藏): Biru (比如), D. D. Tao (陶德定) 11316 (PE);
Gar (噶尔), Qinghai-Xizang Exped. (青藏队) 76-8316, 76-8587 (PE); Gêrzê (改则), F. Z. Li (李法曾) 23
(PE); Gonjo (贡觉), Qinghai-Xizang Exped. (青藏队) 76-12581 (PE); Qamdo (昌都), Qinghai-Xizang Exped.
(青藏队) 76-12694 (PE); Riwoqê (类乌齐), Qinghai-Xizang Exped. (青藏队) 76-12943 (PE); Sog Xian (索
县), D. D. Tao (陶德定) 10894 (PE); Xainza (申扎), D. D. Tao (陶德定) 10712 (PE).
Kashmir Region. Rupshu, Koelz 22886 (GH, LE, S).
No. 4 WANG Wen-Tsai: A revision of Clematis sect. Meclatis (Ranunculaceae) 413
Mongolia. Bogd Somon, Norlindh 10221 (S); South-Gobi: Aimak, Norlindh 10303 (S); Ulaanbaatar,
Lavrenko 19153, Kalinina s.n. (LE); Urga, Krascheninikov 24 (LE); Urzu, Klementz s.n. (LE).
Tadzhikistan. Bai-Kara River, Brohesisky s.n. (LE); Central Pamir: Murgal Valley, Tolmacheva 4367
(G, S, US); Pamir, Fedtschenko s.n. (MO).
Of the specimens cited above, the vast majority are woody vines, and only two, C. Y. Yang
1145 and Geobot. Group Exped. 1393, collected from Qinghai Province, China, are dwarf
erect shrubs. The dwarf shrubby habit appears to be derived from scandent habit due to
adaptation to adverse environmental conditions. In sect. Meclatis, the phenomenon of
derivation from scandent habit to erect habit also occurs in C. tibetana, an ally of C. tangutica.
In the latter case, the dwarf suffrutescent form with sepals puberulous outside represents one
of the varieties of C. tibetana (var. pamiralaica), and might be derived from the woody
scandent variety tibetana with sepals glabrous outside (see below) (Fig. 3)
2b. var. mongolica (Grey-Wilson) W. T. Wang, st. nov.——C. tangutica (Maxim.) Korsh. ssp.
mongolica Grey-Wilson in Kew Bull. 44: 54, fig. 1: W. 1989; M. Johnson, Klematis 355.
1997; Grey-Wilson, Clematis 173. 2000, syn. nov. Type: Mongolia, Tula River, ca. 25 km NE
of Ulaanbaatar, alt. ca. 1450 m, 1970-07-30, Jeffrey 1436 (holotype, K!; isotype, PE!).
C. tibetana Kuntze ssp. tangutica (Maxim.) Brandenb., Meclatis in Clematis 169. 2000,
p.p., quoad syn. C. tangutica ssp. mongolica Grey-Wilson.
多齿甘青铁线莲 Fig. 4: E
This variety differs from var. tangutica in its narrowly lanceolate, long attenuate leaflets
with (5–)8–13 teeth per side.
Woody vine. Leaflets 2.2–5.2×0.6–1.1 cm. Sepals submembranous or thinly papery,
broadly ovate, inside glabrous, apex long acuminate. Fl. Jul. –Aug.
China (C & S Gansu) and N Mongolia. In bushes on slopes; alt. 1450–3100 m.
Additional specimens examined:
China. Gansu (甘肃): in valle Drakana (near Jonê (卓尼)), Hummel 4637 (S); Tianzhu (天祝), X. Z.
Han (韩学俊) 621127 (NAS).
Mongolia. Ulaanbaatar, Gorbunova 4, Ikonnikov-Galtzky 384 (LE); Urga to Lake Ihe-tuhumnol,
Krascheninikov 4 (GH).
The formation of the scattered distribution areas of var. mongolica and var. pubescens
(see below) (Fig. 2) might be caused by the strong impact of the Quaternary glaciers (Wang,
1989).
2c. var. pubescens M. C. Chang & P. P. Li in Fl. Reip. Pop. Sin. 28: 356. 1980; M. Y. Fang in
Fl. Xizang. 2: 89. 1985; L. Q. Li in Vasc. Pl. Hengduan Mount. 1: 523. 1993; L. H. Zhou in Fl.
Qinghai. 1: 347. 1997; W. T. Wang in Acta Phytotax. Sin. 38: 327. 2000; W. T. Wang & Barth.
in Fl. China 6: 364. 2001. Type: China. Xizang (西藏): Zhag’yab (察雅), 1976-09-20,
Qinghai-Xizang Exped. (青藏队) 76-13014 (holotype, PE!).
C. tibetana Kuntze ssp. vernayi (C. E. C. Fisch.) Grey-Wilson var. dentata Grey-Wilson
in Kew Bull. 44: 48, fig. 1: Z. 1989; M. Johnson, Klematis 356. 1997; Grey-Wilson, Clematis
170. 2000. Type: China. Gansu (甘肃): Radja, alt. 3300 m, 1926-06, Rock 14124 (holotype, K;
isotypes, GH!, NAS!, P!, S!).
C. tibetica Kuntze ssp. vernayi auct. non (C. E. C. Fisch.) Grey-Wilson: Brandenb.,
Meclatis in Clematis 175. 2000, p.p., quoad syn. C. tibetana ssp. vernayi var. dentata
Grey-Wilson et C. tangutica var. pubescens M. C. Chang & P. P. Ling.
毛萼甘青铁线莲 Fig. 4: C, D
This variety differs from var. tangutica only in its sepals puberulous inside.
Woody vine. Fl. Jun.–Sept.
China (SW Gansu, S Qinghai, W Sichuan, SW Xinjiang, E Xizang) and Mongolia. On
grassy slopes or gravelly river banks, or in bushes; alt. 3100–3600 m.

Acta Phytotaxonomica Sinica Vol. 44 414


Fig. 4. Clematis tangutica (Maxim.) Korsh. A, B, var. tangutica. A, flowering branch; B, stamen (from D. D. Tao
10894). C, D, var. pubescens M. C. Chang & P. P. Ling. C, flowering branch; D, stamen (from Anonymous 519); E, var.
mongolica (Grey-Wilson) W. T. Wang leaf (from Hummel 4637).
No. 4 WANG Wen-Tsai: A revision of Clematis sect. Meclatis (Ranunculaceae) 415


Fig. 5. A, B, Clematis tibetana Kuntze var. pamiralaica (Grey-Wilson) W. T. Wang. A, habit; B, stamen (from
Tolmatcheva 4367). C, D, C. tangutica (Maxim.) Korsh. var. tangutica. C, habit; D, stamen (from Y. C. Yang 1145).
Acta Phytotaxonomica Sinica Vol. 44 416
Additional specimens examined:
China. Qinghai (青海): Yushu (玉树), Anonymous 519 (PE). Sichuan (四川): Dêgê (德格), S. X. Jia
(贾慎修) s.n., Y. W. Tsui (崔友文) 5146 (PE); Garzê (甘孜), Q. S. Zhao et al. (赵清盛等) 111219 (PE);
Hongyuan (红原), Longriba (龙日坝), C. L. Wu (吴中伦) 32918 (PE); Queer Shan (雀儿山), Y. T. Zhang &
K. Y. Lang (张永田, 郎楷永) 147 (PE); Yajiang (雅江), Y. W. Tsui (崔友文) 4327 (PE). Xinjiang (新疆):
Wuqia (乌恰), Inst. Northwest Bot. Exped. (西北植物所队) 1849, 2073 (PE). Xizang (西藏): Jomda (江达),
Qing hai-Xizang Exped. (青藏队) 76-12463 (PE); Qamdo (昌都), Qinghai-Xizang Exped. (青藏队) 73-70
(PE).
Mongolia. Near Sandensu, Gusev 121 (S).
While var. pubescens, published in 1980, and var. dentata, published in 1989, are proved
to represent the same taxon, the earlier name, var. pubescens, has priority at the varietal rank
regardless of that this taxon is treated as a variety of C. tangutica or of C. tibetana ssp.
vernayi, according to Article 11 of the International Code of Botanical Nomenclature (Greuter
et al., 2000).
3. Clematis intricata Bunge in Mém. Sav. Etrang. Acad. Sci. St.-Petersb. 2: 75. 1833;
Maxim., Prim. Fl. Amur. Suppl. 1. 1858; Kom. in Acta Hort. Petrop. 22: 288. 1903; Kitag.,
Lineam. Fl. Mansh. 217. 1939; Anonymous in Iconogr. Corm. Sin. 1: 740, fig. 1480. 1972;
Anonymous in Fl. Tsinling. 1 (2): 290. 1974; Liou et al., Fl. Pl. Herb. Chinae Bor.-Or. 3: 173,
pl. 75, fig. 5. 1975; P. P. Ling in Fl. Reip. Pop. Sin. 28: 142, fig. 15. 1980; Grubov, Key Vasc.
Pl. Mongol. 112, t. 52, fig. 237. 1982; He, Fl. Beijing, rev. ed., 1: 250. 1984; J. W. Wang in Fl.
Hebei. 1: 474. 1986; Grey-Wilson in Kew Bull. 44: 45. 1989; Y. Z. Zhao in Fl. Intramongol.,
ed. 2, 2: 531. 1990; Y. J. Ling et al. in Fl. Shanxi. 1: 633. 1992; L. H. Zhou in Fl. Qinghai. 1:
346. 1997; M. Johnson, Klematis 337. 1997; Grey-Wilson, Clematis 167, fig. 121. 2000; J. Q.
Fu in Fl. Loess Plat. 1: 440. 2000; Brandenb., Meclatis in Clematis 151. 2000, p.p., excl. syn.
C. hilariae Koval., C. sarezica Ikonn. etc.; Grabovsk. in Grubov, Pl. As. Centr. 12: 81. 2001;
W. T. Wang & Barth. in Fl. China 6: 367. 2001.——C. orientalis L. ssp. thunbergii (Steud.)
Kuntze var. intricata (Bunge) Kuntze in Verh. Bot. Ver. Brand. 26: 125. 1885, p.p., excl. pl.
Himal.——C. orientalis L. var. intricata (Bunge) Maxim. in Bull. Acad. Sci. St.-Petersb. 22:
210. 1877; et in Mél. Biol. 9: 583. 1877; Forbes & Hemsl. in J. Linn. Soc. Bot. 23: 6. 1886;
Maxim., Fl. Tangut. 3. 1889; Enum. Pl. Mongol. 4. 1889; et in Acta Hort. Petrop. 11: 5. 1890.
Type: China borealis, 1831, Bunge s.n. (holotype, LE!; isotypes, G!, P!, PE!).
C. orientalis auct. non L.: Franch., Pl. David. 1: 12. 1884.
C. glauca Willd. var. angustifolia auct. non Ledeb.: Rehd. in J. Arn. Arb. 4: 190. 1923;
Hand.-Mazz. in Acta Hort. Gotob. 13: 217. 1939; Anonymous in Fl. Beijing 1: 305. 1962.
C. orientalis L. var. aethusaefolia Kozlov in Publ. Mus. Hoangho Paiho Tien Tsin 22: 14.
1933, nom. illegit., non C. aethusaefolia Turcz.
黄花铁线莲
This species consists of three varieties widespread on the Loess plateau and adjacent
provinces of China and extending northward to the desert regions of Mongolia.
3a. var. intricata Fig. 3: D, E
Woody vine. Branches shallowly 6–8-sulcate, sparsely puberulous or subglabrous.
Leaves 1–2-pinnate; leaflets often grey-green, papery, lanceolate, linear-lanceolate or linear,
1–4×0.2–2 cm, apex attenuate, base cuneate, margin entire or 1–2-denticulate, undivided or
2–3-lobed, on both surfaces very sparsely puberulous, glabrescent, midrib flat; petioles
1.6–5.5 cm long. Cymes axillary, 1–3(–5)-flowered; peduncles 0.1–3 cm long; bracts petiolate,
lanceolate, undivided or 2–3-lobed. Flower 2–4 cm in diam.; pedicel 2–3.8 cm long, sparsely
puberulous or subglabrous. Sepals 4, yellow, ascending, papery, narrowly ovate, ovate,
suboblong, or lanceolate, 1.2–2.3(–2.5)×0.5–0.8(–1) cm, apex acuminate, on both surfaces
glabrous, occasionally outside above very sparsely puberulous, on margin velutinous. Stamens
5–9.5 mm long; anthers narrowly oblong or linear, 2.5–4 mm long, glabrous, apex obtuse.
No. 4 WANG Wen-Tsai: A revision of Clematis sect. Meclatis (Ranunculaceae) 417
Ovaries puberulous; styles 8–10 mm long, densely villous. Achenes elliptic or rhombic-ovate,
2.5–3.2×2–2.5 mm, puberulous, margin rimmed; persistent styles 2.5–4 cm long, plumose.
Fl. Jun.–Sept.
China (Beijing, Gansu, Hebei, Henan, W Liaoning, Nei Mongol, Ningxia, E Qinghai, N
Shaanxi, Shanxi, W Sichuan) and Mongolia. On slopes or sandy hills, in sandy places, in
bushes, or by streams; alt. 500–2600(–3200) m.
Additional specimens examined:
China. Beijing (北京): David 399 (K, LE, P), 2904 (P), Bretschneider 24 (K, LE), Tatarinow s.n. (LE),
K. K. Tsoong (钟观光) s.n., T. N. Liou (刘慎谔) s.n., T. F. King (金德福) 346 (PE). Gansu (甘肃): Dingxi
(定西), Huanghe Exped. (黄河队) 56-10129 (PE); Huan Xian (环县), C. Hou & P. L. Yang (侯昌, 杨平礼)
154 (PE); Huining (会宁), Huanghe Exped. (黄河队) 56-5800 (PE); Jingyuan (靖远), Licent 6158 (K);
Minqin (民勤), Hexi Exped. (河西队) 2022 (PE); Min Xian (岷县), T. P. Wang (王作宾) 4990 (PE);
Pingliang (平凉), Z. Y. Li (李宗英) 115 (PE); Sunan (肃南), Qinghai-Gansu Exped. (青甘调查队) 60-3292
(PE); Tianzhu (天祝), Y. Q. He (何业祺) 4395 (PE); Wuwei (武威), P. C. Kuo (郭本兆) 3520 (NAS);
Yongchang (永昌), F. M. Xu s.n. (PE); Yuzhong (榆中), P. G. Wu (吴培根) 651043 (NAS); Zhenyuan (镇
原), T. P. Wang (王作宾) 17246 (IBSC, PE); Zhugqu (舟曲), W. Y. Hsia (夏纬瑛) 6411 (PE). Hebei (河北):
Dongling (东陵), Licent 9266 (TIE); Fuping (阜平), K. M. Liou (刘继孟) 3279 (NAS, PE); Huailai (怀来),
Meyer 1047 (K), T. F. King (金德福) 257 (PE, S); Laiyuan (涞源), K. M. Liou (刘继孟) 2573, 2820 (PE);
Neiqiu (内丘), Y. Liou (刘瑛) 13008 (NAS, PE); Pingshan (平山), Chanet 671 (TIE); Xiaowutai Shan (小五
台山), Hancock s.n. (LE), Limpricht 501 (S), H. Smith 297 (S, UPS), H. W. Kung (孔宪武) 1118, T. P. Wang
(王作宾) 617 (PE); Xingtai (邢台), H. F. Chow (周汉藩) 43371 (PE); Xuanhua (宣化), Licent 9711 (G);
Zhangbei (张北), X. L. Huang et al. (黄秀兰等) 2494, 3481 (PE); Zhangjiakou (张家口), Y. W. Tsui (崔友
文) 447 (PE); Zhuolu (涿鹿), C. G. Yang (杨朝广) 1149 (PE). Henan (河南): Yiyang (宜阳), Henan Exped.
(河南队) 59-6351 (IBSC, PE). Liaoning (辽宁): Lingyuan (凌源), C. S. Wang (王崇书) 3022 (IBSC). Nei
Mongol (内蒙古): Dengkou (磴口), T. P. Wang (王作宾) 2404 (PE); Fengzhen (丰镇), Y. W. Tsui (崔友文)
926 (PE); Helan Shan (贺兰山), Przewalski 243 (LE), Y. Q. Ma (马毓泉) 265 (HIMC); Hohhot (呼和浩特),
Y. Q. Ma (马毓泉) s.n. (HIMC); Liangcheng (凉城), Y. Q. Ma (马毓泉) 50 (HIMC); Ordos (鄂尔多斯),
Przewalski s.n. (K, LE), Licent 6920 (K, TIE), 6851, 6873, 6880, 6932 (TIE), W. Y. Hsia (夏纬瑛) 3769
(PE); Urad Qianqi (乌拉特前旗), X. Z. Lang (郎学忠) 323 (PE); Zhuozi (卓资), Y. Q. Ma & Q. R. Wu (马毓
泉, 吴庆茹) 188 (HIMC). Ningxia (宁夏): Guyuan (固原), Huanghe Exped. (黄河队) 56-2356 (PE);
Haiyuan (海原), Huanghe Exped. (黄河队) 56-5501 (PE); Helan Shan (贺兰山), Y. Y. Pai (白银元) 173
(PE); Pingluo (平罗), Z. W. Zhang (张振万) 391 (PE); Tongxin (同心), Huanghe Exped. (黄河队) 56-8666
(PE); Zhongning (中宁), Z. W. Zhang (张振万) 306 (PE). Qinghai (青海): Datong (大通), Przewalski s.n.
(LE), K. M. Liou (刘继孟) 6504 (PE); Guide (贵德), T. N. Ho (何廷农) 1027 (PE); Huangyuan (湟源), P. C.
Tsoong (钟补求) 8921 (PE); Huzhu (互助), K. M. Liou (刘继孟) 5881 (PE); Minhe (民和), T. N. Ho (何廷
农) 715 (PE); Qilian (祁连), P. C. Kuo (郭本兆) 12568 (PE); Regio Tangut, Przewalski s.n. (LE, S); Xining
(西宁), K. M. Liou (刘继孟) 5904 (PE). Shaanxi (陕西): Hengshan (横山), K. T. Fu (傅坤俊) 7105 (IBSC,
PE); Huanglong (黄龙), K. T. Fu (傅坤俊) 3097 (NAS, PE); Jingbian (靖边), K. T. Fu (傅坤俊) 7390 (IBSC,
PE); Suide (绥德), K. T. Fu (傅坤俊) 6748 (PE); Wuqi (吴旗), Huanghe Exped. (黄河队) 56-8253 (PE);
Yulin (榆林), Huanghe Exped. (黄河队) 56-7010 (PE). Shanxi (山西): Datong (大同), Licent 200 (TIE),
Shanxi Exped. (山西考察队) 53-770 (PE); Fenyang (汾阳), K. M. Liou (刘继孟) 2724 (PE); Jiaocheng (交
城), Licent 2163 (IBSC, K); Jiexiu (介休), K. M. Liou (刘继孟) 1340 (PE); Lingchuan (凌川), K. M. Liou
(刘继孟) 7488, 7513 (PE); Lishi (离石), Y. W. Tsui (崔友文) 10310 (NAS); Taiyuan (太原), Licent 10857
(G), Limpricht 680 (S), Yabe s.n. (NAS); Wutai Shan (五台山), Hancock 10, Licent 2036, K. M. Liou (刘继
孟) 2643 (K), T. Tang (唐进) 1053 (NAS, PE), K. C. Kuan & Y. L. Chen (关克俭, 陈艺林) 2239 (PE); Xi
Xian (隰县), T. P. Wang (王作宾) 3156 (PE). Sichuan (四川): Kangding (康定), Zhonggu (中谷), K. C.
Kuan et al. (关克俭等) 389 (PE).
Mongolia. Chalcha, Lisovski s.n. (LE); Gobi, Glajolev 194 (S); Hara Usu, Mashalseba 20 (LE); Lake
Nehaituhim-nor, Polynov & Lebedev 253 (LE); Shabarakh Usu, R. W. Chaney 583 (LE).
3b. var. intrapuberula W. T. Wang, var. nov. Type: China. Gansu (甘肃): Heshui (合水),
Acta Phytotaxonomica Sinica Vol. 44 418
Jiajia Valley (贾家沟), in shady places near road, woody vine, flowers yellow, 1954-07-07,
Huanghe Exped. (黄河队) 54-461 (holotype, PE; isotype, IBSC).
毛萼黄花铁线莲
A var. intricata differt sepalis intus puberulis.
This variety differs from var. intricata by having sepals puberulous inside.
China (E Gansu).
3c. var. purpurea Y. Z. Zhao in Fl. Intramongol. 2: 242, 369, pl. 125, fig. 11. 1978; P. P. Ling
in Fl. Reip. Pop. Sin. 28: 142. 1980; Y. Z. Zhao in Fl. Intramongol., ed. 2, 2: 533. 1990; W. T.
Wang & Barth. in Fl. China 6: 367. 2001. Type: China. Nei Mongol (内蒙古): Daqing Shan
(大青山), Halaqingou (哈拉沁沟), in Betula forest, 1964-09-06, Y. Q. Ma (马毓泉) 28
(holotype, not seen).
紫萼铁线莲
This variety differs from var. intricata by having lanceolate terminal leaflets and elliptic
lateral leaflets, and purple sepals. Fl. Jul.–Sept.
China (SW Nei Mongol). In Betula forests.
An unclear form:
Clematis glauca Willd. var. akebioides (Maxim.) Rehd. & Wils. f. phaeantha Rehd. in J.
Arn. Arb. 1: 195. 1920; Hand.-Mazz. in Acta Hort. Gotob. 13: 218. 1939; Rehd., Man. Cult.
Trees & Shrubs, ed. 2, 219. 1951; Grey-Wilson in Kew Bull. 44: 46. 1989; et Clematis 167.
2000.——C. glauca Willd. var. phaeantha (Rehd.) Rehd., Man. Cult. Trees & Shrubs 229.
1927.——C. intricata Bunge var. phaeantha (Rehd.) M. Johnson, Klematis 840. 1997, p.p.,
excl. Hummel 4137. Type: Plant cultivated in Highland Park, Rochester, New York, U.S.A.,
1918-09-20, W. L. J. Edson s.n. (holotype, GH!).
This form, described from a cultivated plant, is similar to C. intricata in the lanceolate or
linear-lanceolate leaflets and ascending lanceolate sepals, but differs mainly by having “dull
violet” (Rehder, 1920, l.c.) sepals. In the protologue, Rehder (1920) pointed out that “of the
origin of this form nothing is known to” him. After its publication, forma phaeantha was
firstly mentioned by Handel-Mazzetti (1939) in the enumeration of the Chinese species of the
Clematis, but with no specimen being cited for this form and its origin not indicated.
Grey-Wilson (1989, 2000) reduced C. intricata var. purpurea to the synonymy under forma
phaeantha, for which no specimen was cited and the origin was not indicated either. However,
the former with lanceolate terminal leaflets and elliptic lateral leaflets is different from the
latter. Johnson (1997) reported the occurrence of forma phaeantha in Gansu Province, China.
However, the specimen, Hummel 4137, on which his record was based, is in fact C.
akebioides, in which the leaflets are oblong, narrowly ovate, or elliptic in outline, and the
sepals are yellow in colour. In the two books entitled Meclatis in Clematis by Brandenburg
(2000) and An Illustrated Encyclopedia of Clematis by Toomey & Leeds (2001), forma
phaeantha was not included. So, this curious cultivar is only known from the one type
gathering, and its origin still remains unclear.
4. Clematis tibetana Kuntze in Verh. Bot. Ver. Brand. 26: 172. 1885; Grey-Wilson in Kew
Bull. 44: 46. 1989; Naithani, Flow. Pl. India, Nepal & Bhutan 9. 1990; Rau in Sharma et al.,
Fl. Ind. 1: 78. 1993; M. Johnson, Klematis 356. 1997; Grey-Wilson, Clematis 168. 2000;
Brandenb., Meclatis in Clematis 164. 2000, p.p., excl. syn. C. akebioides (Maxim.) Veitch et
C. ladakhiana Grey-Wilson; W. T. Wang & Barth. in Fl. China 6: 365. 2001. Type: India.
Kumaon, Strachey & Winterbottom s.n. (holotype, K; isotype, BM!).
中印铁线莲
This species consists of four varieties occurring in the western Himalayas, the Pamirs and
adjacent mountains.

No. 4 WANG Wen-Tsai: A revision of Clematis sect. Meclatis (Ranunculaceae) 419
4a. var. tibetana Fig. 6: A, B
Woody vine. Branches shallowly 6–10-sulcate, sparsely puberulous, often glabrescent.
Leaves 1–2-pinnate; leaflets green, papery, broadly lanceolate, narrowly ovate, ovate, or
elliptic, 1–4.2×0.5–2.4 cm, apex acute or attenuate, base broadly cuneate or rounded, margin
usually entire, undivided or near base 2–3-lobed, sparsely puberulous on both surfaces or
adaxially subglabrous, basal veins abaxially nearly flat; petioles 1.2–6 cm long. Flowers
solitary, terminal, or also in axillary 1–3-flowered cymes, 2–4.5 cm in diam.; peduncles
0.3–7(–10) cm long; bracts foliaceous or simple, 3-sect; pedicels 2–15(–20) cm long,
puberulous or glabrous. Sepals 4, yellow, yellow-brown or brown-purple, ascending, thickly
papery, narrowly ovate or lanceolate, 2–2.8(–3.5)×0.6–1.1(–1.4) cm, apex long acuminate,
inside puberulous, outside glabrous. Stamens 7–12 mm long; anthers oblong, 1.6–2 mm long,
glabrous, apex obtuse or minutely apiculate. Ovaries puberulous; styles 12–15 mm long,
densely villous. Fl. May–Jul.
China (SW Xinjiang, W Xizang), N India, Kashmir Region, and Nepal. In bushes or on
dry slopes; alt. 2800–5000 m.
Additional specimens examined:
China. Xinjiang (新疆): Pamir (帕米尔), Jarkand, C. Persson 19, Norstedt 4 (S); Taxkorgan (塔什库尔
干), Inst. Northwest Bot. Exped. (西北植物所队) 893 (PE); Wuqia (乌恰), Inst. Northwest Bot. Exped. (西北
植物所队) 1849, 2073 (PE, WUK). Xizang (西藏): Pulan (普兰), Qinghai-Xizang Exped. (青藏队) 76-8422,
76-13271 (PE); Without precise locality, J. D. Hooker & Thomson 1077 (P).
India. Garhwal, Osmaston 745 (GH); Mulapa Gadh, Duthie 5246 (G).
Kashmir Region. Ladak, Schlagintweit s.n., Thuolou 540 (BM).
Nepal. Barbung Khola, Kakkotgon, Polunin, Sykes & Williams 1086 (S, US); Marsidandi Valley,
Lowndes 1078 (G); Muktinath, Stainton et al. 1395 (GH), 5646 (G); Ringmo, Shrestha 5315 (US).
4b. var. vernayi (C. E. C. Fisch.) W. T. Wang in Acta Phytotax. Sin. 36: 164. 1998; W. T.
Wang & Barth. in Fl. China 6: 365. 2001.——C. vernayi C. E. C. Fisch. in Bull. Misc. Inform.
Kew 1937: 95. 1937; Gupta in Bull. Nat. Bot. Gard. Lucknow 80: pl. 45. 1963.——C.
tibetana Kuntze ssp. vernayi (C. E. C. Fisch.) Grey-Wilson in Kew Bull. 44: 47. 1989; M.
Johnson, Klematis 356. 1997; Grey-Wilson, Clematis 169, figs. 122–126. 2000; Brandenb.,
Meclatis in Clematis 175. 2000, p.p., excl. syn. C. tangutica var. pubescens M. C. Chang & P.
P. Ling et C. tibetana ssp. vernayi var. dentata Grey-Wilson et var. laciniifolia Grey-Wilson.
Type: China. Xizang: 12 ml NW of Gyantze, alt. 4150 m, Cutting & Vernay 57 (holotype, K;
isotype, GH!).
C. chrysantha Ulbr. var. brevipes Tamura in Acta Phytotax. Geobot. 23: 30. 1968. Type:
Nepal. Pijehl, 1958-09, Namikawa 124 (holotype, KYO!).
?C. orientalis L. var. uniflora Tamura in Acta Phytotax. Geobot. 37: 156. 1986. Type: Nepal.
Mustang Distr.: near Muktinath, alt. 3300 m, 1976-10-11, Tabata et al. 9269 (holotype, KYO).
C. tibetana Kuntze var. lineariloba W. T. Wang in Acta Phytotax. Sin. 36: 164. 1998; W.
T. Wang & Barth. in Fl. China 6: 365. 2001. Type: China. Xizang (西藏): Nyingchi (林芝),
Nixi, 1965-07-29, Y. T. Zhang & K. Y. Lang (张永田, 郎楷永) 1089 (holotype, PE!).
C. tibetana auct. non Kuntze: Hara in Hara & Williams, Enum. Flow. Pl. Nepal 2: 16.
1979.
C. tenuifolia auct. non Royal: Ling in Fl. Reip. Pop. Sin. 28: 140, pl. 41. 1980; M. Y.
Fang in Fl. Xizang. 2: 90. 1985.
厚萼中印铁线莲 Fig. 6: C, D
This variety differs from var. tibetana in its sepals often thicker, thickly papery or
coriaceous, broadly ovate, ovate, or oblong in outline, with acute apexes, and in its longer
anthers (2.4–3.5 mm).
Woody vine. Leaflets or their terminal lobes lanceolate, narrowly ovate, or linear, margin
entire or subentire. Fl. May–Sept.
Acta Phytotaxonomica Sinica Vol. 44 420


Fig. 6. A, B, Clematis tibetana Kuntze var. tibetana. A, flowering branch; B, stamen. Drawn from Shrestha 5315. C, D,
C. tibetana var. vernayi (C. E. C. Fisch.) W. T. Wang. C, flowering branch; D, stamen. Drawn from Qinghai-Xizang Exped.
76-7153. E, F, C. ladakhiana Grey-Wilson. E, flowering branch; F, stamen. Drawn from Thomson s.n.
No. 4 WANG Wen-Tsai: A revision of Clematis sect. Meclatis (Ranunculaceae) 421
China (SW Xinjiang, S Xizang), Kashmir Region, and W & N Nepal. In bushes, on
slopes or on gravelly river banks; alt. 1850–4800 m.
Additional specimens examined:
China. Xinjiang (新疆): Jarkand-Serek-kol, Norstedt 4 (S). Xizang (西藏): Bainang (白朗), J. W.
Zhang (张经纬) 2405 (PE); Bomi (波密), T. S. Ying & D. Y. Hong (应俊生, 洪德元) 1057 (PE); Changphu
Chu, Wager 228 (K); Gyaca (加查), Qinghai-Xizang Exped. (青藏队) 75-656 (PE); Gyangzê (江孜), Ludlow
75, Walton 128 (BM); Gyirong (吉隆), Qinghai-Xizang Exped. (青藏队) 76-7153 (PE); Khambajary,
Younghusband 70 (G, P); Kongbo: Pasum Tso, Ludlow et al. 13928 (BM); Lhasa (拉萨), Dungbou s.n. (K),
Y. T. Zhang & K. Y. Lang (张永田, 郎楷永) 2662 (PE); Mainling (米林), Med. Pl. Exped. (药植队) 72-4321
(PE); Namling (南木林), Med. Pl. Exped. (中草药队) 72-850 (PE); Nang Xian (郎县), Med. Pl. Exped. (中草
药队) 72-4246 (PE); Ngamring (昂仁), Z. W. Zhang (张振万) 2701 (PE); Nyalam (聂拉木), Y. T. Zhang &
K. Y. Lang (张永田, 郎楷永) 4152 (PE); Nye, Ludlow & Sheriff 1944 (BM); Nyang Chu, Ludlow, Sherriff
& Taylor 6199 (BM, G); Nyingchi (林芝), Y. T. Zhang & K. Y. Lang (张永田, 郎楷永) 1002 (PE); Qushui
(曲水), G. X. Fu (傅国勋) 71 (PE); Rinbung (仁布), Xizang Exped. (西藏队) 61-1519 (PE); Sa’gya (萨迦),
Xizang Exped (西藏队) 61-1647 (PE); Tingri (定日), Y. T. Zhang & K. Y. Lang (张永田, 郎楷永) 3711
(PE); Zayü (察隅), Z. C. Ni et al. (倪志诚等) 1047 (PE); Zogang (左贡), Qinghai-Xizang Exped. (青藏队)
76-12047 (PE).
Kashmir Region. Purig, Koelz 6071 (GH).
Nepal. Barbung Khola, Grey-Wilson & Philips 751 (K); Kali Kandaki, Lange 34 (K); Muktinath, 12000
ft., Stainton, Sykes & Williams 1395 (GH); Mt. Everest, Hingston 48 (K); Shimen, Grey-Wilson & Philips
534 (K).
4c. var. pamiralaica (Grey-Wilson) W. T. Wang, st. nov.——C. pamiralaica Grey-Wilson in
Kew Bull. 44: 54, fig. 5. 1989; M. Johnson, Klematis 347. 1997; W. T. Wang in Acta Phytotax.
Sin. 36: 165. 1998; Grey-Wilson, Clematis 171. 2000; Grabovsk. in Grubov, Pl. As. Centr. 12:
83. 2001; W. T. Wang & Barth. in Fl. China 6: 365. 2001, syn. nov. Type: Tadzhikistan. Central
Pamir, Murghäb, alt. 3350 m, 1958-06-29, Tolmacheva 4367 (holotype, K!).
C. tangutica (Maxim.) Korsh. in Bull. Acad. Sci. St.-Petersb. 9: 399. 1898, p.p. quoad
specim. cit.; Krasch. in Kom., Fl. URSS 7: 322. 1937, p.p.
C. tibetana Kuntze ssp. tangutica (Maxim.) Brandenb., Meclatis in Clematis 169. 2000,
p.p. quoad syn. C. pamiralaica Grey-Wilson.
帕米尔铁线莲 Fig. 5: A, B
This variety differs from var. tibetana in its dwarf suffrutescent habit and outside
puberulous sepals.
Stem 15 to 45 cm tall, indistinctly 6-angulate, not or shallowly 6-sulcate, glabrous or
puberulous, simple or branched. Leaves pinnate or ternate; leaflets coriaceous, narrowly to
broadly ovate, 1–2(–3)×0.5–2 cm, margin entire or 1–4-denticulate per side, below the
middle 2–3-lobed or undivided, on both surfaces sparsely puberulous. Fl. Jun.–Jul.
China (SW Xinjiang) and E Tadzhikistan. In grassy places on slopes or on rocky cliffs;
alt. 3300–4600 m.
Additional specimens examined:
China. Xinjiang (新疆): Akto (阿克陶), Inst. Northwest Bot. Exped. (西北植物所队) 698 (PE, WUK);
Pishan (皮山), B. S. Li & D. Zheng (李勃生, 郑度) 11675 (PE); Suget Karaul, Thundlov 607 (BM);
Taxkorgan (塔什库尔干), Inst. Northwest Bot. Exped. (西北植物所队) 1221 (PE, WUK); Tagdumbash
Pamir, Appleton 41 (LE).
Tadzhikistan. Koung-Koul, Lacoste s.n. (P); Pamir: Kara Kul, Kuschakewicz s.n., S. Hedin s.n. (S);
Pamir Mts., Fedtschenko s.n. (LE); Ak-baisal, Kuschakewicz s.n. (LE).
4d. var. laciniifolia Grey-Wilson in Kew Bull. 44: 48, fig. 1: T. 1989; M. Johnson, Klematis
356. 1997; Grey-Wilson, Clematis 170. 2000. Type: Nepal. Kali Gandaki, Yara, S of Mustang,
alt. ca. 3900 m, Stainton, Sykes & Williams 2130 (holotype, BM!).
C. tibetana ssp. vernayi auct. non (C. E. C. Fisch.) Grey-Wilson: Brandenb., Meclatis in
Clematis 175. 2000, p.p. quoad syn. C. tibetana ssp. vernayi var. laciniifolia Grey-Wilson.
Acta Phytotaxonomica Sinica Vol. 44 422
This variety differs from var. tibetana in the leaflets being rhombic-ovate in outline,
3-lobed to 3-sect, and acutely dentate.
Woody vine. Branches only on nodes sparsely puberulous, elsewhere glabrous. Leaflets
rhombic-ovate, broadly rhombic, or broadly ovate, 1.4–3.8×1.2–4 cm, 3-lobed to 3-sect,
margin sparsely acutely dentate. Sepals thickly papery or subcoriaceous, narrowly ovate, 1.6–2
cm long, inside puberulous, outside glabrous.
W Nepal.
Additional specimens examined:
Nepal. Bheri Valley, Dunaihi, Polunin, Sykes & Williams 231 (BM); Kali Gandaki, Titre, N of Dana,
Stainton, Sykes & Williams 7543 (BM).
5. Clematis ladakhiana Grey-Wilson in Kew Bull. 44: 49. 1989; Rau in Sharma et al., Fl. Ind.
1: 68. 1993; M. Johnson, Klematis 341. 1997; Grey-Wilson, Clematis 170, fig. 127. 2000;
Grabovsk. in Grubov, Pl. As. Centr. 12: 81. 2001. Type: Kashmir Region. Nubra, Thomson s.n.
(holotype, K!; isotype, LE!).
C. orientalis L. var. acutifolia Hook. f. & Thoms., Fl. Ind. 9. 1885; et in Hook. f., Fl. Brit.
Ind. 1: 5. 1872. Type: Kashmir Region. Ladakh, Thomson s.n. (syntype, K—Grey-Wilson,
1989).
C. orientalis ssp. normalis var. daurica (Pers.) Kuntze f. thomsonii Kuntze in Verh. Bot.
Ver. Brand. 26: 124. 1885. Type: China. Xizang (西藏): Without precise locality, Thomson s.n.
(holotype, B; isotypes, GH!, S!).
C. orientalis ssp. normalis var. daurica f. dyeri Kuntze in l.c. Type: Kashmir Region.
Askole, Clarke 30329B (holotype, K).
?C. orientalis L. var. longifoliolata Tamura in Kitamura, Pl. W Pakist. & Afghan. 58.
1964. Type: Karakoram: Askole, alt. 3200 m, 1955-07-24, Nakao s.n. (holotype, KYO).
C. tibetana auct. non Kuntze: Brandenb., Meclatis in Clematis 165. 2000, p.p., quoad
syn. C. ladakhiana Grey-Wilson.
Fig. 6: E, F
Woody vine. Branches 6-angulate or subterete, indistinctly shallowly 6–8-sulcate, on
nodes sparsely puberulous, elsewhere glabrous. Leaves 1–2-pinnate; leaflets chartaceous or
subcoriaceous, lanceolate-linear or narrowly lanceolate, 1.5–5.5×0.2–1 cm, apex attenuate,
margin entire or 1–2-dentate, undivided or above base 2–3-lobed, on both surfaces glabrous,
occasionally adaxially near base with a few hairs, basal veins nearly flat; petioles 3–6.8 cm
long. Cymes axillary, 1–5-flowered; peduncles 0.2–3(–4.7) cm long, glabrous or sparsely
puberulous; bracts pinnate. Flower 1.8–3 cm in diam.; pedicel 2.2–9 cm long, glabrous. Sepals
4, yellow, often tinged with purple-brown outside, ascending, papery, narrowly ovate or
broadly lanceolate, 1.2–2×0.5–0.9 cm, apex attenuate or acuminate, inside puberulous,
outside glabrous, on margin velutinous. Stamens 7–9 mm long; anthers narrowly oblong or
oblong, 2–2.5(–3) mm long, glabrous, apex obtuse. Ovaries pubescent; styles 7–9 mm long,
densely villous. Achenes obovate, ca. 3.5×1.8 mm, densely pubescent; persistent styles ca.
3.5 cm long, plumose. Fl. Jul. –Sept.
China (SW Xizang) and Kashmir Region. In bushes or on rocks by river; alt. 2800–3850
m.
Additional specimens examined:
China. Xizang (西藏): from Kargyay, to Lingti, Stolczka s.n. (K); from Upschi to Tel, Schlagintweit
s.n. (P).
Kashmir Region: Ladakh, Koelz 2537 (GH, LE, S, US), Heber s.n. (US), Stainton 8413 (K); Nubra,
Schlagintweit s.n. (US).
6. Clematis glauca Willd., Berl. Baumz. 65. t. 4, fig. 1. 1796; Poir., Encyc. Suppl. 2: 295.
1811; DC., Syst. 1: 136. 1818; et Prodr. 1: 3. 1824; Ledeb., Fl. Ross. 1: 3. 1841; Prantl in Bot.
Jahrb. 9: 261. 1888; Krasch. in Kom., Fl. URSS 7: 321. 1937; Gamayou. in Fl. Kazakhst. 4:
No. 4 WANG Wen-Tsai: A revision of Clematis sect. Meclatis (Ranunculaceae) 423
72, t. 9, fig. 4. 1961; Anonymous in Iconogr. Corm. Sin. 1: 740. 1972; P. P. Ling in Fl. Reip.
Pop. Sin. 28: 143. 1980, p.p.; Grubov, Key Vasc. Pl. Mongol. 113. 1982; Liou f., Fl. Desert.
Sin. 1: 485. 1985; J. G. Liu in Fl. Xinjiang. 2: 291, pl. 79, figs. 3, 4. 1994; Grabovsk. in
Grubov, Pl. As. Centr. 12: 79. 2001; W. T. Wang & Barth. in Fl. China 6: 366. 2001, p.p.——
Meclatis sibirica Spach, Hist. Nat. Veg. Phan. 7: 273. 1839.——C. orientalis L. var. glauca
(Willd.) Maxim., Fl. Tangut. 3. 1889, quoad nomen tantum. Type: In Herb. Willd., n. 10474,
from plant cultivated in the Botanical Garden, Berlin—Grey-Wilson, 1989; Brandenburg,
2000).
C. daurica Pers., Synop. 2: 99. 1806; DC., Syst. 1: 153. 1818; et Prodr. 1: 7. 1824;
Ledeb., Fl. Ross. 1: 4. 1841.——C. orientalis L. ssp. normalis Kuntze var. daurica (Pers.)
Kuntze in Verh. Bot. Ver. Brand. 26: 123. 1885; Grey-Wilson in Kew Bull. 44: 43. 1989; et
Clematis 165, fig. 118. 2000; M. Johnson, Klematis 345. 1997. Type: Without precise locality,
Patrin 10502 (holotype, P-Ju—Brandenburg, 2000).
C. orientalis ssp. normalis var. daurica f. persoonii Kuntze in Verh. Bot. Ver. Brand. 26:
124. 1885. Type: Specimen from plant collected from Dahuria (syntype, P-Ju); specimen from
plant cultivated in Bot. Gard., Paris (syntype).
C. orientalis auct. non L.: Brandenb., Meclatis in Clematis 130. 2000, p.p., quoad syn. C.
glauca Willd. et C. daurica Pers.
粉绿铁线莲 Fig. 7: A–C
Woody vine. Branches 4–8-sulcate, sparsely puberulous or subglabrous. Leaves 1–2-
pinnate; leaflets blue-green, papery or herbaceous, narrowly ovate, lanceolate, elliptic, or
ovate, (1–)2–5×0.6–1.8 cm, apex acute, base broadly cuneate, margin entire, undivided or
near base 1–2-lobed, adaxially glabrous, abaxially very sparsely puberulous, basal veins
slightly prominent; petioles 2.5–6 cm long, subglabrous. Cymes axillary, 1–7(–many)-
flowered; peduncles 1–4.6 cm long, glabrous; bracts petiolate, ovate or lanceolate, 0.8–1.6 cm
long. Flower 3.5–3.8 cm in diam.; pedicel 2–7 cm long, near apex puberulous. Sepals 4,
ascending or nearly spreading, yellow, thinly papery, narrowly lanceolate or narrowly oblong,
1.4–2.3×0.4–0.7 cm, glabrous or sparsely puberulous on both surfaces, margin velutinous,
apex attenuate or acute. Stamens 7–14 mm long; anthers linear or broadly linear, 2–3.5 mm
long, glabrous, apex obtuse, rarely minutely apiculate. Ovaries pubescent; styles 7–10 mm
long, densely villous. Achenes elliptic or ovate, 1.8–2.2×1.2–1.6 mm, pubescent, rimmed;
persistent styles 3–5 cm long, plumose. Fl. Jul. –Sept.
China (C Gansu, Xinjiang), Kazakhstan, Kyrgyzstan, Mongolia, and Russia (S Siberia).
On slopes or waste lands, or by rivers; alt. 500–2000 m.
Additional specimens examined:
China. Gansu (甘肃): Kangle (康乐), Y. S. Lian (廉永善) 96873 (PE). Xinjiang (新疆): Altay Shan
(阿尔泰山), R. C. Ching (秦仁昌) 2332 (PE); Burqin (布尔津), Y. R. Ling (林有润) 74-1008 (PE); Fukang
(阜康), K. C. Kuan (关克俭) 4191 (PE); Hoxud (和硕), A. J. Li & J. N. Zhu (李安仁, 朱家柟) 7705 (PE);
Korla (库尔勒), A. J. Li & J. N. Zhu (李安仁, 朱家柟) 8641 (PE); Nilka (尼勒克), K. C. Kuan (关克俭)
4046 (PE); Tacheng (塔城), Y. R. Ling (林有润) 74-1360 (PE); Turpan (吐鲁番), Z. M. Zhang (张治明) 285
(PE); Ulastai (乌拉斯台), K. C. Kuan (关克俭) 3816 (PE); Ürümqi (乌鲁木齐), A. J. Li & J. N. Zhu (李安仁,
朱家柟) 10949 (PE); Wenquan (温泉), Hoch & J. R. Chen 86-308 (PE).
Kazakhstan. Kungej-Alatau, Roldugin 5367 (US); Semipalatinsk, Korshinsky s.n. (US).
Kyrgyzstan. ad fluv. Tiup prope Santash, Skvortsov s.n. (GH).
Mongolia. Kobdosk, Baranov s.n. (LE); Near Lake Kirghiz-nor, Potanin s.n. (LE).
Russia. Altai Mountain: Chuya River, Elias, Weber, Tomb & Krasnoborov 4411 (PE); Tuva AR: Kazil,
Elias, Shetler & Murray 7594 (US); Tuvinskaja AR: Erzinsk, Timochia & Mrichin 2340 (LE); Siberia,
Turczaninow s.n. (P).
7. Clematis zandaensis W. T. Wang in Acta Phytotax. Sin. 36: 165, fig. 2: 1–4. 1998; Grey-
Wilson, Clematis 174. 2000; W. T. Wang & Barth. in Fl. China 6: 366. 2001. Type: China.
Acta Phytotaxonomica Sinica Vol. 44 424



Fig. 7. A–C, Clematis glauca Willd. A, flowering branch; B, sepal outside; C, stamen. Drawn from Skvortsov s.n. D–G,
C. zandaensis W. T. Wang. D, flowering branch; E, F, leaflets; G, stamen. Drawn from Qinghai-Xizang Exped. 76-8160.


No. 4 WANG Wen-Tsai: A revision of Clematis sect. Meclatis (Ranunculaceae) 425
Xizang (西藏): Zanda (札达), between Gulang (古浪) and Shibuqi (什布奇), alt. 3500 m,
1976-07-02, Qinghai-Xizang Exped. (青藏队) 76-8160 (holotype, PE!).
C. graveolens auct. non Lindl.: P. P. Ling in Fl. Reip. Pop. Sin. 28: 144. 1980; M. Y. Fang
in Fl. Xizang. 2: 89. 1985.
札达铁线莲 Fig. 7: D–G
Woody vine. Branches shallowly 4–8-sulcate, glabrous. Leaves 1–2-pinnate; leaflets
green, papery, ovate or broadly ovate, 1.8–5×1.8–4 cm, base subcordate, rounded, or broadly
cuneate, 2–3-lobed to 2–3-parted, margin sparsely dentate, terminal lobe larger, ovate-oblong,
lateral lobes smaller, glabrous on both surfaces or abaxially on basal veins sparsely
puberulous, basal veins abaxially slightly prominent; petioles 5.4–7 cm long. Cymes axillary,
1–3-flowered; peduncles 4–9.5 cm long, glabrous; bracts shortly petiolate, ternate. Flower
1.4–2 cm in diam.; pedicel 1.5–10 cm long, glabrous. Sepals 4, yellow, ascending, papery,
ovate or narrowly ovate, 0.9–1.8×0.5–0.8 cm, inside puberulous, on margin velutinous,
outside glabrous, apex acute. Stamens 3.5–10 mm long; anthers narrowly oblong, 1.4–3.2 mm
long, glabrous, apex obtuse. Ovaries pubescent; styles 7–10 mm long, densely villous. Fl. Jun.
–Jul.
China (SW Xizang) and Kashmir Region. In thickets on slopes or on river banks; alt.
3500 m.
Additional specimen examined:
China. Xizang (西藏): Zanda (札达), Qinghai-Xizang Exped. Vegetation Group (青藏植被队) 12914
(PE).
Kashmir Region. Chitral: Tirich Gol, Stainton 2789 (BM).
8. Clematis serratifolia Rehd. in Mitt. Deutsch. Dendr. Ges. 248. 1910; et in Repert. Sp. Nov.
13: 362. 1914; Bailey in Stand. Cyclop. Hort. 1: 798. 1914; Krasch. in Kom., Fl. URSS 7: 321.
1937; Nakai in J. Jap. Bot. 13 (7): 1. 1937; Kitag., Lineam. Fl. Mansh. 218. 1939; Ohwi, Fl.
Japan 442. 1965; Liou et al., Fl. Pl. Herb. Chinae Bor.-Or. 3: 173, pl. 75, figs. 4–7. 1975; T. B.
Lee, Ill. Fl. Korea 342, fig. 1368. 1979; P. P. Ling in Fl. Reip. Pop. Sin. 28: 143, fig. 16. 1980;
Kitamura & Murata, Colour. Ill. Herb. Pl. Japan, rev. ed., 2: 225. 1980; Tamura in Satake et al.,
Wild Flow. Japan 2: 71. 1982; Grey-Wilson in Kew Bull. 44: 59. 1989; Ohwi & Kitag., New
Fl. Japan 680. 1992; Luferov in Pl. Vasc. Or. Extr. Soviet. 7: 97. 1995; Y. N. Lee, Fl. Korea
167, fig. 494. 1996; M. Johnson, Klematis 349. 1997; Grey-Wilson, Clematis 175, figs. 133,
134. 2000; Brandenb., Meclatis in Clematis 160, fig. 221. 2000; W. T. Wang & Barth. in Fl.
China 6: 366. 2001. Type: Korea. Ping Yang, 1905-09-18, Jack s.n. (holotype, GH!).
C. orientalis L. var. serrata Maxim. in Bull. Acad. Sci. St.-Petersb. 9: 21. 1879; Nakai in
J. Coll. Sci. Univ. Tokyo 26: 7. 1909.——C. intricata Bunge var. serrata (Maxim.) Kom. in
Acta Hort. Petrop. 22: 289. 1904.——C. serrata (Maxim.) Kom. in Kom. & Alis., Key Pl. Far.
East Reg. USSR 1: 549. 1931. Type: Manshuria ross. austr., 1872-08-10, Goldenstadt 255
(holotype, LE!).
C. orientalis L. var. wilfordi Maxim. in l.c.; Nakai in l.c.——C. eriopoda Maxim. var.
wilfordi (Maxim.) Kuntze in Verh. Bot. Ver. Brand. 26: 165. 1885; Huth in Bull. Herb. Boiss.
5: 1064. 1897; Matsum., Ind. Pl. Japan 2: 110. 1912.——C. wilfordi (Maxim.) Kom. in Kom.
& Alis., Key Far East Reg. USSR 1: 549. 1931.——C. serratifolia Rehd. f. wilfordi (Maxim.)
Kitag., Neu-Lineam. Fl. Mansh. 299. 1979. Type: China. Without precise locality: Coast of
Manchuria, 1859, Wilford s.n. (holotype, LE!; isotypes, G!, UPS!).
C. sibiricoides Nakai in J. Jap. Bot. 23: 13. 1949. Type: Japan. Jeso: Prov. Kitami
Wekka-nai, 1946-07-30, Hiraoka Tomie s.n. (holotype, TI!).
齿叶铁线莲 Fig. 8: A, B
Woody vine. Branches shallowly 6–8-sulcate, sparsely puberulous or glabrous. Leaves
2-ternate; leaflets green, papery, lanceolate, narrowly ovate, or ovate, 3–6(–8)×1–2.5(–3) cm,
Acta Phytotaxonomica Sinica Vol. 44 426

Fig. 8. A, B, Clematis serratifolia Rehd. A, flowering branch; B, stamen. Drawn from Sato 3039. C, D, C. orientalis L.
var. orientalis. C, flowering branch; D, stamen. Drawn from Sintenis 848.
No. 4 WANG Wen-Tsai: A revision of Clematis sect. Meclatis (Ranunculaceae) 427
apex attenuate, base broadly cuneate or rounded, margin serrate or denticulate, sparsely
puberulous on both surfaces, glabrescent, basal veins abaxially nearly flat; petioles 3.5–6 cm
long. Cymes axillary, 1–3-flowered; peduncles 0.5–1.5 cm long; bracts petiolate or sessile,
lanceolate or linear, 0.4–1.5 cm long. Sepals 4, yellow, ascending, papery or submembranous,
oblong, elliptic-lanceolate, or narrowly ovate, 1.5–2.5×0.5–0.8 cm, inside puberulous, outside
glabrous, margin velutinous, apex attenuate or caudate-cuspidate. Stamens 8–12 mm long;
anthers narrowly oblong, 1.2–2.6 mm long, glabrous, apex minutely apiculate. Ovaries
pubescent; styles 7–9 mm long, densely villous. Achenes elliptic, ca. 3×2 mm, pubescent;
persistent styles ca. 3 cm long, plumose. Fl. Jul.–Sept.
China (E Jilin, Liaoning), N Japan, N Korea, and Russia (Far East Region). In forests, on
slopes or on gravelly river banks; alt. ca. 400 m.
Additional specimens examined:
China. Jilin (吉林): Antu (安图), T. N. Liou (刘慎谔) 3785, 4336 (LE, PE); Changbai Shan (长白山),
J. J. Chien (钱家驹) 799 (PE); Helong (和龙), Yanbian Exped. (延边队) 59-792 (PE); Hunchun (珲春), P. Y.
Fu et al. (傅沛云等) 798 (PE); inter Kirin et Omosa, Komarov 703 (LE, P), 710 (1896-08-28, K); Linjiang (临
江), S. X. Li (李书心等) 1092 (PE); Wangqing (汪清), P. Y. Fu et al. (傅沛云等) 989 (PE). Liaoning (辽宁):
Benxi (本溪), C. Q. Lin (林长清) 1218 (PE); Caohekou (草河口), Yabe s.n. (NAS); Kuandian (宽甸), S. C.
Tsui (崔顺昌) 338 (PE); Youyan (岫岩), W. Wang et al. (王薇等) 1487 (PE).
Korea. Bakuho, Uchiyama s.n. (NAS); He-chun, Mills s.n. (PE); Kangkai, Mills s.n. (PE); Kapsan,
Komarov s.n. (PE); Lyongdam, Mills s.n. (PE); Prov. Hengkyo: Tumen-Yalu divide, E. H. Wilson 8927 (GH,
US); Prov. Kogen: Kongo-san, E. H. Wilson 10711 (GH, K, US); Taebak-san, Hagman 272 (UPS);
Un-czchen-gen, Komarov 710 (1897-07-01, GH, K).
Russia. Prov. Primorje: near Vladivostok, Skvortsov 54 (G, GH, LE), Topping 2436 (US), Ulanova
6884 (G. LE).
9. Clematis hilariae Koval. in Not. Syst. Herb. Inst. Bot. Acad. Sci. Uzbek. 18: 34. 1967;
Grey-Wilson in Kew Bull. 44: 55, fig. 6: A–C. 1989, p.p., excl. syn. C. sarezica Ikonn.; M.
Johnson, Klematis 336. 1997; Grey-Wilson, Clematis 174, fig. 132. 2000. Type: Plants
cultivated in Bot. Gard. Sci. Uzbek. from seeds collected in the Pamir Mts. near Ljangar,
1966-09-17, Kovalevskaja 5541 (holotype, ?; isotypes, G!, LE!, S!).
C. orientalis L. var. roschanica Korsh. in Bull. Acad. Sci. St.-Petersb., ser. 5, 9: 399.
1898. Type: Iter Turkistanicum, 1897-08, Korshinsky 2772, 3790 (syntypes, LE!).
C. chrysantha Ulbr. var. paucidentata Tamura in Kitamura, Add. Corr. Fl. Afghan. 92.
1966. Type: Afghanistan. Badakhshan, Wakhlan, Qasideh Valley, Mt. Noshaq, 1960-07-20,
Yosii 488 (holotype, KYO!).
C. chrysantha Ulbr. var. monantha Tamura in l.c. Type: Afghanistan. Wakhan, Ishkashim,
Yosii 801 (holotype, ?).
C. intricata auct. non Bunge: Brandenb., Meclatis in Clematis 151. 2000, p.p., quoad syn.
C. hilariae Koval. et C. chrysantha Ulbr. var. paucidentata Tamura et var. monantha Tamura.
Fig. 9: C, D
Woody vine. Branches shallowly 8–12-sulcate, puberulous. Leaves 1–2-pinnate; leaflets
green, papery, narrowly ovate or lanceolate, 1–3.5×0.3–1.2 cm, apex attenuate, base cuneate,
undivided or 2–3-lobed, margin regularly or only below sparsely dentate or serrate, adaxially
subglabrous, abaxially on veins sparsely puberulous, basal veins abaxially slightly prominent;
petioles 1.5–4 cm long. Cymes axillary and terminal, 1–7-flowered; peduncles 0.4–6 cm long;
bracts pinnate or simple. Flower 2.8–4 cm in diam.; pedicel 3–10 cm long, puberulous. Sepals
4(–5), yellow, spreading, subcoriaceous, narrowly ovate or lanceolate, 13–22(–25)×5–8(–9)
mm, inside puberulous, outside sparsely puberulous, on margin velutinous, apex attenuate.
Stamens 7–9 mm long; anthers narrowly oblong or linear, 2.5–3(–3.5) mm long, glabrous,
apex obtuse or obscurely apiculate. Ovaries puberulous; styles ca. 9 mm long, densely villous.
Achenes elliptic, ca. 2.5×1.2 mm, puberulous, narrowly rimmed; persistent styles 3–3.8 cm
long, plumose. Fl. Jul.–Sept.
Acta Phytotaxonomica Sinica Vol. 44 428
NE Afghanistan and E Tadzhikistan. On borders of cultivated fields or by rivers; alt. ca.
3000 m.
10. Clematis sarezica Ikonn. in Novit. Syst. Pl. Vasc. 14: 231. 1977; M. Johnson, Klematis
349. 1997; Grey-Wilson, Clematis 174. 2000. Type: Tadzhikistan. Badachshan, Lake Sarez,
alt. 3350 m, 1958-08-23, Ikonnikov 5830 (holotype, LE!; isotypes, PE!, S!).
C. hilariae auct. non Koval.: Grey-Wilson in Kew Bull. 44: 55. 1989, p.p. quoad syn. C.
sarezica Ikonn.
C. intricata auct. non Bunge: Brandenb., Meclatis in Clematis 151. 2000, p.p., quoad syn.
C. sarezica Ikonn.
Fig. 9: A, B
Woody vine. Branches subterete, shallowly 10-sulcate, only on nodes sparsely
puberulous, elsewhere glabrous. Leaves 1–2-pinnate; leaflets green, subcoriaceous or
chartaceous, narrowly lanceolate or lanceolate, 1.6–4.6×0.3–1.2 cm, apex attenuate, base
cuneate, undivided or 2–3-lobed or 2–3-sect (the terminal lobe larger, lanceolate, the lateral
ones smaller, obliquely lanceolate or long elliptic), adaxially below on veins sparsely
puberulous, abaxially glabrous, midrib adaxially impressed, abaxially slightly prominent;
petioles 4–5.4 cm long. Cymes axillary, 1-flowered; peduncles 0.2–8 cm long, glabrous; bracts
petiolate, simple or ternate, 1.8–2.5 cm long. Flower ca. 3 cm in diam.; pedicel 2–4.5 cm long,
glabrous. Sepals 4, brownish-purple, ascending, thickly papery, narrowly ovate or lanceolate,
15–19×5–8 mm, outside below apex shortly corniculate, inside puberulous, outside glabrous,
on margin velutinous. Stamens 7–9.5 mm long; anthers narrowly oblong, ca. 2.5 mm long,
glabrous, apex minutely apiculate. Ovaries pubescent; styles ca. 9 mm long, densely villous.
Achenes ovate, ca. 2.5×1.5 mm, pubescent; persistent styles ca. 3 cm long, plumose. Fl. Aug.
E Tadzhikistan. On grassy banks of lake; alt 3350 m.
11. Clematis orientalis L., Sp. Pl. 1: 543. 1753; DC., Syst. 1: 135. 1818; et Prodr. 1: 3. 1824;
Ledeb., Fl. Ross. 1: 3. 1841; Trautv, in Bull. Soc. Nat. Mosc. 33: 57. 1860; Boiss., Fl. Orient.
1: 3. 1867; Koch, Dendr. 1: 422. 1869; Hook. f. & Thoms. in Hook. f., Fl. Brit. Ind. 1: 5. 1872,
p.p.; Maxim. in Bull. Acad. Sci. St.-Petersb. 22: 211. 1876; Tritram, Fauna & Fl. Palest. 207.
1884; Kuntze in Verh. Bot. Ver. Brand. 26: 123. 1885, p.p.; Maxim., Enum. Pl. Mongol. 3.
1889; Halácsy, Consp. Fl. Graec. 1: 2. 1901; Finet & Gagnep. in Bull. Soc. Bot. France 50:
540. 1903, p.p.; Ostenf. & Pauls in Hendin, S Tibet 6: 3. 1922; Hayek, Prodr. Fl. Penin.
Balcan. 1: 322. 1927; Krasch. in Kom., Fl. URSS 7: 322. 1937; Reching., Fl. Aegaea 184.
1943; Parsa, Fl. Iraq 1: 353. 1952; Gamayou. in Fl. Kazakhst. 4: 72. 1961; Tutin, Fl. Europ. 1:
221. 1964; Davis et al., Fl. Turkey 1: 139. 1965; P. P. Ling in Fl. Reip. Pop. Sin. 28: 140. 1980;
Liou f., Fl. Desert. Sin. 1: 185. 1985; Grey-Wilson in Kew Bull. 44: 36, fig. 2. 1989; Y. Z.
Zhao in Fl. Intramongol., ed. 2, 2: 531. 1990; Riedl & Nasir in Fl. Pakist 193: 93. 1991;
Reching., Fl. Iraq 171: 233. 1992; Rau in Sharma et al., Fl. Ind. 1: 72. 1993; J. G. Liu in Fl.
Xinjiang. 2: 291. 1994; Heller & Heyn, Consp. Fl. Orient. 444. 1994; M. Johnson, Klematis
342. 1997; Grey-Wilson, Clematis 163. 2000; Brandenb., Meclatis in Clematis 130, fig. 2: 10,
11. 2000, p.p.; Grabovsk. in Grubov, Pl. As. Centr. 12: 82. 2001; W. T. Wang & Barth. in Fl.
China 6: 367. 2001.——Meclatis orientalis (L.) Spach, Hist. Nat. Veg. Phan. 7: 274. 1839.—
—Clematis orientalis L. var. vulgaris Trautv. in Bull. Soc. Nat. Mosc. 33: 57. 1860; Maxim.,
Enum. Pl. Mongol. 4. 1889.——Viticella orientalis (L.) W. A. Weber in Phytologia 55: 9.
1984. Type: Dillenius Hort. Eltham. 144, t. 119, fig. 145. 1732 (lectotype, BM!); Dillenium
Herb. n. 2868 (typotype, OXF)—Grey-Wilson, 1989; Brandenburg, 2000.
Clematitis flava Moench, Method. 296. 1794.——Clematis orientalis ssp. normalis
Kuntze var. flava (Moench) Kuntze in Verh. Bot. Ver. Brand. 26: 123. 1885. Type: No type
specimen designated.
No. 4 WANG Wen-Tsai: A revision of Clematis sect. Meclatis (Ranunculaceae) 429


Fig. 9. A, B, Clematis sarezica Ikonn. A, flowering branch; B, stamen. Drawn from Ikonnikov 5830. C, D, C. hilariae Koval. C,
flowering and fruiting branch; D, stamen. Drawn from Kovalevskaja 5541.
Acta Phytotaxonomica Sinica Vol. 44 430
Clematis longecaudata Ledeb., Fl. Ross. 1: 3. 1841.——C. orientalis ssp. wightiana
(Wall.) Kuntze var. longecaudata (Ledeb.) Kuntze in l.c. 125. Type: Without field notes
(holotype, LE!).
C. glauca Willd. β. angustifolia Ledeb., Fl. Ross. 1: 3. 1841. Type: in Mt. Altay. vel fl.
Katunja (syntypes).
东方铁线莲
This species consists of seven varieties occurring in southeastern Europe (Aegean Islands
of Greece) and western and central Asia.
11a. var. orientalis Fig. 8: C, D
Woody vine. Branches shallowly 6–10-sulcate, puberulous, sometimes subglabrous.
Leaves 1–2-pinnate; leaflets grey-green, thickly papery or subcoriaceous, variable in shape
and division, ovate, narrowly ovate, or lanceolate, 1.5–5×0.5–5 cm, appressed-puberulous on
both surfaces, sometimes subglabrous, usually below 2–3-lobed to 2–3-sect, sometimes
undivided, terminal lobe larger, narrowly ovate, lanceolate, linear-lanceolate, or narrowly
obovate, 0.4–1.8 cm broad, margin entire or 1–2-dentate per side, midrib adaxially flat,
abaxially slightly prominent, lateral lobes smaller; petioles 2.8–6.5 cm long. Cymes axillary
and terminal, few- to many-flowered, often panicle-like; peduncles 1.4–6.5 cm long, 1–1.2
mm thick; bracts petiolate, leaflet-like. Flower 1.4–2.8 cm in diam.; pedicel 1–2.5 cm long,
appressed-puberulous, usually above densely hairy. Sepals 4, yellow, spreading or reflexed,
oblong-lanceolate or narrowly oblong, 6.5–15×3–5 mm, puberulous on both surfaces, on
margin velutinous, apex acute. Stamens 5–9 mm long; anthers linear or narrowly oblong, 2–4
mm long, glabrous, apex obtuse, sometimes minutely apiculate. Ovaries pubescent; styles 6–8
mm long, densely villous. Achenes narrowly elliptic, ca. 2.8×1.2 mm , puberulous, slightly
rimmed; persistent styles 2.5–5.5 cm long, plumose. Fl. Jun.–Sept.
Greece (Aegean Islands), Afghanistan, Armenia, Azerbaijan, China (NW Gansu, SW Nei
Mongol, Xinjiang), Georgia, Iran, N Iraq, Kashmir Region, Kazakhstan, Kirghizstan,
Mongolia, N Pakistan, Palestine, Russia (?SW Siberia), Tadzhikistan, Turkey, Turkmenistan,
and Uzbekistan. On slopes, in bushes or thickets, or by streams or rivers; alt. 20–2600 m.
Additional specimens examined:
Afghanistan. Bamian, Podlech 18872 (G); Doab, Pabot 1230 (G); Helmund Valley, Long 443 (US);
Kunar, Podlech 16638 (G); Lashkari Basar, Frumkin 54 (G).
Armenia. Amassia, Manissadjian 639 (P); Without precise locality, Safier s.n. (K), Schischkin s.n. (LE),
Sintenis 2998 (G, P), 3390 (LE).
Azerbaijan. Kuba, Karjajev s.n. (S).
China. Gansu (甘肃): Anxi (安西), Anonymous 308 (PE); Jiuquan (酒泉), Qinghai-Gansu Exped. (青
甘调查队) 60-2966 (PE); Minqin (民勤), Y. Q. He (何业祺) 3332 (PE). Xinjiang (新疆): Alataw Shan (阿拉
套山), Xinjiang Exped. (新疆队) 78-3835 (PE); Altay Shan (阿尔泰山), R. C. Ching (秦仁昌) 2813 (PE);
Aksu (阿克苏), Z. N. Feng (冯肇南) 417 (PE); Fuyun (富蕴), R. C. Ching (秦仁昌) 1892 (PE); Gongliu (巩
留), Xinjiang Exped. Inst. Northwest Bot. (西北植物所新疆队) 2667 (PE); Hami (哈密), R. C. Ching (秦仁
昌) 122 (PE); Hoxud (和硕), A. J. Li & J. N. Zhu (李安仁, 朱家柟)7705 (PE); Korla (库尔勒), A. J. Li & J.
N. Zhu (李安仁, 朱家柟) 8641 (PE); Lop Nur (罗布泊), Xinjiang Exped. (新疆队) 59-10120 (PE); Manas
(玛纳斯), K. C. Kuan (关克俭) 1339 (PE); Nilka (尼勒克), K. C. Kuan (关克俭) 3885 (PE); Qira (策勒),
Xinjiang Exped. (新疆队) 56-129 (PE); Shanshan (鄯善), A. J. Li & J. N. Zhu (李安仁, 朱家柟) 6692 (PE);
Shawan (沙湾), G. L. Zhu (朱格麟) 5517 (PE); Tacheng (塔城), Y. R. Ling (林有润) 74-1363 (PE); Tian
Shan (天山), T. N. Liou (刘慎谔) 2689 (PE); Toksun (托克逊), A. J. Li & J. N. Zhu (李安仁, 朱家柟) 7288
(PE); Turpan (吐鲁番), Z. M. Zhang (张智敏) 294 (PE); Ürümqi (乌鲁木齐), T. N. Liou (刘慎谔) 2891, R.
C. Ching (秦仁昌) 3372 (PE); Wenquan (温泉), K. C. Kuan (关克俭) 4560 (PE); Yecheng (叶城),
Qinghai-Xizang Exped. (青藏队) 87-764 (PE); Yining (伊宁), Z. M. Zhang (张智敏) 206, Y. X. Liou (刘瑛
心) 712 (PE); Yiwu (伊吾), Q. R. Wang (王庆瑞) 4522 (PE); Zhaosu (昭苏), Xinjiang Exped. Inst. Northwest
Bot. (西北植物所新疆队) 2593 (PE).
Georgia. Caucasus, Vasak s.n. (G).
No. 4 WANG Wen-Tsai: A revision of Clematis sect. Meclatis (Ranunculaceae) 431
Iran. Amol, Hewer 1492 (LE); Khorasan, Koelz 16822 (US), Rechinger 1424 (S); Lusistan, Bornmuller
s.n. (LE); Nischapur, Bunge s.n. (LE); Ostan, Schmid 6289 (G); Salukhi, Sintenis 1108 (G); Techeran, Acher
Eloy 4025 (P).
Kashmir region. Jacquemont 1162 (P), Stewart 2015 (G).
Kazakhstan. Dzhambul, Raikova 2915 (G, GH, K, PE, S); Kungej-Alatau, Roldugin 5367 (K); Western
Tianshan, Mekerov 400 (PE).
Kirghizstan. Chatkal Range, Skvortsov s.n. (GH); Kurdjich, Petrova s.n. (GH).
Mongolia. Shabarakh Usu, Chaney 583 (US).
Pakistan. Karakorum, Polunin 6414 (G).
Tadzhikistan. Achty, Alexenko s.n. (LE).
Turkey. Camardi, Nydegger 15449 (G); Kalecik, Guichard s.n. (LE); Malatya, Godfrey & Taysi
SH-386 (US); Mrgrp, Kotte s.n. (K); Paphlagonia: Wilajet Kastanbuli, Sintenis 4881 (S); Tortum, Davis
47565 (K), Winter 369 (US).
Turkmenistan. Ashabad, Litwinow 23 (G), Sintenis 848 (US); ad fl. Daino-Ssu, Litwinow 411 (P);
Farab, Androsov 2916 (K).
Uzbekistan. Taskent, Ellas, Murray & Newcomba 9873 (PE), Korovina s.n. (G).
11b. var. albida (Klotzsch) Kuntze in Verh. Bot. Ver. Brand. 26: 124. 1885.——C. albida
Klotzsch in Klotzsch & Garke, Bot. Ergeb. Reise Pr. Waldemar 131, t. 40. 1862. Type: W.
Himalaya, Hoffmeister s.n. (holotype, B).
This variety differs from var. orientalis by having leaflets ovate or broadly ovate,
2–3-lobed to 2–3-sect and dentate.
Branches puberulous. Leaflets grey-green, ovate or broadly ovate, 1.2–3.8×1–4 cm,
2–3-lobed to 2–3-sect, terminal lobe rhombic, often 3-lobulate, margin dentate or
incised-dentate, lateral lobes smaller. Cymes 3–9-flowered; peduncles 1.5–5.5 cm long, 1 mm
in diam. Sepals oblong-lanceolate, ca. 14 mm long, puberulous on both surfaces.
N Pakistan and probably SW Afghanistan and Kashmir region.
Additional specimens examined:
Pakistan. Kurram Valley, Aitchison 415 (K, P), 455 (GH).
11c. var. latifolia Hook. f. & Thoms., Fl. Ind. 1: 9. 1855; et in Hook. f., Fl. Brit. Ind. 1: 5.
1875; Grey-Wilson in Kew Bull. 44: 42. 1989; et Clematis 165. 2000; M. Johnson, Klematis
346. 1997. Type: India. Himachal Pradesh, Piti, Munro s.n. (syntype, K!—Grey-Wilson,
1989).
C. globosa Royle, Ill. Bot. Himal. Mts. 51. 1834; Lauener in Not. R. Bot. Gard. Edinb.
36: 128. 1978.——C. orientalis L. var. globosa (Royle) Mukerjee in Bull. Bot. Surv. Ind. 1:
140. 1959; Kapoor in Bull. Nat. Bot. Gard. Lucknow 124: 57. 1966. Type: Kashmir Region.
Soongnum, Royle s.n. (holotype, LIV).
This variety differs from var. orientalis by having leaflets green, ovate, 3-parted, and
dentate,and sepals glabrous outside. It shows striking resemblance to var. albida, but differs by
having green leaflets and sepals glabrous outside.
Branches puberulous or glabrous. Leaflets green, ovate, 1.5–3×0.7–3 cm, 2–3-parted,
terminal lobe rhombic or rhombic-ovate, often 3-lobulate, margin dentate, lateral lobes
smaller. Cymes 3–many-flowered; peduncles 1.5–6(–9) cm long, 1–1.8 mm in diam. Sepals
oblong-lanceolate, 10–13(–16) mm long, inside puberulous, outside glabrous.
SE Afghanistan, N India, and Kashmir Region.
Additional specimens examined:
Afghanistan. Kunar, Bashgal-Tal, Podlech 32143 (S).
India. Kunawar, Thomson s.n. (GH).
11d. var. robusta Grey-Wilson in Kew Bull. 44: 40. 1989; et Clematis 164, fig. 119. 2000; M.
Johnson, Klematis 346. 1997. Type: Afghanistan. Badakhshan Province: Wakham, between
Urgun and Qazideh, alt. 2800 m, 1971-07, Grey-Wilson & Hewer 1688 (holotype, K!).
C. orientalis L. var. hindukushensis Grey-Wilson in Kew Bull. 44: 40. 1989; et Clematis
164. 2000; M. Johnson, Klematis 345. 1997. Type: Afghanistan. Ghorat Province, 1 mile E of
Minaret of Djam, alt. 2000 m, 1971-06, Grey-Wilson & Hewer 1224 (holotype, K!).
Acta Phytotaxonomica Sinica Vol. 44 432
This variety differs from var. orientalis by having larger leaflets and sepals, and longer
peduncles and persistent styles.
Branches puberulous. Leaflets grey-green, ovate, 3–7×1–4.5 cm, usually above base
3-parted, terminal lobe lanceolate or rhombic-obovate, 0.7–2 cm broad, 1-dentate per side or
entire, sparsely puberulous on both surfaces, lateral lobes usually much shorter. Cymes
7–many-flowered; peduncles 6–12 cm long, 1–1.8 mm in diam. Sepals lanceolate, 16–21 mm
long, puberulous on both surfaces. Persistent styles up to 8 cm long.
NE Afghanistan and probably Kashmir region.
Additional specimen examined:
Afghanistan. Badakhshan Province: Wakham, Grey-Wilson & Hewer 1634 (K).
?Kashmir region. Songono, 1851, Tatarinov s.n. (S).
11e. var. tenuifolia (Royle) Grey-Wilson in Kew Bull. 44: 42. 1989; et Clematis 165. 2000; M.
Johnson, Klematis 347. 1997.——C. tenuifolia Royle, Ill. Bot. Himal. Mts. 51. 1834. Type:
Kashmir Region. Without precise locality, Royle s.n. (holotype, LIV).
?C. baltistanica Qureshi & Chaudhri in Pakist. Syst. 2: 11. 1978. Type: unknown.
光枝东方铁线莲
This variety differs from var. orientalis by having glabrous branches and leaflets, and
sepals glabrous or subglabrous outside.
Branches glabrous. Leaflets up to 4 cm long, glabrous, undivided or 2–3-parted, terminal
lobe linear-lanceolate, 3–6 mm broad, entire or 1-dentate per side, lateral lobes much smaller.
Cymes 3–many-flowered; peduncles 1.5–6.7 cm long, 0.8–1 mm in diam. Sepals
oblong-lanceolate, 13–16 mm long, inside puberulous, outside glabrous or subglabrous.
Persistent styles 2.5–3.8 cm long.
Afghanistan, China (SW Xinjiang), and Kashmir region.
Additional specimens examined:
Afghanistan. Without precise locality, Griffith 1381, 1382, 1383 (K).
China. Xinjiang (新疆): Pamir (帕米尔), C. Persson 228, 522 (S).
Kashmir region. Karakoram, Polunin 6453 (BM).
11f. var. baluchistanica Grey-Wilson in Kew Bull. 44: 40. 1989; et Clematis 165. 2000; M.
Johnson, Klematis 345. 1997. Type: Pakistan. Baluchistan, Urak Zarghoun, alt. 2350 m,
1952-06, Crookshank 373 (holotype, K!).
This variety differs from var. orientalis by having leaflets with terminal lobe linear or
linear-lanceolate, 1–4(–5) mm broad.
Branches sparsely puberulous. Leaflets up to 3 cm long, 2–3-sect, terminal lobe linear or
linear-lanceolate, 1–4(–5) mm broad, entire or 1-dentate per side, lateral lobes much smaller.
Cymes 3–9-flowered; peduncles 1–6.4 cm long, 0.8–1 mm in diam. Sepals narrowly oblong,
8–11 mm long, puberulous on both surfaces. Persistent styles 2–4 cm long.
N Pakistan and possibly SW Afghanistan.
Additional specimen examined:
Pakistan. Without precise locality, Aitchison 772 (GH).
11g. var. sinorobusta W. T. Wang in Acta Phytotax. Sin. 36: 167. 1998.——C. orientalis L.
var. robusta W. T. Wang in l.c. 29: 466. 1991, non Grey-Wilson, 1989; M. Johnson, Klematis
346. 1997. Type: China. Xinjiang (新疆): Yecheng (叶城), Yiliki (伊力克), alt. 3800 m,
1987-09-15, B. S. Li et al. (李勃生等) 11557 (holotype, PE!).
This variety differs from var. orientalis by having 1-flowered axillary cymes with short,
thick, robust peduncles.
Branches only on nodes sparsely puberulous, elsewhere glabrous. Leaf-lets
linear-lanceolate, 3–7 cm long, 0.4–0.8 cm wide, entire, undivided or above base 3-lobed,
terminal lobe much longer than lateral ones, subglabrous. Cymes axillary, 1-flowered;
peduncles 2–5 mm long, up to 2 mm in diam. Persistent styles 2–2.8 cm long.
China (SW Xinjiang).
No. 4 WANG Wen-Tsai: A revision of Clematis sect. Meclatis (Ranunculaceae) 433


Fig. 10. A, B, Clematis corniculata W. T. Wang. A, upper part of flowering branch; B, stamen. Drawn from K. Guo
12158. C, D, C. caudigera W. T. Wang. C, upper part of flowering branch; D, stamen. Drawn from B. S. Li 11741.


An unclear variety:
Clematis orientalis L. var. obtusifolia Hook. f. & Thoms., Fl. Ind. 9. 1855. Type: no type
specimen designated.
While visiting the Herbarium K in 1999, I failed to find out any specimen identified as
this variety by Hooker & Thomson. Hence the identity of this variety cannot be yet
determined.
12. Clematis caudigera W. T. Wang in Acta Phytotax. Sin. 36: 165, fig. 2: 5, 6. 1998;
Grey-Wilson, Clematis 165. 2000; W. T. Wang & Barth. in Fl. China 6: 365. 2001. Type:
Acta Phytotaxonomica Sinica Vol. 44 434
China. Xinjiang (新疆): Yutian (于田), Fulu (甫鲁), 1988-06-25, B. S. Li (李勃生) 11741
(holotype, PE!).
尾尖铁线莲 Fig. 10: C, D
Woody vine. Branches shallowly 4-sulcate, glabrous. Leaves pinnate, 5-foliolate; leaflets
grey-green, thickly papery or coriaceous, triangular in outline, 0.8–2×0.4–1.6 cm, base
cordate-truncate or subcordate, near base 3-parted or 3-sect, terminal lobe narrowly
rhombic-lanceolate or linear-lanceolate, 1–6 mm broad, margin 1–2-denticulate per side,
seldom entire, lateral lobes smaller, obliquely cuneate, unequally 2-lobulate, adaxially
glabrous, abaxially sparsely appressed-puberulous, basal veins nearly flat, inconspicuous;
petioles 2.2–4.5 cm long, sparsely puberulous. Flowers solitary, terminal, 3.5–6 cm in diam.;
pedicels 9–21.5 cm long, glabrous or sparsely puberulous. Sepals 4, yellow, ascending, papery,
oblong-lanceolate, 2.3–4.2×0.8–1.1 cm, inside densely appressed-puberulous, on margin
velutinous, outside glabrous, apex attenuate with tail-like projections 3–6 mm long. Stamens
8–12 mm long; anthers oblong or narrowly oblong, 2.4–3 mm long, glabrous, apex obtuse.
Ovaries pubescent; styles 11–13 mm long, densely villous. Fl. Jun.
China (SW Xinjiang). On slopes in valley; alt. 3000–3700 m.
13. Clematis corniculata W. T. Wang in Acta Phytotax. Sin. 29: 466, fig. 3: 1, 2. 1991; M.
Johnson, Klematis 334. 1997; Grey-Wilson, Clematis 165. 2000; W. T. Wang & Barth. in Fl.
China 6: 366. 2001. Type: China. Xinjiang (新疆): Yecheng (叶城), Supikiya (苏皮克牙), alt.
2800 m, 1987-09-01, D. Zheng (郑度) K102 (holotype, PE!).
角萼铁线莲 Fig. 10: A, B
Woody vine. Branches shallowly 6-sulcate, glabrous. Leaves pinnate, glabrous; leaflets
grey-green, coriaceous, lanceolate, narrowly ovate or narrowly lanceolate, 1.5–3×0.3–0.8 cm,
base broadly cuneate or cuneate, margin usually entire, sometimes 1-denticulate, undivided or
2–3-lobed or 2–3-sect, terminal lobe larger, linear-lanceolate, 2–4 mm broad, lateral lobes
much smaller, narrowly ovate, 0.3–1 cm long, midrib abaxially slightly prominent; petioles ca.
7 cm long. Flowers solitary, terminal, ca. 4 cm in diam.; pedicels 14–20 cm long, glabrous.
Sepals 4, yellow, ascending, papery, oblong, 2.3–2.6×0.7–0.9 cm, outside below apex
corniculate (hornlike projections ca. 2.5 mm long), glabrous on both surfaces, outside on
margin velutinous. Stamens 8–9 mm long; anthers oblong, ca. 2 mm long, glabrous, apex
minutely apiculate. Ovaries pubescent; styles ca. 10 mm long, densely villous. Fl. Aug. –Sept.
China (SW Xinjiang). On grassy slopes; alt. 2800–2930 m.
Additional specimen examined:
China. Xinjiang (新疆): Qira (策勒), K. Guo (郭柯) 12518 (PE).
Acknowledgements I would like to express my sincere thanks to the directors and curators
of BM, G, GH, HIMC, K, LE, MO, NAS, P, S, TI, TIE, UPS, US, and WUK for kindly
inviting me to visit their herbaria and/or sending herbarium material on loan; to David E.
Boufford & Zhu Guang-Hua for sending me the book An Illustrated Encylopedia of Clematis
as a gift; to LI Liang-Qian, QIN Hai-Ning, ZHANG Zhi-Yun, ZHU Xiang-Yun, WANG
Zhong-Tao, BAN Qin, and XIE Lei for various kinds of help in the course of research; and to
SUN Ying-Bao for making the drawings.
References
Baillon H. 1867. Clematites. Histoire des plantes. Paris. 1: 52–62.
Brandenburg W A. 2000. Meclatis in Clematis: Yellow flowering Clematis species. Systematic studies in
Clematis L. (Ranunculaceae), inclusive of cultonomic species. Wageningen: Wageningen Universiteit.
De Candolle A P. 1818. Regni Vegetabilis Systema Naturale. Paris. 1: 131–167.
Essig F B. 1991. Seedling morphology in Clematis (Ranunculaceae) and its taxonomic implications. Sida 14:
377–390.
No. 4 WANG Wen-Tsai: A revision of Clematis sect. Meclatis (Ranunculaceae) 435
Grabovskaya A E. 2000. Ranunculaceae. In: Grubov V I ed. Catalogue of the Type Specimens of Central Asian
Vascular Plants in the Herbarium of the V. L. Komarov Botanical Institute (LE). St. Petersburg: St.
Petersburg University Press. 189–200.
Greuter W, McNeill J, Barrie F R, Burdet H M, Demoulin V, Filgueiras T S, Nicolson D H, Silva P C, Skog J
E, Trehane P, Turland N J, Hawksworth D L. 2000. International Code of Botanical Nomenclature (St.
Louis Code). Königstein: Koeltz Scientific Books.
Grey-Wilson C. 1989. Clematis orientalis (Ranunculaceae) and its allies. Kew Bulletin 44: 33–60.
Grey-Wilson C. 2000. Clematis the Genus. Oregon, Portland: 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.
Johnson M. 1997. Släktet Klematis. Södertälje: M Johnsons Plantskola AB.
Kuntze O. 1885. Monographie der Gattung Clematis. Verhandlungen des Botanischen Vereins für
Brandenburg (Abhandlungen) 26: 83–203.
Ling P-P (凌苹苹). 1980. Clematis sect. Meclatis. In: Flora Reipublicae Popularis Sinicae (中国植物志).
Beijing: Science Press. 28: 139–147.
Linnaeus C. 1753. Species Plantarum. Stockholm.
Prantl K. 1888. Beiträge zur Morphologie und Systematik der Ranunculaceen. Botanische Jahrbücher 9:
225–273.
Rehder A. 1920. New species, varieties and combinations from the herbarium and collections of the Arnold
Arboretum. Journal of the Arnold Arboretum 1:191–210.
Rehder A. 1951. Clematis. Manual of Cultivated Trees and Shrubs. 2nd ed. New York: The Macmilan
Company. 206–220.
Rehder A, Wilson E H. 1913. Clematis. In: Sargent C S ed. Plantae Wilsonianae. Cambridge: The University
Press. 1: 319–343.
Schneider C K. 1906. Clematis L. Illustriertes Handbuch der Laubholzkunde. Jena: Verlag von Gustav Fisher.
1: 273–294.
Spach E. 1839. Trib. Clematideae. Histoire des Naturelle Végetaux. Phanérogarmes. Paris. 7: 257–284.
Tamura M. 1955. Systema Clematidis Asiae orientalis. Osaka University Science Reports. 4: 43–55.
Tamura M. 1967. Morphology, ecology and phylogeny of the Ranunculaceae. Vll. Osaka University Science
Reports 16: 21–43.
Tamura M. 1987. A classification of genus Clematis. Acta Phytotaxonomica et Geobotanica 38: 33–44.
Tamura M. 1995. Clematis L. In: Hiepko P ed. Engler’s Die Natürlichen Pflanzenfamilien. Zweite Aufl.
Berlin: Dunker & Humbolt. 17a (4): 368–387.
Toomey M, Leeds E. 2001. An illustrated encyclopedia of Clematis. Oregon, Portland: Timber Press.
Wang W-T (王文采). 1989. Notes on disjunction in the flora of China. Bulletin of Botanical Research (植物研
究), Harbin 9: 1–16.
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). 2004. A revision of Clematis sect. Brachiatae (Ranunculaceae). Acta Phytotaxonomica
Sinica (植物分类学报) 42: 289–332.
Wang W-T (王文采), Bartholomew B. 2001. Clematis L. In: Wu Z-Y, Raven P H eds. Flora of China. Beijing:
Science Press; St. Louis: Missouri Botanical Garden Press. 6: 333–386.
Wang W-T (王文采), Li L-Q (李良千). 2005. A new system of classification of the genus Clematis
(Ranunculaceae). Acta Phytotaxonomica Sinica (植物分类学报) 43: 431–488.
铁线莲属黄花铁线莲组修订
王文采
(系统与进化植物学国家重点实验室, 中国科学院植物研究所 北京 100093)

摘要 对毛茛科铁线莲属Clematis的黄花铁线莲组sect. Meclatis进行了全面修订, 确定此属含13种和13
变种(包括1新变种和2新变种等级); 写出了此组的分类学简史和地理分布, 并对其在铁线莲属中的系
Acta Phytotaxonomica Sinica Vol. 44 436
统位置和组内诸种的亲缘关系进行了讨论; 还写出了此组的分种、分变种检索表, 以及各种植物的形态
描述、地理分布、生长环境等, 并附有各种的插图。此组的花构造与对枝铁线莲组sect. Brachiatae的近
似, 与后者在亲缘关系上相近, 区别在于此组的萼片通常斜上方开展, 呈黄色, 被毛的花丝下部变宽,
呈狭披针形, 而在对枝铁线莲组, 萼片水平开展, 呈白色, 被毛的雄蕊花丝呈狭条形, 下部不变宽; 二
组可能均起源于欧洲铁线莲组的sect. Clematis subsect. Clematis, 因此, 均应是隶属欧洲铁线莲亚属
subgen. Clematis的成员。根据对此组植物形态特征的分析, 观察到以下演化趋势: (1)叶的颜色由于适应
干旱气候, 由绿色变为灰绿色; (2)卵形或宽卵形、掌状分裂、边缘具齿的小叶可能是原始的特征, 而披
针形或条形、不分裂、全缘的小叶是衍生的特征; (3)单独、顶生、只是花梗的花是由具花序梗和二苞
片的聚伞花序发生减化(reduction)而衍生的; (4)萼片形状的演化趋势与小叶形状的演化趋势近似, 也由
卵形演变到披针形或条形; (5)萼片内面无毛是原始现象, 而被毛则是衍生现象; (6)萼片顶端无突起是
原始现象, 出现突起则为衍生现象; (7)花药形状由长圆形演变到狭长圆形和条形。根据上述演化趋势,
推测具较多原始特征的甘川铁线莲C. akebioides和甘青铁线莲C. tangutica为此组的原始种, 而具较多
衍生特征的尾尖铁线莲C. caudigera和角萼铁线莲C. corniculata为此组的进化种。组成世界屋脊的青藏
高原西缘、帕米尔高原和邻近山地集中分布有此组10种(包括7特有种), 当是此组的分布中心; 而甘川
铁线莲和甘青铁线莲二种分布区的主要重叠部分所在的青藏高原东缘则可能是此组的起源中心。过去,
一些铁线莲属专家将属于欧洲铁线莲组的C. ispahanica Boiss.和属于对枝铁线莲组的C. graveolens
Lindl.误置于黄花铁线莲组中, 对此, 本文予以纠正。
关键词 铁线莲属; 黄花铁线莲组; 分类学修订