全 文 ：植 物 分 类 学 报 45 (4): 425–457(2007) doi:10.1360/aps06114
Acta Phytotaxonomica Sinica http://www.plantsystematics.com
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Received: 18 July 2006 Accepted: 30 November 2006
Supported by the National Natural Science Foundation of China, Grant No. 30470126.
* In this paper, the second section (Pollen morphology) is written by XIE Lei, and the other four sections by WANG Wen-Tsai.
A revision of Clematis sect. Tubulosae (Ranunculaceae)*
WANG Wen-Tsai XIE Lei
(State Key Laboratory of Systematic and Evolutionary Botany, Institute of Botany, the Chinese Academy of Sciences,
Beijing 100093, China)
Abstract Clematis sect. Tubulosae is revised in this paper. Nine species, two varieties, and
three forms are recognized and classified into two subsections. An identification key is
provided, and each species is described and illustrated. Brief taxonomic history is given,
along with a summary of pollen morphology and geographical distribution. The relationships
among the infrasectional groups are also discussed: Subsect. Pinnatae, characterized by its
scandent habit, bisexual flowers, white or pinkish, at length spreading, obovate-oblong sepals,
and tricolpate pollen, is regarded as the more primitive group, whereas subsect. Tubulosae,
characterized by its erect habit, usually polygamous flowers, blue or purple, erect, usually
narrowly oblong sepals, and usually pantoporate pollen, is regarded as the more advanced
group of the section. Subsect. Pinnatae is believed to have originated from sect. Clematis in
Central or East China, and subsect. Tubulosae might be derived from subsect. Pinnatae.
Key words Clematis, sect. Tubulosae, Ranunculaceae, taxonomic review.
1 Brief taxonomic history
The first species of Clematis L. sect. Tubulosae Decne. known to science is C.
heracleifolia DC., which was described in 1818 by de Candolle on the basis of a flowering
specimen collected by an English officer, G. L. Staunton, from the north of Beijing, China in
1793. This species was placed in the large and heterogeneous sect. Flammula by de Candolle.
Afterwards, Turczaninow (1837) and Siebold and Zuccarini (1845) described two new species
closely allied to C. heracleifolia, C. tubulosa Turcz. and C. stans Sieb. & Zucc., from Beijing,
China and Honshu, Japan, respectively.
In 1877, in a paper dealing with the genus Clematis of China and Japan, Maximowicz
described two new species, C. tatarinowii Maxim. and C. pinnata Maxim., on the basis of two
gatherings collected from Beijing by A. A. Tatarinow. Maximowicz placed C. tatarinowii near
C. tubulosa based on the fact that they both have erect sepals, and placed C. pinnata near C.
brevicaudata DC. and C. grata Wall., two members of sect. Clematis subsect. Clematis (Wang,
2003), according to the fact that all three species have spreading sepals. Maximowicz (1877)
did not, however, notice that the stamen filaments of his two new species bear a few hairs near
apex. Besides, in his enumeration of the Asian Clematis species, C. heracleifolia was not
included, and in an annotation under C. tubulosa, Maximowicz cast doubt upon the identity of
this species with C. tubulosa.
In 1867, Verlot published a new species, Clematis davidiana, named by M. J. Decaisne,
on the basis of flowering specimens collected by P. A. David from Beijing in 1863. In 1881,
Decaisne provided a revision of the species of the C. tubulosa group, and established a new
section to accommodate them, sect. Tubulosae. In it, seven species were recognized (viz. C.
tubulosa, C. stans, C. davidiana, and four new species, C. hookeri Decne., C. kousabotan
Decne., C. lavallei Decne., and C. savatieri Decne.); C. heracleifolia was not included. In fact,
however, C. heracleifolia was misidentified by Verlot (1867) as C. tubulosa, and in 1937, each
of the five new species named by Decaisne was reduced into synonymy by Kitagawa under
Acta Phytotaxonomica Sinica Vol. 45 426
one of three species, C. heracleifolia, C. tubulosa, and C. stans.
In a paper dealing with four Chinese species of Clematis, Forbes (1884) concluded that
C. heracleifolia and C. tubulosa were the same entity, as previously suspected by
Maximowicz (1877), and relegated C. tubulosa to the synonymy under C. heracleifolia. This
treatment was subsequently adopted by many authors, including Forbes and Hemsley (1886),
Finet and Gagnepain (1903), Rehder and Wilson (1913), Handel-Mazzetti (1939), Anonymous
(1972), Fang (1980), and Wang and Bartholomew (2001), sustaining a taxonomic confusion
that has lasted for more than one hundred and twenty years.
In his monograph of the genus, Kuntze (1885) for the first time associated C. pinnata
with C. heracleifolia based on the fact that the sepals of these two species at the beginning of
anthesis are erect and closely connivent, and during anthesis gradually separate from top
downwards. These two species were placed in his sect. 3 Escandentes. Kuntze (1885) also
treated C. tatarinowii as a variety under C. pinnata, and recognized several other previously
described species as infraspecific taxa within C. heracleifolia, including four treated as
subspecies (C. davidiana, C. stans, C. lavallei, and C. savatieri) and one as a variety (C.
tubulosa).
Prantl (1888), in his classification of Clematis, followed Kuntze by continuing the
association of C. heracleifolia and C. stans with C. tatarinowii, placing them in sect. Viorna 4
Tubulosae, although he did not mention C. pinnata. At the same time, in the infrasectional
group Tubulosae, Prantl misplaced the species of the C. connata group, which are now
regarded as close allies of the C. viorna group (Wang & Li, 2005).
In 1907, Makino described a new species, C. takedana Makino, based on a specimen
collected from Shinano, Japan by H. Takeda, regarding this new species as a hybrid between
C. apiifolia DC., a member of sect. Clematis (Wang, 2003), and C. stans, a member of sect.
Tubulosae. Makino’s viewpoint was accepted by Kitagawa (1937), Ohwi (1965), and Tamura
(1982). Unfortunately, Makino was unaware of the fact that C. takedana is a close ally of the
Chinese C. pinnata (Wang, 2001; Wang & Li, 2005).
In 1937, Kitagawa published the second revision of the C. tubulosa group, in which he
recognized seven species, C. speciosa (Makino) Makino, C. stans, C. heracleifolia, C.
tubulosa, C. tsugetorum Ohwi, C. urticifolia Nakai ex Kitagawa and C. psilandra Kitagawa,
the latter two of which were described as new. In his revision, the morphological differences
between C. heracleifolia and C. tubulosa were clearly given, and the confusion about the
identities of these two species created by Decaisne (1881), Forbes (1884), Forbes & Hemsley
(1886), Finet & Gagnepain (1903), and Rehder & Wilson (1913) was clarified. As to the
relationships of the C. tubulosa group, Kitagawa (1937: p. 344) stated that “this group is
phyletically old and its evolution had either stopped already or is just stopping. One can hardly
trace back its phyletical connection with other sections, nor can make out the same of each
species”. Kitagawa’s above point of view appears to show that he was ignorant of the presence
of a group closely related to C. tubulosa group and consisting of C. pinnata and C. takedana,
which he followed Makino to treat as a hybrid between C. apiifolia and C. stans as mentioned
above.
Two years later, in his revision of the Chinese Clematis, Handel-Mazzetti (1939) adopted
Prantl’s treatment of the C. heracleifolia group, and placed C. heracleifolia and C. tatarinowii
in sect. Viorna subsect. Tubulosae. However, he erroneously referred C. pinnata to sect.
Flammula DC. subsect. Vitalbae Prantl (=sect. Clematis subsect. Clematis—Wang, 2003). His
misplacement of C. pinnata was later adopted by Ting (1980), Johnson (1997), and
Grey-Wilson (2000). Moreover, Handel-Mazzetti (1939) did not agree with Kitagawa (1937)
in recognizing C. heracleifolia and C. tubulosa as distinct species, and instead followed
Rehder & Wilson (1913) by maintaining the reduction of C. tubulosa into the synonymy under
C. heracleifolia, as mentioned above.
No. 4 WANG & XIE: A revision of Clematis sect. Tubulosae (Ranunculaceae) 427
In the classification of Clematis published by Tamura (1995), sect. Tubulosae, placed near
the C. connata group (sect. Campanella Tamura), comprises only the species of the C.
tubulosa group, and does not include C. pinnata or C. takedana. Tamura’s sect. Tubulosae s.s.
was adopted by Snoeijer (1992) and elevated to subgeneric rank by Yang (Yang & Moore,
1999).
In the monograph of Clematis published by Johnson (1997), sect. Tubulosae was likewise
placed near the C. connata group, as had been done by Tamura (1995), and was circumscribed
to contain C. tatarinowii as well as the seven species recognized by Kitagawa (1937); C.
pinnata was erroneously referred to sect. Clematis as Handel-Mazzetti (1939) had done.
Johnson’s treatment of sect. Tubulosae was adopted by Grey-Wilson (2000), although the
group was elevated to subgeneric rank.
In 2001, Wang classified the species of sect. Tubulosae into two subsections, subsect.
Pinnatae, comprising C. pinnata, C. takedana, and C. tatarinowii, and subsect. Tubulosae,
including the seven species recognized by Kitagawa (1937).
In 2003, in the course of field studies in the hilly regions of Beijing, Shi① found that C.
pinnata is highly variable in leaf division, and can not be distinguished from C. tatarinowii. In
her master degree dissertation, this author reduced C. tatarinowii Turcz. and C. pinnata var.
ternatifolia W. T. Wang into the synonymy under C. pinnata. In 2005, in a paper by Xie et al.,
this reduction was formally published.
2 Pollen morphology
2.1 Materials and methods
Pollen grains from seven species and two varieties of Clematis sect. Tubulosae were
investigated using Scanning Electron Microscopy (SEM) (Table 1), of which four species and
two varieties were examined for the first time, i.e. Clematis takedana, C. urticifolia, C.
speciosa, C. tubulosa var. tubulosa, C. tubulosa var. ichangensis, and C. stans var.
austrojapanensis. Pollen samples were obtained from the following collections: the Herbarium
of Institute of Botany, the Chinese Academy of Sciences (PE), the Herbarium of Kagoshima
University Museum (KAG), and the Herbarium of Kyoto University (KYO). Pollen grains
were prepared from herbarium material without special treatment. Anthers were broken to
release the pollen directly onto aluminium stubs, sputtered with gold, and then observed and
photographed using a Hitachi S-800 SEM unit. The values of P (polar axis length) and E
(equatorial diameter) were measured, and means were calculated based on examination of 20
pollen grains. Descriptive terminology follows Clarke et al. (1991).
2.2 Results
The pollen characters of all taxa examined are summarized based on observations from
SEM (Table 1).
Pollen grains of taxa assigned to sect. Tubulosae are of two types, one tricolpate and the
other pantoporate. The pollen is radially symmetrical, isopolar, and spheroidal to prolate with
broad or relatively narrow poles. The grains are small to medium-sized, P (polar axis length)
20.1–24.8 μm×E (equatorial diameter) 15.4–22.8 μm for tricolpate pollen, with a diameter
from 18.7 μm to 22.6 μm for pantoporate pollen. They are elliptical to circular in equatorial
outline, and their surface is uniformly ornamented with microechinate (conical spinules) that
are evenly distributed on the microperforate tectum. The apertures are tricolpate or
pantoporate, and the colpi are narrow or broad with distinct and straight, but incised margins,
sunken or open in tricolpate grains. The colpus membrane is often covered with granular or
microechinate elements usually more or less of the same size or larger than those on the

① Shi J-H (史京华). 2003. A preliminary study of the hybrid origin of Clematis pinnata Maxim. (Ranunculaceae). Master’s Degree
Dissertation. Beijing: Institute of Botany, the Chinese Academy of Sciences.
Acta Phytotaxonomica Sinica Vol. 45 428
Table 1 Pollen characteristics in Clematis sect. Tubulosae
Taxon Type of
aperture
Size (polar ×
equatorial) (μm)
Shape* Colpus (in
width) or
pore (in
diam.) (μm)
End of
corpus
Colpus or
pore
membrane
Figure Locality Voucher
C. pinnata tricolpate (21.3–24.8) 22.9×
(20.2–22.8) 21.6
SP 2–3
(in width)
acute dense conical
spinules
1: A, B Baihuashan,
Beijing, China
S. Y. Ho
15037 (PE)
C. takedana tricolpate (20.3–23.7) 21.6×
(20.1–21.8) 20.2
SP 2–3
(in width)
acute dense conical
spinules
1: C, D Mt.Shirouma-
yama,
Shinano,
Japan
G.
Koidzumi
s.n. (KYO)
C.
heracleifolia
tricolpate (20.1–24.5) 23.6×
(15.4–19.1) 17.3
P 3–4
(in width)
acute dense conical
spinules
1: E, F Ji Xian,
Tianjin, China
T. N. Liou
4521 (PE)
C. tubulosa
var.
tubulosa
pantoporate (18.5–22.4) 20.6 S 3–5
(in diam.)
– dense conical
spinules
2: C, D Dalian,
Liaoning,
China
Z. Wang &
Y. X. Liu
889 (PE)
C. tubulosa
var.
ichangensis
pantoporate (19.5–22.1) 21.0 S 3–5
(in diam.)
– dense conical
spinules
2: A, B Hua Shan,
Shaanxi,
China
K. S. Hao
3973 (PE)
C. urticifolia pantoporate (19.7–22.3) 21.8 S 4–7
(in diam.)
– dense conical
spinules
2: E, F Toji-Myun,
Mt. Chiri,
Chollanamdo,
Korea
Boufford et
al. 25833
(PE)
C. speciosa pantoporate (18.7–22.1) 20.9 S 4–7
(in diam.)
– dense
hemispherical
spinules or
conical
spinules
1: K, L Mt. Dodake,
Hyuga, Japan
S Hatusima,
et al. 22497
(KAG)
C. stans var.
stans
pantoporate (20.5–21.8) 21.1 S 3–4
(in diam.)
– dense conical
spinules
1: G, H Shizuoka,
Japan
F. Chikata
15778 (PE)
C. stans var.
austrojapo-
nensis
pantoporate (19.4–22.6) 21.4 S 4–5
(in diam.)
– dense conical
spinules
1: I, J Mt. Shiraiwa,
Hyuga, Japan
S Hatusima
& S Sako
26198
(KYO)
* S, spheroidal; SP, subprolate; P, prolate.

tectum. On pantoporate grains, the pores are irregular in shape and distribution, often obscure.
The membrane of the pores is often covered with granular or microechinate elements usually
the same size or larger than those on the tectum (Figs. 1, 2).
Among the members of sect. Tubulosae, all except C. pinnata, C. takedana and C.
heracleifolia have pantoporate pollen grains, which represent the relatively advanced pollen
type in Clematis (Xie, 2005),whereas C. pinnata, C. takedana and C. heracleifolia show a
primitive pollen type. Pollen morphology is thus, at least to some extent, systematically
significant in sect. Tubulosae.
3 Relationships among the infrasectional groups
The species of sect. Tubulosae fall distinctly into two groups that are easily
distinguishable on the basis of habit and floral structure.
The first group, subsect. Pinnatae, consists of two closely allied species, the Chinese C.
pinnata and the Japanese C. takedana, in which the stems are woody, scandent or prostrate, the
leaves are 1–2-ternate or 1–2-pinnate, the flowers are bisexual, arranged in panicle-like
compound cymes, the sepals are white or pinkish in colour, obovate-oblong in outline, at
___________________________________________________________________________________________________
→
Fig. 1. A, B, Clematis pinnata. A, Equatorial view. B, Showing microechinate ornamentation. C, D, C. takedana. C,
Equatorial view. D, Details of ornamentation. E, F, C. heracleifolia. E, Equatorial view. F, Details of ornamentation.
G, H, C. stans var. stans. G, Whole pollen grain view. H, Details of ornamentation. I, J, C. stans var. austrojapanensis.
I, Whole view of pollen grain. J, Details of ornamentation. K, L, C. speciosa. K, Whole view of pollen grain. L, Details
of ornamentation.
Scale bar: A, C, E, I, K, 8.6 μm; B, D, F, H, J, L, 3 μm; G, 10 μm.
No. 4 WANG & XIE: A revision of Clematis sect. Tubulosae (Ranunculaceae) 429

Acta Phytotaxonomica Sinica Vol. 45 430

Fig. 2. A, B, C. tubulosa var. ichangensis. A, Whole view of pollen grain. B, Details of ornamentation. C, D, C.
tubulosa var. tubulosa. C, Whole view of pollen grain. D, Details of ornamentation. E, F, C. urticifolia. E, Whole view of
pollen grain. F, Details of ornamentation.
Scale bar: A, C, E, 8.6 μm; B, D, F, 3 μm.

apex, and the pollen grains are tricolpate① (Xie, 2005).
The seven species of the second group, subsect. Tubulosae, are erect perennial herbs,
small subshrubs or shrubs, with usually ternate, rarely pinnate leaves. Their floral structure is
essentially similar to that of subsect. Pinnatae, differing in the usual abortion of one whorl of
sexual organs, in their sepals being usually blue or purple in colour, always erect, usually
narrowly oblong in outline, more closely connivent during anthesis, and only being recurved
above after anthesis. Moreover, their pollen grains are usually pantoporate, rarely tricolpate①
(Yang & Huang, 1992; Nowicke & Skvarla, 1995; Xie, 2005), as shown in Figs. 1, 2 and Table
1.
In subsect. Tubulosae, most species are perennial herbs or subshrubs, and have ternate
leaves and large, terminal, many-flowered panicle-like compound cymes with clustered
flowers. Only one species, C. tsugetorum, an endemic of Taiwan Island, China, is a small
shrub with ternate or pinnate leaves and solitary terminal flowers (the solitary terminal flowers
may result from the reduction of the many-flowered compound cymes due to the adaptation to
the harsh alpine climate); most species have polygamous or unisexual flowers and pantoporate

① Shi J-H (史京华). 2003. A preliminary study of the hybrid origin of Clematis pinnata Maxim. (Ranunculaceae). Master’s
Degree Dissertation. Beijing: Institute of Botany, the Chinese Academy of Sciences.
Zhang Y-L (张镱锂). 1987. Pollen morphology of some sections of Clematis in China and its taxonomic significance. Master’s
Degree Dissertation. Beijing: Beijing Normal University.
No. 4 WANG & XIE: A revision of Clematis sect. Tubulosae (Ranunculaceae) 431
pollen grains, only C. heracleifolia has bisexual flowers and tricolpate pollen grains. Also,
most species have narrowly oblong sepals that are slightly dilated near apex after anthesis,
whereas only C. urticifolia has lanceolate undilated sepals and only C. tubulosa var. tubulosa
has sepals strongly dilated above after anthesis.
From the morphological and palynological characters mentioned above several
evolutionary trends can be seen within sect. Tubulosae. A scandent habit, ternate leaves,
many-flowered cymes, bisexual flowers, white, undilated or slightly dilated sepals, and
tricolpate pollen grains may represent the primitive states, while an erect habit, 1–2-pinnate
leaves, solitary, terminal, polygamous or unisexual flowers, blue or purple, strongly dilated
sepals, and pantoporate pollen grains may represent the derived states.
Clematis pinnata, a member of subsect. Pinnatae, was misplaced in sect. Clematis by
Handel-Mazzetti (1939) and other botanists, as mentioned above. In fact, C. pinnata shows a
striking resemblance to C. grata and C. brevicaudata of sect. Clematis in habit, floral
structure, and pollen morphology (Xie, 2005), differing from them only in its sepals being
suberect at the beginning of anthesis and in having pilose stamen filaments. The misplacement
of C. pinnata in sect. Clematis and the resemblance of C. pinnata to species of sect. Clematis
may thus imply that subsect. Pinnatae is the more primitive group of sect. Tubulosae, and
might have been derived from sect. Clematis, and that the two close allies of this subsection,
C. pinnata and C. takedana, might be sister taxa derived from a species (probably C. apiifolia
DC.) of sect. Clematis. This speculation regarding the derivation of sect. Tubulosae from sect.
Clematis is supported by a recent palynological study of the genus (Xie, 2005). As to the
systematic position of subsect. Tubulosae, it is believed to be the more advanced group of sect.
Tubulosae in having unisexual flowers, blue or purple, narrowly oblong sepals, pantoporate
pollen grains, and other advanced features, and might be derived from subsect. Pinnatae
(Wang, 2001; Wang & Li, 2005).
Clematis takedana was regarded as a hybrid between C. apiifolia (Sect. Clematis) and C.
stans (sect. Tubulosae) by Makino (1907) and several later botanists, as mentioned above.
Interestingly, this interpretation is not unique and has other counterparts within the genus. For
example, in 2003, Shi① suggested that C. pinnata might be a hybrid between C. brevicaudata
(sect. Clematis) and C. heracleifolia (sect. Tubulosae). As indicated above, C. pinnata and C.
takedana have similar habit and floral structure, which clearly shows that they are closely
related. It is thus difficult to interpret why these two closely related species represent hybrids
produced from different parentages occurring in different regions, and further studies are
needed to resolve this issue.
4 Geographical distribution
As indicated above, Clematis sect. Tubulosae, widespread in eastern Asia, is here defined
to consist of nine species, two varieties, and three forms.
The more primitive group within the section, subsect. Pinnatae, consists of two species.
Regarding C. pinnata, most populations are concentrated to the northern and western mountains
surrounding the city of Beijing and in several neighboring counties of Hebei Province and
Tianjin, with two outliers in Shenyang, Liaoning Province and in Harbin, Heilongiang
Province, respectively (Fig. 3). This curious distribution pattern appears to imply that the past
range of C. pinnata might have been larger than at present, being widespread in North and
Northeast China, and that following the Quaternary glacial periods it was strongly reduced,
with the northern populations remaining in two refugia. The second species, C. takedana,

① Shi J-H (史京华). 2003. A preliminary study of the hybrid origin of Clematis pinnata Maxim. (Ranunculaceae). Master’s Degree
Dissertation. Beijing: Institute of Botany, the Chinese Academy of Sciences.
Acta Phytotaxonomica Sinica Vol. 45 432

Fig. 3. Map showing the distribution of the two species of subsect. Pinnatae.

is restricted to central and northern Honshu Island, Japan (Ohwi, 1965). Given that C. pinnata
and C. takedana are close allies and that the islands of Japan have been separated from the
Asian continent since the Miocene (Byльф, 1944), the senior author of this paper surmises that
these two taxa may be sister groups and might have originated in Central or East China before
the Miocene from a common ancestor belonging to sect. Clematis, whose present center of
distribution is situated in Southwest China (Wang, 2003).
The more advanced subsect. Tubulosae consists of seven species, two varieties, and one
form, ranging from the middle and lower reaches of the Changjiang and Huanghe Rivers,
southern Nei Mongol, Liaoning and Taiwan Provinces of China eastward via Korea to Japan
(Fig. 4). The most primitive species of this subsection, C. heracleifolia, is widespread in the
middle and lower parts of the Huanghe River, southern Nei Mongol, and Liaoning Province of
China and in northern Korea. Elsewhere in Asia two other species occur, C. tubulosa (with var.
tubulosa occurring in North China and northern Korea, and var. ichangensis widespread in
middle and lower reaches of the Changjiang River and Huanghe River) and C. urticifolia
occurring in Korea. All three of these species inhabit hills or mountains at low to moderate
altitudes. On the other hand, the two species endemic to Taiwan Island are found in higher
montane or alpine regions: C. psilandra occurs from 1000 to 2500 m above sea level, and the
most advanced species of subsect. Tubulosae, C. tsugetorum, grows from 2400 to 3600 m.
Three species occur in Japan, C. urticifolia (Sado Island—Ohwi, 1965), and two endemics, C.
speciosa and C. stans (Fig. 4). The latter, with its unisexual flowers and dioecious individuals,
may be one of the advanced species in subsect. Tubulosae.
On the basis of a rececnt palynological study of the genus Clematis, Xie (2005) pointed
out that sect. Tubulosae might have originated in eastern Asian and subsequently extended
eastward to Japan. The senior author of this paper would agree with this idea.
No. 4 WANG & XIE: A revision of Clematis sect. Tubulosae (Ranunculaceae) 433

Fig. 4. Map showing the distribution of the species and varieties of Clematis subsect. Tubulosae. The distribution of C.
tubulosa var. tubulosa, C. urticifolia, C. speciosa, and C. stans in Korea and Japan is based on the data given by Kitagawa
in 1937.
5 Taxonomic treatment
Clematis L. sect. Tubulosae Decne. in Nouv. Arch. Mus. Hist.: Nat. Paris, ser. 2, 4: 203.
1881; H. Eichler in Bibl. Bot. 124: 18. 1958, excl. syn.; M. Johnson, Klematis 265. 1997; W.
T. Wang in Acta Phytotax. Sin. 39: 6. 2001.——Sect. Viorna (Reichb.) Prantl subsect.
Tubulosae (Decne.) Prantl in Bot. Jahrb. 9: 258. 1888, p.p. excl.c. acuminata DC. et al.;
Hand.-Mazz. in Acta Hort. Gotob. 13: 191. 1939; M. Y. Fang in Fl. Reip. Pop. Sin. 28: 93.
1980.——Subgen. Tubulosae (Decne.) Grey-Wilson, Clematis 30, 189. 2000. Lectotype: C.
heracleifolia DC. (Eichler, 1958).
Woody vines, erect perennial herbs, low subshrubs or low shrubs. Seedling leaves
alternate (only known from C. heracleifolia—Essig, 1991). Leaves 1–2-ternate or 1–2-pinnate;
leaflets dentate. Flowers bisexual or unisexual. Sepals 4, valvate, suberect or erect, white, blue
or purple, obovate-oblong, narrowly oblong, or lanceolate. Stamen filaments linear, usually
pilose near apex. Pollen grains tricolpate or pantoporate. Achenes compressed but not
Acta Phytotaxonomica Sinica Vol. 45 434
flattened; persistent styles elongate, plumose.
Nine species, widespread in eastern Asia.
Key to infrasectional taxa
1. Woody vines; leaves 1–2-ternate or 1–2-pinnate; flowers in axillary or terminal panicles, bisexual; sepals
suberect at beginning of anthesis, thereafter spreading, white or pinkish, obovate-oblong; pollen grains
tricolpate…………………………………………………………………………………Subsect. 1. Pinnatae
2. Sepals white, apex acute or shortly cuspidate; leaves 1–2-ternate or 1–2-pinnate……………1. C. pinnata
2. Sepals pinkish, apex subtruncate or truncate; leaves ternate, rarely pinnate………………2. C. takedana
1. Erect perennial herbs, or low subshrubs or shrubs; leaves ternate, rarely pinnate; flowers usually
polygamous, rarely bisexual; sepals always erect, blue, purple, or pink, usually narrowly obovate- oblong,
rarely lanceolate or above strongly dilated and elliptic, firmly connivent during anthesis, near apex or above
recurved; pollen grains pantoporate, rarely tricolpate (C. heracleifolia)……Subsect. 2. Tubulosae
3. Perennial herbs or low subshrubs; leaves ternate; flowers in terminal many-flowered panicle-like
compound cymes; pollen grains pantoporate, rarely tricolpate…………………………Ser. 1. Tubulosae
4. Calyx urceolate or tubular-urceolate; sepals lanceolate or narrowly ovate, not dilated, apex recurved;
pedicels short, 0.5–3 mm long …………………………………………………………5. C. urticifolia
5. Sepals violet …………………………………………………………………………5a. f. urticifolia
5. Sepals flesh-coloured or rose-pink……………………………………………………… 5b. f. rosea
5. Sepals white……………………………………………………………………………5c. f. albiflora
4. Calyx tubular; sepals narrowly oblong or narrowly obovate-oblong, sometimes lanceolate-linear, rarely
elliptic, near apex slightly dilated, rarely strongly dilated above after anthesis.
6. Leaflet teeth strongly reduced in size, usually only represented by mucrons; sepals lanceolate-linear,
18–22×2–3 mm, slightly dilated near apex ………………………………………… 7. C. speciosa
6. Leaflets distinctly dentate.
7. Pedicels more or less slender, densely puberulous, rarely velutinous; sepals only near apex
slightly dilated after anthesis.
8. Persistent styles villous from base to near apex.
9. Flowers bisexual; pedicels densely puberulous; sepals 16–23(–30) mm long; pollen grains
tricolpate …………………………………………………………………3. C. heracleifolia
9. Flowers polygamoous; pedicels velutinous; sepals 14–20 mm long; pollen grains pantopo-
rate ……………………………………………………………………………6. C. psilandra
8. Persistent styles glabrous at base ………………………………………………… 8. C. stans
10. Anthers 4–5 mm long ……………………………………………………… 8a. var. stans
10. Anthers 3–3.8 mm long ……………………………………… 8b. var. austrojaponensis
7. Pedicels robust, densely velutinous …………………………………………… 4. C. tubulosa
11. Sepals strongly dilated above after anthesis, elliptic, 8–15(–20)×4–7(–12) mm, linear below,
8–12×2–3.5 mm……………………………………………………………4a. var. tubulosa
12. Sepals purple or blue………………………………………………………4ai. f. tubulosa
12. Sepals white………………………………………………………………4aii. f. albicalyx
11. Sepals narrowly obovate-oblong or narrowly oblong, 12–28×2.2–7 mm, after anthesis
slightly dilated only near apex ………………………………………… 4b. var. ichangensis
3. Small shrubs; leaves ternate or 5-foliolately pinnate; flowers terminal, solitary, rarely in 2–3-flowered
cymes, polygamous; pollen grains pantoporate…………………………………………Ser. 2. Uniflorae
9. C. tsugetorum
Subsect. 1. Pinnatae (W. T. Wang ) W. T. Wang in Acta Phytotax. Sin. 39: 9. 2001; et 43:
977. 2005.——Sect. Clematis subsect. Pinnatae W. T. Wang in Acta Phytotax. Sin. 36: 159.
1998.——Subsect. Pinnatae ser. Pinnatae W. T. Wang in Acta Phytotax. Sin. 39: 9. 2001.
Type: C. pinnata Maxim.
Subsect. Pinnatae ser. Tatarinowianae W. T. Wang in Acta Phytotax. Sin. 39: 9. 2001.
Type: C. tatarinowii Maxim.
No. 4 WANG & XIE: A revision of Clematis sect. Tubulosae (Ranunculaceae) 435
Woody vines. Leaves 1–2-ternate or 1–2-pinnate. Inflorescences axillary and terminal,
usually many-flowered. Flowers bisexual. Sepals suberect or ascending at beginning of
anthesis, thereafter spreading, white or pinkish, obovate-oblong, not connivent during anthesis.
Pollen grains tricolpate.
Two species, one occurring in China, the other in Japan.
1. Clematis pinnata Maxim. in Bull. Acad. Sci. St.-Petersb. 22: 216. 1877; et in Mél. Biol. 9:
591. 1877; Kuntze in Verh. Bot. Ver. Brand. 26: 182. 1885; Forbes & Hemsl. in J. Linn. Soc.
Bot. 23: 7. 1886; Limpr. in Repert. Sp. Nov. Beih. 12: 376. 1922; Rehd. in J. Arn. Arb. 4: 188.
1923; Kozlov in Publ. Mus. Hoangho Paiho Tien Tsin 22: 14. 1933; Hand.-Mazz. in Acta Hort.
Gotob. 13: 217. 1939; Ting in Fl. Reip. Pop. Sin. 28: 180. 1980; He, Fl. Beijing, rev. ed., 1:
252. 1984; J. W. Wang in Fl. Hebei 1: 477, fig. 486. 1986; M. Johnson, Klematis 448. 1997;
W. T. Wang in Acta Phytotax. Sin. 36: 160. 1998; et 39: 9, fig. 2: 1–3. 2001; W. T. Wang &
Barth. in Fl. China 6: 370. 2001; S. C. Jiang in Fl. Tianjin 181. 2004; Xie et al. in Ann. Bot.
Fennici 42: 305. 2005.——C. pinnata var. normalis Kuntze in l.c. Type: China. Near Beijing
(北京), ca. 1845, Tatarinow s.n. (holotype, LE!; isotype, PE!).
C. tatarinowii Maxim. in Bull. Acad. Sci. St.-Petersb. 22: 215. 1877; et in Mél. Biol. 9:
590. 1877; Decne. in Nouv. Arch. Mus. Hist. Nat. Paris, ser. 2, 4: 212. 1881; Forbes & Hemsl.
in J. Linn. Soc. Bot. 23: 7. 1886; Hand.-Mazz. in Acta Hort. Gotob. 13: 391. 1939; M. Y. Fang
in Fl. Reip. Pop. Sin. 28: 95, pl. 26, figs. 3, 4. 1980; He, Fl. Beijing, rev. ed., 1: 248. 1984; M.
Johnson, Klematis 276. 1997; Grey-Wilson, Clematis 190. 2000; W. T. Wang in Acta Phytotax.
Sin. 39: 9. 2001; W. T. Wang & Barth. in Fl. China 6: 370. 2001. ——C. pinnata Maxim. var.
tatarinowii (Maxim.) Kuntze in Verh. Bot. Ver. Brand. 26: 182. 1885. Type: China. Near
Beijing (北京): ca. 1845, Tatarinow s.n. (holotype, LE!; isotype, PE!).
C. pinnata var. ternatifolia W. T. Wang in Acta Phytotax. Sin. 39: 331. 2001. Type: China.
Beijing (北京): Pinggu (平谷), Nanji Shan (南吉山), 1972-06-13, Pinggu Exped. 224
(holotype, PE!).
羽叶铁线莲 Fig. 1: A, B; Fig. 5; Fig. 6: A–C
Woody vine. Branches shallowly 4–6-sulcate, appressed-puberulous, often glabrescent.
Leaves usually 5-foliolately pinnate, sometimes 1–2-ternate or 2-pinnate; leaflets papery or
herbaceous, ovate, broadly ovate, or rhombic, (2.5–)5–13×(1.5–)3–7.5 cm, apex acuminate,
base rounded or subcordate, margin dentate, 2–3-lobed or undivided, sparsely
appressed-puberulous on both surfaces, abaxial surface laxely reticulate, basal veins abaxially
prominent; petioles 3–11 cm long. Cymes axillary or terminal, usually many-flowered;
peduncles 3–8 cm long, puberulous; bracts triangular, long elliptic or lanceolate, 3–10 mm
long, sometimes ternate, 3 cm long. Flower 1.5–2 cm in diam.; pedicel 0.5–3 cm long, densely
puberulous. Sepals 4, white, obovate-oblong, 12–19×3–5 mm, glabrous inside, densely
puberulous outside, velutinous on margin, apex acute or shortly cuspidate. Stamens 7–10 mm
long; filaments pilose or glabrous near apex; anthers linear or narrowly oblong, 2.5–3 mm
long, glabrous or pilose, apex minutely apiculate. Ovaries puberulous; styles ca. 5 mm long,
densely villous. Fl. Jul.–Aug.
China (Beijing, W & N Hebei, S Heilongjiang, S Liaoning, NW Tianjin). On slopes, in
bushes, or at field edges; alt. 700–1600 m.
Additional specimens examined:
China. Beijing (北京): Haidian (海淀), Jin Shan (金山), S. Y. He (贺士元) s.n. (BNU); Jiufeng (鹫峰),
J. H. Shi et al. (史京华等) 2002037 (PE); Huairou (怀柔), Shayu (沙峪), Anonymous 621 (PE); Mentougou
(门头沟), Baihua Shan (百花山), S. Y. He (贺士元) 15037, J. H. Shi et al. (史京华等) 2002055 (PE); Miyun
(密云), Wuling Shan (雾灵山), PE Exped. (植物所标本馆队) 56-2276 (PE), Q. R. Liu (刘全儒) 978047,
987039 (BNU); Pingu (平谷), Nanshan Cun (南山村), S. Y. He (贺士元) s.n. (BNU); Shijingshan (石景山),
Acta Phytotaxonomica Sinica Vol. 45 436


Fig. 5. Clematis pinnata Maxim. A, flowering branch; B, flower; C, stamens. Drawn from Pinggu Exped. 224.

No. 4 WANG & XIE: A revision of Clematis sect. Tubulosae (Ranunculaceae) 437
Xi Shan (西山), Anonymous 735 (PE); Yanqing (延庆), Licent 9831 (PE); Without precise locality,
Bretschneider 25 (GH, K, LE), Hemeling s.n. (K), Lechman s.n. (K). Hebei (河北): Qinglong (青龙),
Chengde Exped. (承德队) 71-1775 (PE); Xiaowutai Shan (小五台山), Limpricht 627 (S), K. S. Hao (郝景盛)
2057 (PE); Yi Xian (易县), Sanggang (桑岗), Yi Xian Exped. (易县队) 71-140 (PE); Zhuolu (涿鹿),
Yangjiaping (杨家坪), Licent 9081 (TIE); Without precise locality, Chanet & Serre 2886 (P), Serre 9831
(TIE). Heilongjiang (黑龙江): Harbin (哈尔滨), Licent 9221 (TIE). Liaoning (辽宁): Shenyang (沈阳),
Dongling (东陵), Yabe s.n. (NAS). Tianjin (天津): Ji Xian (蓟县), Pan Shan (盘山), S. Y. He (贺士元)
17407 (BNU).
2. Clematis takedana Makino in Bot. Mag. Tokyo 21: 87. 1907; Matsum., Ind. Pl. Japon. 2:
113. 1912; Kitagawa in J. Jap. Bot. 13: 359. 1937; Ohwi, Fl. Jap. 443. 1965; Tamura in Satake
et al., Wild Flow. Jap. 2: 73, pl. 2, fig. 2. 1982; W. T. Wang in Acta Phytotax. Sin. 39: 9, fig. 2:
4–6. 2001. Type: Japon. Honshu: Shinano, 1905-08-28, Takeda s.n. (holotype, not seen).
Figs. 1: C, D; Fig. 6: D–F
Woody vine. Branches shallowly 6–8-sulcate, appressed-puberulous. Leaves ternate,
rarely 5-foliolately pinnate; leaflets papery, broadly ovate or ovate, 3–8.2×1.4–8 cm, apex
acuminate, base rounded or subtruncate, margin irregularly dentate, 2–3-lobed, adaxial surface
sparsely appressed-puberulous, abaxial surface puberulous on veins, basal veins abaxially
prominent; petioles 4.5–5 cm long. Cymes axillary or terminal, usually many-flowered,
panicle-like; peduncles 0.4–4 cm long; bracts triangular, ca. 2 mm long, densely puberulous.
Flower ca. 1.4 cm in diam.; pedicel slender, 6–16 mm long, densely appressed-puberulous.
Sepals 4, pinkish or purplish, narrowly obovate-oblong, 9–15×2–3.6 mm, glabrous inside,
densely appressed-puberulous outside, velutinous on margin, apex subtruncate or truncate.
Stamens 6–9(–11) mm long; filaments pilose near apex; anthers linear, 2–3 mm long, below
pilose, apex acute or minutely apiculate. Ovaries puberulous; styles ca. 6 mm long, densely
villous. Fl. Aug.–Sept.
Japan (C & N Honshu).
Additional specimens examined:
Japan. Akita: Kazunogun Hachimantai, Matsumura s.n. (KYO); Nagano: Mt. Shiroumayama, Makino
95711, Koidzumi s.n. (KYO), Shishu, Koidzumi s.n. (KYO); Nagaro, Hisauchi & Kimura 2145 (TI).
Subsect. 2. Tubulosae (Decne.) W. T. Wang in Acta Phytotax. Sin. 39: 9. 2001; et 43:
477. 2005.——Sect. Tubulosae Decne. in Nouv. Arch. Mus. Hist. Nat. Paris, ser. 2, 4: 203.
1881; Kitagawa in J. Jap. Bot. 13: 342. 1937; Tamura in Sci. Rep. Osaka Univ. 16 (2): 32.
1967; et in Acta Phytotax. Geobot. 38: 39. 1987; et in Hiepko, Nat. Pflanzenfam., Zwei. Auf.,
17a (4): 372. 1995; Snoeijer in Clematis 1992: 10. 1992.——Subgen. Tubulosa T. Y. A. Yang
in Syst. Georgr. Pl. 68: 301. 1999; et in Chiu & Peng, Proc. Cross-str. Symp. Fl. Divers. &
Cons. 92. 1998. Lectotype: C. heracleifolia DC.
Sect. Viorna subsect. Tubulosae auct. non (Decne.) Prantl: Schneid., Ill. Handb. Laubh. 1:
28. 1906; Tamura in Acta Phytotax. Geobot. 16: 79. 1956.
Erect perennial herbs or low subshrubs. Inflorescences terminal. Flowers usually
polygamous, rarely bisexual. Sepals erect, blue, purple, or pink, usually narrowly
obovate-oblong or narrowly oblong, rarely lanceolate or elliptic, after anthesis still firmly
connivent for a long time. Pollen grains pantoporate, rarely tricolpate.
Seven species, distributed in E China, Korea, and Japan.
Ser. 1. Tubulosae Rehd. & Wils. in Sarg., Pl. Wils. 1: 320. 1913; Rehd., Man. Cult. Trees
& Shrubs, ed. 2, 209. 1951, p.p. excl.c. ranunculoides Franch.; W. T. Wang in Acta Phytotax.
Sin. 39: 10. 2001; et 43: 478. 2005. Lectotype: C. heracleifolia DC.
Erect perennial herbs or low subshrubs. Leaves ternate. Flowers in terminal, many-
flowered, panicle-like compound cymes. Pollen grains pantoporate, rarely tricolpate (C.
heracleifolia DC.).
Acta Phytotaxonomica Sinica Vol. 45 438



Fig. 6. A–C, Clematis pinnata Maxim. A, flowering branch; B, sepal outside; C, stamen. Drawn from S. Y. He 15037.
D–F, C. takedana Makino. D, flowering branch; E, sepal outside; F, stamen. Drawn from Hisauchi & Kimura 2145.


No. 4 WANG & XIE: A revision of Clematis sect. Tubulosae (Ranunculaceae) 439
Six species, distributed in E China, Korea, and Japan.
3. Clematis heracleifolia DC., Syst. 1: 138. 1818, ut “heracleaefolia”; et Prodr. 1: 3. 1824;
Forbes in J. Bot. 2: 265. 1884, p.p. excl. syn. C. tubulosa Turcz., C. davidiana Verlot. et C.
stans Sieb. & Zucc.; Forbes & Hemsl. in J. Linn. Soc. Bot. 23: 4. 1886, p.p. excl. syn. C.
tubulosa; Finet & Gagnep. in Bull. Soc. Bot. France 50: 545. 1903, p.p. excl. syn. C. tubulosa,
C. stans etc.; Kom. in Acta Hort. Petrop. 22: 285. 1903, p.p. excl. syn. C. tubulosa; Rehd. &
Wils. in Sarg., Pl. Wils. 1: 320. 1913, p.p. excl. syn. C. tubulosa et C. davidiana; Rehd., Man.
Cult. Trees & Shrubs 219. 1927, excl. syn.; Kitagawa in Rep. First Sci. Exped. Manch., sect. 4,
4: 17. 1936; et in J. Jap. Bot. 13: 354. 1937; et Lineam. Fl. Mansh. 217. 1939; Hand.-Mazz. in
Acta Hort. Gotob. 13: 191. 1939, p.p.; Liou et al., Clav. Pl. Chinae Bor.-Or. 18, pl. 13, fig. 4.
1959; Noda, Fl. N-E Prov. China 522. 1970; Anonymous in Iconogr. Corm. Sin. 1: 735. 1972,
p.p. excl. fig. 1469; Anonymous in Fl. Pl. Herb. Chinae Bor.-Or. 3: 175, pl. 76, figs. 1–4. 1975;
T. B. Lee, Ill. Fl. Korea 342, fig. 1366. 1979; M. Y. Fang in Fl. Reip. Pop. Sin. 28: 94. 1980,
p.p. excl. syn. C. tubulosa et fig. 3; Ding et al., Fl. Henan. 447. 1981, p.p.; He, Fl. Beijing, rev.
ed., 1: 248. 1984, p.p.; J. W. Wang in Fl. Hebei 1: 471, fig. 475. 1986, p.p.; S. H. Li, Fl.
Liaoning. 1: 520. 1988; Y. Z. Zhao in Fl. Intramongol., ed. 2, 2: 520, pl. 260, figs. 6, 7. 1990;
Y. J. Ling et al. in Fl. Shaanxi. 1: 627. 1992, p.p.; Y. N. Lee, Fl. Korea 165. 1996, p.p. excl.
syn. C. urticifolia Nakai ex Kitagawa; Y. J. Zheng in Fl. Shandong. 1: 22, 1997, p.p.; M.
Johnson, Klematis 269. 1997; T. Y. A. Yang & Moore in Syst. Geogr. Pl. 68: 294. 1999, p.p.;
Grey-Wilson, Clematis 190, fig. 149. 2000, p.p.; W. T. Wang & Barth. in Fl. China 6: 370,
2001, p.p.; S. C. Jiang in Fl. Tianjin 179. 2004, p.p. Type: China. “inter Peking et Jehol”, 1793,
Staunton s.n. (holotype, BM!).
C. hookeri Decne. in Nouv. Arch. Mus. Hist. Nat. Paris, ser. 2, 4: 206, pl. 11. 1881.——
C. tubulosa Turcz. var. hookeri (Decne.) Hook. f. in Curtis, Bot. Mag. 111: t. 6801. 1885.——
C. heracleifolia DC. var. hookeri (Decne.) Makino in Bot. Mag. Tokyo 21: 87. 1907, excl.
specim. cit. et syn. C. heracleifolia var. speciosa. No type specimen designated.
C. tubulosa auct. non Turcz.: Hemsl. in J. Linn. Soc. Bot. 13: 75. 1872; Decne. in Nouv.
Arch. Mus. Hist. Nat. Paris, ser. 2, 4: 204, pl. 9. 1881; Lavall., Clemat. 82. 1884; Schneid., Ill.
Handb. Laubh. 1: 281, figs. 184h, 185a–c. 1906.
大叶铁线莲 Fig. 1: E, F; Fig. 7
Perennial herb or subshrub. Stems 20–70 cm tall, longitudinally sulcate, appressed-
puberulous. Leaves ternate; terminal leaflets chartaceous, broadly ovate, rhombic, or elliptic,
6–16×4.5–13.5 cm, apex acute or acuminate, base subtruncate, rounded, or broadly cuneate,
margin irregularly dentate or incised-dentate, 3-lobed, rarely undivided, adaxial surface
subglabrous, abaxial surface reticulate, puberulous on veins; lateral leaflets smaller, slightly
unsymmetrical; petioles 6–16 cm long. Terminal secondary panicles 9–30 cm long, with 2–6
nodes and with flowers 2–5 fascicled in each bract axil; lateral secondary panicles 2.5–12 cm
long, with 1–2 nodes; bracts ternate or simple, ovate, those of fascicled flowers
linear-lanceolate, 2–6 mm long, densely puberulous. Flowers bisexual, 1.5–2 cm in diam.;
pedicels more or less slender, 1.2–3.4(–4.5) cm long, densely puberulous. Sepals 4,
blue-purple, narrowly obovate-oblong or linear, 16–23(–30)×(2–)3–6 mm, slightly dilated
and recurved near apex, glabrous inside, densely appressed-puberulous outside, velutinous on
margin, apex acuminate or acute. Stamens ca. 16, 9–12.5 mm long; filaments 5–8 mm long,
sparsely puberulous or glabrous near apex; anthers linear, 4–6.8 mm long, on connective
pilose; pollen grains tricolpate. Carpels ca. 4 mm long, densely villous. Achenes compressed,
rhombic-elliptic, 3–4.5×1.5–2.2 mm, appressed-puberulous; persistent styles 1.2–2.5 cm
long, plumose. Fl. Jul.–Sept.
China (Beijing, Hebei, Henan, Liaoning, Nei Mongol, Shandong, Shanxi, NW Tianjin)
and N Korea. In grassy places on slopes or in sparse forests; alt. 85–1600 m.
Acta Phytotaxonomica Sinica Vol. 45 440

Fig. 7. Clematis heracleifolia DC. A, upper part of flowering stem; B, sepal inside; C, stamen. Drawn from Y. Liu
11893. D, achene. Drawn from Changping Exped. 121.

No. 4 WANG & XIE: A revision of Clematis sect. Tubulosae (Ranunculaceae) 441
Additional specimens examined:
China. Beijing (北京): Changping (昌平), Baihecun (百合村), Changping Exped. (昌平队) 121 (PE),
Duibaiyu (堆白峪), K. C. Kuan et al. (关克俭等) 12 (PE), Nankou (南口), H. F. Chow (周汉藩) 41588 (PE),
Xiakou (下口), PE Exped. (植物所标本馆队) 56-1518 (PE); Fangshan (房山), Shangfang Shan (上方山), H.
F. Chow (周汉藩) 41707, T. N. Liou (刘慎谔) 8326, W. Y. Hsia (夏纬瑛) 3217 (PE); Haidian (海淀), Dajuesi
(大觉寺), Read 727 (GH); Wofosi (卧佛寺), Cowdry 1009 (BNU, GH), Z. Y. Cao (曹子余) 39 (PE);
Mentougou (门头沟), Baihua Shan (百花山), T. F. King (金德福) 599 (PE, S), C. G. Yang (杨朝广) 526
(PE), Miaofeng Shan (妙峰山), PE Exped. (植物所标本馆队) 56-2771 (PE), Qianjuntai (千军台), W. T.
Wang (王文采) 3092 (PE), Xiaolongmen (小龙门), Z. Y. Mi (米志勇) s.n. (BNU); Shijingshan (石景山), Xi
Shan (西山), T. N. Liou (刘慎谔) 1395 (K, PE), 1396 (PE); Yanqing (延庆), Badaling (八达岭), Z. Y. Cao
(曹子余) 92 (PE), Song Shan (松山), D. Z. Fu & Q. Y. Xiang (傅德志, 向秋云) 163 (PE). Hebei (河北):
Neiqiu (内邱), S. Y. He (贺士元) 2289 (BNU); Wuan (武安), S. Y. He & R. T. Yin (贺士元, 尹汝棠) 21138
(BNU); Xiaowutai Shan (小五台山), Y. Liu (刘瑛) 11349, 11893 (PE), Hers 2181 (GH); Xinglong (兴隆), T.
N. Liou & P. Y. Fu (刘慎谔, 傅沛云) 4594 (LE, PE); Xingtai (邢台), H. F. Chow (周汉藩) 43430 (PE); Yi
Xian (易县), X. L. Huang et al. (黄秀兰等) 868 (PE); Zhuolu (涿鹿): Yangjiaping (杨家坪), H. Smith 312
(GH); Zunhua (遵化): Dongling (东陵), T. N. Liou (刘慎谔) 1397, Y. Liu (刘瑛) 11893 (PE). Henan (河南):
Dengfeng (登封), Henan Exped. (河南队) 59-50705 (PE); Jigong Shan (鸡公山), K. K. Tsoong (钟观光)
3620 (PE); Tongbai (桐柏), Henan Exped. 59-56016 (PE); Yuzhou (禹州), Henan Exped. 59-55011 (PE).
Liaoning (辽宁): Caohekou (草河口), Yabe s.n. (PE); Qian Shan (千山), T. N. Liou et al. (刘慎谔等) 6821
(PE), Y. L. Zhou et al. (周以良等) 2542 (LE, NAS, PE); Shenyang (沈阳), T. N. Liou et al. (刘慎谔等) 359
(IFP); Xiongyuecheng (熊岳城), Y. L. Zhou et al. (周以良等) 2650 (PE). Nei Mongol (内蒙古): Chifeng (赤
峰), Y. M. Zhu (朱亚民) 421 (HIMC); Without precise locality, Y. W. Tsui (崔友文) 2818 (PE). Shandong
(山东): Changqing (长清), C. Y. Guo (郭成勇) 55187-10 (PE); Jinan (济南), Longdong (龙洞), C. Y. Chiao
(焦启源) 3073 (GH, K, NAS); Pingyi (平邑), Y. T. Hou et al. (侯元同等) 98001-1, 98001-2, 98001-3,
98001-7 (PE); Tai Shan (泰山), T. Y. Chou et al. (周太炎等) 7182, 7321 (NAS, PE); Yantai (烟台), Forbes
s.n. (BM), K. M. Liou (刘继孟) 1593 9PE). Shanxi (山西): Fenyang (汾阳), T. P. Wang (王作宾) 2774 (PE);
Jiaocheng (交城), T. Tang (唐进) 1526 (GH, PE); Jiexiou (介休), H. Smith 7764 (GH, UPS); Licheng (黎城),
Licent 11366 (PE); Pingding (平定), K. M. Liou (刘继孟) 3972 (PE); Qinyuan (沁源), K. C. Kuan & Y. L.
Chen (关克俭, 陈艺林) 1001 (PE); Ruicheng (芮城), S. Y. Bao (包士英) 962 (PE); Wutai Shan (五台山), K.
C. Kuan & Y. L. Chen (关克俭, 陈艺林) 2675 (PE); Yuanqu (垣曲), S. Y. Bao (包士英) 744 (PE);
Yuncheng (运城), Huanghe Exped. (黄河队) 57-582 (PE); Without precise locality, Licent 12777 (PE).
Tianjin (天津): Ji Xian (蓟县), T. N. Liou et al. (刘慎谔等) 4521 (PE).
Korea. Kyong Gi: Paekryong Island, Yinger et al. 2849 (K).
The flowers of Clematis heracleifolia are bisexual and protandrous, i.e. when the anthers
of a flower of it mature, the carpels of that flower are still very young and very small, and after
the anthers withered, they began to develop.
4. Clematis tubulosa Turcz. in Bull. Soc. Nat. Mosc. 10 (7): 148. 1837; Walp., Rep. Bot. Syst.
1: 5. 1842; Hook. f. in Curtis, Bot. Mag. 72: t. 4269. 1846; Lamaire, Fl. Serres, ser. 1, 3: 195,
pl. 4. 1847; Paxton in Mag. Bot. 14: 31. 1848; Maxim. in Bull. Acad. Sci. St.-Petersb. 22: 214.
1877, p.p.; et in Mél. Biol. 9: 589. 1877, p.p.; Franch. in Nouv. Arch. Mus. Hist. Nat. Paris,
ser. 2, 5: 165. 1883; et Pl. David. 1: 13. 1884; Kitagawa in J. Jap. Bot. 13: 356. 1937; et
Lineam. Fl. Mansh. 219. 1939; Liou et al., Clav. Pl. Chinae Bor.-Or. 78, pl. 13, fig. 4. 1959;
Noda, Fl. N-E. Prov. China 520. 1970; M. Johnson, Klematis 277. 1997; Grey-Wilson,
Clematis 192, fig. 150. 2000; W. T. Wang in Acta Phytotax. Sin. 44: 334. 2006.——C.
heracleifolia DC. ssp. normalis Kuntze var. tubulosa (Turcz.) Kuntze in Verh. Bot. Ver. Brand.
26: 183. 1885.——C. heracleifolia var. tubulosa Turcz. ex Nakai in J. Coll. Sci. Univ. Tokyo
26: 12. 1909, quoad nomen tantum. Type: North China. Without precise locality, Kirilow s.n.
(holotype, LE).
C. davidiana Decne. ex Verlot in Rev. Hort. 90 cum icon. 1867; Decne. in Nouv. Arch.
Mus. Hist. Nat. Paris, ser. 2, 4: 205, pl. 10. 1881; Lavall., Clemat. 82. 1884; Schneid., Ill.
Acta Phytotaxonomica Sinica Vol. 45 442
Handb. Laubh. 1: 281, figs. 184k, 185f–g. 1906; Kitagawa in Rep. First Sci. Exped. Manch.,
sect. 4, 4: 17. 1936.——C. tubulosa Turcz. var. davidiana (Decne. ex Verlot) Franch. in Nouv.
Arch. Mus. Hist. Nat. Paris, ser. 2, 5: 165. 1883; et Pl. David. 1: 13. 1884.——C. heracleifolia
DC. ssp. davidiana (Decne. ex Verlot) Kuntze in Verh. Bot. Ver. Brand. 26: 183. 1885;
Anonymous in Fl. Pl. Herb. Chinae Bor. -Or. 3: 175, pl. 76, figs. 5, 6. 1975, ut var.; S. H. Li,
Fl. Liaoning. 1: 522. 1988; et in P. Y. Fu, Clav. Pl. Chinae Bor.-Or., ed. 2, 193. 1995, ut var. —
—C. heracleifolia var. davidiana (Decne. ex Verlot) Forbes & Hemsl. in J. Linn. Soc. Bot. 23:
4. 1886; T. B. Lee, Ill. Fl. Korea 342, fig. 1367. 1979; Y. N. Lee, Fl. Korea 165, fig. 483. 1996;
Toomey & Leeds, Ill. Encycl. Clematis 219. 2001.——C. heracleifolia var. davidiana Nakai
in J. Coll. Sci. Univ. Tokyo 26: 12. 1909. Type: China. Beijing (北京): Without precise
locality, 1863, David 417, 423 (syntypes, P!).
C. heracleifolia auct. non DC.: Forbes in J. Bot. 22: 265. 1884, p.p.; Forbes & Hemsl. in
J. Linn. Soc. Bot. 23: 4. 1886, p.p.; Finet & Gagnep. in Bull. Soc. Bot. France 50: 545. 1903,
p.p.; Kom. in Acta Hort. Petrop. 22: 285. 1903, p.p.; Rehd. & Wils. in Sarg., Pl. Wils. 1: 320.
1913, p.p.; Hand.-Mazz. in Acta Hort. Gotob. 13: 191. 1939, p.p.; Anonymous in Iconogr.
Corm. Sin. 1: 735. 1972, p.p. quoad fig. 1469; M. Y. Fang in Fl. Reip. Pop. Sin. 28: 94. 1980,
p.p. quoad fig. 3; Anonymous in Fl. Jiangsu. 2: 167. 1982; He, Fl. Beijing, rev. ed., 1: 248.
1984, p.p.; J. W. Wang in Fl. Hebei 1: 471. 1986, p.p.; Y. J. Zheng in Fl. Shandong. 1: 22.
1997, p.p.; T. Y. A. Yang & Moore in Syst. Geogr. Pl. 68: 294. 1999, p.p.; W. T. Wang & Barth.
in Fl. China 6: 370. 2001, p.p.; S. J. Fan et al., Fl. Laoshan. 1: 238. 2003, p.p. excl. fig. 246; S.
C. Jiang in Fl. Tianjin 179. 2004, p.p.; J. C. Zhao et al., High. Pl. Cat. Hebei Prov. China 50.
2005, p.p. quoad fig. 44.
卷萼铁钱莲
4a. var. tubulosa Fig. 2: C, D; Fig. 8
4ai. f. tubulosa
Perennial herb or subshrub. Stems 50–150 cm tall, longitudinally sulcate, densely
puberulous. Leaves ternate; terminal leaflets chartaceous, broadly ovate, elliptic, or obovate,
6.5–19×5–16 cm, apex acuminate or acute, base subtruncate, rounded, or broadly cuneate,
margin irregularly dentate or incised-dentate, 3-lobed or undivided, adaxial surface
subglabrous, abaxial surface puberulous on prominent veins; lateral leaflets smaller, slightly
unsymmetrical; petioles 4.5–16 cm long. Terminal secondary panicles 10–50 cm long, with
1–4 nodes and with flowers 2–7 fascided in each bract axil, lateral secondary panicles 1.5–18
cm long, with 1–3 nodes, or absent; panicle bracts ternate or simple, ovate; bracts of fascicled
flowers triangular or linear-lanceolate, 3–9 mm long, abaxially densely puberulous. Flowers
polygamous; pedicels robust, 0.3–2 cm long, densely velutinous. Sepals 4, blue-purple, elliptic
or oblong-elliptic above, limb-like, 8–15(–20)×4–7(–12) mm, recurved, linear below,
claw-like, 8–12×2–3.5 mm, glabrous inside, appressed-puberulous outside, velutinous on
margin. Stamens 12–20, 9–12 mm long; filaments 3–5 mm long, pilose near apex; anthers
linear, 5–8 mm long, on connective pilose, apex minutely apiculate; pollen grains pantoporate.
Carpels 20–30, 5–7 mm long, densely villous. In staminate flowers there are no sterile carpels.
Achenes compressed, elliptic, 3×1.8–2 mm, puberulous; persistent styles 1.4–2 cm long,
plumose. Fl. Jul.–Sept.
China (Beijing, N Hebei, NE Jiangsu, Liaoning, E Shandong, NW Tianjin) and N Korea.
In bushes or forests, on slopes, or by streams; alt. 80–1400 m.
Additional specimens examined:
China. Beijing (北京): Changping (昌平), Ming Tombs (十三陵), K. M. Liou (刘继孟) 666 (PE),
Nankou (南口), H. F. Chow (周汉藩) 40471, 40513 (PE), Xiakou (下口), H. F. Chow (周汉藩) 41986, J.
Zhang (张敬) 2023, 2045 (PE), Baihe Cun (百合村), Changping Exped. (昌平队) 41 (PE); Fangshan (房山),
No. 4 WANG & XIE: A revision of Clematis sect. Tubulosae (Ranunculaceae) 443



Fig. 8. Clematis tubulosa Turcz. var. tubulosa. A, upper part of a bisexual flowering plant; B, sepal inside; C, stamen.
Drawn from C. G. Yang 197. D, achene. Drawn from W. Wang 3544.

Acta Phytotaxonomica Sinica Vol. 45 444
Shangfang Shan (上方山), W. Y. Xia (夏纬瑛) 3217a (PE); Haidian (海淀), Beijing Zoo (北京动物园), T. N.
Liou (刘慎谔) 1399 (PE), Qinglongqiao (青龙桥), Z. T. Wang (王忠涛) 156 (PE), Summer Palace (颐和园),
W. T. Wang (王文采) 0601 (PE), Wofosi (卧佛寺), Cowdry 41 (BNU), Jin Shan (金山), S. Y. He (贺士元)
s.n. (BNU), Dajuesi (大觉寺), Anonymous 8 (PE), Xiang Shan (香山), Bartholomev & Boufford 2038 (GH);
Huairou (怀柔): Yunmeng Shan (云蒙山), S. Y. He (贺士元) 33088 (BNU); Mentougou (门头沟), Baihua
Shan (百花山), W. Y. Hsia (夏纬瑛) 2005, 2583 (PE); Jietaisi (戒台寺), Bartholomev & Boufford 2060 (GH),
Jiulong Shan (九龙山), Hancock s.n. (K); Xiaolongmen (小龙门), T. F. King (金德福) 237 (PE); Miyun (密
云): Wuling Shan (雾灵山), S. Y. He (贺士元) 13589 (BNU); Shijingshan (石景山), Xi Shan (西山), H. F.
Chow (周汉藩) 42046 (PE); Yanqing (延庆), Song Shan (松山), D. Z. Fu & Q. Y. Xiang (傅德志, 向秋云)
83012 (PE); Without precise locality, 1843, Kirilow s.n. (LE), Bretschneider 1212 (BM). Hebei (河北):
Changli (昌黎), W. Y. Hsia (夏纬瑛) 1944 (PE); Chengde (承德), Nankai Univ. Exped. (南开大学队) 59-276
(PE); Fengning (丰宁), Z. T. Yin (尹祖棠) 19898 (BNU); Huailai (怀来), Licent 2494 (K, PE); Laiyuan (涞
源), Baishi Shan (白石山), S. Y. He (贺士元) 20318 (BNU); Pai Ta, Licent & Serre 2885 (P); Qinglong (青
龙), Chengde Exped. (承德队) 71-558 (PE); Shanhaiguan (山海关), Licent 1580 (P); Xiaowutai Shan (小五
台山), X. L. Huang et al. (黄秀兰等) 2357, 5622, 6041 (PE); Xinglong (兴隆), T. N. Liou (刘慎谔) 4604
(PE); Yi Xian (易县), J. C. Liu (刘汝强) 1161 (BNU); Zhuolu (涿鹿), Xiling Shan (西灵山), T. F. King (金德
福) 237 (PE, S), Yangjiaping (杨家坪), H. Smith 291, 302 (GH, UPS), Schindler 49 (BM); Zunhua (遵化),
Dongling (东陵), K. M. Liou (刘继孟) 425 (PE). Jiangsu (江苏): Ganyu (赣榆), T. Y. Chou et al. (周太炎等)
21242 (HHBG, NAS); Lianyungang (连云港): Yuntai Shan (云台山), T. Y. Chou et al. (周太炎等) 20909
(NAS). Liaoning (辽宁): Beining (北宁), Z. S. Qin et al. (秦忠时等) 531 (IFP); Chaoyang (朝阳), C. R.
Liang (梁慈荣) 6 (IFP); Dalian (大连), Z. Wang et al. (王战等) 889 (IFP, LE, PE); Dalu Island (大鹿岛), W.
Wang et al. (王薇等) 1286 (IFP); Dandong (丹东), W. Wang et al. (王薇等) 1047 (IFP); Faku (法库), S. Z.
Liu (刘淑珍) 133 (IFP); Fengcheng (凤城), Z. Wang et al. (王战等) 1628 (LE, PE); Huanren (桓仁), C. S.
Wang (王崇书) 4054 (PE); Jianchang (建昌), C. S. Wang (王崇书) 3221 (IFP); Jinzhou (锦州), Z. Wang et
al. (王战等) 3544 (PE); Kelaqin (喀喇沁), C. S. Wang (王崇书) 3705 (IFP); Lingyuan (凌源), S. X. Li (李书
馨) 334 (IFP, NAS, PE); Qian Shan (千山), S. X. Li (李书馨) 5916 (IFP); near Shenyang (沈阳), Komarov
708 (GH); Suizhong (绥中), S. X. Li (李书馨) 542 (IFP); Tieling (铁岭), H. W. Kung (孔宪武) 640 (PE);
Wafangdian (瓦房店), J. Wei (魏均) 38 (IFP). Shandong (山东): Huang Xian (黄县), Lai Shan (莱山), Z. X.
Zhao (赵子孝) 16128 (CAF); Muping (牟平), Kunyu Shan (昆嵛山), T. N. Liou & K. M. Liou (刘慎谔, 刘
继孟) 1391, 1485 (PE), Q. R. Liu (刘全儒) 977176 (BNU); Lao Shan (崂山), C. Y. Chiao (焦启源) 2740
(GH, K), 2840 (IBSC, NAS), T. Y. Chou et al. (周太炎等) 1226 (NAS, PE). Tianjin (天津): Ji Xian (蓟县),
Pan Shan (盘山), PE Exped. (植物所标本馆队) 56-2002 (PE).
Korea. Hamgyong-namdo, Koidzumi s.n. (KYO); Kangwon-do: Mt. Odae National Park, Beyer,
Erskine & Cowley 166 (K); Kyongyi-do, Koidzumi s.n. (KYO).
The type specimen of Clematis tubulosa Turcz. was collected from North China by P.
Kirilow (Turczaninow, 1837). While working in LE in 2001, I failed to find it, but found two
authentic specimens of that species, one flowering and the other fruiting, which were collected
also by Kirilow in 1843 from Beijing (Peking). If the type specimen was destroyed or lost, the
flowering specimen collected by Kirilow just mentioned, I think, may be designated as the
neotype of Clematis tubulosa Turcz.
Var. tubulosa was introduced into cultivation from Beijing by the French plant collector P.
A. David in 1863 under the name of Clematis davidiana, and has been involved with other
species belonging to sect. Tubulosae or to sect. Clematis to produce several cultivars
(Grey-Wilson, 2000).
4aii. f. albicalyx (D. Z. Lu) W. T. Wang, comb. nov.——C. heracleifolia DC. f. albicalyx D.
Z. Lu in J. Beijing Agr. Coll. 17 (3): 19. 2002. Type: China. Beijing (北京): Miyun (密云),
Potou (坡头), alt. 700 m, 2001-07-11, D. Z. Lu (路端正) 01007006 (holotype, BJFC!).
白花卷萼铁线莲
Sepals white. Fl. Jul.
No. 4 WANG & XIE: A revision of Clematis sect. Tubulosae (Ranunculaceae) 445
China (Beijing: Miyun). By stream in valley; alt. 700 m.
This form with short robust pedicels and white sepals which are strongly dilated above
obviously belongs to C. tubulosa var. tubulosa, and should be transferred to the latter from C.
heracleifolia.
4b. var. ichangensis (Rehd. & Wils.) W. T. Wang in Acta Phytotax. Sin. 44: 335. 2006.——C.
heracleifolia DC. var. ichangensis Rehd. & Wils. in Sarg., Pl. Wils. 1: 321. 1913; Rehd. in J.
Arn. Arb. 4: 183. 1923; et Man. Cult. Trees & Shrubs 219. 1927; P. C. Tsoong in Contr. Inst.
Bot. Nat. Acad. Peiping 4: 119. 1936; Kitagawa in J. Jap. Bot. 13: 354. 1937; Hand.-Mazz. in
Acta Hort. Gotob. 13: 191. 1939, p.p.; Grey-Wilson, Clematis 191. 2000. Type: China. Hubei
(湖北): Yichang (宜昌), foot of limestone cliff, alt. 50–600 m, 1907-08, E. H. Wilson 763
(holotype, GH!). Hubei: Yichang, E. H. Wilson 2596a (paratypes, GH!, K!); Ou-tan-shan,
1907-08, Silvestri 632, 633 (paratypes, GH!, K!); without precise locality, Henry 3053, 4478
(paratype, GH!), 4359 (paratypes, G!, G-Boiss!, GH!). Shaanxi (陕西): Taibai Shan (太白山),
Purdom 1010 (paratypes, GH!, K!); Huan-tuo-san, 1897-10-10, Giraldi s.n. (paratype, GH!).
C. heracleifolia auct. non DC.: Hand.-Mazz. in Acta Hort. Gotob. 13: 191. 1939, p.p.
quoad H. Smith 5952; Hsu in Obs. Fl. Huangshan. 115. 1965; Anonymous in Iconogr. Corm.
Sin. 1: 735. 1972, p.p.; Anonymous in Fl. Tsinling. 1 (2): 287, fig. 246. 1974; Anonymous in
Fl. Hupeh. 1: 362, fig. 506. 1976; M. Y. Fang in Fl. Reip. Pop. Sin. 28: 94. 1980, p.p. excl. fig.
3; Ding et al., Fl. Henan. 1: 447. 1981, p.p.; X. W. Wang in Fl. Anhui 2: 331. 1986; Z. H. Lin
in Fl. Zhejiang 2: 283. 1992; Y. J. Ling et al. in Fl. Shanxi. 1: 627. 1992, p.p.; W. T. Wang in
Keys Vasc. Pl. Wuling Mount. 168. 1995; Y. Z. Zheng in Fl. Shandong 2: 22. 1997, p.p.; T. Y.
A. Yang & Moore in Syst. Geogr. Pl. 68: 294. 1999, p.p.; K. M. Li in Fl. Hunan. 2: 679. 2000;
Z. H. Peng et al., Encycl. Pl. Three Gorg. Yangtze Riv. China 196. 2005.
狭卷萼铁线莲 Fig. 2: A, B; Fig. 9
This variety differs from var. tubulosa in its narrowly obovate-oblong or narrowly oblong
sepals, which are slightly dilated and slightly recurved near apex.
Sepals 12–18×2.2–7 mm. Stamens 8–11 mm long; filaments longer or shorter than
anthers, 3.5–6 mm long; anthers 4–6 mm long, apex not or slightly apiculate. Fl. Jun.–Sept.
China (Anhui, E Guizhou, SW Hebei, Henan, W Hubei, W Hunan, S Shaanxi, Shandong,
Shanxi, NW Zhejiang). On grassy slopes, in bushes, by streams, or on cliffs; alt. 50–2000 m.
Additional specimens examined:
China. Anhui (安徽): Huang Shan (黄山), R. C. Ching (秦仁昌) 3032 (K), T. N. Liou & P. C. Tsoong
(刘慎谔, 钟补求) 2809 (PE, WUK), M. J. Wang (王名金) 3712 (IBSC, NAS, PE); Huoshan (霍山), Pl. Res.
Exped. (植物资源队) Da0480 (PE); Jinzhai (金寨), X. S. Shen (沈显生) 1596 (ANUB); She Xian (歙县), X.
P. Zhang (张小平) 931 (ANUB); Shucheng (舒城), Z. W. Xie (谢中稳) 97227 (PE). Guizhou (贵州): Shibing
(施秉), Wulingshan Exped. (武陵山队) 88-2570 (PE). Hebei (河北): Ci Xian (磁县), K. C. Kuan (关克俭)
6117 (PE); Neiqiu (内邱), X. Y. Liu (刘心源) 651 (PE); Xingtai (邢台), Y. Liu & X. Y. Liu (刘瑛, 刘心源)
13133 (PE); Zhengding (正定), Zhengding Exped. (正定队) V622 (PE). Henan (河南): Jigong Shan (鸡公
山), X. Q. Zhang (张祥卿) 20348 (PE); Lingbao (灵宝), J. Q. Fu (傅竞秋) 87 (KUN); Lushi (卢氏), K. M.
Liou (刘继孟) 4885 (PE); Shangcheng (商城), Henan Exped. (河南队) 59-10452 (PE); Song Xian (嵩县), K.
C. Kuan & T. L. Dai (关克俭, 戴天伦) 2293 (PE); Tongbai (桐柏), Henan Exped. 59-56261 (河南队) (PE);
Xixia (西峡), K. C. Kuan & T. L. Dai (关克俭, 戴天伦) 1144 (PE); Yichuan (奕川), Henan Exped. (河南队)
59-20523, 59-20768 (PE); Yiyang (宜阳), Xinxiang Norm. Coll. Exped. (新乡师院队) 22657 (PE). Hubei (湖
北): Shiyan (十堰), C. S. Wang (王崇书) 231 (PE); Wudang Shan (武当山), K. R. Liu (刘克荣) 135 (PE);
Without precise locality, Henry 7494, E. H. Wilson 2596 (K). Hunan (湖南): Fenghuang (凤凰), F. Z. Tian
(田凤珍) 251 (HUTM); Yuanling (沅陵), G. C. Zhang (张桂才) 564 (PE). Shaanxi (陕西): Baoji (宝鸡),
Hugh s.n. (BM); Feng Xian (凤县), Z. Y. Zhang (张志英) 281 (IBSC, KUN, WUK); Foping (佛坪), X. M.
Zhang (张襄明) 336 (IBSC, WUK); Hu Xian (户县), P. C. Kuo (郭本兆) 542, 640 (PE); Hua Shan (华山), T.
P. Wang (王作宾) 19794 (PE, WUK); Hua Xian (华县), T. P. Wang (王作宾) 15895 (KUN, PE); Lantian (蓝
Acta Phytotaxonomica Sinica Vol. 45 446

Fig. 9. Clematis tubulosa Turcz. var. ichangensis (Rehd. & Wils.) W. T. Wang. A, upper part of a bisexual flowering
plant; B, sepal outside; C, stamen. Drawn from T. P. Wang 15895. D, achene. Drawn from T. P. Wang 16601.

No. 4 WANG & XIE: A revision of Clematis sect. Tubulosae (Ranunculaceae) 447
田), Z. Y. Zhang (张志英) 281 (WUK); Luonan (洛南), T. P. Wang (王作宾) 15847 (PE, WUK); Mt.
Ngo-san, 1899-08, Hugh s.n. (PE); Shangnan (商南), P. C. Kuo (郭本兆) 4436 (IBSC, KUN); Shangzhou (商
州), T. P. Wang (王作宾) 16122 (IBSC, PE); Shanyang (山阳), T. P. Wang (王作宾) 16367, 16601 (PE);
Taibai Shan (太白山), J. X. Yang (杨金祥) 2188 (PE, WUK), Hugh s.n. (BM); Zhongnan Shan (终南山), T.
P. Wang (王作宾) 2164 (PE); Zhouzhi (周至), X. M. Zhang (张襄明) 243 (IBSC, KUN, WUK), Hugh s.n.
(BM). Shandong (山东): Fei Xian (费县), Meng Shan (蒙山), T. Y. Cheo (周太炎) & L. Yan 200 (G, P);
Mengyin (蒙阴), T. Y. Cheo et al. (周太炎等) 6372 (NAS, PE); Tai Shan (泰山), S. C. Cui (崔顺昌) 59 (PE);
Yantai (烟台), 1890-02, Faber 253 (K, LE). Shanxi (山西): Jincheng (晋城), S. Y. Bao (包世英) 339 (PE);
Linfen (临汾), Licent 2245 (BM); Lingchuan (陵川), S. Y. Bao (包世英) 398 (PE); Ruicheng (芮城), S. Y.
Bao (包士英) 845 (PE); Yuncheng (运城), H. Smith 5952 (PE, UPS); Wutai (五台), K. C. Kuan & Y. L. Chen
(关克俭, 陈艺林) 2515 (PE). Zhejiang (浙江): Xitianmu Shan (西天目山), Zhejiang Exped. (浙江队)
59-29559 (NAS, PE); Changhua (昌化), X. Y. He (贺贤育) 23934 (HHBG, NAS, PE).
The flowers of populations of Clematis tubulosa var. ichangensis occurring in most
provinces of its distribution area are bisexual, and only those of populations occurring in
Shandong Province are unisexual and staminate.
5. Clematis urticifolia Nakai (in Tyosen Sanrin Kaiho no. 122-5: 23 et 31, cum diagn. Jap., ut
urticiflora. 1935) ex Kitagawa in J. Jap. Bot. 13: 346, fig. 1. 1937; Chong, Ill. Man. Korean
Trees & Shrubs 202, fig. 302. 1943; Ohwi, Fl. Jap. 441. 1965; Kitamura & Murata, Colour. Ill.
Herb. Pl. Jap., rev. ed., 2: 224. 1980; Ohwi & Kitagawa, New Fl. Jap. 679. 1992; M. Johnson,
Klematis 285. 1997; Grey-Wilson, Clematis 191. 2000. Type: Korea. Prov. Tyu-nan: in
dumosis montis Zoku-ri-san, 1934-08-12, Nakai 14932 (holotype, TI).
C. heracleifolia DC. var. tubulosa sensu Nakai in J. Coll. Sci. Univ. Tokyo 26: 12. 1909,
non C. tubulosa Turcz.
C. tubulosa auct. non Turcz.: Nakai, Tyosen Syokubutu 39. 1914.
C. heracleifolia auct. non DC.: Y. N. Lee, Fl. Korea 165. 1996, p.p. quoad syn. C.
urticifolia; T. Y. A. Yang & Moore in Syst. Geogr. Pl. 68: 294. 1999, p.p. quoad Okamoto s.n.
5a. f. urticifolia Fig. 2: E, F; Fig. 10
Subshrub. Stems up to 2 m tall, longitudinally sulcate, puberulous. Leaves ternate;
terminal leaflets papery, broadly rhombic, rhombic, broadly ovate, elliptic, or obovate, 7–16×
4–11 cm, apex acuminate, base broadly cuneate or rounded, margin irregularly dentate,
3-lobed or 3-lobulate, adaxial surface appressed-puberulous, abaxial surface reticulate,
pubescent; lateral leaflets smaller, obliquely ovate; petioles 8.5–16 cm long. Panicles terminal,
3–40 cm long, 2–3 times branched, 10–many-flowered; panicle bracts foliiform or simple,
obovate; bracts of flowers linear-lanceolate or triangular, 1.2–5 mm long, densely puberulous.
Flowers polygamous; pedicels robust, short, 0.5–3 mm long, 1.2–1.8 mm in diam., velutinous.
Calyx urceolate; sepals 4, violet, oblong-lanceolate or narrowly ovate, 12–15×3–6 mm,
glabrous inside, appressed-velutinous outside, often 3-costate, velutinous on margin, apex
acuminate, recurved. Stamens 12–16, 9–12 mm long; filaments lanceolate-linear, 5–9 mm
long, 1-veined, pilose near apex; anthers linear, 4–5 mm long, apex apiculate; pollen grains
pantoporate. Carpels ca. 12, 4 mm long, densely villous. Achenes compressed, broadly ovate
or broadly elliptic, 3–3.2×2.8–3.2 mm, pilose, not or slightly rimmed; persistent styles ca. 2.5
cm long, plumose. Fl. Jul.–Sept.
S Korea. At forest edges or in sparse forests on slopes; alt. 700–1300 m.
Additional specimens examined:
Korea. Cholla-namdo: Kurge-gun, Mt. Chiri, Boufford et al. 25833 (PE); Kangwon-do: Mt.
Jeombong-san, Qin et al. 18016 (PE), Mt. Kumgang, Koidzumi s.n. (KYO), Kondo 9135 (PE); Kogen: Mt.
Kumgansan, Kitamura s.n. (KYO); Kyongsannamdo: Mt. Tii, Koidzumi s.n. (KYO); the same locality,
1937-04-23–28, Okamoto s.n. (four sheets, KYO).

Acta Phytotaxonomica Sinica Vol. 45 448

Fig. 10. Clematis urticifolia Nakai ex Kitagawa f. urticifolia. A, upper part of a bisexual flowering plant; B, sepal
inside; C, stamen. Drawn from Boufford et al. 25833. D, achene. Drawn from H. N. Qin et al. 18016.

No. 4 WANG & XIE: A revision of Clematis sect. Tubulosae (Ranunculaceae) 449
5b. f. rosea (Nakai) Nakai ex Kitagawa in J. Jap. Bot. 13: 350. 1937; M. Johnson, Klematis
286. 1997; Grey-Wilson, Clematis 191. 2000. ——C. tubulosa Turcz. var. rosea Nakai in Bot.
Mag. Tokyo 31: 4. 1917; et Rep. Veget. Diamond Mt. 172, 195. 1918; Chong, Ill. Man. Korean
Tr. & Shr. 202, fig. 301. 1993. Type: Korea. Mt. Unzyong-ryong, alt. 720 m, collector name
and collection number both not given (holotype, TI).
C. urticifolia Nakai var. carnea Nakai, Tyuosen Sanrin Kaiho no. 122–125: 23, 31. 1935,
nom. illegit.
Sepals flesh-coloured or rose-pink.
Korea.
5c. f. albiflora (Y. Lee) W. T. Wang, comb. nov. ——C. heracleifolia DC. f. albiflora Y. Lee,
Fl. Korea 165, 1157, fig. 484. 1996. Type: Korea. Kangwon-do: Osaek, 1983-08-10, Y. Lee s.n.
(holotype, not seen).
Sepals white.
C Korea.
In Flora of Korea written by Lee (1996), the plant in the photograph (fig. 482) of
Clematis heracleifolia DC. has flowers with very short pedicels and purple urceolate calyxes,
distinctly different from those of C. heracleifolia, and is actually a plant of C. urticifolia f.
urticifolia. Moreover, the photograph (fig. 484) of C. heracleifolia f. albiflora Y. Lee in that
book shows that this new form with white calyxes also has very short pedicels and narrowly
urceolate calyxes, and thus should belong to C. urticifolia rather than to C. heracleifolia.
6. Clematis psilandra Kitagawa in J. Jap. Bot. 13: 352, fig. 2. 1937; S. T. Liu & Hsieh in Fl.
Taiwan 2: 489. 1976; T. Y. A. Yang & T. C. Huang in Taiwania 37: 47, pl. 12. 1992; et 40: 238.
1995; et in Fl. Taiwan, ed. 2, 2: 536, pl. 252. 1996; M. Johnson, Klematis 272. 1997; T. Y. A.
Yang & Moore in Syst. Geogr. Pl. 68: 298. 1999; Grey-Wilson, Clematis 191. 2000; W. T.
Wang & Barth. in Fl. China 6: 371. 2001. ——C. heracleifolia DC. var. taiwanica Suzuki &
Hosokawa in Trans. Nat. Hist. Soc. Formos. 23: 96. 1933. Type: China. Taiwan (台湾):
Pingdong (屏东), Mt. Mutoo, Hosakawa 5405 (holotype, TAI; photograph, PE!).
光蕊铁线莲 Fig. 11: E–H
Small shrub. Stems 50–120 cm tall; branches puberulous, glabrescent. Leaves ternate;
terminal leaflets thickly papery, broadly ovate, pentagonous, ovate, or rhombic, 7–10×4–10
cm, apex acuminate, base subtruncate or broadly cuneate, margin dentate or denticulate, often
3–5-lobed, adaxial surface glabrous, abaxial surface sparsely puberulous on veins, basal veins
abaxially prominent; leteral leaflets smaller, obliquely ovate, usually 1–3-lobulate; petioles
5–10 cm long, velutinous, glabrescent. Compound cymes terminal, panicle-like, usually
many-flowered; peduncles 3.5–8.5 cm long, velutinous; bracts foliiform or small, narrowly
ovate, 3–6 mm long, velutinous. Flowers polygamous, ca. 1 cm in diam.; pedicels 0.7–2 cm
long, velutinous. Sepals 4, pinkish, narrowly oblong, 14–20×4–10 mm, glabrous inside,
velutinous outside, apex mucronate and recurved. Stamens ca. 14, 4–10 mm long; filaments
abaxially above with a few short hairs; anthers narrowly oblong, 2.2–3.2 mm long, glabrous,
apex minutely apiculate; pollen grains pantoporate (Yang & Huang, 1992). Carpels numerous,
ovaries pubescent, styles ca. 8 mm long, villous. Abortive carpels several in a staminate
flower, 0.6–1 mm long. Achenes convex on both sides, elliptic or ovate, 3–4×1.6–2 mm,
puberulous; persistent styles ca. 2 cm long, plumose. Fl. Jul.–Sept.
China (Taiwan). On open slopes or on cliffs; alt. 1000–2500 m.
Additional specimens examined:
China. Taiwan (台湾): Hualian (花莲), C. C. Liao 482 (GH), Tamura, Shimizu & Kao 24737 (KYO);
Pingdong (屏东), C. C. Liao 677 (GH), Y. R. Lin 123 (GH).

Acta Phytotaxonomica Sinica Vol. 45 450

Fig. 11. A–D. Clematis tsugetorum Ohwi. A, a pistillate flowering plant; B, sepal inside; C, staminode. Drawn from J. C.
Wang et al. 3752. D, stamen (after Yang & Huang, 1996). E–H. C. psilandra Kitagawa. E, upper part of a staminate
flowering plant; F, sepal outside; G, stamen. Drawn from C. C. Liao et al. 677. H, achene. Drawn from Y. R. Lin 123.

No. 4 WANG & XIE: A revision of Clematis sect. Tubulosae (Ranunculaceae) 451
7. Clematis speciosa (Makino) Makino in J. Jap. Bot. 1: 39. 1918; Makino & Nemato, Fl. Jap.
971. 1925; Kitagawa in J. Jap. Bot. 13: 345. 1937; Ohwi, Fl. Jap. 441. 1965; Tamura in Satake
et al., Wild Flow. Jap. 2: 73, pl. 71: 1. 1982; Ohwi & Kitagawa. New Fl. Jap. 679. 1992; M.
Johnson, Klematis 273. 1997; Grey-Wilson, Clematis 192. 2000. ——C. heracleifolia DC.
var. speciosa Makino in Bot. Mag. Tokyo 6: 50, 170. 1892; et 11: 332. 1897. Type: Japan.
Tosa: Aso, Makino s.n. (holotype, not seen).
C. heracleifolia DC. var. hookeri Makino in Bot. Mag. Tokyo 21: 27. 1907, nec C.
hookeri Decne., nec C. tubulosa Turcz. var. hookeri (Decne.) Hook. f.; Matsum., Ind. Pl. Jap. 2
(2): 111. 1912.
C. tubulosa auct. non Turcz.: Koidz., Fl. Symb. Or.-Asiat. 47. 1930.
Fig. 1: K–L; Fig. 12
Subshrub or perennial herb. Stems 10–80 cm tall, few-branched; hornotinous branches
2.2–11 cm long, terete or 6-angulate, shallowly 6–10-sulcate, puberulous. Leaves ternate;
terminal leaflets herbaceous, broadly ovate, broadly elliptic, rhombic, or pentagonous, 5–17×
2–14 cm, apex acuminate or acute, base broadly cuneate, subtruncate, or subcordate, margin
denticulate with teeth usually strongly reduced in size and their mucrones remained, usually
3-lobed, adaxial surface sparsely puberulous, abaxial surface more or less reticulate, sparsely
puberulous on veins; lateral leaflets smaller, obliquely ovate, undivided or 1–2-lobulate;
petioles 3.5–11.5 cm long, puberulous. Inflorescences terminal, 11–24 cm long, few- to
many-flowered, with 1–4 nodes and with fascicled flowers; peduncles usually absent; bracts
ovate, 4–7 mm long, abaxially velutinous. Flowers polygamous, 1.8–2.4 cm long; pedicels
0.6–2.2 cm long, densely puberulous or velutinous. Sepals 4, lanceolate-linear, 18–24×2–5
mm, glabrous inside, densely appressed-puberulous outside, velutinous on margin, apex
slightly dilated and recurved. Stamens 8–11 mm long; filaments oblanceolate-linear, near apex
pilose; anthers lanceolate-linear, ca. 3 mm long, abaxially sparsely pilose, apex often apiculate.
Ovaries densely puberulous; styles ca. 4.5 mm long, densely villous. Achenes compressed,
ovate or narrowly ovate, 2–3×1.2 mm, densely puberulous; persistent styles ca. 1.6 cm long,
plumose. Fl. Aug.–Oct.
Japan (Kyushu, Shikoku). On limestone or in gravelly places; alt. 450–1100 mm.
Additional specimens examined.
Japan. Hinataku, Tashiro s.n. (KYO); Hyuga: Mt. Dodake, Sako s.n., Sako & Kawanabe 22497 (KAG);
Kochi: Hatagun, Yamamoto s.n. (KYO); Kumamoto, Shimizu 5230, 6023 (KYO); Miyazaki: Mt. Dodake,
Shimizu 3365 (KYO); Nishigomura, Nadani 24806 (KYO); Oita: Saiki, Hatushima 19719 (KAG); Tosa,
Yamamoto s.n. (KYO).
8. Clematis stans Sieb. & Zucc. in Abh. Bayer. Akad. Wiss. Math.-Phys. 4 (2): 177. 1845;
Walp. in Ann. Bot. Syst. 1: 953. 1848–1849; Miq., Ann. Mus. Bot. Lugd.-Bat. 3: 1. 1867;
Regel in Gartenfl. 19: 203, t. 357. 1870; Franch. & Savat., Enum. Pl. Jap. 1: 2. 1875; Maxim.
in Bull. Acad. Sci. St.-Petersb. 22: 215. 1877; Decne. in Nouv. Arch. Mus. Hist. Nat. Paris, ser.
2, 4: 407, pl. 12. 1881; Lavall., Clemat. 83. 1884; Hook. f. in Curtis, Bot. Mag. 111: t. 6810.
1885; Schneid., Ill. Handb. Laubh. 1: 281, fig. 1841, fig. 185h–i. 1906; Kitagawa in J. Jap.
Bot. 13: 350. 1937; Rehd., Man. Cult. Trees & Shrubs, ed. 2, 210. 1951; Makino, Ill. Fl. Jap.,
rev. ed., 552, fig. 1656. 1953; Ohwi, Fl. Jap. 441. 1965; Kitamura & Murata, Colour. Ill. Herb.
Pl. Jap., rev. ed., 2: 223, pl. 51: 436. 1980; Tamura in Satake et al., Wild Flow. Jap. 2: 73, pl.
70: 2, 3. 1982; Ohwi & Kitagawa, New Fl. Jap. 679. 1992; M. Johnson, Klematis 273. 1997;
T. Y. A. Yang & Moore in Syst. Geogr. Pl. 68: 299. 1999, p.p. excl. syn. C. hookeri Decne.;
Grey-Wilson, Clematis 191. 2000.——C. heracleifolia DC. ssp. stans (Sieb. & Zucc.) Kuntze
in Verh. Bot. Ver. Brand. 26: 183. 1885.——C. heracleifolia var. stans (Sieb. & Zucc.) Huth in
Bull. Herb. Boiss. 5: 1062. 1897; Matsum., Ind. Pl. Japon. 2: 112. 1912.——C. stans var.
typica Schneid. l.c. Type: Japan. Without precise locality, Siebold s.n. (lectotype, LE!——
Yang & Moore, 1999).

Acta Phytotaxonomica Sinica Vol. 45 452


Fig. 12. Clematis speciosa (Makino) Makino. A, upper part of a bisexual flowering plant; B, stamen. Drawn from
Dadani 24806. C, achene. Drawn from Shimizu 3365.

No. 4 WANG & XIE: A revision of Clematis sect. Tubulosae (Ranunculaceae) 453
C. kousaboton Decne. in Nouv. Arch. Mus. Hist. Nat. Paris, ser. 2, 4: 208, pl. 13.
1881.——C. heracleifolia DC. ssp. lavalei var. kousaboton (Decne.) Kuntze in l.c., cum f.
monoica Kuntze et f. affinis Kuntze.——C. stans var. kousaboton (Decne.) Schneid., l.c. No
type specimen designated.
C. lavallei Decne. in l.c. 209, pl. 14. 1881; Lavall., Clemat. 83. 1884.——C. heracleifolia
ssp. lavallei (Decne.) Kuntze in l.c. 183. 1885.——C. heracleifolia var. lavallei (Decne.) Huth
in Bull. Herb. Boiss. 5: 1062. 1897; Matsum., Ind. Pl. Japon. 2: 112. 1912.——C. stans var.
lavallei (Decne.) Schneid., l.c. No type specimen designated.
C. lavallei var. foliosa Decne. in l.c. 210, pl. 15. 1881. No type specimen designated.
C. savatieri Decne. in l.c. 211, pl. 16. 1881; Lavall., Clemat. 83. 1884.——C.
heracleifolia ssp. savatieri (Decne.) Kuntze in l.c. 184. 1885.——C. heracleifolia var.
savatieri (Decne.) Huth in Bull. Herb. Boiss. 5: 1062. 1897; Matsum., Ind. Pl. Japon. 2: 112.
1912. No type specimen designated.
C. stans var. monoica Lavall., Clemat. 83. 1884. Type unknown.
C. heracleifolia ssp. stans var. decaisneana Kuntze, var. maximowicziana Kuntze, et var.
savatieroides Kuntze in l.c. 183. 1885. Type unknown.
C. heracleifolia ssp. lavallei var. lanceolata Kuntze in l.c. 183. 1885. Type unknown.
C. maximowicziana Decne. ex Rehd & Wils. in Sarg., Pl. Wils. 1: 321. 1913, pro syn.
C. heracleifolia auct. non DC.: Forbes in J. Bot. 22: 265. 1884, p.p. quoad syn. C. stans
Sieb. & Zucc.; Huth in Bull. Herb. Boiss. 5: 1061. 1897; Finet & Gagnep. in Bull. Soc. Bot.
France 50: 545. 1903, p.p. quoad syn. C. stans; Matsum., Ind. Pl. Japon. 2: 111. 1912.
8a. var. stans Fig. 1: G, H (see page 429); Fig. 13
Perennial herb or subshrub. Stems 50–100 cm tall, shallowly 10-sulcate, sparsely
appressed-puberulous or subglabrous. Leaves ternate; terminal leaflets chartaceous, broadly
rhombic-obovate, obovate, elliptic, or pentagonous, 4–12×4–10 cm, apex acuminate, base
broadly cuneate or subtruncate, margin dentate, often 2–3-lobed, adaxial surface subglabrous,
abaxial surface reticulate, on veins puberulous; lateral leaflets smaller, obliquely ovate or
rhombic; petioles 1.5–12 cm long. Panicles usually pedunculate, 2–3 times branched,
many-flowered; bracts folii form or small, linear, undivided or 3-sect, 3–12 mm long,
puberulous. Flowers unisexual; pedicels 4–18 mm long, velutinous. Sepals 4, purplish,
narrowly oblong or narrowly obovate-oblong, 12–16(–20)×1.8–3 mm, glabrous inside,
appressed-puberulous outside, velutinous on margin, apex slightly narrowed or slightly
dilated, recurved, rarely strongly recurved or circinate. Stamens 15–18, 8–10 mm long;
filaments 2.2–4.8 mm long, pilose; anthers linear, 4–5 mm long, connective pilose; pollen
grains pantoporate. Staminodes of pistillate flower ca. 18, 5–7 mm long; abortive anthers
subulate, 1.5–3 mm long, pilose or subglabrous. Carpels ca. 20; ovaries densely pubescent;
styles 4–5 mm long, densely villous. Abortive carpels of staminate flower several, 1–3 mm
long. Achenes compressed, elliptic, 2–3.5×1.2–2.2 mm, puberulous, slightly tumidly rimmed;
persistent styles 1.2–2.2 cm long, plumose, but with glabrous bases. Fl. Jul.–Sept.
Japan (Honshu, S Hokkaido). On slopes, in grassy places, at thicket edges, or by lake; alt.
100–2000 m.
Additional specimens examined:
Japan. Honshu: Aomori, Mt. Hakkoda, Faurie 868 (P), Naito 1297 (GH, MO); Chiba: Mt. Kiyosomi,
Makino 76714 (KAG); Echigo, Kobayashi 12853, 14191 (S); Hyogo, Boufford 19527 (GH); Ikao, Bisset 3243
(BM); Ishikawa, Deguchi 16719 (GH), Tsugaru 7199 (GH, MO); Iwate, Suzuki s.n., Koyama s.n. (GH);
Kanagawa: Sagami, Furuse 34671 (GH, PE), Mizushima 144 (GH), Yamazaki 2622 (S); Karuizawa, Fox s.n.
(BM); Miyagi, Kurasawa 821 (GH); Mt. Echigo-Koma, Ohwi s.n. (K); Mt. Juji, Kubo 213 (UPS); Musashi:
Nippara, Togashi 1219 (BM, G, GH, K, KAG, LE, P, PE); Mutsu, Tamura 9309 (S); Nagana, Boufford 23495
(GH, MO), Furuse 36064 (PE, S), Shimizu 5401 (S), Tateishi & Togushi 513 (BM, K, P); Nakauonuma,
Kobayashi 12853 (S); Saitama, Shimizu 13672 (S); Senano, Tschnoski s.n. (BM, K); Shiga, Tateishi 12025
(GH); Shizuoka: Tashiro, Konda 15778 (PE); Tochigi, Furuse 28035 (PE); Tokyo, Suzuki 273 (GH); Totomi:
Senzu, Ohwi & Koyama 1108 (G, GH, K, KAG, LE, P, PE, S, UPS); Ugo, Ikeda s.n. (BM); Yamagata,
Acta Phytotaxonomica Sinica Vol. 45 454

Fig. 13. Clematis stans Sieb. & Zucc. var. stans. A, upper part of a staminate flowering plant; B, sepal outside; C,
stamen. Drawn from Furuse 36064. D, achene. Drawn from Furuse 34671.

No. 4 WANG & XIE: A revision of Clematis sect. Tubulosae (Ranunculaceae) 455
Boufford 19883 (MO), Takahashi 309 (GH); Yamanashi, Furuse 9721 (K, PE), 29800 (PE, S), 36136 (GH,
PE), Tamura & Hotta 3751 (KYO, PE), Togashi 512 (BM, GH, P); Yokohama, Maximowicz s.n. (K, P, S);
Yokoska, Savatier 5 (P). Hokkaido. Hakodake, Albrecht s.n. (G, GH, K, LE, UPS), Maximowicz s.n. (BM),
Yatabe s.n. (UPS).
8b. var. austrojaponensis (Ohwi) Ohwi, Fl. Jap. 441. 1965; Kitamura & Murata, Colour. Ill.
Herb. Pl. Jap., rev. ed., 2: 223. 1980; Tamura in Satake et al., Wild Flow. Jap. 2: 73. 1982; M.
Johnson, Klematis 276. 1997; Grey-Wilson, Clematis 192. 2000.——C. austrojaponensis
Ohwi in Acta Phytotax. Geobot. 7: 45. 1938. Type: Japan. Kyushu: Hyuga, Mt. Dogatake,
1915-08-23, Tashiro s.n. (holotype, KYO!). Kyushu: Higo, Mt. Nokieboshi, Tashiro s.n.
(paratype, KYO!), Mt. Naidaijin, Tashiro s.n. (paratype, KYO!), Mt. Siroiwa, Tashiro s.n.
(paratype, KYO!).
Fig. 1: I, J (see page 429)
This variety differs from var. stans in its shorter anthers 3–3.8 mm long.
Japan (Kyushu, Shikoku). On limestone cliffs; alt. 900–1600 m.
Additional specimens examined:
Japan. Higo, Gokanosyo, Hatushima & Sako 27817, 32255, Sako 1047 (KAG); Hintaku, Tashiro s.n.
(KAG, KYO); Hyuga: Mt. Shiraiva, Hatushima & Sako 26198 (KAG, KYO); Kumamoto, Hatushima 31886
(KAG), Shimizu 5046 (KYO); Tokushima: Naka, Takato 957, 1166 (KYO).
Ser. 2. Uniflorae W. T. Wang in Acta Phytotax. Sin. 39: 10. 2001; et 43: 478. 2005. Type:
C. tsugetorum Ohwi.
Small shrubs. Leaves ternate or 5-foliolately pinnate. Flowers terminal, solitary, rarely in
2–3-flowered cymes, polygamous. Pollen grains pantoporate.
One species, endemic to Taiwan, China.
9. Clematis tsugetorum Ohwi in Acta Phytotax. Geobot. 2: 153. 1933; T. S. Liu & Hsieh in
Fl. Taiwan 2: 493. 1976; M. Y. Fang in Fl. Reip. Pop. Sin. 28: 93, fig. 2. 1980; T. Y. A. Yang &
T. C. Huang in Taiwania 37: 48, pl. 13. 1992; et 40: 247. 1995; et in Fl. Taiwan, ed. 2, 2: 541,
pl. 256. 1996; M. Johnson, Klematis 276. 1997; T. Y. A. Yang & Moore in Syst. Geogr. Pl. 68:
299. 1999; Grey-Wilson, Clematis 189. 2000; W. T. Wang & Barth. in Fl. China 6: 371. 2001.
Type: China. Taiwan (台湾): Taizhong (台中), Nenggao Shan (能高山), 1933-06-15, Ohwi
3277 (holotype, KYO!; isotype, UPS!).
高山铁线莲 Fig. 11: A–D
Small shrub. Stems 30–60 cm tall; branches shallowly 6-sulcate, puberulous. Leaves
ternate or 5-foliolately pinnate; leaflets papery, broadly ovate, orbicular-ovate, or ovate, 1–3.8
×1–3 cm, apex acute or shortly acuminate, base truncate, rounded, or broadly cuneate, margin
unequally dentate, undivided or 3-lobed, adaxial surface sparsely puberulous, abaxially
puberulous on veins, basal veins abaxially prominent; petioles 1–5 cm long. Flowers
polygamous, terminal, solitary, rarely in 2–3-flowered cymes; pedicels 2–6 cm long, densely
puberulous. Sepals 4, bluish or purple, narrowly oblong, 10–20×3–7 mm, slightly dilated
toward apex, glabrous inside, densely puberulous outside, velutinous on margin, apex
mucronate and recurved. Stamens 5–10 mm long; filaments pilose near apex; anthers narrowly
oblong, ca. 2.5 mm long, on connective pilose, apex minutely apiculate. Staminodes of
pistillate flower several, 3–4 mm long, with broadly linear filaments and thinly subulate
abortive anthers. Carpels numerous; ovaries pubescent; styles 6 mm long, densely villous.
Abortive carpels of staminate flower several. Achenes compressed, elliptic, 3–5×2–2.5 mm,
pubescent; persistent styles 1.8–2.5 cm long, plumose. Fl. Jul.–Sept.
China (Taiwan). In limestone montane regions; alt. 2400–3600 m.
Additional specimens examined:
China. Taiwan (台湾): Hualian (花莲), Ohwi 3162 (KYO); Yilan (宜兰), en route from Mt.
Chong-Yang-Chien to Nan-shan, Tamura & Koyama 23654 (KYO, S); Nantou (南投). Mt. Neng-kao, Tamura
& Koyama 23330 (KYO, S); Taizhong (台中), Nanhu Dashan (南湖大山), J. C. Wang 3752, 3753 (GH).
Acta Phytotaxonomica Sinica Vol. 45 456
Acknowledgements The senior author of this paper is most grateful to the directors and
curators of ANUB, BJFC, BM, BNU, CAF, G, GH, HHBG, HIMC, HUTM, IBSC, IFP, IMD,
K, KAG, KUN, KYO, LE, MO, NAS, P, S, TI, TIE, UPS, and WUK for kindly inviting him to
visit their herbaria or/and sending herbarium material on loan; to F. Jacquemoud (G), S.
Landrein (K) and JIN Xiao-Feng (HZU) for kindly providing some literature; to P. P. Lowry II
for critical reading of the manuscript and improving the English; to LI Liang-Qian, LI
Zhen-Yu, ZHU Xiang-Yun, ZHANG Shu-Ren, WANG Zhong-Tao, and BAN Qin for various
kinds of help during the preparation of the present revision; and to SUN Ying-Bao for making
the drawings.
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铁线莲属大叶铁线莲组修订
王文采 谢 磊
(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Ranunculaceae铁线莲属Clematis的大叶铁线莲组sect. Tubulosae进行了修订, 确定此组
含9种2变种和3变型, 对此组的分类简史和地理分布做了介绍, 研究了此属大多数种的花粉形态, 写出
此组组下分类群的形态特征、地理分布等, 并附有全部种的插图。此组9种被划分为2亚组, 其中原始群
羽叶铁线莲亚组subsect. Pinnatae(有2种, 1种分布于中国河北和东北, 另1种产日本)在木质藤本习性、花
构造、花粉形态(具3沟)等方面与威灵仙组sect. Clematis颇为近似, 区别主要在于萼片在开放初期近直
立, 以后平展, 雄蕊花丝被毛, 此亚组可能源于威灵仙组。进化群大叶铁线莲亚组subsect. Tubulosae(有
7种, 分布于我国东部、北部和台湾, 朝鲜和日本)为直立多年生草本、小亚灌木或小灌木, 花通常杂性,
萼片直立(花萼呈筒状, 稀呈坛状), 顶端或上部反曲, 雄蕊常有毛, 花粉通常具散孔, 只在1种(原始种
大叶铁线莲C. heracleifolia)具3沟, 此亚组可能由羽叶铁线莲亚组演化而来。
关键词 铁线莲属; 大叶铁线莲组; 毛茛科; 分类学修订