全 文 ：热带亚热带植物学报　2015， 23(2)： 123 ~ 132Journal of Tropical and Subtropical Botany
Received: 2014–09–29　　　　Accepted: 2014–12–04
This study was supported by the Science and Technology Basic Work of China (2013FY112100).
* Corresponding author. E-mail: qeyang@scib.ac.cn
涪陵银莲花(毛茛科)的名实订正
张煜1,2, 杨亲二1*
(1. 中国科学院华南植物园， 中国科学院植物资源保护与可持续利用重点实验室， 广州 510650； 2. 中国科学院大学， 北京， 100049)

摘要： 通过标本检查，发现毛茛科涪陵银莲花(Anemone fulingensis W. T. Wang & Z. Y. Liu)与川西银莲花(A. prattii Huth ex Ulbr.)
属于同一种植物，故将前者处理为后者的异名。揭示川西银莲花的花粉具 6~10带沟。澄清了川西银莲花的地理分布，讨论了
与其近缘种滇川银莲花(A. delavayi Franch.)的形态区别。
关键词： 银莲花属； 新异名； 毛茛科； 分类学
doi: 10.11926/j.issn.1005–3395.2015.02.002
The Identity of Anemone fulingensis (Ranunculaceae)
ZHANG Yu1,2, YANG Qin-er1*
(1. Key Laboratory of Plant Resources Conservation and Sustainable Utilization, South China Botanical Garden, Chinese Academy of Sciences,
Guangzhou 510650, China; 2. University of Chinese Academy of Sciences, Beijing 100049, China)
Abstract: Examination of herbarium specimens has shown that Anemone fulingensis W. T. Wang & Z. Y. Liu
(Ranunculaceae) is conspecifi c with A. prattii Huth ex Ulbr. We therefore reduce A. fulingensis to the synonymy
of A. prattii. Pollen grains in A. prattii are revealed to be 6–10-zonocolpate. The geographical range of A. prattii is
clarifi ed, and its morphological differentiation from A. delavayi Franch. is discussed.
Key words: Anemone; New synonymy; Ranunculaceae; Taxonomy
Anemone fulingensis W. T. Wang & Z. Y. Liu
(Ranunculaceae) was described on the basis of two
collections, Z. Y. Liu & M. B. Ren 2060366 (PE) and
205007 (PE) (Fig. 1), from Fuling, Chongqing, China,
with the former being designated as the holotype[1].
In the protologue, the authors stated that the species
was a member of A. sect. Stolonifera (Ulbr.) Juz. by
having sessile involucral bracts, 4–5 sepals, filiform
fi laments, and pantocolpate pollen, but distinguishable
from all other species of the section by its moniliform
rhizome consisting of 2–10 tubers. They stressed that
the moniliform rhizome of the species was not only
unique in A. sect. Stolonifera but also in the whole
genus Anemone L.
Judging from its external morphology and pollen
morphology (for the correct description of the pollen,
see below), Anemone fulingensis is indeed readily
referable to A. sect. Stolonifera. However, its moniliform
rhizome is far from a unique character in the section.
Our examination of ample material in major Chinese
herbaria has shown that such rhizome occurs at least
in three species of the section, i.e., A. baicalensis Turcz.
(Fig. 2), A. delavayi Franch. (Fig. 3), and A. prattii
Huth ex Ulbr. (Fig. 4). In literature, the rhizomes of
these species are often described as having long and
slender internodes with the nodes remote from each
124 第23卷热带亚热带植物学报
Fig. 1 Specimens of Anemone prattii. A: Z. Y. Liu and M. B. Ren 2060366 (holotype of A. fulingensis, PE), Fuling, Chongqing, China; B, C: Z. Y. Liu
and M. B. Ren 2060366 (isotypes of A. fulingensis, PE); D: Z. Y. Liu 2050007 (paratype of A. fulingensis, PE), Fuling, Chongqing, China.
第2期 125
Fig. 2 Specimens of Anemone baicalensis. A: Baishuijiang Exped. 5765 (PE), Wenxian, Gansu, China; B: Temperate Forest Exped. 45 (PE), Changbai
Shan, Jilin, China; C: Qinling Exped. 10257 (PE), Taibai Shan, Shaanxi, China; D: Farges 386 (P), Chengkou, Chongqing, China. Arrows indicates
tuber-like nodes of the rhizome.
张煜等：涪陵银莲花(毛茛科)的名实订正
126 第23卷热带亚热带植物学报
Fig. 3 Specimens of Anemone delavayi. A: T. T. Yu 19291 (PE), Deqen, Yunnan, China; B: Delavay 1504 (lectotype, here designated, P), Heqing,
Yunnan, China; C: Inst. Bot. Hengduan Shan Exped. 135 (PE), Lushui, Yunnan, China; D: T. T. Yu 11557 (PE), Zhongdian, Yunnan, China. Arrows
indicate tuber-like nodes of the rhizome.
第2期 127
Fig. 4 Specimens of Anemone prattii. A: Pratt 773 (lectotype, P), Ta-tsien-lou (= Kangding), Sichuan, China; B: K. C. Kuan et al. 440 (PE), Emei
Shan, Sichuan, China; C: Veget. Exped. 41688 (CDBI), Luding, Sichuan, China; D: Y. B. Yang 21715 (CDBI), Tianquan, Sichuan, China. Arrows
indicate tuber-like nodes of the rhizome.
张煜等：涪陵银莲花(毛茛科)的名实订正
128 第23卷热带亚热带植物学报
other.
Among the three species just mentioned, Anemone
prattii caught our special attention because of its great
resemblance with A. fulingensis in general aspect.
Ulbrich[2] described A. prattii on the basis of a collection,
Pratt 773, from the neighborhood of Kangding, western
Sichuan, China. He described the rhizome of the
species as having very strongly elongated internodes,
with the tuber-like nodes 3–5 cm distant from each
other. The rhizome was also clearly illustrated in the
accompanying line drawing in the paper. The original
material of A. prattii deposited in the Botanical Garden
and Botanical Museum Berlin-Dahlem (B) was most
likely destroyed in World War II. Fortunately, we found
a duplicate of Pratt 773 (Fig. 4: A) from the National
Natural History Museum, Paris (P). This specimen
had been previously identifi ed as A. baicalensis, but it
matches perfectly the original description of A. prattii
and thus we designate it as the lectotype herein. In
describing A. fulingensis, Wang et al.[1] must have not
examined the type material and original description of
A. prattii, and thus did not compare their A. fulingensis
with A. prattii. As shown in Figures 1, 4 and Table 1,
A. fulingensis is not different from A. prattii in any
essentials. In the protologue, A. fulingensis was described
as having only one pedicel, but actually it has one or
two pedicels as illustrated in Figure 1.
In pollen morphology, Anemone fulingensis is also
Table 1 Comparison of morphological characters for Anemone fulingensis and A. prattii
A. fulingensis A. prattii
Plant height (cm) 15–25 11–30
Habit Erect herb with 1–2 basal leaves Erect herb with 1–3 basal leaves
Perennating structure Rhizomes, tubers Rhizomes, tubers
Leaf shape and dissection Cordate-pentagonal; 3-sect, central segment 3-lobed,
rhombic, base cuneate, margin incised serrate, lateral
segments 2-parted, obliquely fl abellate
Cordate-pentagonal; 3-sect, central segment 3-lobed,
rhombic, base cuneate, margin incised serrate, lateral
segments 2-parted, obliquely fl abellate
Leaf size (cm) 3–5 × 5.5–7.6 3–3.6 × 4.8–5.5
Leaf pubescence Sparsely puberulent adaxially, subglabrous abaxially Sparsely puberulent adaxially, subglabrous abaxially
Petiole Glabrous Glabrous
Scape Glabrous Glabrous or distally sparsely puberulent
Involucral bracts Rhombic, 3-partite or 3-lobed , 2.6–4 cm × 1.2–4.8 cm Rhombic, 3-partite or 3-lobed, 2–5 cm × 0.8–4 cm
Pedicel number (fl ower number) 1–2 1–3
Sepal 4–5, white; 9.5–12 mm × 3.2–7 mm 5, white; 8–10 mm × 4 –7 mm
Carpel number 5–6 4–7
Achene Obovoid, densely puberulent Obovoid, densely puberulent
not essentially different from A. prattii. Wang et al.[1]
reported A. fulingensis to be 6-colpate. Xi and Chang[3]
reported A. prattii to be 8(–10)-colpate, and Fang
and Yang[4] reported it to be (6–)7(–10)-colpate. The
material of A. prattii that we examined had 8-zonocolpate
pollen (Fig. 5). Obviously A. prattii is somewhat
variable in the number of pollen aperture. It is to be
noted that Wang et al.[1] was wrong to have described
the pollen of A. fulingensis as “pantocolpate”. The six
apertures of pollen in A. fulingensis are situated at the
equator and thus the pollen should be referred to as “6-
zonocolpate”[5] or “stephanocolpate”[6]. In their studies
of the pollen in Anemone, Xi and Chang[3], and Huynh[7]
described such pollen as “polycolpate”. In pantocolpate
pollen grains, the apertures are more or less regularly
distributed over the whole surface[6], such as those in
Anemone griffithii Hook. f. & Thoms.[3–4,8]
From the above analyses it is necessary to reduce
Anemone fulingensis to the synonymy of A. prattii.
第2期 129
Fig. 5 Pollen grains of Anemone prattii. [Scanning electron microscopy, both same scale; voucher: L. X. Zhou 12132 (IBSC), Mabian, Sichuan, China],
8-zonocolpate. A: Polar view; B: Equatorial view.
Anemone prattii Huth ex Ulbr. in Bot. Jahrb.
Syst. 36 (Beibl. 80): 4. 1905; W. T. Wang in Fl.
Reipubl. Popularis Sin. 28: 18. 1980, p.p.; L. Q. Li in
Vas. Pl. Hengduan Mount. 1: 510. 1993, p.p.; W. T.
Wang in Fl. Yunnan. 11: 188. 2000, p.p.; W. T. Wang
et al. in Fl. China 6: 312. 2001, p.p.; Ziman et al. in
J. Jpn. Bot. 79: 296. 2004, p.p. Type: China. Sichuan:
Tachienlu (= Kangding), alt. 9000–13500 ft, 1890,
Pratt 773 (B, most likely destroyed; lectotype, here
designated, P!).
A. fulingensis W. T. Wang & Z. Y. Liu in Acta
Phytotax. Sin. 45: 290. 2007. syn. nov. Type. China.
Chongqing: Fuling, Damu Xiang, in shady moist
forest on slope in ravine, alt. 1420 m, 22 Apr. 2006, Z.
Y. Liu & M. B. Ren 2060366 (holotype, PE!; isotypes,
PE!).
Notes. Ziman et al.[9] have wrongly cited Ducloux
5678 (P; Fig. 6: A) from Qiaojia, northeastern Yunnan,
China as the holotype of Anemone prattii. Ulbrich[1]
did not cite this specimen when he described A. prattii,
with his description based on only Pratt 773. In fact,
Ducloux 5678 was collected in 1908 whereas A. prattii
was described in 1905. It was Handel-Mazzetti[10] who
referred Ducloux 5678 (P) to A. prattii for the fi rst
time. This specimen, in our opinion, should belong
to A. delavayi. Handel-Mazzetti, however, correctly
identifi ed Ducloux 6170 (P; Fig. 6: B) from Qiaojia
as A. prattii on the determinavit slip of the specimen,
although he did not mention this collection in any of
his works.
Anemone pratti has long been recorded to occur
in Deqen, northwestern Yunnan, China[11–14]. This wrong
record is obviously attributed to Handel-Mazzetti’s
misidentifi cation of his own collection, Handel-Mazzetti
8881 (WU; Fig. 6: C)[15]. This specimen, just like Ducloux
5678 from Qiaojia, should be referred to A. delavayi.
He also misidentified Ducloux 4592 (P; Fig. 6: D)
from Binchuan, northwestern Yunnan as A. pratti on
the determinavit slip; this specimen also should be
identifi ed as A. delavayi. Another collection from
northwestern Yunnan, Forrest 21495 (not shown here),
which Handel-Mazzetti first considered to represent
the transition from A. baicalensis to A. prattii[15], and
then to be most likely A. prattii[10], has been correctly
identified as A. delavayi at PE. All these specimens
have a slender rhizome with more or less tuber-like
nodes, indicating that Handel-Mazzetti regarded the
presence of tubers as an important diagnostic character
of A. prattii.
Morphologically, Anemone delavayi is indeed
somewhat confused with A. prattii owing to their
similarity in rhizome structure. The involucral bracts
张煜等：涪陵银莲花(毛茛科)的名实订正
130 第23卷热带亚热带植物学报
Fig. 6 Specimens of Anemone delavayi. (A, C, D; all misidentifi ed as A. prattii by Handel-Mazzetti) and A. pratti (B). A: Ducloux 5678 (P), Qiaojia,
Yunnan, China; B: Ducloux 6170 (P), Qiaojia, Yunnan, China; C: Handel-Mazzetti 8881 (WU), Deqen, Yunnan, China; D: Ducloux 4592 (P),
Binchuan, Yunnan, China. Arrows indicate tuber-like nodes of the rhizome.
第2期 131
in A. delavayi, just like those in A. baicalensis, are highly
variable in size, as pointed out by Handel-Mazzetti[15]
in his remarks on A. delavayi and as shown in Figures
3, 6. Some plants of the species have bracts much
smaller than the basal leaves, while some have very
large ones similar to the basal leaves in shape. Those
with large leaf-like bracts are, in particular, easily
confused with A. prattii, in which the bracts are almost
always as large as the basal leaves. It seems that A.
delavayi is different from A. prattii mainly in the basal
leaves solitary or absent (vs. 1–3), and the leaf segments
undivided or only shallowly divided (vs. deeply divided).
Ziman et al.[9,16] classifi ed A. baicalensis, A. prattii and A.
delavayi in different series under A. sect. Stolonifera,
with the former two in ser. Stolonifera Ziman et al.
and the latter one in ser. Flaccidae Juz. It seems to us
that these three species are morphologically so closely
similar to each other that they may better be placed
together in a series.
Additional specimens examined. Chongqing:
Fuling, Z. Y. Liu & M. B. Ren 205007 (PE); Sichuan:
Baoxing, Anonymous 00247 (PE); Dujiangyan, F.
T. Wang 20845 (NAS, PE), Q. E. Yang & M. Feng
95028 (PE); Ebian, T. T. Yu 656 (NAS, PE); Emei,
Anonymous 54210 (PE), K. C. Kuan et al. 373 (CDBI,
PE), 440 (PE), G. H. Yang 54266 (PE), T. T. Yu 361
(PE), 461 (NAS, PE); Luding, Sichuan Veget. Exped.
25118 (CDBI)，41688 (CDBI), 42059 (CDBI);
Mabian, W. P. Fang 457 (NAS, PE); Shimian, C. J.
Xie 40035 (PE), 40059 (PE); Tianquan, T. Naito et al.
153 (PE), X. L. Jiang 33722 (IBSC, PE), Y. B. Yang
21715 (CDBI); Yunnan: Qiaojia, Ducloux 6170 (P).
Distribution and habitat. Anemone prattii is
distributed in Chongqing (Fuling), western Sichuan,
Fig. 7 Distribution of Anemone prattii (●).
and northeastern Yunnan (Qiaojia), China. It grows in
shady places in mixed forests or thickets at altitudes
of 1400–2400 m above sea level.
Acknowledgments　　We are grateful to the curators of CDBI,
IBSC, NAS, P, PE and WU for the permission to use their scanned
images of specimens and for research facilities.
References
[1]　 Wang W T, Liu Z Y, Xie L. Anemone fulingensis W. T. Wang & Z.
Y. Liu, a new species of Ranunculaceae from Chongqing, China
[J]. Acta Phytotax Sin, 2007, 45(3): 289–292. (in Chinese)
[2]　 Ulbrich E. Über einige neue Ranunculaceen ostasiens [J]. Bot
Jahrb Syst, 1905, 36(80): 19–38.
[3]　 Xi Y Z, Chang K T. Studies on pollen morphology of Anemone L.
[J]. Acta Bot Sin, 1964, 12(1): 19–38. (in Chinese)
张煜等：涪陵银莲花(毛茛科)的名实订正
132 第23卷热带亚热带植物学报
[4]　 Fang M Y, Yang M E. A study on pollen morphology and evolution
of the genus Anemone from Sichuan [J]. J Sichuan Univ (Nat Sci),
1994, 31(2): 246–258. (in Chinese)
[5]　 Nowicke J W, Skvarla J J. Pollen morphology [M]// Die Natürlichen
Pflanzenfamilien, Band 17 a IV. Berlin: Duncker & Humblot,
1995: 129–159.
[6]　 Hesse M, Halbritter H, Zetter R, et al. Pollen Terminology [M].
Wien: Springer-Verlag, 2009: 1–261.
[7]　 Huynh K L. Le pollen du genre Anemone et du genre Hepatica
(Ranunculaceae) et leur taxonomie [J]. Pollen et Spores, 1970, 12
(3): 329–364.
[8]　 Zhang Y, Yang Q E. Notes on the geographical range of Anemone
davidii (Ranunculaceae) with special reference to pollen morphology
[J]. J Trop Subtrop Bot, 2015, 23(1): 25–30. (in Chinese)
[9]　 Ziman S N, Keener C S, Kadota Y, et al. A taxonomic revision
of Anemone L. subgenus Anemonanthea (DC.) Juz. sensu lato
(Ranunculaceae) III [J]. J Jpn Bot, 2004, 79(5): 281–310.
[10]　 Handel-Mazzettii H. Plantae sinensis: XXXIII. Ranunculaceae
[J]. Acta Horti Gothob, 1939, 13: 37–219.
[11]　 Wang W T. Anemone L. [M]// Flora Reipublicae Popularis Sinicae,
Tomus 27. Beijing: Science Press, 1980: 1–56. (in Chinese)
[12]　 Wang W T. Anemone L. [M]// Flora Yunnanica, Tomus 11. Beijing:
Science Press, 2000: 183–204. (in Chinese)
[13]　 Li L Q. Anemone L. [M]// Vascular Plants of the Hengduan Moun-
tains, Vol. 1. Beijing: Science Press, 1993: 506–516. (in Chinese)
[14]　 Wang W T, Ziman S N, Dutton B E. Anemone L. [M]// Flora
of China, Vol. 6. Beijing: Science Press & St. Louis: Missouri
Botanical Garden Press, 2001: 307–328.
[15]　 Handel-Mazzetti H. Symbolae Sinicae, Teil. 7(2) [M]// Wien:
Verlag von Julius Springer, 1931: 211–448.
[16]　 Ziman S N, Keener C S, Kadota Y, et al. A taxonomic revision
of Anemone L. subgenus Anemonanthea (DC.) Juz. sensu lato
(Ranunculaceae) I [J]. J Jpn Bot, 2004, 79(1): 43–71.