全 文 ：植 物 分 类 学 报 45 (5): 742–750(2007) doi:10.1360/aps06028
Acta Phytotaxonomica Sinica http://www.plantsystematics.com
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Received: 24 February 2006 Accepted: 12 September 2006
Supported by the National Natural Science Foundation of China, Grant No. 30370111.
* Author for correspondence. E-mail: .
Comparison and assessment of three East-Asian species of
the genus Scapania (Hepaticae: Scapaniaceae)
1, 2, 3ZUO Ben-Rong 2, 1CAO Tong* 2GUO Shui-Liang
1 (Institute of Applied Ecology, the Chinese Academy of Sciences, Shenyang 110016, China)
2 (College of Life & Environmental Sciences, Shanghai Normal University, Shanghai 200234, China)
3 (Graduate University of the Chinese Academy of Sciences, Beijing 100049, China)
Abstract Three East-Asian species of the hepatic genus Scapania, namely, S. parvidens
Steph., S. parvitexta Steph., and S. glaucoviridis Horik., which were recently treated as one
species (S. parvidens) by Potemkin, are described and typified. Examination and comparison
of the types as well as representative specimens showed that these three should be treated as
separate species. Descriptions with figures for each species are presented and differences
among the three species are discussed in detail.
Key words Hepaticae, Scapaniaceae, Scapania, Scapania parvidens, Scapania parvitexta,
Scapania glaucoviridis, East-Asia, assessment.
Scapania (Dumort.) Dumort. is the largest genus of the hepatic family Scapaniaceae, and
is distributed mainly in temperate regions and on high mountain areas in tropical and
subtropical regions. About 230 species have been described in the world (Müller, 1905; Long,
1990) and more than 110 species were recognized by Potemkin in 1998. Later in 2002,
Potemkin put forward a phylogeny and classification of the genus Scapania and recognized
87 recent and one fossil species in the genus. He also proposed a number of new synonyms in
the genus. Among them, there are three East-Asian species, namely, S. parvidens Steph. and
S. parvitexta Steph. in Japan and China, as well as S. glaucoviridis Horik. in Japan, China and
Himalayas. Potemkin (2002) treated S. parvidens and S. glaucoviridis as new synonyms of S.
parvitexta because of considerable overlap in variability and some unstable characters among
them.
During our study of Chinese Scapania, we examined the types and other specimens from
China and Japan of these three species. After examination and comparison of types and some
additional specimens, we concluded that the three are different and should be treated as
separate species. Comparison and assessment of these species are presented here.
1. Scapania parvitexta Steph., Bull. Herb. Bossier 5: 107. 1897. Type: “Kattasan, Towada,
Hakodate”, Faurie 151123 (paratype, G!), 14262 (paratype, G). Fig. 1
Plants median in size, 0.5–1.2 cm long, 0.6–1.9 mm wide with leaves, yellow-green.
Stems ascending above, unbranched or rarely forked, in transverse section cortical cells in 3–5
layers, small, thick-walled, interior cells large, thin-walled. Rhizoids scattered on stems, long,
colorless. Leaves densely imbricate, unequally divided 2/5–3/5 their lengths. Keel 0.3–0.6 the
length of the ventral lobe, slightly concave dorsally, in cross section 2–4 cells thick, wing
wanting. Ventral lobe nearly transversely inserted on the stem, oval to obovate, distinctly
convex backwards, more or less decurrent at base, apex rounded to obtuse, sometimes
subacute with a point, margins densely ciliate-dentate to dentate, teeth often acute at tip, 1–4
No. 5 ZUO et al.: Comparison and assessment of three East-Asian species of the genus Scapania 743

Fig. 1. Scapania parvitexta Steph. A–D, leaves; E, F, leaf margin teeth; G, leaf cells near basal margin; H, median leaf
cells; I, basal leaf cells; J, perianth; K, perianth mouth pattern; L, cells of perianth mouth; M, gemmae; N, cross section of
leaf; O, cross section of leaf keel; P, cross section of stem; Q, part of plants with perianths, dorsal view; R, leaf insertion
pattern, dorsal view; S, leaf insertion pattern, ventral view. Drawn by B. R. Zuo from Faurie 151123, paratype of Scapania
parvitexta Steph.
Line scales: 150 µm for A–D, J, R, S; 100 µm for K; 75 µm for Q; 50 µm for E, F, L, P; 25 µm for G–I, M–O.
Acta Phytotaxonomica Sinica Vol. 45 744
cells long, 1–3 cells wide. Dorsal lobe transversely inserted and appressed to the stem, arching
beyond the farther edge of the stem, not decurrent at base, rectangular to obovate, 0.5–0.7 the
ventral lobe in size, apex rounded to obtuse, rarely with a point, margins irregularly dentate to
ciliate-dentate, teeth similar to those of dorsal lobe. Wings absent. Leaf cells along margin
8–10 μm, median cells 12–15 μm wide×17–20 μm long, basal cells 12–15 μm wide×20–34
μm long, walls usually equally thickened, trigones moderately large to small. Cuticle coarsely
verrucose, sometimes with distinct papillae. Dioicous. Perianth terminal, oblong, truncate,
dorsiventrally compressed, the upper part often bent towards the ventral side, mouth with
numerous lobes, lobe with irregular teeth. Gemmae ellipsoid, green to reddish, 1-celled.
Habitat: On granite or in shaded habitats on soil-covered surfaces of rock, and sometimes
on bark of trees or rotten wood.
Distribution: China, Japan.
Stephani (1897) described S. parvitexta based on Japanese specimens. Later, this species
was recorded from Japan by Müller (1905), Stephani (1910), Hattori (1944), Amakawa &
Hattori (1954), etc. It was reported from Anhui (Chen & Wu, 1965; Guo et al., 1988; Piippo,
1990; Zhang et al., 1993; Hu & Liang, 1996; Gao & Cao, 2000), Jiangxi (Zhang et al., 1993;
Fang et al., 1998), Yunnan (Gao & Cao, 2000) and Zhejiang (Liu, 1985; Zhang et al., 1993;
Wang & Hu, 1995; Zhu et al., 1998) provinces in China before the present study. It seems that
S. parvitexta is one of the most common species of Scapania in China. According to the
present study, it is distributed in East China (Anhui, Fujian, Zhejiang, and Jiangxi), South
China (Guangxi), Southwest China (Guizhou, Yunnan, and Xizang (Tibet)), Northeast China
(Liaoning), and Taiwan.
S. parvitexta is sometimes confused with S. parvidens. But comparison of types and
additional specimens from China and Japan showed that these two species differ in size of the
dorsal lobe and leaf margin teeth, as well as in plant size. S. parvitexta has dorsal lobes
distinctly smaller than the ventral lobes; margin teeth large, consisting of numerous cells; and
plants about 1–3 cm long, while S. parvidens has dorsal lobes nearly equal to the ventral lobes
in size; margin teeth small, mostly 1-celled, sometimes 2-celled; and plants about 0.5–1 cm
long.
S. parvitexta differs from S. glaucoviridis in having smaller dorsal lobes, about 0.7–0.8
of the ventral lobes in size, and shorter keels without wings, while S. glaucoviridis has dorsal
lobes nearly equal to the ventral lobes and long, straight keels with distinct wings.
Specimens examined.
China. Anhui (安徽): Mt. Huangshan (黄山), D. K. Li (李登科) 00839 (SHM); B. R. Zuo & G. Y.
Song (左本荣, 宋国元) 0552650-2, 0551379-1, 0551376, 05429175, 05429273, 050503607 (SHNU).
Fujian (福建): Mt. Wuyishan (武夷山), B. J. Lin (林邦娟) 2823 (IFSBH), Training Class of Bryology (苔藓
植物培训班), 978, 57 (PE); Chong’an (崇安), D. K. Li (李登科) 01837 (SHM); Jiangle (将乐), Mt. Longxi
(陇西), M. Z. Wang (汪楣芝) 48822, 48823 (PE). Guangxi (广西): Xing’an (兴安), Mao’ershan (猫儿山),
C. Gao & G. C. Chang (高谦, 张光初) 1447, 1635 (IFSBH). Guizhou (贵州): Guiyang (贵阳), Mt.
Bijieguanmen (毕节关门山), Z. W. Li (李之文) 501 (PE). Jiangxi (江西): Mt. Lushan (庐山), Training
Class of Bryology (苔藓植物培训班) 98, 111 (PE). Liaoning (辽宁): Kuandian (宽甸), Baishilazi (白石砬
子), T. Cao & S. Z. Wang (曹同 , 王淑芝) 38321 (IFSBH). Taiwan (台湾): Taibei (台北), Mt.
Yangmingshan (阳明山), C. Gao & T. Cao (高谦 , 曹同) 980841 (IFSBH); Taizhong (台中), Mt.
Hehuanshan (合欢山), T. Cao & C. Gao (曹同, 高谦) 980169 (IFSBH), Anmashanzhuang (鞍马山庄), T.
Cao & C. Gao (曹同, 高谦) 981433 (IFSBH); Xinzhu (新竹), Yuanyanghu Lake Nature Reserve (鸳鸯湖自
然保育区), T. Cao & C. Gao (曹同, 高谦) 980671, 980332 (IFSBH). Xizang (西藏): Yadong (亚东), Mt.
Asangxia (阿桑下山), Zangmu (臧穆) 245, 241 (KUN, IFSBH). Yunnan (云南): Dulongjiang (独龙江),
Zangmu (臧穆) 1644, 1225, 2846, 2616, 2846, 1701 (KUN, IFSBH); Lijiang (丽江), Mt. Yulongxueshan (玉
龙雪山), T. Cao & G. Y. Song (曹同, 宋国元) 030695 (SHNU). Zhejiang (浙江): Mt. Jiulong (九龙山), Z.
No. 5 ZUO et al.: Comparison and assessment of three East-Asian species of the genus Scapania 745
L. Liu (刘仲苓) 1484, 249a, 573 (IFSBH).
Japan. Hakodate: Yezo, Faurie March 1899 (BM). Nagasaki, Mt. Tarudake, Faurie 10 June 1899
(BM); Kittasan, Faurie 14098 (M); Sanze, Tsuruoka-shi, Yamagata, S. Ono Nov. 5, 1978 (NY); Kuman Oto,
Mt. Ichifusa, K. Mayebara (NY); S. Hattori (NY); Fuka-yabakei, Ooita Prefecture, A. Noguchi, July 13, 1950
as S. Parvitexta var. minor (NY). Higo: Kiushiu, Hitoyoshi, K. Mayebara 1381 (NY); Kiushiu, Y. Kuwahara
1280, 132, 4225 (NY); Kyushu, Y. Kuwahara 5166 (NY).
2. Scapania parvidens Steph., Hedwigia 44: 15. 1904. Type: Japan. Without precise locality,
Faurie 1296 (holotype, G!). Fig. 2
Plants small in size, 0.5–1.0 cm long, 1–1.5 mm wide with leaves, green to yellow-green.
Stem ascending at upper part, unbranched or scarcely branched, in transverse section cortical
cells in 2–3 layers, small, thick-walled, interior cells large, thin-walled. Leaves densely
imbricate, nearly equally divided 2/3 their lengths. Keel long, straight, about 0.6 the length of
the ventral lobe. Ventral lobe transversely inserted, somewhat concave backwards, hardly
decurrent at base, obovate, 110–180 μm wide×140–240 μm long, width equal to or 3/4 the
length, apex rounded to obtuse, margins denticulate towards tip, sometimes nearly entire,
teeth mostly 1-celled, rarely 2-celled, usually acute. Dorsal lobe subequal to slightly smaller
than ventral lobe, 3/4 the ventral lobe in size, nearly transversely inserted, arching beyond the
farther edge of the stem, not decurrent at base, oblong-oval, apex rounded to obtuse, margins
of upper part with small, 1-celled teeth, entire below. Wings absent. Leaf cells small, rounded
to subrounded, cells near margins 7–10 μm, median cells 12–15 μm, basal cells 15 μm
wide×30 μm long, walls slightly thickened, trigones not distinct. Cuticle smooth to finely
roughened. Dioicous. Perianth terminal, oblong, dorsiventrally compressed, truncate above,
upper part often bent towards the ventral side, mouth with small teeth. Gemmae rare, green,
oval to ellipsoid, 1-celled.
Habitat: This species is common on granite at high altitudes (usually up to 1000 m), and
also occurs on the thin soil over rocks.
Distribution: China, Japan.
S. parvidens was described as a species new to science from Japan by Stephani (1904).
Later, the species was recorded in Japan by Müller (1905), Stephani (1910), Amakawa and
Hattori (1954), etc. It was reported from Anhui by Chen and Wu (1965), Jilin by Koponen et
al. (1983) and Yunnan Province by Gao and Cao (2000) in China before the present study.
Based on checked specimens, new records of S. parvidens in China include Liaoning,
Sichuan, Taiwan and Zhejiang Provinces, and Guangxi Autonomous Region.
S. parvidens is similar to S. parvitexta. The former differs from the latter by (1) smaller
plants, only 0.5–1.0 cm long; (2) dorsal lobes nearly equal to the ventral lobes in size, with
longer keels; and (3) teeth of leaf margins small, mostly consisting of 1-cell (rarely 2-celled).
S. parvidens and S. glaucoviridis both have large dorsal lobes nearly equal to the ventral
lobes, but S. parvidens has mostly 1-celled small marginal teeth and keels without wings,
while S. glaucoviridis has larger marginal teeth consisting of numerous cells and keels with
distinct wings.
Specimens examined:
China. Anhui (安徽): P. C. Chen (陈邦杰) 6947 (IFSBH); Mt. Huangshan (黄山), C. Gao (高谦),
1982-05-20 (IFSBH). Guangxi (广西): Xing’an (兴安), Mt. Mao’ershan (猫儿山), C. Gao & G. C. Zhang
(高谦, 张光初) 1405 (IFSBH). Jilin (吉林): Antu (安图), Mt. Changbai (长白山), T. Koponen 36688 (H,
IFSBH, PE), 36857, 36882 (H, IFSBH); T. Cao & Yan Baoying (曹同, 闫宝英) 396, 407 (IFSBH); T. Cao &
B. R. Zuo (曹同, 左本荣) 040515, 040576, 040568, 040571, 040572 (SHNU). Liaoning (辽宁): Kuandian
(宽甸), Baishilazi (白石砬子), T. Cao & S. Z. Wang (曹同, 王顺忠) 38321 (IFSBH). Sichuan (四川):
Jiangjin (江津), Mt. Simianshan (四面山), T. Cao & Q. Li (曹同, 李乾) 41695 (IFSBH); Mt. Emeishan (峨
眉山), T. Cao & G. Y. Song (曹同 , 宋国元) 031030 (SHNU). Taiwan (台湾): Xinzhu (新竹),
Yuan-yang-hu Lake Nature Reserve (鸳鸯湖自然保育区), T. Cao & C. Gao (曹同, 高谦) 980340 (IFSBH).
Acta Phytotaxonomica Sinica Vol. 45 746

Fig. 2. Scapania parvidens Steph. A–G, leaves; H–J, leaf margin teeth; K, leaf cells near margin; L, median leaf cells;
M, basal leaf cells; N, perianth; O, P, cross section of leaf keel; Q, cross section of leaf; R, cells of perianth mouth; S,
gemmae; T, cross section of stem; U, V, parts of plants, dorsal view; W, leaf insertion pattern, ventral view; X, leaf
insertion pattern, dorsal view. Figs. N, R were drawn from C. Gao & G. C. Zhang 1405, others were all drawn from Faurie
1296, holotype of Scapania parvidens, by B. R. Zuo and T. Cao.
Line scales: 0.3 mm for W, X; 150 µm for A–G, N; 75 µm for U, V; 50 µm for H–J, R, T; 25 µm for K–M, O–Q.
No. 5 ZUO et al.: Comparison and assessment of three East-Asian species of the genus Scapania 747
Yunnan (云南): Gongshan (贡山), Dulongjiang (独龙江), Zangmu (臧穆) 937 (KUN, IFSBH). Zhejiang
(浙江): Taishun (泰顺), Wuyanling Nature Reserve (乌岩岭自然保护区), T. Cao & S. L. Guo (曹同, 郭水
良) 010269 (SHNU).
Japan. Hokkaido: Kawakami, Y. Kuwahara 5520 (NY); Kawakawi, Y. Kuwahara 5521 (NY);
Daisetsuzan National Park, Y. Kuwahara 5502, 5511 (NY); Kyushu: Pref. Nagasaki, Y. K. 1985 (NY);
Chikugo, Y. Kuwahara 4340 (NY). Honshu: Hiroshima-ken, Mt. Gokurakuji, H. Ando E8810 (NY). Kiushiu:
Kirishima, Y. Kuwahara 117 (NY); Hiuga, Mt. Osuzu, Y. Kuwahara 194 (NY).
3. Scapania glaucovidis Horik., Sci. Hiroshima Univ. Ser. B, Div. 2, 2: 221, 1934. Type:
China. Taiwan. Tainan and Taichu, Mt. Morrison, Niitaka-Tozanguchi-Tataka-summit-
Hattsukwan, 1932-08-20, Y. Horikawa 9222 (holotype, HIRO!). Mt. Taihezhan (Mururoafu-
Kyôgetsu), prov. Taikoku (S. IWAMASA, Aug. 1932). (paratype, not seen). Fig. 3
Scapania okamurana Steph. ex Amak. & Hatt., J. Hatt. Bot. Lab. 12: 106. 1954. cf.
Inoue 1972.
Plants medium in size, about 2 cm long, 2.1 mm wide with leaves, pale-green, reddish
above, often in dense tufts on ground. Stems dark-brown, 0.38 mm in diam., strong, usually
unbranched. In cross section of stem, cortical cells in 3–4 layers, small, thick-walled, interior
cells large, thin-walled. Leaves densely imbricate, often falcate towards one direction. Keel
0.7–0.8 the length of the ventral lobe, nearly straight, often with distinct wing. Ventral lobe
nearly transversely inserted on the stem, concave, broad-ovate, 1.33 mm wide, about 1.55 mm
long, not decurrent at base, apex rounded to obtuse, margins densely toothed towards the tip,
teeth triangular, consisting of numerous cells, acute, nearly entire below. Dorsal lobe nearly
equal to or slightly smaller than ventral lobe, transversely inserted and often appressed on the
stem, arching beyond the farther edge of the stem, concave, broad-oval, 1.44 mm wide×1.3
mm long, width longer than length, margins of upper part of lobe toothed, teeth similar to
those of ventral lobe. Wings present, 1–6-celled. Leaf cells near margins 15 μm, median cells
18 μm wide×22 μm long, cells near base 22 μm wide×48 μm long, walls thickened, trigones
distinct, large. Cuticle smooth. Dioicous. Perianth terminal, ovate, dorsiventrally compressed,
upper part often bent towards the ventral side, mouth truncate, toothed, teeth consisting of
several thick-walled cells.
Habitat: On banks and moist rocks at high altitudes, alt. 2000–3000 m.
Distribution: China, Japan, and Himalayas.
S. glaucoviridis was described as a new species by Horikawa from Taiwan, China in
1934. Inoue (1972) treated S. okamurana Steph. ex Amak. & Hatt. (Amakawa & Hattori,
1954; Amakawa, 1964) as a synonym of S. glaucoviridis and recorded this species from
China (Taiwan), Japan, and Himalayas. According to present study, S. glaucoviridis also
occurs in Mt. Huangshan, Anhui Province and is newly recorded from mainland China.
S. glaucoviridis is similar to S. parvitexta in having leaf margin teeth irregular, large,
consisting of numerous cells, whereas S. glaucoviridis has large dorsal lobes, nearly equal to
or slightly smaller than ventral lobes in size, and long, straight keels with distinct wings. S.
parvitexta has the dorsal lobe distinctly smaller, about 0.7–0.8 of the ventral lobe in size, and
slightly concave keels without wings.
Specimens examined:
China. Anhui(安徽): Mt. Huangshan (黄山), P. C. Chen et al. (陈邦杰等) 6362, 7442 (PE). Taiwan
(台湾): Taizhong (台中), Mt. Xiaoxueshan (小雪山), T. Cao & C. Gao (曹同, 高谦) 980960, 980948,
980984 (IFSBH, SHNU); Hualian (花莲), between Mt. Tailukota Shan and Mt. Patolu Shan, C. C. Chuang
5675 (PE); Ilan (宜兰), C. C. Chuang 6043 (PE).
In conclusion, comparisons of three Eastern-Asian species indicate that S. parvitexta, S.
parvidens, and S. glaucoviridis differ in several features (Table 1) and can be distinguished
practically (key to the species). Therefore, the three should be treated as separate species.
Acta Phytotaxonomica Sinica Vol. 45 748


Fig. 3. Scapania glaucovidis Horik. A–D, leaves; E, perianth mouth pattern; F, G, cells of perianth mouth; H, leaf
margin teeth; I, J, cross section of leaf keel; K, median leaf cells; L, basal leaf cells; M, parts of plants with perianth and
sporophyte, dorsal view; N, cross section of stem. Drawn by B. R. Zuo from Horikawa 9222, holotype of S. glaucovidis.
Line scales: 150 µm for A–D; 100 µm for E; 75 µm for M; 50 µm for F–H, N; 25 µm for I–L.
No. 5 ZUO et al.: Comparison and assessment of three East-Asian species of the genus Scapania 749
Table 1 Comparison of Scapania glaucoviridis, S. parvidens and S. parvitexta in some criteria
Criteria S. glaucoviridis S. parvidens S. parvitexta
Wing of the leaf-keel wing developed no no
Dorsal and ventral lobes
in size
dorsal lobe nearly equal to
ventral lobe
dorsal lobe nearly equal to
ventral lobe
dorsal lobe small equal to
ventral lobe
Leaf-margin teeth large, with numerous cells small, 1-celled, rarely
2-celled
large, with numerous cells
Cuticle of leaves smooth smooth to finely roughened coarsely verrucose

Key to the three species
1. Dorsal lobe nearly equal to ventral lobe.
2. Leaf margin teeth small, mostly 1-celled (sometimes 2-celled); keel without wing………………………
………………………………………………………………………………………Scapania parvidens
2. Leaf margin teeth large, consisting of numerous cells, keel with distinct wing…………S. glaucoviridis
1. Dorsal lobe distinctly smaller than ventral lobe ……………………………………………… S. parvitexta

Acknowledgements Thanks are due to the curators and directors of the herbaria of BM, G,
H, HIRO, IFSBH, KUN, M, NY, PE, and SHM for their loan of the specimens, especially the
type specimens examined for the present study. Special thanks go to Dr. Jonathan Shaw of
Duke University for his help with English. Appreciations are also given to the first author’s
good friend Dr. XU Ye-Ping of Berlin Freedom University with his help on some old
literatures.
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东亚分布合叶苔属(苔纲: 合叶苔科)
三种植物的比较和评估
1, 2, 3左本荣 2, 1曹 同* 2郭水良
1(中国科学院沈阳应用生态研究所 沈阳 110016)
2(上海师范大学生命与环境科学学院 上海 200234)
3(中国科学院研究生院 北京 100049)

摘要 对最近被Potemkin作为同一种处理的合叶苔属Scapania中东亚分布的三种即细齿合叶苔S.
parvidens Steph.、弯瓣合叶苔S. parvitexta Steph.和灰绿合叶苔S. glaucoviridis Horik.的模式标本及相关
标本进行了比较研究。结果表明, 它们仍应该作为三个独立不同的种来处理。对三个种的模式标本进
行了绘图和描述, 并详细讨论了三种之间的区别。
关键词 苔类植物; 合叶苔科; 合叶苔属; 细齿合叶苔; 弯瓣合叶苔; 灰绿合叶苔; 东亚; 评估