全 文 ：　中国卷柏科植物的研究(Ⅰ):卷柏属同形叶亚属 　
张 宪 春
(中国科学院植物研究所　北京 100093)
Studies on the Chinese species of Selaginellaceae(Ⅰ):
Selaginella subgenus Tetragonostachys Jermy
ZHANG Xian_Chun
(Institute of Botany , the Chinese Academy of Sciences , Beijing 100093)
Abstract　The Chinese species of Selaginella subgenus Tetragonostachys are revised.Three spe-
cies are distinguished in the present study , i.e.S .sibirica , S.indica and S .vardei.Illustra-
tions of the leaves and strobili , distribution maps of the subgenus and the species occurring in China
are made to facilitate easy identification of the species.The distribution of the species in China
shows an interesting southwestern China and northeastern China distribution pattern.
Key words　Selaginellaceae;Selaginella subgenus Tetragonostachys;China;Distribution and
ecology
The Selaginellaceae are an ancient group of fern allies composed of about 700 ～ 750 species
and are presently regarded as monotypic and monophyletic (Korall et al., 1999;Jermy , 1990 ,
1986).
Many classifications of the Selaginellaceae have been proposed.Traditionally , leaf dimorphism
was regarded as a primary criterion in the subgeneric classification of Selaginella in many classifica-
tions(Jermy , 1986;Hieronymus , 1901;Spring , 1850).Jermy (1986)recognized five subgenera ,
three of which , Selaginella , Ericetorum and Tetragonostachys , are isophyllous and tow , Stachyg-
ynandrum and Heterostachys , are anisophyllous.
The typical isophyllous Selaginella rupestris and its allies form a distinctive group in Selaginel-
laceae.It is recognized as a section(sect.Tetragonostachys)by Hieronymus in 1901 and subse-
quently by Tryon in 1955 , and treated as a subgenus by Jermy (1990 , 1986).In 1992 , Sojàk
(1992)proposed Bryodesma , a new generic name for this group of plants with decussate uniform
sporophylls , and isophyllous , spirally arranged leaves.
Korall et al.(1999)made a cladistic analysis based on rbcL gene sequences of selective repr-
esentative species in the framework of Jermy s classification.The result of their molecular study
strongly supports monophyly of Selaginellaceae.Within Selaginellaceae , the morphologically distinc-
tive subgenus Selaginella is resolved as sister group to a clade composed of all other species.A
close relationship for all isophyllous species , as hypothesized in many classifications , is not support-
ed by the rbcL data.
The taxonomy of the isophyllous Selaginella species from China was confusing in the past.In
1934 , Alston recorded only one isophyllous Selaginella , S.vardei from China , without a mention
　Received 13 December 2000;accepted 7 April 2001
This work was supported by the National Natural Science Foundation of China(NSFC_39800011), and partly by the projects“Tax-
onomy and Floristic Special Grant” and the “Floristic and Biogeographic Studies of the Plants from Qinghai_Xizang(Tibet)Plateau
and Adjacent Regions” (No.KSCX 2_1_06B)from the Chinese Academy of Sciences.
植 物 分 类 学 报　39(4):345～ 355(2001)　[ 创刊 50 周年纪念特刊]
Acta Phytotaxonomica Sinica　[ Golden Jubilee issue]
of the well_known S .sibirica.Tryon recorded S .sibirica and S.vardei from China in his mono-
graphic study in 1955.In 1983 , Ching and Wu recognized S.vardei and a new variety (var.gra-
cilis)from Xizang.
After a long time of study on Selaginella with an access of the type materials kept in European
and American herbaria and collections from China and the Himalayan countries , I realized that there
are three distinct isophyllous Selaginella species in Chinese pteridophyte flora.S .indica , the Hi-
malayan isophyllous Selaginella , has not been recognised from China , and so far it was only record-
ed from India , Nepal and Bhutan.It was misidentified as S.vardei in China.The fact tells us that
the taxonomy of the Himalayan plants can not be made without references to the floras of China and
India at both sides of the Himalayas and access of the type materials usually kept in European her-
baria.
This is the first of a series of articles contributed to the study of the Chinese Selaginellaceae ,
presenting a taxonomic revision and geographical analysis of Selaginella subgenus Tetragonostachys
in China.
1　Taxonomy
Selaginella subgen.Tetragonostachys Jermy
Jermy in Fern Gaz.13(2):118.1986;in Kramer &Green , Fam.Gen.Vascul.Pl.1:43.
1990;Valdespino in Fl.N.Amer.2:41.1993.——— Tetragonostachyae A.Br.in Ind.Sem.
Hport.Bot.Berol.Append.11.1857 , without rank.——— Selaginella sect.Tetragonostachyae
(A.Br.)Hieron.in Engler &Prantl , Nat.Pflanzenfam.1(4):669.1901;R.M.Tryon in
Ann.Miss.Bot.Gard.42(1):17.1955.——— Selaginella subgen.Homoeophyllum (Spring)
Hieron.in Engler &Prantl , Nat.Pflanzfam.1(4):669.1901 , pro parte.——— Bryodesma Sojàk
in Preslia 64(2):155.1992.TYPE:Selaginella rupestris(L.)Spring.
Plants creeping , stems much branched , prostrate , and often mat_forming or with short erect
branches;vessel elements present.Rhizophores borne on the angles of branches from the upperside
of stems , throughout stem length or confined to base of stem.Leaves isophyllous , tightly appressed
and spirally arranged , or upperside and underside leaves slightly differentiated , all leaves linear or
linear_lanceolate , thick or fleshy (seldom thin);margins dentate , serrate , or ciliate;abaxial groove
with stomates along vein;axillary leaves absent.Strobili quadrangular.Sporophylls differentiated
from vegetative leaves;megasporophylls and micorsporophylls same size , in 4 alternating ranks , ap-
pressed.
About 50 species ranging from southern North America through the tropics of South America ,
Africa including Madagascar , and the Indian subcontinent to China , Siberia and Far East of Russia ,
northern Korea and northern Japan.In China , three species are distinguished as follows.
Key to the Chinese species of Selaginella subgenus Tetragonostachys Jermy
1.Leaf apex keeled , obtuse to truncate;margins densely ciliate 1.S.sibirica…………………………………
1.Leaf apex plane , attenuate;margins sparsely ciliate 2…………………………………………………………
2.Stems and branches strongly dorsiventral in position , about 2 mm wide including leaves;apex
bristle about 1 5 of the leaf length 2.S.indica……………………………………………………………
2.Stems and branches not dorsiventral in position , about 1 mm wide including leaves;apex bristle
about 1 3 of the leaf length 3.S.vardei…………………………………………………………………
1.1　西伯利亚卷柏　图 1:1 ～ 2
Selaginella sibirica(Milde)Hieron.Fig.1:1 ～ 2
Hieron.in Hedwigia 39:290.1900;in Engler &Prantl , Nat.Pflanzenfam.1(4):671.
346　 植　物　分　类　学　报 39 卷
Fig.1　1～ 2.Selaginella sibirica (T.N.Liou et al.8780);3～ 4.S.indica(W.M.Chu 2598);
5～ 6.S.vardei(X.C.Zhang et al.903).
1899;Kom.&Alis , Key Pl.Far East.Reg.USSR 1:106.t.23.f.5 ～ 8.1931;Iljin in
Kom., Fl.URSS 1:124.t.6.f.21 a～ c.1934;R.M.Tryon in Ann.Miss.Bot.Gard.42:
71.f.48.map 47.1955;W.Wang in N.T.Liou , Fl.Herb.N.E.China 1:9.f.10.1958;
Noda , Fl.N.E.Prov.China 1:6.pl.49.f.8.1971;C.J.Wu &X.L.Bai in Y.C.Ma ,
Fl.Intramongolica 1:57.pl.2.f.10 ～ 14.1985;C.W.Aur &G.F.Li , Pterid.Heilongjiang
36.f.10.1990;Valdespino in Fl.N.Amer.2:51.1993;P.Y.Fu , Clav.Pl.Chin.Bor._
Orient.sec.ed.22.pl.3.f.1.1995.——— Selaginella rupestris(L.)Spring f.sibirica Mil-
de , Filic.Eur.&Atl.262.1867.——— Bryodesma sibiricum (Milde)Sojàk in Preslia 64(2):
155.1992;R.J.Johns , Ind.Fil.Suppl.7:18.1997.TYPE:USA.Alaska , Unalaska ,
Chamisso (lectotype , proposed by Tryon , B , NY);Ajan , Tilling , Dahuria ad flumen Ingoda ,
Pallas(paratype , n.v.).
Selaginella rupestris f.amurensis Milde , Filic.Eur.&Atl.262.1867.TYPE:Russia.
Amur , Maximowicz (holotype , n.v.).
Selaginella rupestris f.manchuriensis Milde , Filic.Eur.&Atl.262.1867.TYPE:China.
Manchuria , Khalkyli , Maack (holotype , n.v.).
Selaginella schmidtii Hieron.in Hedwigia 39:292.1900.——— Selaginella rupestris f.
4 期 张宪春:中国卷柏科植物的研究(Ⅰ):卷柏属同形叶亚属 347　
schmidtii (Hieron.)Clute in Fern Bull.16:52.1908.TYPE:Russia.Sakhalin , Schmidt(lecto-
type , B;isolectotypes , A , K;fragment , NY).
Selaginella schmidtii var.krauseorum Hieron.in Hedwigia 39:293.1900.TYPE:Russia.
Tschuktschen peninsula , Emma_harbor , Aurel&Arthur Krause 53(lectotype , B;fragment , NY).
Plants on rock or terrestrial , forming cushion_like mats.Stems prostrate , or decumbent;main
stems obvious , with many short lateral branches;lateral branches ascending , about 2 mm wide in-
cluding leaves , short , 0.5 ～ 1 cm long , with 1 ～ 3 short branchlets.Rhizophores borne on upper-
side of stems , throughout stem length.Leaves thick , monomorphic , in alternate pseudowhorls of 5 ,
tightly appressed , deep green , linear_lanceloate , 1.9 ～ 2.2 mm×0.3 ～ 0.4mm(length excluding
the apex bristle);underside and lateral leaves ascending and turning to the upperside of stem , up-
perside leaves ascending;abaxial ridges very prominent;base cuneate and decurrent to rounded and
adnate , margins densely ciliate , cilia transparent , spreading to ascending;apex keeled , obtuse to
truncate;apex bristle white , about 1 3 to 1 5 of the leaf length , straight , puberulent.Strobili solit-
ary , 0.5 ～ 1(～ 2.5)cm×1.5 ～ 2mm;sporophylls deltate_ovate to ovate_lanceolate , margins cili-
ate , apex truncate in profile , bristled.Megasporangia in basal sporophylls , microsporangia in upper
sporophylls.
Distribution and Ecology:NE.China;also in N.Korea , N.Japan , E.Siberia , Bering
Straits and Far East of Russia , Alaska , Canada(Yukon district).On dry rocky slopes , or in bare
open grassy tundra.
China.Heilongjiang:Da HingganMts., T.N.Liou et al.8780(PE);Heihe , 185m , Y.
C.Chu et al.82(PE);ibid., J.Sato 6734(PE);Shangzhi , G.Z.Wang et al.250 (PE);
Inner Mongolia:Horqin Youyi Qianqi , Wuchagou , ca.1200m , Inner Mongolia Univ.Med.Pl.
Exp.893(PE).
1.2　印度卷柏　图 1:3 ～ 4
Selaginella indica(Milde)R.M.Tryon.Fig.1:3 ～ 4
R.M.Tryon in Ann.Miss.Bot.Gard.42:52.f.27.map 32.1955;R.D.Dixit , Selag-
inellaceae of India 32.f.1:A ～ G.pl.1.1992.———Bryodesma indicum (Milde)Sojàk in
Preslia 64(2):154.1992(publ.1993);R.J.Johns , Ind.Fil.Suppl.7:17.1997.——— Se-
laginella rupestris f.indica Milde , Filic.Eur &Atl.262.1867.TYPE:India.Khasia , Hook.f.
&Thomson s.n.(lectotype , BM , here designated;photograph , PE).
Selaginella wightii auct.non Hieron.:Panigrahi &R.D.Dixit in Proc.Nat.Inst.Sci.In-
dia 34B(4):206.f.11.1968.
Selaginella vardei auct.non H.Lév.:Ching &S.K.Wu in C.Y.Wu , Fl.Xizangica 1:
19.1983 , pro parte , excl.figs.;H.S.Kung , Fl.Sichuanica 6:58.1988 , pro parte , excl.
figs;Shing in W.T.Wang et al., Vascul.Pl.Hengduan Mts.1:8.1993 , pro parte.
Selaginella longipila auct.non Hieron.:Alston in Proc.Nat.Inst.Sci.India 11:214.
1945.
Plants on rock or terrestrial , forming discrete long_spreading mats.Stems and branches strongly
dorsiventral in position , prostrate , or decumbent;main stems obvious , with many lateral branches;
lateral branches 1 ～ 2_pinnate , spreading , about 2 mm wide including leaves , 1.5 ～ 2.5 cm long ,
with 3 ～ 7 short branchlets.Rhizophores borne on upperside of stems , throughout stem length.
Leaves monomorphic , herbaceous , in alternate pseudowhorls of 5 , loosely appressed , linear_lanceo-
late , 1.8 ～ 2.3 mm×0.3 ～ 0.5 mm (length excluding the apex bristle);underside leaves of main
348　 植　物　分　类　学　报 39 卷
Fig.2　Histogram of alt itudinal ranges of the individual taxa.Horizontally:percentage of records(%);vertically:
altitudinal zone(m).1.Selaginella indica;2.S.vardei.
stems brown in color , lateral leaves ascending and turning to the upperside of stem , upperside leav-
es ascending or spreading;abaxial ridges prominent;base cuneate and decurrent to rounded and ad-
nate , margins sparsely ciliate;apex plane , attenuate;apex bristle white , about 1 5 of the leaf
length.Strobili solitary , 0.5 ～ 2.5 cm×1.5 ～ 2 mm;sporophylls deltate_ovate to ovate_lanceolate ,
margins ciliate , apex truncate in profile , bristled.Megasporangia in basal sporophylls , microspor-
angia in upper sporophylls.
Distribution and Ecology:India , Nepal , Bhutan and China (Xizang , Yunnan , Sichuan).In
dry valley or on open slopes , usually on dry exposed rocks.1500 ～ 2800 m.
China.Xizang(Tibet):Cona , Nyam Jang Chu , 27°47′N , 91°50′E , 2743 m(9000 ft), F.
Ludlow , G.Sherriff &G.Taylor 6677(BM , PE);Gyirong , 2200 m , Qinghai_Xizang Exp.75 ～
6842(PE).Yunnan:Anning , 2200 m , J.J.He 8438 (PYU);Dali , T.N.Liou , 17577 ,
22992(PE);ibid., 2600 m , W.H.Hsu 300 (PE , PYU);ibid., C.Y.Wu 11242(PE);
ibid., 2000 ～ 2500 m , W.M.Chu 2598(PE , PYU);ibid., 2100 m , Kunming Inst.Bot.NW.
Yunnan Jinshajiang Exp.4377 (PE);ibid., 2100 m , Yunnan Univ.W.Yunnan Exp.4333
(PE , PYU);ibid., 2500 m , Q.Xia &Y.L.Ma 112(PE);ibid., 2000 ～ 2300 m , M.Kato
131(KUN);Eryuan , Mo_so_yn , Pee_cha_he , Delavay 3035(P , photograph , PE);ibid., 2150 ～
2200 m , W.M.Chu et al.29033(PYU);Shuangbai , 1500 m , W.M.Chu &Y.M.Feng
4605(PE , PYU);Yangbi , 1700 ～ 1850 m , W.M.Chu et al.9502 (PYU);ibid., H.C.
Wang 1942;Yongsheng , 1500 m , Y.K.Yang 108 (PE , PYU).Sichuan:Dechang , H.S.
Kung s.n.(PE), VI_1971.
India.Khasia , 1524 m(5000 ft), J.D.Hooker &T.Thomson s.n.(A , BM;photograph ,
PE);Kali Valley , 2133 ～ 2438 m(7000 ～ 8000 ft), J.F.Duthie 3727(BM;photograph , PE).
Bhutan.Griffith 2871(BM);near Trongsa Bridge , 2133m(7000 ft), exposed on rocks , F.
Ludlow , G.Sherriff &J.H.Hicks 17039(BM;photograph , PE);Kinga Rapden , Maugie Chu ,
4 期 张宪春:中国卷柏科植物的研究(Ⅰ):卷柏属同形叶亚属 349　
1524 m (5000 ft), F.Ludlow , G.Sherriff &J.H.Hicks 18606(BM;photograph , PE).
Nepal.Syabrubensi , Trisuli Khda , 1524 m (5000 ft), on rocks , O.Polunin 1478(BM ,
photograph , PE);Langtang , between Lama Hotel and Sharpugaon , 2600 ～ 2800 m , Toshiyuki Na-
kaike 1325(PE).
On the type specimen of S .indica preserved in BM , bearing the determination label “Proba-
bly an isotype of S .rupestris f.indica Milde , Fil.Eur.Atlant.262.1867.Leg.Hook.fil.&
Thomson , Khasia , 5000 ft” of Tryon(1953), the original label reads “Coll.Rawson W.Rawson ,
—No.2286 , Spec.951 , Selaginella rupestris Spr., E.India , Khasia.Ded.Hooker 1861” .Ex-
cept the type material , another two collections , Griffith 2871 and Duthie 3727 are also mounted on
this sheet.In the Khasia type material , a fragment of S.vaginata is found among the dense
branches.This is also the phenomenon in Yunnan that S.indica and S.vaginata Spring (syn.
S .compta Hand._Mazz.)are sometimes found growing together on rather dry rocks.
S .indica is rather common in Cangshan range in Yunnan , at medium elevations.Collections
of this species were all identified as S .vardei in Chinese herbaria , and were believed to be the typ-
ical form of S .vardei.Only after a check of the type materials borrowed from BM , I realized that
the Chinese plants identified as S.vardei by us are identical to those of S.indica from the Himal-
ayas , and what we called S .vardei var.gracilis is the real typical form of S .vardei.
The nomenclature of S .indica was also complicated by the misapplication of the name S .
longipila Hieron., because Hieronymus type materials were wrongly labeled as from the Himala-
yas.Tryon (1955)clarified the erroneous and found out that S .longipila Hieron.is a synonym of
S .densa Rydb., a species from North America.
1.3　细瘦卷柏　图 1:5 ～ 6
Selaginella vardeiH.Lév.Fig.1:5 ～ 6.
H.Lév., Cat.Pl.Yunnan 172.f.41.1915 ～ 1917;Alston in Bull.Fan Mem.Inst.Biol.
Bot.5(6):267.1934;R.M.Tryon in Ann.Miss.Bot.Gard.42(1):61.1959;Ching &S.
K.Wu in C.Y.Wu , Fl.Xizangica 1:19.f.4:6 ～ 10.1983 , pro parte , quoad figs.;H.S.
Kung , Fl.Sichuanica 6:58.pl.17.f.1 ～ 4.1988 , pro parte;Shing in W.T.Wang et al.,
Vascul.Pl.Hengduan Mts.1:8.1993 , pro parte.——— Bryodesma vardei (H.Lév.)Sojàk in
Preslia 64(2):155.1992(publ.1993);R.J.Johns , Ind.Fil.Suppl.7:18.1997.TYPE:
China.Yunnan , vicinity of Huize , 7 km from the town , Xiaowulong (Siao_ou_long), 2900 m ,
Maire 56(holotype , E;isotype , BM;photograph , PE).
Selaginella vardei var.gracilis Ching in C.Y.Wu , Fl.Xizangica 1:19.1983.TYPE:
China.Xizang , Mainling , F.Ludlow , G.Sherriff &G.Taylor 5494 (holotype , PE;isotype ,
BM).
Selaginella wightii auct.non Hieron.:Hand._Mazz., Symb.Sin.6:4.1929.
Selaginella longipila auct.non Hieron.:O.C.Schmidt in Act.Hort.Gothob.5:51.1929
(1930).
Plants on rocks or terrestrial , forming long_spreading mats.Stems and branches not dorsiven-
tral in position , creeping or decumbent , long slender , much branched;lateral branches about 1mm
wide including leaves , 1 ～ 5 cm long or more , with 1 ～ 4 short branchlets.Rhizophores borne on
upperside of stems , throughout stem length.Leaves monomorphic , herbaceous , in alternate pseud-
owhorls of 5 , tightly appressed , underside leaves same as the upper leaves , green , ascending , lin-
ear_lanceolate to narrowly lanceolate , 1.8 ～ 2.0 mm×0.3 ～ 0.4 mm (length excluding the apex
350　 植　物　分　类　学　报 39 卷
bristle);abaxial ridges prominent;base decurrent to adnate , often pubescent , margins sparsely cil-
iate;apex plane , attenuate;apex bristle white , about 1 3 of the leaf length , straight or curled , pu-
berulent.Strobili solitary , 0.5 ～ 1.5(～ 2)cm ×1 ～ 1.5 mm;sporophylls ovate_lanceolate , mar-
gins short_ciliate , bristled;Megasporangia in basal sporophylls , microsporangia in upper sporo-
phylls , sometimes monoecious , microsporangia occurring throughout the strobili.
Distribution and Ecology:Endemic to China(Xizang , Yunnan , Sichuan and Gansu).In crev-
ices under shrubs , or on mossy rocks;(950 ～ )2700 ～ 3800 m.
China.Xizang(Tibet):Kongpo , Tsangbo Valley , 3236 m(9300 ft), F.Ludlow , G.Sher-
riff &H.H.Elliot 14233(BM;photograph , PE);Nyingchi , 3650 m , C.Y.Wu et al.75_1171
(PE);Mainling , Dr.Tsela , Tsangpo Valley , 2895 m(9500 ft), F.Ludlow , G.Sherriff &G.
Taylor 5494 (BM , PE);ibid., Between Rongchaka and Lhagyari , Tsangpo Valley , 3505 m
(11500 ft), F.Ludlow , G.Sherriff &G.Taylor 4116(BM , PE);Markam , 3000 m , H.S.
Kung 6470(PE);Nyingchi , 3680m , C.Y.Wu et al.75_1171(KUN);Nüdong , Tsetang , 3700
m , 29°15′N , 91°45′E , Xizang Ecology Team M_7407a , M_74106 (PE);Qamdo , 3100 m , X.
Y.Zhu &Y.F.Du 20069 (PE);Zayǜ , 1800 ～ 2000 m , Qinghai_Xizang Exp.7722 , 10920
(PE);Zhag yab , Yéndum , 30°41′N , 97°32′E , Y.W.Tsui 5753(PE).Yunnan:Düqün ,
2700 ～ 3400 m , C.W.Wang 64723 , 69195 (PE);ibid., 2150 m , Qinghai_Xizang Exp.2878
(PE);ibid., 2050 ～ 2210m , W.M.Chu et al.17272(PE), 21047(PYU);Jingshajiang , 900
～ 1500 m , Handel_Mazzetti 824 (W;photograph , PE);Ludian , 1350 m , W.M.Chu 2830
(PE , PYU);Luquan , Wumeng Mt., 900 ～ 2700 m , W.M.Chu 2388(PE , PYU);Nangjiang ,
2100 ～ 2200 m , Handel_Mazzetti 8544(W;photograph , PE);Mekong Valley , G.Forrest 20349
(E , PE);Midu , 1800 m , W.M.Chu 6232(PE , PYU);Qiaojia , Chongxi , 2200 m , W.M.
Chu &H.Z .Yan 24417(PE , PYU);ibid., Loufang , 2500m , W.M.Chu &H.Z.Yan 24548
(PE , PYU);Yangtze ferry Lakatiang , Schneider 488(K);Yiliang , 1800 m , Q.S.Ran s.n.
(PYU);Yongsheng , H.J.Li s.n.(KUN).Sichuan:Barkam , 2750 m , The Eighth Forestry
Exp.Team 2432(PE);ibid., 2520 m , K.Y.Lang et al.2119 (PE);Batang , 3200 m , Si-
chuan Med.Pl.Exp.1555(PE);ibid., 2700m , Sichuan Vegetation Exp.3928(PE);Danba ,
K.L.Chǜ 7884 (PE);ibid., C.T.Kuan 5021(PE);ibid., 1700 m , The Eighth Forestry
Exp.Team 5021 (PE);Dujiangyan (Guanxian), Yinxiuwan (Yin_hsiu_wan), 1500 m , Harry
Smith 2394(PE);Hsǜ_Tsing , ca.2200 m , Harry Smith 4573(UPS , photograph PE);Huili , S.
K.Wu s.n.(PE);Jiuzhaigou(Nanping), 2000 m , X.C.Zhang et al.1336(PE);Kangding ,
ca.2000 m , Harry Smith 13330(UPS;photograph PE);H.C.Chow 583(PE);ibid., 1500 m ,
H.S.Kung 6098(PE , PYU);ibid., 3500 m , F.Y.Zou 4819;ibid., Paoma Mt., 2550 m ,
X.C.Zhang et al.901 , 903 (PE);ibid., Bamei (Qianning), 2800 m , Q.Q.Wang 5710;
Jinchuan , 2700 m , Sichuan Pl.Exp.9874 (PE), 13_IX_1975;Lixian , 2700 m , W.P.Fang
44881(SZ), D.P.He 1822(SZ);Luding , K.C.Kuan et al.1524(PE);Maoxian , 1800 m ,
X.C.Zhang et al.623 (PE);vicinity of Wenchuan , 1600 m , s.coll.8483 (PE);W.Si-
chuan , E.H.Wilson 5411 (K , P).Gansu:South part , Bailongjiang area , Zhugqu (?), s.
coll.s.n.(Tüwo County Forestry Herbarium);Wenxian , vicinity of Wenxian , by the river , alt.
950 m , Z.Y.Zhang 8123(WUK).
Different from Selaginella indica in that its stem and branches are slender , leaves smaller ,
shorter , the leaf apex bristle about 1 3 of the leaf length.
This species is uncommonly encountered in the field usually at higher elevations and north of
4 期 张宪春:中国卷柏科植物的研究(Ⅰ):卷柏属同形叶亚属 351　
352　 植　物　分　类　学　报 39 卷
4 期 张宪春:中国卷柏科植物的研究(Ⅰ):卷柏属同形叶亚属 353　
the range of S.indica (Fig.2;Fig.4).It was recognized only as a variety of S .vardei in herbar-
ia and concerned floras , while the plant with dorsiventral branches is regarded as the typical form of
S .vardei.The mistake continued until a check of the type of S .vardei , collected by Maire(no.
56)from Siao_ou_long , Yunnan , which was kindly sent on loan from BM recently.Maire s plant is
of slender form , the same as type of S .vardei var.gracilis and quite different from the dorsiventral
form , common in Cangshan range of Dali , NW.Yunnan.The Chinese name “Cangshan Juanbai”
for S.vardei contributed somewhat to the mystery.Since the specie is very common in Cangshan
range , circumscribed here to be S.indica , the Chinese name for S .vardei is better to be changed
to “Xishou Juanbai” (means slender Selaginella), once named for S.vardei var.gracilis.
This species is found growing with S .sanguinolenta and S.pulvinata in the dry_warm_valley
areas in Sichuan.They are all xeric plants.
In “Flora Xizangica” , the long apex bristle of the sterile leaf is not illustrated , because it is
easy to be broken off.This species represents a new record to the pteridoflora of Gansu Province.
Besides the Wenxian collection , I found another one in the local forestry herbarium of Tüwo (Tebbu
by Rock)County in 1998 during our floristic explorations in Bailongjiang region of South Gansu.I
was informed that the specimen was probably collected from Zhugqu.
2　Phytogeograph and Ecology
Subgenus Tetragonostachys (Fig.3)has a wide distribution in America , Asia and Africa (in-
cluding Madagascar).This subgenus is notably absent from the Malaysia_Australian region , the Pa-
cific islands and Europe (Tryon 1955).The modern distribution and differentiation centre of this
subgenus is in America , with over half of its species(Valdespino , 1993).
There are few representatives of this subgenus in Asia.S.vardei and S .indica are confined
to the Himalayas and southwestern China , S.wightii from southern India , Sri Lanka with a variety
from eastern Africa;S .sibirica from northeastern China , northern Korea and Japan through the Si-
beria and Far East via Bering Sea to North America;S.shakotanensis is distributed in Sakhalin ,
Japan and Korea.All of the above five species are endemic to Asia except S.wightii and S.sibir-
ica , the former has a disjunct distribution from South Asia to East Africa , the latter has a continued
extension via the Bering Sea to North America.The patterns of the distribution of those species indi-
cate that there once existed migration routes between Asia and Africa in the southwest and between
Asia and America in the northeast in geological time.
In China , the distribution of the three species of subgenus Tetragonostachys shows an interest-
ing southwestern and northeastern China distribution pattern(Fig.4).S.sibirica is only found in
Inner Mongolia and Heilongjiang in China.S.indica and S .vardei are nearly allopatric in distri-
bution.S .vardei , although a Chinese endemic species , is more widely distributed and common
than S .indica in China.Its distribution extends northwards from Yunnan and Xizang to Sichuan
andGansu , the north limit of its range being on the south side of Bailongjiang River in Southern
Gansu.
Species of subgenus Tetragonostachys are plants living in cold alpine , seasonally dry regions ,
characterized by thick cutinized leaves with fine hair points which protect the growing shoot apex.In
China , the species of subgenus Tetragonostachys are restricted to the open , xeric habitats , more
often the plants are found growing with mosses on rocks.The leaves on the old stems and branches
become dry dead leaves , and green leaves are only at the top part of branches and stems , which
354　 植　物　分　类　学　报 39 卷
greatly reduces due to loss of water during the dry seasons.The roots are hairy and form extensive
network and thus enables the plants to utilize rapidly even small amount of moisture.
Acknowledgements　I wish to express my sincere thanks to the directors and curators the following herbaria , BM ,
PYU , WUK for sending specimens on loan , A , BM , K , KUN , NY , P , PYU and SZ for their hospitality during my
visit , UPS and W for help with photographing some important specimens.My thanks go to Prof.Wei_Ming Chu for
sending duplicates of Marire s collection and pointing out its site , to Dr.T.Nakaike for sending duplicates from Ne-
pal , to Profs.Ming_Li Zhang and Yu_Fei Wang for taking photographs from UPS and W respectively , to Miss.Ping
Yang and Mr.Chao_Zheng Ji for drawing the distribution maps and illustrations , to Miss.Alison Paul and Prof.M.
Kato for checking names in Reed s Index Selaginellarum .
References
Alston , 1934 .An enumeration of the Chinese species of Selaginella .Bull Fan Mem Inst , 5:261～ 294.
Ching R_C (秦仁昌), Wu S_K(武素功), 1983.Selaginellaceae.In:Wu C_Y ed.Flora Xizangica.Vol 1.Bei-
jing:Science Press
Hieronymus G , 1901.Selaginellaceae.In:Engler A und Prantl K eds.Die Natǜurlichen Pflanzenfamilien.Band 1
(4).Leipzig:Verlag von Wilhelm Engelmann
Jermy A C , 1986.Subgeneric names in Selaginella.Fern Gaz , 13(2):117～ 118
Jermy A C , 1990.Selaginellaceae.In:Kramer K U , Green P S eds.The Families and Genera of the Vascular Plants
I:Pteridophytes and Gymnosperms.Berlin , Heidelberg:Springer_Verlag
Korall P , Kenric P , Therrien J P , 1999.Phylogeny of Selaginellaceae:evaluation of generic subgeneric relationships
based on rbcL gene sequence.Int J Plant Sci , 160(3):585～ 594
Sojàk J , 1992(publ.1993).Generische Problematik der Selaginellaceae.Preslia , 64(2):151～ 158
Spring A , 1850.Monographie de la famille des Lycopodiacé es.Mém Acad Sci Belg , 24:1～ 358
Tryon RM Jr , 1955.Selaginella rupestris and its allies.AnnMO Bot Gard , 42(1):1 ～ 99
Valdespino I A , 1993.Selaginellaceae.Flora of North America.Vol 2.New York:Oxford Univ Press
摘要　对中国卷柏科卷柏属同形叶亚属进行了分类学修订 , 首次承认中国分布有 3 种 ,即西伯利亚卷柏
S.sibirica ,印度卷柏 S.indica 和细瘦卷柏S .vardei。绘制了每种的叶形 , 孢子叶穗线条图;并绘有同形
叶亚属植物的地理分布图。该亚属在中国的分布呈西南和东北分布格局。
关键词　卷柏科;卷柏属同形叶亚属;中国;地理分布和生态
(责任编辑 郭延平)　
4 期 张宪春:中国卷柏科植物的研究(Ⅰ):卷柏属同形叶亚属 355