全 文 ：植　　　物　　　研　　　究
BULLETIN OF BOTAN ICAL RESEARCH
第 18卷　第 3期 1998年　 7月
Vol. 18　 No. 3 July , 　 1998
西伯利亚嵩草及其相关类群的植物分类学及植物地理学研究
张树仁
(中国科学院植物研究所 ,北京　 100093)
摘　要　研究了产于西伯利亚的西伯利亚嵩草 Kobresia sibirica ( Turcz. ex
Ledeb. ) Boeck. , 产于北美的 K . hyperborea Po rsild和 K . macrocarpa Clokey。三
者之间无显著差异 ,并且它们的雌花都具有由一至三枚小鳞片组成的花被。因此 ,它
们为同种植物 , K . hyperborea包括其变种 va r. alaskana Duman和 va r. lepagei
Duman及 K . macrocarpa被降为西伯利亚嵩草的异名。西伯利亚嵩草分布于亚洲
和北美洲的北极和亚北极地区 ,可能起源于亚洲 ,由东西伯利亚通过白令海峡散布
到北美。
关键词　西伯利亚嵩草 ;分类学 ;植物地理学
TAXONOMY AND PHYTOGEOGRAPHY OF
KOBRESIA SIBIRICA ( TURCZ. EX LEDEB. )
BOECK. AND RELATED TAXA ( CYPERACEAE)
Zhang Shu- ren
( Institute o f Bo tany, Chinese Academy of Sciences, Beijing 100093)
Abstract　 Kobresia sibirica ( Turcz. ex Ledeb. ) Boeck. f rom Siberia , K . hyper-
borea Porsild and K . macrocarpa Clokey f rom No rth America w ere studied. There
a re no signi ficant dif ferences among the three species, and thei r female f low ers a re
all prov ided wi th a perianth of one to three small scales. Therefo re, they are con-
specific. Kobresia hyperborea , including varieties alaskana Duman and lepagei Du-
man, and K . macrocarpa are reduced to synonyms of K . sibirica. The species is
dist ributed in the a rctic and subarctic Asia and NW North America. It might have
evo lv ed in Asia, and mig rated f rom eastern Siberia through the Bering Strait to
No rth America.
Key words　Kobresia sibirica; Taxonomy; Phy tog eog raphy
Th e project supported by a g rant for sys tematic and evolutionary biology, CAS( No. 9701) .　收稿日期: 1998- 3- 10
1. Introduction
In the revision by Iv anova ( 1939) , the Kobresia schoenoides complex ( series Sibiricae)
contains 5 species: K . sibirica ( Turcz. ex Ledeb. ) Boeck. ; K . smirnovii Iv anova; K . ti-
betica Maxim. ; K . pamiroalaica Iv anova, and K . schoenoides ( C. A. Mey. ) Steud. All
these species exist in Asia , f rom the Caucasus and Urals to the Himalayas, SW China,
Mongo lia , and arctic Asia. Raymond ( 1965) considered that tw o Nor th American species,
K . hyperborea Po rsild and K . macrocarpa Clokey should be added to the complex. In addi-
tion, there a re still fiv e mo re Asian species should be refer red to the complex , they a re K .
deasyi C. B. Clarke ( Clarke 1908) ; K . septatonodosa Koyama ( Koyama 1956) ; K .
yushuensis Y. C. Yang and K . maquensis Y. C. Yang ( Yang 1984) ; and K . lacustris P.
C. Li ( Li 1991) . The K . schoenoides complex is ex t remely va riable and dif ficult to be di-
vided ( Dickoré 1995, Kukkonen 1996) .
The identi ty o f K . sibirica and the exact taxonomic po sition o f the tw o American
species ( K . hyperborea and K . macrocarpa ) has still to be investig ated ( Raymond 1965,
Dickoré 1995) . They are resembling each o ther morpho logically ( Raymond 1965) . The
geographical dist ribution of K . sibirica ex tends f rom the Ura ls th rough arctic Asia to the
Chukchi Peninsula and south to M ongolia. While K . hyperborea is distributed in NW
North America , wi th two va rieties, K . hyperborea var. alaskana Duman and va r. lepagei
Duman in Alaska ( Duman 1956) . K . macrocarpa occurs in the mountains of Colorado,
W yoming and M ontana. The mo rpholo gical similari ty and dist ribution pat tern of the three
species is interesting. In this paper, I w ill discuss thier tax onomy and phytog eog raphy.
2. Taxonomy
The Kobresia schoenoides complex is ha rd to be treated and needs a tho rough and de-
tai led study. Whi le K . sibirica is comparatively easy to be identified because of the pres-
ence of a“ perianth", i t is superficially simi lar to o ther members of the complex. Af ter ex-
amining the species bo rrow ed from BM, CAN, CM , K and NY, I found that Ivanova
( 1939) co rrect ly described it as " Flo res ♀ ad basin pistilii foliolis perianthii squami-
formibus, tenui ter membranaceis, anguste lanceo latis v el t riangulari- lanceola tis , cum
achenio deciduis" ( female flower wi th thin membranaceous, narrowly lanceo late or t rian-
gula r- lanceolate sma ll scales o f perianth at the base of pistil falling o ff wi th the achene) .
This characteristic is diagno stic of the species and also interesting in i tsel f. Ivanova
( 1939) believ ed that the small scales w ere the perianth and represented an ancestral sta-
tus, and that the arctic zone o f Angara Land, w hich is now occupied by K . sibirica, w as
an early dif ferentiation center of the genus Kobresia. It is doubtful w hether the scales a re
the perianth. Up to now their true na ture has not been understo od. In this paper, howev-
er, the scales are stil l called The “ perianth " in accordance w ith Ivanova ( 1939) . The
st ructure is signi ficant for classif ica tion no mat ter w hether o r no t i ts homology is co rrect ly
interpreted. The st ructure makes K . sibirica distinct f rom closely rela ted taxa. The reason
3733期　　　张树仁:西伯利亚嵩草及其相关类群的植物分类学及植物地理学研究 (英 )
that severl authors ( Kǜ kenthal 1909, Sergiev skaya 1935) had sunk K . sibirica under K .
schoenoides as a synonym might be the resul t of neg lect of the presence o f the“ perianth".
Kobresia hyperborea was first published il legi tima tely by A. E. Po rsi ld ( 1943) as“K .
arct ica ", w hich is a later homonym of K . arctica Ivanova (= Carex nardina Fries) . Por-
si ld ( 1951) reali zed the problem and co rrected the error. Porsi ld ( 1943) considered tha t
K . hyperborea was mo re slender than K . schoenoides , and no ticed tha t one sheet of speci-
mens co llected f rom w estern Siberia seems clo ser to K . hyperborea. He w as corret , but ig-
nored one important character of his new species- the presence o f the perianth. I have no t
seen the type of K . hyperborea , but I have examined several pa ra types. The spikelet struc-
ture is just the same as tha t of K . sibirica, and the female flowers have the same perianth.
Other characters o f the tw o species are identical. They are conspecific and can be easi ly
separated from K . schoenoides by i ts slender appea rance and perianth. Kobresia hyperborea
should be reduced to a synonym o f K . sibirica. The two va rieties o f K . hyperborea in
Alaska w ere described as having a“ la teral spikelet containing 1- 3 steri le scales" ( Duman
1956) , w hich are actually the "perianth ", and o ther dif ferences prov ided by the author a re
v ery subt le. Scoggan ( 1978) once had reduced K . hyperborea and i ts tw o varieties as syn-
onyms of K . sibirica, but no t mentioned the presence of perianth. The ev idence tha t all
these taxa po ssess perianth suppo rt such tax onomic treatment.
Kobresia macrocarpa Clokey w as inco rrect ly described when i t w as fi rst published by
sta ting that it s spike w as “ clo sely 8- 15- f low ered, the terminal flow ers stamina te, the
la teral pistillate" ( Mackenzie 1931) . Af ter ex amining the specimens o f this species ( in-
cluding types) , I found that it s“ lateral f low ers” are actually spikelets, who se structure is
equal to tha t o f K . sibirica and K . hyperborea. The basal flower of spikelet is female and
upper 1- 4 are male, and also the female flowers have 1- 3 perianth scales at the base o f
the pistil. M oreover, o ther characters of these three species a re all simi lar and are no t dis-
tinguishable. They should be merged into one species:
Kobresia sibirica ( Turcz. ex Ledeb. ) Boeck. in Linnaea 39: 7. 1875; Iv anova in Bot.
Zhurnal 24: 480. 1939; Ego rova in Novi t. Sy st. Pl. V asc. 20: 71. 1983. - Elyna sibirica
Turcz. ex Ledeb. Fl. Ross. 4: 262. 1853; Steud. Synops. Cyperac. 245. 1855. - Kobre-
sia arctica A. E. Po rsild in Sarg entia 4: 15. 1943, non Ivanova ( 1939) . - K . hyperborea
A. E. Po rsi ld in Bull. Nat. M us. Canada , 121: 103. 1951. - K . hyperborea var.
alaskana Duman in Bull. To rrey Bot. Club, 83: 194. 1956. - K . hyperborea va r. lepagei
Duman. l. c. — K . macrocarpa Clokey ex Mackenzie in North America Fl. 18: 5. 1931,
syn. nov.
3. Phytogeography
Kobresia sibirica show s an arctic and subarctic Asia and NW North America distribu-
tion that ex tends f rom the Urals through arctic Asia to the Chukchi Peninsula, south to
Mongo lia , and across the Bering Strai t s to Alaska and NW Canada and along Rocky
Mountains south to central Colo rado ( fig. 1) . Iv anova ( 1939) believ ed that the presence o f
374 植　　物　　研　　究　　　　　　　　　　　　　　　 18卷
the“ perianth " in K . sibirica represent an ancest ral stage in the evolution of the genus Ko-
bresia. In her opinion, series Sibiricae, w hose representativ e is K . sibirica , o ccupies a cen-
tral posi tio n in subgenus Elyna. She believ ed that the series di fferentiated in the a rctic re-
gion of Angara land; o ther groups in the subgenus deriv ed f rom the series th rough more or
less considerable reduction. During the la te Tertia ry ei ther wide mig ration o f the oro-
phy tes o f no rthern Asia to Himalayas o r rev erse happened. The types of sections and the
majo ri ty o f series o f the genus Kobresia were fo rmed, and in the Pleistocene, the represen-
tativ es of the genus mig rated down to Europe and North America ( Iv anova 1939) . The
dist ribution center o f the genus Kobresia is loca ted in the Himala yas and SW China. Every
g roup o f the genus has representativ es in the areas, and the genus most probably devel-
oped there ( Zhang et al 1995) . It is dif ficul t to be veri fied that the tax a of subgenus Elyna
in central Asia , the Himalayas, and China came from the arctic Asia. The most closely re-
lated species to K . sibirica exist in Al tai , cent ral Asian highlands, the Himalayas, and SW
China. The K . schoenoides complex is composed of these species. In the complex , whether
K . sibirica i s the most primi tiv e species is sti ll uncertain, because the nature and phy log e-
netic significance of i ts “ perianth " has no t been adequately explained. There is no convinc-
ing evidence to suppo rt the conclusion tha t the complex originated in Arctic Asia as sug-
gested by Ivanova ( 1939) . Nevertheless, K . sibirica and i ts clo se Asian relativ es migh t
have o rigina ted in Asia. The species could mig ra te to North America acro ss the Bering
Strai t.
Fig . 1　 Geog raphical distribution of Kobresia sibirica ( Turcz. ex Ledeb. ) Boeck*

Acknowledgments
I am g rateful to Pro f. Tang Yan- cheng and Dr. Bruce Bar tholomew , who revised
the manuscript. I am also indebted to the keepers o f BM, CAN, CM , K and NY for the
lo ans of specimens.
3753期　　　张树仁:西伯利亚嵩草及其相关类群的植物分类学及植物地理学研究 (英 )
Some references of geog raphical di s tribu tion are f rom Ivanova ( 1939) , Porsild ( 1943 ) , Duman ( 1965 ) and
Kozhevnikov ( 1981) .
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