全 文 ：中国西南六种广义拂子茅属 (禾本科) 植物的染色体数目*
马海英1, 2 , 刘振稳1, 2 , 彭 华1**
( 1中国科学院昆明植物研究所, 云南 昆明 650204; 2 中国科学院研究生院, 北京 100039)
摘要: 广义拂子茅属 ( Calamagrostis) 是一个世界温带广布的大属, 有些作者又分为拂子茅属和野青茅属, 但
近期的研究表明处理为一个属较为合适。中国共有 37 种广义拂子茅属植物, 但至今没有任何染色体的研究。
本文报道了其中产于中国西南 6 种野青茅的染色体数目, 其中 Deyeuxia petelotii 4 个居群, D1 diffusa, D1moup-
inensis , D1 nivicola 和D1f lavens 各一个居群都是四倍体 ( 2n= 4x= 28) , D1 neglecta 为六倍体 ( 2n= 6x= 42)。根据
广义拂子茅属植物染色体倍性特征, 该属植物中至今未发现二倍体, 四倍体是该属中倍性最低和最普遍的, 广
义拂子茅属的演化很可能是在四倍体的水平上进行的。由于以上几个四倍体种均是狭域分布的类群, 所以可能
是由四倍体的祖先隔离分化形成的。
关键词: 野青茅属; 拂子茅属; 染色体数目; 四倍体; 六倍体
中图分类号: Q 943 文献标识码: A 文章编号: 0253- 2700( 2006) 01- 022- 07
Chromosome counts in the Genus Calamagrostis s1l ( Poaceae)
from Southwestern China
*
MA Ha-i Ying
1, 2
, LIU Zhen-Wen
1, 2
, PENG Hua
1* *
(1 Kunming Insti tute of Botany , Chinese Academy of Sciences, Kunming 650204, China;
2 Graduate School of the ChineseAcademy of Sciences, Beijing 100039, China)
Abstract: Calamagrostis s1l. is a large genus widespread throughout the world in temperate and cold regions. It is sometimes
divided into two genera: Calamagrostis s. str . and Deyeuxia, and both have been accepted in China, while recent studies
show it should be treated as one genus. There are 37 species of Calamagrostis s1 l. occuring in China, but no cytological study
was conducted before. In this paper the chromosome numbers of 9 populations representing 6 species of Deyeuxia ( recognized as
a part of Calamagrostis s1l. ) from southwestern China are reported for the first time. Most of them are tetraploids and no diploid
species is found. All 4 populations of D1 petelotii and each population of D1 diff usa, D1moupinensis , D1 nivicola, and D1f-
laven are tetraploids ( 2n= 4x= 28) , and the only population of D1 neglecta is hexaploid ( 2n= 6x= 42) . On the basis of these
counts and previous reports, tetraploid is again confirmed primitive in Calamagrostis s1l. and the speciation of the genus is
principally occurred at tetraploid level in this region. As these five tetraploid species are restricted to Himalayan-Hengduan
Mountains, they probably have been formed by isolation from tetraploid ancestors.
Key words: Deyeuxia; Calamagrostis; Chromosome number; Tetraploid; Hexaploid
Calamagrostis Adans. s1l. belongs to the tribe
Aveneae, subfamily Pooideae of Poaceae, and it con-
tains about 270 species ( Clayton and Renvoize, 1986) .
The genus is widespread throughout the world in temperate
and cold regions and on tropical mountains ( Clayton and
Renvoize, 1986) . Central Asia, eastern Australia, and
the Andes Mountains of South America are three distribu-
t ion centers of the group (Tateoka, 1974) .
Calamagrostis s1l. has been divided on the basis of
morphological features into sect ions (Koch, 1837; Torg-
云 南 植 物 研 究 2006, 28 ( 1) : 22~ 28
Acta Botanica Yunnanica

*
** 通讯作者: Author for correspondence
收稿日期: 2005- 04- 28, 2005- 08- 02接受发表
作者简介: 马海英 ( 1971- ) 女, 博士研究生, 从事植物系统学研究工作。
基金项目: 国家自然科学基金项目 (30070051) , 云南省自然科学基金 ( 2000C0069M)
es, 1898; Rozhevitz, 1934; Tsvelev, 1983) or sub-
genera ( Wasiljew, 1960) . Some authors, following the
generic delimitat ion of Beauvois ( 1812) , treat it as two
genera: Calamagrostis s. str. and Deyeuxia ( R’golo,
1978; Edgar, 1995; Renvoize, 1998) . In fact, Dey-
euxia makes up the majority of Calamagrostis s1l. with
over 250 out of total 270 species in Calamagrostis s1l.
(Clayton and Renvoize, 1986) , and so for Deyeuxia spe-
cies, Deyeuxia and Calamagrostis are two interchangeable
names. For Chinese species, Keng ( 1959) , Lu ( 1987)
and Chen ( 2001) all recognized two genera, only Yang
( 1983, 1988) recognized Calamagrostis s1l. . As all pre-
vious cytological studies outside China were carried out on
the basis recognizing Calamagrostis s1l. , and the split of
two genera is not supported by our molecular phylogenetic
study (Ma et al . , unpubl. data) , it must be addressed
that the species in the present study, though using the ge-
neric name Deyeuxia as conventionally did in China, are
regarded as species of Calamagrostis s1l. ( this will be
abbreviated as Calamagrostis in the following parts) .
52 species of Calamagrostis have previously been
studied cytologically in the world. Chromosome numbers
of Calamagrostis have been recorded from Europe ( e1g.
LÊve and LÊve, 1961, 1974; Fedorov, 1969; Arohonka,
1982; Micieta, 1986; Petrova and Stoyanova, 1998;
LÊvkvist and Hultg¼rd, 1999) , the far east of Russia
( Zhukova, 1967, 1969; Zhukova and Tikhonova, 1971,
1973; Zhukova and Petrovsky, 1971, 1975; Guzik, 1984;
Rudyka, 1988, 1990; Sorokin, 1991, 1993; Probatova
et al . , 1996) , Kashmir (Koul and Gohil, 1991) , Hima-l
ayas (Mehra and Sharma, 1975) , Japan (Tateoka, 1976,
1978, 1984) , North America ( Greene, 1984; Aiken et
al . , 1989) , and Costa Rica ( Pohl and Davidse, 1971) .
Overall, the genus has been studied in Northern Hemisphere
except China but not studied in Southern Hemisphere. Pre-
vious studies show that species in Calamagrostis have large
Pooid type chromosomes, which is characteristic for Pooid
grasses and unique in Poaceae ( Clayton and Renvoize,
1986) , and the chromosome base number is 7. Various lev-
els of ploidy have been found in the genus: tetraploid,
hexaploid, octoploid, decaploid, dodecaploid, and aneu-
ploid, and different polyploidy in one species is common in
the genus (Tateoka, 1976) . No diploid has been found in
any of the previous studies, and some specialists ( Nygren,
1962; Tateoka, 1976) proposed that the tetraploid is
primitive in the genus.
There are 37 species of Calamagrostis in China, in-
cluding 6 species of Calamagrostis s. str. ( Lu, 1987)
and 31 species of Deyeuxia ( Chen, 2001) , and especia-l
ly rich in southwestern China ( abbreviated as SW China
below) along Himalaya-HengduanMountains. No cytolog-i
cal work on the species from China has been conducted be-
fore. The present work is to invest igate chromosome num-
bers of some species in SW China so as to provide data for
further systematic work for the genus.
Material and Methods
Seeds or living plants were collected from field for further ex-
periment in various localities in SW China. Plants from one popula-
tion were studied for each species, but four populations were studied
for Deyeuxia petelotii because it used to have a controversial system-
atic position. About 3- 10 individuals were used for each popula-
tion. The localities and the herbarium voucher specimens are listed
in Table 1. Herbarium voucher specimens are deposited in KUN
( Herbarium of Kunming Institute of Botany, Chinese Academy of
Sciences, Kunming, China) .
Root tips were used for chromosome count. Root tips were
mostly obtained from germinating seeds, and only in two populations
of Deyeuxia petelotii, one in Kunming ( H. Y . Ma 001) and one
in Bijie ( H. Y. Ma 154) , the root tips were obtained from the
transplanted plants.
Table 1 Material of Calamagrostis s1l . studied and chromosome numbers determined, localities and voucher specimens
Species 2n Locality Voucher
Deyeuxia petelotii (Hitchcock) S. M. Phillips& W. L. Chen 28 Heilongtan, Kunming, Yunnan, 1900 m H. Y. Ma 001
28 Lidiping, Weixi County, Yunnan, 3000m H. Y. Ma 008
28 Mt . Cangshan, Dali , Yunnan, 3000m H. Y. Ma 144
28 Yang Hill, Bijie County, Guizhou, 1450m H. Y. Ma 154
D1 f lavens Keng 28 Mt . Haba, Shangrila County, Yunnan, 2700m H. Y. Ma 119
D1diffusa Keng 28 Xiaoshao, Kunming, Yunnan, 2000m H. Y. Ma 159
D1 nivicola Hook. f 28 Xiangcheng County, Sichuan, 4300m H. Y. Ma 63
D1moupinensis ( Franch. ) Pilger 28 Ganyanggou, Baoxing County, Sichuan, 2600m H. Y. Ma 256
D1 neglecta ( Ehrh. ) Kunth 42 Mt . Zhegu, Ma. erkang County, Sichuan, 4300 m H. Y. Ma 240
231 期 马海英等: 中国西南六种广义拂子茅属 (禾本科) 植物的染色体数目
The root tips were pretreated in a mixture of 01002 mmolPL 8-
hydroxyquinoline and saturated 1, 4-dichlorobenzene ( 1B1 in vo-l
ume) for 5- 6 hours at room temperature, then fixed with Carnoy. s
fluid at 4 e for at least one hour. Before staining, the root tips
were hydrolysed with 1 mmolPL HCl for 15min at 60 e . Finally,
samples were stained with carbolfuchsin and squashed. From each
population at least ten cells were counted. Photographs were taken
with Olympus BX 51.
Results
D1petelotii (Hitchcock) S1M1Phillips & W1L1Chen
The species is special in Calamagrostis because its
glumes are usually slightly shorter than the lemma, while
most species in Calamagrostis have glumes longer than or
equal to the lemma. For this unique feature, the species
has been described as new several times in different gen-
era: Aulacolepis petelotii Hitchcock, Agrostis continen-
talis Hande-l Mazzetti, and Anisachne gracilis Keng, and
several combinations have been made: Aniselytron graci-
lis ( Keng ) N. X. Zhao, A1petelotii ( Hitchcock )
Soj‚k, Calamagrostis petelotii ( Hitchcock ) Govaerts,
Deyeuxia continentalis ( Hande-l Mazzetti ) L. Liou, and
Neoaulacolepis petelotii ( Hitchcock) Rauschert ( Phillips
and Chen, 2003) . The latest revision identif ies it as
Deyeuxia petelotii ( Hitchcock) S. M. Phillips & W.
L. Chen ( Phillips and Chen, 2003) .
This species appears quite widespread from north-
eastern India through southern China to northern Vietnam,
at elevations of 1000 - 3400 m ( Phillips and Chen,
2003) . In China, it is distributed in Guizhou and Yun-
nan and grows at grassy places on sandy acid soils at alt-i
tudes of 1400- 2000m ( Chen, 2001) . Our result shows
that the species is a stable tetraploid in China since all
four populations from Yunnan and Guizhou have chromo-
some number 28 ( 2n= 4x= 28) ( Fig. 1: 1) .
Fig. 1 Mitotic metaphases in species of Deyeuxia
1. Deyeuxia petelotii ( 2n= 28) ; 2. D1 nivicola ( 2n= 28) ; 3. D1 f lavens ( 2n= 28) ; 4. D1diffusa ( 2n= 28) ;
5. D1moupinensis ( 2n= 28) ; 6. D1 neglecta ( 2n= 42) Scale bar= 5Lm
24 云 南 植 物 研 究 28卷
D1nivicola Hook. f
The species is distributed in Qinghai, Sichuan, X-i
zang, and Yunnan of China, as well as Nepal and Sik-
kim ( Chen, 2001) . It grows on grassy and stony moun-
tain slopes at altitudes of 3000- 5000 m. We collected
seeds from Xiangcheng County, Sichuan Province, and it
is a tetraploid ( 2n= 4x= 28) ( Fig. 1: 2) .
D1flavens Keng
The species is distributed relat ively wide in Qingha-i
Tibet Plateau, Gansu, Qinghai, Sichuan, Xizang and
Yunnan ( Chen, 2001) . It grows in alpine meadows,
grassy slopes, open woodland or shrubland, especially
along riverbanks, at alt itudes of 2800- 4500m. We co-l
lected seeds from Mt. Haba in Shangrila County, Yun-
nan, and it is determined a tetraploid ( 2n = 4x= 28)
(Fig. 1: 3) .
D1diff usa Keng
It is restricted to Guizhou, Sichuan, and Yunnan
Provinces of China and grows among shrubs and on waste-
land at altitudes of 1900- 3800 m ( Chen, 2001) . The
seeds were collected from Kunming, Yunnan and it is a
tetraploid ( 2n= 4x= 28) ( Fig. 1: 4) .
D1moupinensis ( Franch. ) Pilger
The species is restricted to a few counties in S-i
chuan, and grows in very moist grassy places in montane
forests at altitudes of 2000- 2600m ( Chen, 2001) . We
collected seeds from the locality of its holotype specimen,
Baoxing County in Sichuan Province. It is a tetraploid
( 2n= 4x= 28) (Fig. 1: 5) .
D1neglecta ( Ehrh. ) Kunth
This is a widespread, highly polymorphic species,
growing at grassy places in forests and damp ground near
ditches at altitudes of 1200- 4000m. In China, it is dis-
tributed in Gansu, Hebei, Heilongjiang, Liaoning, In-
ner Mongolia, Shanxi, Sichuan, and Xinjiang ( Chen,
2001) . It is also distributed in Japan, Kyrgystan, Rus-
sia, Mongolia, Tajikistan, Europe and North America.
We collected seeds from one population in Sichuan Prov-
ince and it is a hexaploid (2n= 6x= 42) ( Fig. 1: 6) .
Discussion
This is the first report of chromosome numbers of
Calamagrostis in China. In accord with the previous
studies on Calamagrostis , species in our study all have
large size chromosomes and have a stable basic number 7,
which are called / Festucoid0 ( Stebbins, 1956) or / Poo-
id0 ( Clayton and Renvoize, 1986) type chromosomes and
characteristic in Pooid grasses.
Including the present study, altogether 57 species of
Calamagrostis have been studied cytologically. Early re-
searchers ( Nygren, 1962; Tateoka, 1976) had found
that diploid plants with 2n= 14, as well as triploids and
pentaploids, are complete lacking while tetraploids are
dominant in Calamagrostis. Meanwhile, studies on re-
production of Calamagrostis showed that tetraploid plants
(2n = 28) are sexual while hexaploids ( 2n = 42) are
apomictic, indicat ing that tetraploid plants behave like a
diploid plant while hexaploid plants like a triploid
(Nygren, 1962) .
Stebbins ( 1956) pointed out that polyploid complex
may pass through a series of stages. In polypoid complex-
es in which no diploid species is found, the presen-t day
polypoids could not have formed from diploids, for if
that, diploids, at least some relics, should still be
found as narrow endemics. In this kind old polyploid
complex, nearly all of the diploid species have become
extinct, and new cycles of polyploidy have arisen, and
tetraploids has become actual diploids ( Stebbins, 1956) .
Based the reproductive mode of Calamagrostis , Nygren
( 1962) thus regarded tetraploid as primitive in the genus.
Based on intensive study of chromosome numbers of Japa-
nese species, in combination with data from Europe,
Tateoka ( 1976) assumed that a tendency towards the ba-
sic number of X= 14 have been developed in earlier stage
and presen-t day Japanese species have been formed from
tetraploid ancestral taxa, and furtherhe postulated that
this should also be applicable to Europe species.
After these studies, many new chromosome counts
for species in Northern Hemisphere have been reported in
1980s and 1990s, but still no diploid plant with 2n= 14
was found, all species have chromosome numbers above
28. The lacking of diploid in Calamagrosits is still ques-
tionable since that no species in tropics and Southern
Hemisphere has been studied, but the chance to f ind dip-
loid plant in those areas is very small. In genera related
to Calamagrostis, such as Agrostis, which has very sim-
251 期 马海英等: 中国西南六种广义拂子茅属 (禾本科) 植物的染色体数目
ilar distribution pattern as Calamagrostis does, diploid
plants with 2n= 14 are not scarce, but all from Northern
Hemisphere, none of species in Southern Hemisphere is
found diploid ( Frey, 1997) .
Greene ( 1984) studied reproduction of some eastern
North American species. Tetraploids and polyploids are
found again in that region. His work also confirmed that
tetraploids are sexual, while hexaploids ( 2n = 42) are
apomictic in seed formation. Moreover, tetraploids a-l
ways have distinct morphology easy to dist inguish from
other species whereas hexaploids are not distinct in mor-
phology and often show somewhat intermediate characters.
This again confirmed that tetraploids actually behave like
a diploid while hexaploids behave like a triploid, thus
agreeing to Tateoka. s ( 1976) postulat ion that the evolu-
tion of the genus occurred at tetraploid level.
In our investigation on Chinese species, in accord
with other parts of the Northern Hemisphere, no diploid
is found. Of the 6 species in our study, 5 are tetra-
ploids, and only one species D1 neglecta is a hexaploid.
Therefore all previous studies and our current study con-
firmed again that tetraploid is dominant in the genus, and
then it is reasonable to suggest that the speciation of Ca-
lamagrostis in China has occurred at the tetraploid level.
Then it is possible to consider the speciat ion mode of
these species from their chromosome numbers and distr-i
bution patterns. All 5 tetraploid species are restricted to
Qingha-i T ibet Plateau or Himalayan-Hengduan regions,
so they probably have been formed by isolation from wide-
spread tetraploid ancestors during the uplift of the Himala-
yas. As no ploidy level above tetraploid has been found,
it seems that no polyploidization has involved during the
process of speciation. This is in accord with the recent
summery on chromosomal studies on plants in Hengduan
mountains ( Nie et al . , 2005) , which shows that the
polyploidy is not as common as people have assumed and
the speciation in the region have principally occurred at
diploid level, only to note that the tetraploids in Ca-
lamagrostis are in fact diploids.
Among these 5 species, D1petelotii used to have
controversial systemat ic position. It has now been widely
recognized as a species of Calamagrostis ( Clayton and
Renvoize, 1986; Chen, 2001; Phillips and Chen,
2003) , and our recent molecular study also show that it
is a member of Calamagrostis ( Ma et al . , unpubl. da-
ta) . In this study, plants from four localities in Yunnan
and Guizhou are all tetraploids. So it also has probably
been formed from widespread tetraploid Calamagrostis
species. Its short glumes should have derived from equal
glumes of Calamagrostis, not from different origin.
The case of D1neglecta is much more complicated.
D1 neglecta is recognized by Soreng ( 2003) as a synonym
of C1 stricta (T imm) Koeler but as a different species by
Chen ( 2001) and Lu et al . ( unpublished manuscript for
Flora of China, English edition) . Some cytological stud-
ies have been done under the name D1 neglecta and
C1stricta respectively. For D1neglecta, tetraploids
were found in North Europe ( Engelskjon, 1979) , octo-
ploids ( 2n = 56) , decaploids ( 2n= 70) , and dodeca-
ploids ( 2n= 84) were found in Russia ( Sokolovskaya and
Probatova, 1977) . For C1stricta, tetraploids ( 2n =
28) were found in southwestern Finland ( Arohonka,
1982) , northern England ( Crackles, 1994) , and west
Greenland ( Dalgaard, 1988) , dodecaploids ( 2n = 12x
= 84) were found in Japan (Tateoka, 1976) , and aneu-
ploids ( 2n= c. 104, 114, 116, 119, 120, 123) were
found in Eastern North America (Greene, 1984) . If D1-
neglecta is a synonym of C1 stricta, the species is distrib-
uted in Eurasia and Americas. From the chromosomal
studies, we can assume that it has been formed in Europe,
where many tetraploids have been found, and has spread
through Asia to North America. In high mountains or cold
regions of Asia to North America, polypoidization and hy-
bridization may have occurred in the species to adapt the
environmental or climate changes, therefore hexaploids,
dodecaploids and aneuploids are found in these areas. If
D1neglecta is a different species from C1 stricta, D1-
neglecta is only distributed in Eurasia and the chromosomal
reports of it are not enough to make a conclusion. There-
fore careful morphological study for D1 neglecta and C1str-
icta is needed to elucidate their relationship. For this
widespread species, this study is only to add a report for
the species and no further conclusion can be made on the
hexaploid plants we studied. However, it is not iceable
that the populat ion we studies in China, though a
hexaploid, exhibits typical morphology of D1neglecta,
26 云 南 植 物 研 究 28卷
not integrated with characters of other species, so it is
certain that it is from the tetraploid and octoploid plants of
the species, not a hybrid with other species.
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