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1990, Vol.28, No.3 Previous Issue    Next Issue
Studies on the Karyotype of 5 Samples of Allium Sect. Bromatorrhiza Ekberg
Yan Yi-Xiang, Huang Rui-Fu, Wei Rong-Cheng, Xu Jie-Mei
1990, 28 (3): 177-184. 
Abstract ( 0 )
The karyotypes of 5 samples in Allium Sect.  Bromatorrhiza Ekberg were
analysed in this paper.
      In Allium wallichii Kunth, the first sample is a diploid, with genome formula is AA and
karyotype formula is K(2n)=2x=14=2m(SAT)+2m+10sm.  The second is an autotetraploid,
with genome formula AAAA, karyotype formul K(2n)=4x=28=2m(SAT)+6m+20sm. These
two karyotypes belong to “3A”.  The two karyotypes of A. wallichii Kunth are similar in
morphology, though different in ploidy.
      In Allium hookeri Thwaites, the first sample is a dibasic autoallotriploid.   Its genome
formula is AAB1; the basic number of the genome A is 7 and that of the genome B1 is 8. The
karyotype formula is K(2n)=2x+x'=22=(12sm+2t)+(1m+4sm+1st+2t).  The second is
also an autoallotriploid.  The genomes in pairs are similar to those in the first sample in size
and morphology of chromosomes.  However, the unpaired genome differs from the first one
apparently.  Therefore, its genome formula is AAB2, and karyotype formula is K(2n)=2x+
x'=22=(12sm+2t)+(3m+1sm+2st+2t). The third is doubling of the first karyotype.  It is
 an autoallohexaploid, with genome formula AAAAB1 B1 and karyotype formula K(2n)=4x+
2x'= 44= (24sm+4t) + (2m+8sm+2st+4t). These three karyotypes belong to “3A”.
Report on Karyotypes of 6 Species in 4 Genera of Polygonateae from China
Hong De-Yuan, Zhu Xiang-Yun
1990, 28 (3): 185-198. 
Abstract ( 0 )
Cytotaxonomically  investigated  in this work were 6  species in 4 genera  of
Polygonateae (sensu Krause, 1930).   Each species was karyotypically analysed using 5 so-
matic metaphase cells with well-spread chromosomes.  The chromosome classification follows
Levan et al. (1964) and the karyotype classification is according to Stebbins (1971). The ma-
terials used are listed in the Appendix and the vouchers are deposited in PE. The chromosome
numbers and karyotypes of Disporum megalanthum and Disporopsis aspera are reported here
for the first time, and those of Chinese Maianthemum bifolium are also reported for the first
time.  The results are shown as follows.
       (1) Disporum  Salisb.  D.  megalanthum  Wang et Tang from tthe Wolong
Nature Reserve, Sichuan, is found to have a karyotype   2n=16=2m(1SAT)+6sm(1SAT)+8st
(3SAT) (Plate I, A).  The parameters of chromosomes are listed in Table 1 and the idiogram
is shown in Fig. 1, A.  The chromosomes range in length from 8.5 to 29.3 μm, with the ratio
of the longest to the shortest 3.45.  The karyotype belongs to Stebbins' (1971) 3B.  In a somatic
chromosome complement the 2nd, 4th, 6th and 7th pairs each have one chromosome carrying
a satellite, showing heterozygosity.
      Another material from the Qinling Range, Shaanxi, is shown to have 2n=16=2m(1SAT)
+8sm(3SAT)+6st (Plate 1, B).  The parameters of chromosomes are listed in Table 1 and
the idiogram is presented in Fig. 1, B.  The chromosomes range in length from 6.3   to
22.6μm, with the ratio of the longest to the shortest 3.61, and thus the karyotype belongs to
3B.  The karyotype shows clear heterozygosity (Fig. 1, B).  The two chromosomes of the first
pair have arm ratios 2.38 and 1.82 respectively, but they are equal in length, 22.6 μm.  It seems
to us that a pericentric inversion has taken place in one of the two chromosomes.  Moreover,
the 3rd and 4th pairs each have one chromosome carrying a satellite attached to the long arm.
     These two materials are of the basically same karyotype, the major difference between
them being that the 3rd pair in the former consists of two st chromosomes with the arm ratio
3.15, while the corresponding pair in the other is of two m chromosomes with an arm ratio
1.67.
     Seven East-Asian species of the genus Disporum are reported to have 2n=14, 16 and 18
(or 16+2B?), but 2n=16 is common to all the species, and therefore the basic number of the
group is x=8.  For the North American group of the genus, however, 3 species (D. hookeri,
D. lanuginosum, D. oreganum) are of 2n=18, D. smithii is of 2n=16, and D.   maculatum
2n=12.  Chromosome numbers are more variable in the North American group, but x=9 seems
to be a dominant basic number.   Even more striking difference  in  karyotype  between  the
two groups exists in size of chromosomes, 2.0-4.9μm.for the North American group, while 4.0-
16.0 μm for the East-Asian counterpart (Therman, 1956) (Our result shows 6.3-22.6 μm and
8.5-29.3 μm for the two materials).  This remarkable contrast in karyotype is clearly correlat-
ed with the differentiation in gross morphology. The East-Asian species have calcarate tepals
but no reticulate veins of leaves, whereas the North American ones have reticulate veins  but
spurless tepals.  The evidence from karyotype and morphology seems to justify the restoration
of the genus Prosartes for the Nortth American species (Conover, 1983, cf. Dahlgren et al.
1985).
      (2) Disporopsis Hance D. pernyi (Hua) Diels from Mapien, Sichuan, is of
2n = 40 = 23m(2SAT)+13sm(2SAT) + 2st+ 2t(2SAT) (Plate 1, C). The parame-
ters of chromosomes are listed in Table 2, and the idiogram is shown in Fig. 2,  A.
The chromosomes range in length 5.2-16.2μm, with the ratio of the longest to  the
shortest 3.11, and thus the karyotype belongs to 2B.
      D. aspera     (Hua) Engl. ex Krause also from Mapien, Sichuan, is found to have
2n=40=30m+8sm(2SAT)+2t(2SAT) (Plate 1,D).  The parameters of chromosomes are listed
in Table 2, and the idiogram is shown in Fig. 2, B.  The chromosomes range in length 5.2-
14.7 μm, with the ratio of the longest to the shortest 2.84.  Therefore, the karyotype belongs to
2B.  Another material from the same locality but different population was also examined and
found to have 2n=40=30m+6sm+2st(2SAT) (Fig. 2, C).
     D. arisanensis (=D. pernyi) from Taiwan is reported to have 2n=40=26m+12sm+2st
(Chang and Hsu, 1974), D. fusco-picta from the Philippines 2n=40=22m+16sm+2st(2SAT)
(Kumar and Brandham, 1974), and D. longifolia from Thailand 2n=40 (Larsen, 1963). Thus,
the species in the genus, except the newly described D. jingfushanensis Z. Y. Liu (1987) with
no chromosome data, are all of 2n = 40, and the basic number of the genus is x = 20.  From
the karyotype formulae, asymmetry of the karyotypes increases from D. aspera to D. fusco-picta
through D. pernyi, which may be correlated with the increasing specialization of gross morpho-
logy.
      (3) Maianthemum Web. M. bifolium (L.) F. W. Schmidt from the Qinling
Range, Shaanxi, is found to have 2n = 36 = 20m + 10sm + 4st + 2t (2SAT) (Plate
1, H).  The parameters of the chromosomes are listed in Table 3, and the idiogram
is shown in Fig. 3, D.  The chromosome lengths range 2.4-8.2μm, with  the  ratio
of the longest to the shortest 3.43.  The karyotype thus belongs to 2B, and is sligh-
tly bimodal: the first 10 pairs and the pair of sat chromosomes are larger  than the
rest 7 pairs, the ratio of the shortest in the former group to the longest in the latter
group being 1.24.
      (4) Polygonatum Mill.  P. humile Fisch. ex Maxim. from Chicheng County,
Hebei, is shown to have a karyotype 2n= 20= 10m(2SAT)+6sm(2SAT)+ 4st
(Plate 1, G).  The parameters of chromosomes are listed in Table 4, and the  hap-
loid idiogram is shown in Fig. 3, C.  The chromosome lengths range from 3.0 to
10.0μm with the ratio of the longest to the shortest  3.3.  The  karyotype  therefore
belongs to 2B.
      P. odoratum  (Mill.) Druce Two materials in this species were examined. One from
Chicheng County, Hebei, has 2n=20=10m+10sm(3SAT)  (Plate 1, E).  The parameters of
chromosomes are presented in Table 4 and the somatic idiogram in Fig. 3, A. The chromoso-
mes range in length 3.1-8.8 μm, with the ratio of the longest to the shortest 2.8. The karyo-
type is thus of 2B.  The other from the Qinling Range, Shaanxi, is found to have 2n=20=
12m(4SAT)+8sm(2SAT) (Plate 1, F).  The parameters of chromosomes are listed in Table
4, and the haploid idiogram is shown in Fig. 3, B.  The chromosomes range in length 4.2-
10.9 μm, with the ratio of the longest to the shortest 2.6.  The karyotype is also of 2B.
     P. odoratum is widely distributed in Eurasian temperate region and its cytological reports
are frequently seen.  All the materials outside of China, from Portugal to Japan, are reported
to have 2n=20, except one material from east Sayan in SE Siberia, which is reported to have
2n=30 (Krogulevich, 1978).  In China, however, three chromosome numbers have so far been
reported under the name P. odoratum, 2n=20 from the Changbai Mountains, Jilin Province
(Fang, 1989), Qinlong County, Hebei Province (Wang et al. 1987), the Jinfo Mountains, Si-
chuan Province (in cultivation), besides the two materials used in this work; 2n=22 from Mt.
Jinshan in Beijing (Li, 1980), Wuhan in Hubei Province, Yixin in Jiangsu Province and Mt.
Emei in Sichuan Province (Fang, 1989); 2n=18 from Yixin in Jiangsu Province and the Dabien
Mountains in Anhui Province (Fang, 1989).  It is, therefore, rather evident that the species
under discussion is variable in chromosome number only in the southern part of its distribution
area.  Karyotypical morphology is also variable in this species.  The 2n=20 group is found to
have following karyotypes: 12m(4SAT)+8sm (in Austria, Hong et al. unpubl.), 14m+6sm
(Jilin): 12m+8sm (Qinlong, Hebei): 10m+10sm (3SAT) (Chicheng, Hebei): 12m(4SAT)+
8sm(2SAT) (Shaanxi) and 10m+6sm+4st(Mt. Jinfo, Sichuan). For the 2n=18 group, 10m+
8sm (Anhui) and 8m+10sm (Jiangsu) have been found.  In the 2n=22 group these karyotype
formulae so far reported are all 10m+8sm+4st. Comparing the karyotypes in the three groups
we find that 4st chromosomes are always present in the 2n=22 group, while in the other two
groups, except the karyotype 10m+6sm+4st found from the Jinfo Mountains in Sichuan, all
the karyotypes consist of m and sm chromosomes.
      Based on the correlation between karyotypical data and cryptic morphological differences
Wang et al. (1988) consider Polygonatum odoratum as a complex, which consists of  three
species: Polygonatum odoratum (s. str. 2n=20), P. macropodium Turcz. (2n=22) and P. simi-
zui Kitag. (2n=18).  But in this complex biosystematic problems, such as relationship between chromosome number and chromosome structure, evolutionary relationship of the different chromosome numbers, relationship between means of reproduction (extent of vegetative propagation) and karyotype variation are still unresolved and deserve further studies.
Studies on Karyotypes of 5 Species of Rohdea and Tupistra
Yang Di-Qing, Zhu Xie-Fu
1990, 28 (3): 199-206. 
Abstract ( 0 )
Five species of the genera Rohdea and Tupistra of Liliaceae were cytota-
xonomically investigated in this work.  The materials were collected from Jiangxi, Zhejiang,
and Sichuan provinces.  The results are shown as follows.
      1.  Rohdea Roth There are two forms of Rohdea japonica (Thunb.) Roth.  The karyo-
type formula of the cultivated form is 2n=38=28m+8sm+2st (Plate, 1), while that of the
wild one is 2n=38=26m+10sm+2st (Plate, 1). They both belong to Stebbins' (1971) karyotype
classification 2 B.
      2. Tupistra Ker-Gawl. The karyotype of Tupistra chinensis Baker from Mt. Lu-
shan, Jiangxi,is 2n=38=24m+14sm(Plate, 2), which differs from that of Campylandra
watanabei(=T. chinensis) from Taiwan, which is 2n = 38 = 32m + 4sm + 2st (Ch-
ang et Hsu, 1974).  The chromosome numbers and karyotypes of the three  species
from Nanchuan, Sichuan, are reported here for the first time.  The karyotype for-
mulas of T. delavayi Franch. and T. jinshanensis Z. L. Yang et X. G. Luo are both
2n= 38 = 26m + 12sm (Plate, 2-3), and that of T. wattii (C. B. Clarke) Hook.
f. is 2n= 38= 28m + 10sm (Plate, 3).  They all belong to 2B.
     3.  In Convallarieae (s. 1.) the chromosome numbers of all the genera, except for Thero-
pogon, are 2n=38 or both 2n=38 and 2n=36 in Aspidistra.  The karyotypes of Rohdea and
Tupistra are most similar, the differenc in karyotype between two genera is only presence or
absence of a pair of subterminal chromosomes. The karyotype of Rohdea is similar to that of
Convallaria rather than to those of Aspidistra and Speirantha, while the karyotype of Tupistra
is similar to that of Reineckia rather than to those of the other genera.
Report on Karyotypes of Smilacina tatsienensis and Ophiopogon japonicus
Wang Shu-Fen, Xu Jie-Mei, Yu Shu-Hua
1990, 28 (3): 207-210. 
Abstract ( 0 )
 In the present paper the karyotypes of Smilacina tatsienensis (Franch.) Wang
et Tang and Ophiopogon japonicus (L. f.) Ker.- Gawl. in Sichuan were analysed.  The karyo-
types of the two species are reported for the first time.  The results are shown as follows.
      Smilacina tatsienensis (Franch.) Wang et Tang is a dipoiid.  Its karyotype formula is
2n=2x=36=16m+10sm+10st(4SAT) (Plate 1: Fig. 1, 3).  The karyotype is bimodal with ten
large and eight small chromosome pairs and the length ratio of the tenth pair to the eleventh
being 1.33.  The length ratio of the largest chromosome and the smallest one is 4.33.
       Ophiopogon japonicus (L.f.) Ker.-Gawl. is a mixoploid, with diploid, triploid and tetra-
ploid cells in a single plant.  The karyotype formula  of the diploid is  2n=2x=36=18m
(4SAT)+18sm(Plate 1: Fig. 2, 4).  The species is of a bimodal karyotype with eight large and
ten small chromosome pairs and the length ratio to the eighth pair and the ninth being 1.10.
There are nine metacentric pairs (two pairs of sat-chromosomes) and nine submetacentric pairs.
A Chromosomal Study on 7 Species of Smilax L.
Fu Cheng-Xin, Hong De-Yuan
1990, 28 (3): 211-222. 
Abstract ( 0 )
The chromosome numbers and karyotypes of 7 species of Smilax L. in Lilia-
ceae (s. 1.) are cytotaxonomically studied in this work.  Their karyotypic characters, distinc-
tion between the species and the chromosomal basis of sexual differentiation are discussed. The
karyotypes of most species are first reported. The results are shown as follows (see Tables
1-4 for the chromosome parameters and the karyotype constitution; Fig. 1 for their idio-
grams):
       1. Smilax nipponica Miq.  The species is one of the herbaceous species distri-
buted in East Asia.  Two karyotypes, 2n = 26(type A) and 2n = 32 (type B),  are
found in the species (Plate 1: 1-7).  The karyotype of No. 88032 (uncertain of
                                        -L-      -M-         -S-
sexuality) is 2n = 26 = 2m + 6st + 6m + 4sm + 6sm + 2st.  The karyotype has 4
pairs of L chromosomes, of which the first three pairs are subterminal, and the 4th
is median.  The karyotype belongs to 3B.  No. 88045 (the male) and No. 88046
(the female) have 2n = 32.  Their karyotypes are basically uniform, and both are
                     -L-       -M-       -  S
2n=32= 2m+4sm+ 2st+ 2m+4sm+ 6m+ 10sm + 2st, also with 4 pairs of L
chromosomes, but the 2nd pair is median, and thus different from the type A.  The
karyotype belongs to 3B.  The first pair of chromosomes of the male  are  distinctly
unequal in length, with the D. V. (0.93) of relative length between them obviously
greater than that of the female (0.1).  The pair seems  to  be  of  sex-chromosomes.
Sixteen bivalents (n= 16) were observed at PMCs MI of No. 88045 (Plate 1: 4).
The major difference between the  karyotypes A and B are  greater relative  length
of L chromosomes in the type A than in the type B, and the increase of chromoso-
me number in the karyotype B mainly due to the increase of  st  chromosomes.  Na-
kajima (1937)reports 2n= 30 for S. hederacea var. nipponica  (=S. nipponica,
Wang and Tang, 1980).
      2. S. riparia A. DC.  This species is also herbaceous, distributed in East Asia.
Thirty chromosomes were found in root-tip cells (uncertain of sexuality).  The kar-
                          -L-     -M-      -S
yotype is 2n = 30 = 8st + 6sm + 2st + 6m + 6sm + 2st  (Plate 3: 1, 5), consisting
mainly of sm and st chromosomes.  There are 4 pairs of L chromosomes which are
all subterminal and the m chromosomes  appear  to fall all into S category.  Though
the karyotype belongs to 3B, it is less symmetrical than that of S.  nipponica.  The
species is karyologically rather different from S. nipponica, therefore. The first pair
of chromosomes of this material are unequal in length, and it may be a male.  The
karyotype of this species is first reported.
     3. S. sieboldii Miq. The species is a thorny climbing shrub, distributed in
East Asia.  At PMCs All, 16 chromosomes (n= 16) were found (Plate 2: 6), in
accordance with Nakajima's (1933) report for a Japanese material.
     4. S. china L.  This species, a thorny climbing shrub, is of a wide distribu-
tion range mainly in East Asia and Southeast Asia.  Two karyotypes  were  observed
in different populations.  (1) The population from Xikou has 2n = 96(6x) = 20st+
L-       -M-
    6t + 6sm + 12st + 52(S) (Plate 3:7), of which the first three pairs of chromo-
somes are terminal, different from those in the  other  species.  The arm  ratios  of
both L and M chromosomes  are larger than 2.0, which resembles those of  S. davi-
diana.  (2) PMCs MI of the population from Shangyu shew 15 chromosomes  (n
15).  The hexaploid of the species is recorded for the first time.  Hsu (1967,1971)
reported 2n = 30 from Taiwai and Nakajima (1937) recorded n = 30 from Japan,
which indicates that the karyotype  of  the  species varies  not  only  in  ploidy,  but
also in number.
      5. S. davidiana A. DC.  The somatic cells were found to have 32 chromoso-
mes, and PMCs MI shew 16 bivalents (Plate 2: 1-5).  The karyotype is 2n = 32=
-L-    -M-       -S
 8st + 4sm + 4st + 8sm + 8st.  The karyotype belongs to 3B, and is less symmetri-
cal than those in herbaceous species.  The D. V. (0.20) of relative length between
the two homologues of the first pair is slightly larger in the male than  in the  fe-
male (0.14), and it is thus difficult to determine whether they are sexual  chromos-
omes or not.
       6. S. glabra Roxb.  The species is a non-thorny climbing shrub,  distributed
in East Asia and Southeast Asia. 32 chromosomes were found in somatic cells. The
                                -L-   -M-     - S-
karyotype is 2n= 32=  8st  + 10st+6sm+8st (Plate 3: 2, 6),with only 3 pairs
of sm chromosomes  (12, 13 and 16th).   The  karyotype  is  more  asymmetric  than
that of S. davidiana, although it is also of 3B  (Table 1).  The karyotype is first
reported for the species.
      7. S. nervo-marginata Hay. var. liukiuensis (Hay.) Wang et Tang The variety
has a relatively narrow distribution range, mainly occurring in eastern  China.  The
chromosomal number of somatic cells is 2n= 32 (Plate 3: 3-4).  The karyotype is
                  -L-       -M-      -S
2n = 32 = 2sm + 6st + 2sm + 2st + 2m + 6sm + 12st, evidently different from that
of S. glabra.  The first pair of chromosomes are submedian, and much longer  than
the 2nd to 4th pairs. The ratio in length of the largest chromosome to the smallest
one  is  4.3.  The symmetric degree is of 3C, a unique type.  The karyotype of the
species is reported for the first time.
      In Smilax, the known basic numbers are 13, 15, 16 and 17.  The two herbaceous species
distributed in East Asia have three basic numbers: 13, 15 and 16, while the woody species studi-
ed mainly have 16, with no 13 recorded.  Mangaly (1968) studied 8 herbaceous species in North
America and reported 2n=26 for them except S. pseudo-china with 2n=30.  Mangaly consi-
dered that a probably ancestral home of Smilax, both the herbaceous and woody, is in Southeast
Asia and the eastern Himalayas, and speculated that the ancestral type of Sect.  Coprosman-
thus is possibly an Asian species, S. riparia.  The karyotypes of the two herbaceous species in
East Asia consist mostly of sm and m chromosomes, whereas those for the North American
species are all of st chromosomes.  Based on the general rule of karyotypic evolution, i.e. from
symmetry to asymmetry, his speculation seems reasonable.
      Researches on sex-chromosomes of Smilax have been carried out since 1930 (Lindsay, 1930;
Jensen, 1937; Nakajima, 1937; Mangaly, 1968), and they are generally considered to be the
largest pair, but there is still no adequate evidence. The result of our observation on S. nipponica
may confirm that the first pair of chromosomes of this species is XY type of sex-chromosomes.
      Chromosomes of the genus are small and medium-sized, varying between 1-6 μm, slightly
larger in herbaceous species than in woody ones, larger in the karyotype of 2n=26 than in
that of 2n=32.   Based on karyotype constitution of the above 5 species, the karyotype in the
genus is characterized by 4 pairs of L chromosomes and 2-5 pairs of M chromosomes, and
mostly st and sm chromosomes, and by rather asymmetrical 3B type.  The degree of symmetry
in the above 5 species is from Sect. Coprosmanthus to Sect. Coilanthus, and herbaceous species to
woody ones.
Investigation on Embryology of Anemarrhena asphodeloides
Chen Zu-Keng, Wang Fu-Hsiung, Li Zhi-Li
1990, 28 (3): 223-227. 
Abstract ( 0 )
 Anemarrhena  asphodeloides  Bunge is the only species  of Anemarrhena in Li-
liaceae, which possesses three stamens.  The flowers in this species have following features:
(1) Crystalliferous cells are present in the perianth and the filament. (2) Epidermal cells of
filaments and the inner perianth appears verruciform. (3) In longitudinal section, a number
of the multicellular hairs were found in the apex of the inner perianth. The above characteris-
tics of Anemarrhena are possibly important and differ from those of the  other  genera  in
Liliaceae.
       The main aim of the present paper is to deal with the female gametophyte and embryo-
genesis in Anemarrhena.  The development of embryo sac is similar to that of Ornithogalum
(Tilton et al., 1981), belonging to the Polygonum type, but there is a short embryo sac hausto-
rium at the antipodal end.  Before fertilization the two polar muclei fuse into a secondary nu-
cleus. The filiform apparatus was found in the synergid.
       The early development of proembryo in Anemarrhena is similar to that of Najas  (Hu,
1982).   After fertilization the zygote has a short stage of dormancy.   When the endosperm
has 12-16 free nuclei, the first division of the zygote takes place, forming an apical cell and
a basal cell.  Then the apical cell undergoes transversal divisions 2 or 3 times, forming a line
of three to four cells.  The basal cell usually does not further divide.
       The endosperm formation in Anemarrhena is the Helobial type.  The small chalazal cham-
ber is usually ephemeral and 2-4-nucleate, while the large micropylar one may be a multi-
nucleate before wall formation.
Pollen Morphology and Epidermal Characters of Leaves in Convallarieae (s. l.)
Ma Li-Ming, Hong De-Yuan
1990, 28 (3): 228-236. 
Abstract ( 0 )
 The tribe Convallarieae (sensu Krause 1930) consists of 7 genera, i.e. Con-
vallaria, Speirantha, Reineckia, Theropogon, Tupistra, Rohdea and Aspidistra, but now gen-
erally recognized as two tribes, Convallarieae (the former 4 genera) and Aspidistreae (the
rest).  Observed in this work were pollen morphology of 17 species and epidermal characters
of leaves of 12 species.  All the 7 genera are covered in observations.
     Pollen grains in Convallarieae (s. str.) are all monosulcate and boat-shaped (Plate 1: A-
F). The exine is rather uniformly microperforate (Plate 1: A-F); only Theropogon is ex-
ceptional in this respect: it has rugulate exine (Plate 1: O, P). Tang and Zhang (1985) have
pointed out the heterogeneity of Theropogon in this tribe.   Pollen morphology in the  tribe
Aspidistreae is widely variable.  The genera Tupistra and Rohdea were shown to have mono-
sulcate and boat-shaped pollen grains.   Their exine is perforate or reticulate (Plate 1: G-
N).  Pollen grains in the genus Aspidistra, however, are nonaperturate and spheroidal.  The
exine in the genus varies from crass-rugulate, variously gemmate to tuberculate-baculate (Plate
2; A-H).  The pollen morphology of Aspidistra is therefore distinctly different from that of
Tupistra and Rohdea, which supports the Nakai's (1936) establishment of the tribe Rohdeae
for Tupistra and Rohdea.  Therefore, Krause's Convallarieae is reasonably divided into at least
three tribes, Convallarieae (Speirantha, Convallaria, Reineckia and Theropogon), Aspidistreae
(Aspidistra) and Rohdeae (Rohdea and Tupistra).  The pollen characters of all the 7 genera
are shown in Table 1. The evolutionary trends of pollen morphology (aperture and exine) in
the three tribes are discussed and our major view-points are shown in Fig. 1.
     Observations on epidermal characters of leaves show that in the Convallarieae (s. 1.) sto-
matal apparatuses are all anomocytic; cuticular layer on the upper epidermis is mainly striate-
thickened or rather uniformly thickened (Plate 2: J--P; Plate 3: A-C, F-N), whereas in
the genus Convallaria the cuticular layer is squamosely thickened (Plate 2: I; Plate 3: D, E).
The epidermal characters of leaves in the 7 genera are summarized in Table 2.
A Conspectus and Phytogeography of the Genus Carex Subgen. Vignea (P. Beauv.) Kirsch. in China
Liang Song-Yun, Tang Yan-Cheng
1990, 28 (3): 237-260. 
Abstract ( 0 )
In spite of different views on the classification of the genus Carex, the su-
bgenus Vignea (P. Beauv.) Kirsch. in it is relatively natural subgroup adopted by most mo-
dern caricologists.  The total number of species in this subgenus is about 330, particularly abun-
dant in the subarctic and temperate regions of the Northern Hemisphere. (Fig. 1. Tab. 1.).
The conspectus of Chinese subgenus Vignea, consisting of 48 species, 7 subspecies and 1 variety,
is given in this article.  These 54 taxa (with species, subspecies and variety treated equally at
the same rank and with one widely distributed species and one uncertain species excluded for
floristic analysis here) can be grouped into 4 types of floristic elements according to the
floristic regions of the world suggested by Takhtajan (1986).
       1.  The elements of the Circumboreal Region (Fig. 2.): Carex diandra, C. vulpina, C.
 stipata, C. otrubae, C. curaica, C. disperma, C. bohemica, C. angustior, C. loliacea, C. tenui-
flora and C. lachenalii.  They constitute 20.4% of the total and are principally distributed in
the Eurasian Forest Subkingdom of China. Wu's scheme (1979) for the Chinese floristic division
is adopted here.
       2.  The elements of the Eastern Asiatic Region (Figs. 3, 4, 5): Carex echinochloaeformis,
 C. enervis subsp. chuanxibeiensis, C. rochebruni subsp. remotispicula, C. ovatispiculata, C. neu-
 rocarpa, C. nubigena subsp. pseudo-arenicola, C. nubigena subsp. albata, C. paxii, C. leiorhyn-
 chya, C. laevissima, C. pseudocuraica, C. pallida, C. yamatsutana, C. lithophyla, C. kobomugi,
 C. gibba, C. remotiuscula, C. rocheruni subsp. rochebruni, C. rochebruni subsp. reptans, C.
 alta, C. maackii, C. omiana, C. pallida var. angustifolia, C. earistata, C. thompsonii, C. larice-
 torum, C. maorshanica, C. dailingensis, C. unisexualis   C. heilongjingensis.   They constitute
 55.5% of the total taxa.  Wu (1979) considers that the Eastern Asiatic Region is better divid-
 ed into the Sino-Himalayan Forest Subkingdom and the Sino-Japan Forest Subkingdom. Among
the taxa mentioned above, only the first four species occur in the Sino-Himalayan Forest Sub-
kingdom and the remaining ones are of the Sino-Japanese Forest Subkingdom.  In fact, the
elements of the Sino-Japanese Forest Subkingdom constitute 48.1% of the total, obviously higher
than in the other regions.   Moreover, of these taxa the latter eight are endemic to   Sino-
Japanese Subkingdom and constitute 61.5% of the endemics of China.  It comes to a conclusion
that speciation of Chinese subgenus Vignea is more rapid here than elsewhere.
     3.  The elements of the Irano-Turanian Region (Fig. 6.): Carex duriuscula subsp, durius-
cula, C. duriuscula subsp, rigescens, C. duriuscula subsp, stenophylloides, C. reptabunda,   C.
 pycnostachya, C. enervis, C. pseudofoetida, C. sagaensis and C. physodes.   They constitute
16.7% of the total and are mainly distributed in Asiatic Desert Subkingdom, Eurasian Steppe
Subkingdom and Qinghai-Xizang (Ching-Tibet) Plateau Subkingdom of China.
     4.  The elements of the Indo-Chinese Region and the elements of Indian Region (Fig. 7.):
Carex thomsonii, C. fluviatilis, C. craspedotricha and C. nubigena.  They constitute 7.4% of the
total taxa and mostly occur in the Malaysian Subkingdom of China.  Of these taxa C. thom-
sonii with higher culm-nodes and C. nubigena with inflorescence of basal compound branch.
are regarded by us as primitive ones in the subgenus Vignea.  It is interesting to note at this
point that in the Indo-Malaysian Region not only is the Indocaricoid group, a primitive one
of Carex, more concentrated, but also the primitive ones of the subgenus Vignea, the   most
advanced group of Carex, are present.  The fact supports Nelmes' view (1951) that the genus
Carex had its origin in Indo-Malysian region.