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Cytology of Dicranostigma and Glaucium (Papaveraceae) from China

中国秃疮花属和海罂粟属的细胞学研究



全 文 :中国秃疮花属和海罂粟属的细胞学研究∗
向春雷1ꎬ 董洪进2ꎬ 胡国雄1ꎬ 纳迪亚􀅰扎赫拉3ꎬ 船本常男4∗∗
(1 中国科学院东亚植物多样性与生物地理学重点实验室ꎬ 昆明  650201ꎻ 2 国家林业局昆明勘察
设计院ꎬ 昆明  650216ꎻ 3 伊斯兰堡真纳大学生物技术系ꎬ 巴基斯坦 伊斯兰堡  45320ꎻ
4 昭和药科大学药学基础教育与研究中心生物研究所ꎬ 日本 东京  194 ̄8543)
摘要: 报道了秃疮花属 3种ꎬ 即苣叶秃疮花、 秃疮花和宽果秃疮花共 11 个居群及其姐妹类群海罂粟属 1
种 (海罂粟) 一个居群的染色体数目ꎮ 研究结果表明ꎬ 秃疮花属全部三种的染色体数目均为 2n= 12ꎻ 海罂
粟的染色体数目为 2n= 10ꎬ 并且是该属中新发现的最小数目ꎮ
关键词: 秃疮花属ꎻ 海罂粟属ꎻ 罂粟科ꎻ 染色体数目
中图分类号: Q 942              文献标志码: A              文章编号: 2095-0845(2015)06-721-06
Cytology of Dicranostigma and Glaucium (Papaveraceae) from China
XIANG Chun ̄lei1ꎬ DONG Hong ̄jin2ꎬ HU Guo ̄xiong1ꎬ Nadia B. Zahra3ꎬ Tsuneo FUNAMOTO4∗∗
(1 Key Laboratory for Plant Diversity and Biogeography of East Asiaꎬ Kunming Institute of Botanyꎬ Chinese Academy of Sciencesꎬ
Kunming 650201ꎬ Chinaꎻ 2 China Forest Exploration and Design Institute of Kunmingꎬ Kunming 650216ꎻ 3 Department of
Biotechnologyꎬ Quaid ̄i ̄Azam Universityꎬ Islamabadꎬ 45320 Pakistanꎻ 4 Biological Instituteꎬ Fundamental Education
and Research Center of Pharmaceutical Sciencesꎬ Showa Pharmaceutical Universityꎬ 194 ̄8543ꎬ Tokyoꎬ Japan)
Abstract: The cytological studies were performed in 11 populations of Dicranostigma lactucoidesꎬ D􀆰 leptopodumꎬ
and D􀆰 erectum and one population for Galucium fimbrilligerum. All three species of Dicranostigma possessed chromo ̄
some number 2n= 2x= 12ꎬ of which chromosome numbers for D􀆰 erectum and D􀆰 lactucoides are reported here for the
first time. Glaucium fimbrilligerum presented the number 2n = 2x = 10ꎬ which is not only new to the genus but also
the known lowest number for Glaucium.
Key words: Dicranostigmaꎻ Glauciumꎻ Papaveraceaeꎻ Chromosome number
  There are approximately 40 genera and 800
species in family Papaveraceae (Judd et al.ꎬ 1999).
This family has gained special attention for its phar ̄
maceuticalꎬ ornamentalꎬ and alimentary importance
due to the presence of high content of alkaloids and
a number of species which are of horticultural inter ̄
est (Wu and Chuangꎬ 1999). There are 440 species
of Papaveraceae belonging to 19 genera presently re ̄
ported from China. This large occurrence has made
China as an important distribution center of this fam ̄
ily (Wu and Chuangꎬ 1999ꎻ Zhang et al.ꎬ 2008).
Chromosome numbers in Papaveraceae vary
greatly along with basic numbersꎬ and with cytologi ̄
cal variability in different members of the family.
Howeverꎬ basic chromosome numbers are usually the
multiples of x= 6ꎬ 7ꎬ 8ꎬ 9ꎬ 10 and 11 and rarely of
other numbers (Kadereitꎬ 1993). Chromosome num ̄
bers for some Chinese genera are reported by differ ̄
植 物 分 类 与 资 源 学 报  2015ꎬ 37 (6): 721~726
Plant Diversity and Resources                                    DOI: 10.7677 / ynzwyj201515104

∗∗
Funding: Youth Innovation Promotion Associationꎬ Chinese Academy of Sciences (2013253)ꎻ Key Laboratory for Plant Diversity and Bio ̄
geography of East Asiaꎬ Kunming Institute of Botanyꎬ Chinese Academy of Sciences (KLBB201309)
Author for correspondenceꎬ E ̄mail: funamoto@ac􀆰 shoyaku􀆰 ac􀆰 jp
Received date: 2015-06-23ꎬ Accepted date: 2015-08-31
作者简介: 向春雷 (1984-) 男ꎬ 博士ꎬ 副研究员ꎬ 主要从事植物分类与系统进化研究ꎮ E ̄mail: xiangchunlei@mail􀆰 kib􀆰 ac􀆰 cn
ent studies (Krahulcovaꎬ 1982ꎻ Li and Hsuꎬ 1986ꎻ
Huang et al.ꎬ 1989ꎻ Safonovaꎬ 1989ꎬ 1991ꎬ 1994ꎻ
Zhang and Fuꎬ 1993ꎻ Xu and Weiꎬ 1994ꎻ Meng et
al.ꎬ 2006). Howeverꎬ the relevant cytological infor ̄
mation for Dicranostigma J􀆰 D. Hooker & Thomson
and its sister group Glaucium Miller is very scarce.
Dicranostigma is a small genus in subfamily
Chelidonioideae. As currently circumscribed (Zhang
et al.ꎬ 2008)ꎬ the genus consists of three speciesꎬ of
which D􀆰 lactucoides Hook. f. & Thomson is found in
Himalayas and D􀆰 erectum K. ̄F. Günther ( =D􀆰 platy ̄
carpum C􀆰 Y. Wuꎻ Dong and Xiangꎬ 2015) and D􀆰 lep ̄
topodum (Maxim.) Fedde are present in the Hengd ̄
uan Mountains and Losses Plateau region. Safonova
(1991) and Wu and Wang (1997) reported chro ̄
mosome number 2n=12 for D􀆰 leptopodum and D􀆰 hen ̄
anense S􀆰 Y. Wang & L􀆰 H. Wuꎬ respectively. How ̄
everꎬ the latter was suggested to be a synonym of the
former species ( Zhang et al.ꎬ 2008). Thereforeꎬ
chromosome data is only available for D􀆰 leptodpodum
to dateꎬ and the variability for chromosome number
within the genus is still unknown.
Glaucium Miller has been shown as sister to Di ̄
cranostigma (Brücknerꎬ 1992ꎻ Hoot et al.ꎬ 1997).
The genus consists of 23 speciesꎬ which occur around
the Mediterranean. In Chinaꎬ only three species were
recorded from Xinjiang ( Chuangꎬ 1993ꎻ Wu and
Chuangꎬ 1999). The chromosome numbers for seven
species belonging to the genus Glaucium have already
been reported by different authors ( Aminꎬ 1979ꎻ
Moryꎬ 1979ꎻ Ceschmedzievꎬ 1983ꎻ Ghaffari and
Chariat ̄Panahiꎬ 1985ꎻ Ghaffari and Kelichꎬ 2006ꎻ
Ruíz de Clavijoꎬ 1990ꎻ Safonovaꎬ 1991ꎬ 1993ꎬ 1994ꎻ
Runemarkꎬ 2000). These are G􀆰 corniculatum (L.)
Rudolphꎬ G􀆰 elegans Fisch. & C􀆰 A. Mey.ꎬ G􀆰 flavum
Crantzꎬ G􀆰 fimbrilligerum Boiss.ꎬ G􀆰 leiocarpum Boiss.ꎬ
G􀆰 oxylobum Boiss. & Buhseꎬ and G􀆰 squamigerum Kar.
& Kir. Unexceptionallyꎬ the chromosome number for
all species is 2n = 12 ( Table 1). The chromosome
numbers seemed to be constant in Glauciumꎬ but
there is not enough cytological data reported in Di ̄
cranostigma. Thus further cytological studies as well
as morphological and molecular data are needed.
As part of our studies for taxonomicalꎬ system ̄
atics and population genetics of the genus Dicranos ̄
tigma and related generaꎬ we investigated the cyto ̄
logical study of 11 populations of three Dicranostig ̄
ma species and one population of Glaucium fimbril ̄
ligerum from China and the results are presented
here. The aims of the present study were to deter ̄
mine chromosome numbers in Dicranostigma and
Glaucium in Chinaꎬ and to test the variability of
chromosome numbers within different populations of
these genera. This will contribute our knowledge for
future systematic study of this group.
1  Materials and method
The seeds of Dicranostigma and Glaucium fim ̄
brilligerum for the chromosome study were collected
from field in 2014 (Table 2)ꎬ and voucher specimens
are kept at the Herbarium of Kunming Institute of
Botanyꎬ Chinese Academy of Sciences (KUN). Seeds
were sown on wet filter paper in Petri dishes for ger ̄
mination and incubated at 23-25 ℃ in Temperature
Table 1  List of taxaꎬ location and chromosome number reported previously
No. Taxon Chromosome number (2n) References
1 Dicranostigma leptopodum 12 Safonova (1991ꎬ 1994)
2 Glaucium corniculatum 12 Amin (1979)ꎻ Ruíz de Clavijo (1990)
3 G􀆰 elegans 12 Mory (1979)ꎻ Ghaffari and Chariat ̄Panahi (1985)
4 G􀆰 flavum 12 Baltisberger and Voelger (2006)ꎻ Runemark (2000)
5 G􀆰 fimbrilligerum 12 Safonova (1991)ꎻ Ghaffari and Kelich (2006)
6 G􀆰 leiocarpum 12 Ceschmedziev (1983)
7 G􀆰 oxylobum 12 Safonova (1991)
8 G􀆰 squamigerum 12 Safonova (1991)
227                                  植 物 分 类 与 资 源 学 报                            第 37卷
Table 2  List of taxaꎬ location and chromosome number examined in this study
No. Taxon Chromosome number (2n) Collection data
A Glaucium fimbrilligerum 10 Tongdeꎬ Qinghaiꎻ Xiang Chunlei et al. 1067 (KUN)
B Dicranostigma leptopodum 12 Rangtangꎬ Sichuanꎻ Xiang Chunlei et al. 1045 (KUN)
C D􀆰 leptopodum 12 Dangchangꎬ Gansuꎻ Xiang Chunlei et al. 1127 (KUN)
D D􀆰 leptopodum 12 Tianshuiꎬ Gansuꎻ Xiang Chunlei et al. 1120 (KUN)
E D􀆰 leptopodum 12 Maerkangꎬ Sichuanꎻ Xiang Chunlei et al. 1042 (KUN)
F D􀆰 leptopodum 12 Kangdingꎬ Sichuanꎻ Xiang Chunlei et al. 1012 (KUN)
G D􀆰 leptopodum 12 Maoxianꎬ Sichuanꎻ Xiang Chunlei et al. 1159 (KUN)
H D􀆰 leptopodum 12 Xiaojinꎬ Sichuanꎻ Xiang Chunlei et al. 1036 (KUN)
I D􀆰 leptopodum 12 Shangri ̄Laꎬ Yunnanꎻ Xiang Chunlei et al. 969 (KUN)
J D􀆰 leptopodum 12 Lijiangꎬ Yunnanꎻ Xiang Chunlei et al. 928 (KUN)
K D􀆰 erectum 12 Lijiangꎬ Yunnanꎻ Xiang Chunlei et al. 959 (KUN)
L D􀆰 luctucoides 12 Nielamuꎬ Xizangꎻ Dong Hongjin and Hu Guoxiong 992 (KUN)
Gradation Chamber (TG100ADꎬ Nippon Medical &
Chemical Instruments Co.ꎬ LTD). Chromosome ob ̄
servations were made in meristematic cells of freshly
growing root tip cells. Freshly growing root tips were
obtained from germinated plants. They were pretrea ̄
ted in 2 mmol􀅰L-1 8 ̄hydroxyquinoline for 4 hrs at
ca. 20 ℃ and then fixed 45% acetic acid for 10 min
at ca. 2 ℃ . Finally they were macerated in a mixture
of 45% acetic acid and 1 mol􀅰L-1 hydrochloric acid
(1∶1) for 17-18 s at ca. 60 ℃. The macerated seeds
were stained in 2% aceto ̄orcein for over 30 min at
room temperature in a moist chamber with 45% ace ̄
tic acid (see Funamoto and Ogawaꎬ 2014ꎻ Xiang et
al.ꎬ 2014).
Slides were prepared by the conventional squa ̄
sh method. The slides were faintly heated under an
alcohol flame for 1 - 2 s before observation. Photo ̄
graphs were taken using OLYMPUS Digital Camera
(DP72SETA2) System. Classification of somatic meta ̄
phase chromosomes of centromeric position for Glau ̄
cium followed Levan et al. (1964): Arm ratio was
calculated by long arm / short armꎬ M=median (1􀆰 0)ꎬ
m=median (1􀆰 0<m<1􀆰 7)ꎬ sm=submedian (1􀆰 7≤
sm<3􀆰 0)ꎬ st = subterminal (3􀆰 0≤st<7􀆰 0)ꎬ t = ter ̄
minal (≤7􀆰 0).
2  Results and discussion
The chromosome number of Glaucium fimbril ̄
ligerum from population of Tongdeꎬ Qinghaiꎬ Chinaꎬ
is 2n= 2x= 10 (Fig􀆰 1ꎬ A). This number is reported
here for the first time for this genus. The length of
Glaucium fimbrilligerum chromosome ranged from
2􀆰 7 to 4􀆰 0 μm longꎬ and haploid chromosome length
was 31􀆰 8 μm long. The chromosome complement at
mitotic metaphase consisted of 6 median regionꎬ 4
submedian region chromosomes. No satellite chromo ̄
somes were observed in this population.
Safonova (1993) first reported that chromosome
number for Glaucium fimbrilligerum is 2n = 2x = 12.
Recentlyꎬ the parallel findings have been found by
Ghaffari and Kelich (2006) in their cytological stud ̄
ies for Iranian angiosperm plants. Besidesꎬ all previ ̄
ous studies showed that chromosome numbers for
Glaucium species are 2n = 2x = 12 (Ceschmedzievꎬ
1983ꎻ Ghaffari and Chariat ̄Panahiꎬ 1985ꎻ Ruíz de
Clavijoꎬ 1990ꎻ Safonovaꎬ 1993ꎻ Runemarkꎬ 2000).
Incorporating these findings with our result hereꎬ it
is evident that two different base numbers occur in
Glaucium: x= 5 and 6ꎬ but all these previous results
revealed that the base chromosome number x = 6 is
more frequent in Glaucium. It′ s also worth noting
that the basic chromosome number x = 5 is rare in
Papaveraceae. To our knowledgeꎬ this number was
only reported from Chelidonium asiaticum Krahulco ̄
va (Krahulcovaꎬ 1982ꎻ Shatalovaꎬ 2000)ꎬ C􀆰 majus
L. (Starodubtsevꎬ 1989ꎻ Sha et al.ꎬ 1995ꎻ Yan et
al.ꎬ 2000)ꎬ Macleaya cordata (Willd.) R. Br. and
M􀆰 microcarpa (Maxim.) Fedde (Safonovaꎬ 1989).
3276期          XIANG Chun ̄lei et al.: Cytology of Dicranostigma and Glaucium (Papaveraceae) from China           
Fig􀆰 1  Selected photomicrographs of somatic chromosomes at metaphase of Dicranostigma and Glaucium from China
A. Glaucium fimbrilligerum (Tongdeꎬ Qinghai)ꎻ B-J. Dicranostigma leptopodum (Bꎬ Rangtangꎬ Sichuanꎻ C. Dangchangꎬ Gansuꎻ
D. Tianshuiꎬ Gansuꎻ E. Maerkangꎬ Sichuanꎻ F. Kangdingꎬ Sichuanꎻ G. Maoxianꎬ Sichuanꎻ H. Xiaojinꎬ Sichuanꎻ I. Shangri ̄Laꎬ
Yunnanꎻ J. Lijiangꎬ Yunnan)ꎻ K. D􀆰 erectum (Lijiangꎬ Yunnan)ꎻ L. D􀆰 luctucoides (Nielamuꎬ Xizang) (bar = 5 μm)
427                                  植 物 分 类 与 资 源 学 报                            第 37卷
  Most Glaucium species are distributed in Medi ̄
terranean area. In China Glaucium species are con ̄
sidered endemic to Xinjiangꎬ here our collection of
Glaucium fimbrilligerum from Qinghai constitutes the
new record of the genus for Qinghai. In Xinjiangꎬ
these species grow in arid habitatꎬ while in Tongdeꎬ
Qinghaiꎬ the plants grow on high altitude and the
habitat is cooler and wetter than in Xinjiang. The
populations thus react to changes in their environment
by adaptive modifications which may cause the chro ̄
mosome number change. Hereꎬ the lower base num ̄
ber x= 5 has possibly developed through descending
dysploidy as an adaption to cool ̄wet habitatꎬ al ̄
though the process behind this phenomenon remains
hypothetical in the absence of in ̄depth studies.
Chromosome number for all the three species of
Dicranostigma is 2n = 2x = 12 ( Fig􀆰 1ꎬ B -L) and
chromosome numbers for D􀆰 erectum and D􀆰 lactucoides
have never been reported earlier. The length of chro ̄
mosome in all populations varied from 2 to 3 μmꎬ
which is too small to conduct a karyotype analysis.
For Dicranostigmaꎬ the chromosome numbers
reported here corroborates with previous study (Saf ̄
onovaꎬ 1991). In Chinaꎬ our three Dicranostigma
species have a disjunct distribution and occupy dif ̄
ferent habitats. Dicranostigma leptopodum and D􀆰 lac ̄
tucoides grow in dry ̄warm valley but D􀆰 erectum sur ̄
vives in high latitude and grows in alpine meadows
and / or rock crevices. Considering polyploidization
may play an important role in Papaveraceae distribu ̄
ted in the high altitude areaꎬ D􀆰 erectum is supposed
to be a polyploid species. Howeverꎬ there are no
differences in chromosome numbers among the Di ̄
cranostigma populations and all of them are diploid
species. According to some investigationsꎬ polyploid ̄
ization may play an important role in Meconopsis Vig.
(Xie et al.ꎬ 2014) and Corydalis DC. in Papaver ̄
aceae and some other plants ( Hong and Zhangꎬ
1990ꎻ Yangꎬ 2002ꎻ Zhou et al.ꎬ 2012). Howeverꎬ
polyploidy is not common in the alpine flora of the
Hengduan Mountains Hotspotꎬ and the frequency of
polyploidy is recorded to be only 22% in the area
(Nie et al.ꎬ 2005). Hereꎬ our study shows that
polyploidization has not played an important role in
the evolutionary history of the genus Dicranostigma.
Evidences from molecular data showed that Di ̄
cranostigma is sister to Glaucium ( Gleissberg and
Kadereitꎬ 1999ꎻ Ruiz ̄Sanchez et al.ꎬ 2012). Addi ̄
tionallyꎬ morphological similarities between them
such as growth formꎬ sepalsꎬ leaf and fruit shape
and seed morphology also shows that these genera
are closely related. The similar chromosome numbers
were reported for both generaꎬ i􀆰 e. 2n = 12 in the
previous investigations (Table 1). Thereforeꎬ Vent
and Mory (1973) even though these two genera are
congeneric. Althoughꎬ our findings do not provide
further support for their close relationship because
they have different chromosome numbers. Henceꎬ in
future there is a need to conduct cytological studies
including increased populations of other Glaucium
species.
References:
Amin Aꎬ 1979. Chromosome numbers of ten flowering plants of Egypt
[J] . United Arab Republic Journalꎬ 22: 73—75
Baltisberger Mꎬ Voelger Mꎬ 2006. IAPT / IOPB chromosome data 1
[J] . Taxonꎬ 55: 444
Brückner Cꎬ 1992. A tool for better identification of Dicranostigma
species (Papaveraceae) [ J] . Feddes Repertoriumꎬ 103: 399—
409
Ceschmedziev IVꎬ 1983. In IOPB chromosome number reports LXXX
[J] . Taxonꎬ 32: 506
Chuang Hꎬ 1993. The taxonomic and evolution and distribution of Pa ̄
paveraceae [J] . Acta Botanica Yunnanicaꎬ 15: 137—148
Dong HJꎬ Xiang CLꎬ 2015. Dicranostigma platycarpumꎬ a new syno ̄
nym of Dicranostigma erectum (Papaveraceae) [J] . Phytotaxaꎬ
230: 198—200
Funamoto Tꎬ Ogawa Mꎬ 2014. Karyomorphology of Perillula reptans
Maxim.ꎬ Lamiaceaeꎬ endemic species in Japan [ J] . Chromo ̄
some Botanyꎬ 9: 69—71
Ghaffari SMꎬ Chariat ̄Panahi MSꎬ 1985. Chromosome counts of some
angiosperms from Iran [J]. Iranian Journal of Botanyꎬ 3: 67—73
Ghaffari SMꎬ Kelich Kꎬ 2006. New or rare chromosome counts of
some angiosperm species from Iran II [ J] . Iranian Journal of
Botanyꎬ 12: 81—86
Gleissberg Sꎬ Kadereit JWꎬ 1999. Evolution of leaf morphogenesis:
evidence from developmental and phylogenetic data in Papaverace ̄
ae [J] . International Journal of Plant Scienceꎬ 160: 787—794
5276期          XIANG Chun ̄lei et al.: Cytology of Dicranostigma and Glaucium (Papaveraceae) from China           
Hong DYꎬ Zhang DZꎬ 1990. Observations on chromosomes of some
plants from west Sichuan [J] . Cathayaꎬ 2: 191—197
Hoot SBꎬ Kadereit JWꎬ Blattner FR et al.ꎬ 1997. Data congruence
and phylogeny of the Papaveraceae s. l. based on four data sets:
atpB and rbcL sequencesꎬ trnK restriction sitesꎬ and morphologi ̄
cal characters [J] . Systematic Botanyꎬ 22: 575—590
Huang SFꎬ Zhao ZFꎬ Chen ZY et al.ꎬ 1989. Chromosome counts on
one hundred species and infraspecific taxa [ J] . Acta Botanica
Austro Sinicaꎬ 5: 161—176
Judd WSꎬ Campbell CSꎬ Kellogg EA et al.ꎬ 1999. Plant Systematics:
A Phylogenetic Approach [M]. Sinauerꎬ Sunderland.
Kadereit JWꎬ 1993. Papaveraceae [A] / / Kubitzki K ed.ꎬ The Fam ̄
ilies and Genera of Vascular Plants [M]. Berlin: Springer Ver ̄
lagꎬ 2: 494—506
Krahulcova Aꎬ 1982. Cytotaxonomic study of Chelidonium majus L.
s􀆰 l. [J] . Folia Geobotanica Phytotaxonomicaꎬ 17: 237—268
Levan Aꎬ Fredga Kꎬ Sandberg AAꎬ 1964. Nomenclature for centro ̄
meric position on chromosomes [J] . Hereditasꎬ 52: 201—220
Li LCꎬ Hsu PSꎬ 1986. Chromosome observations of eight species en ̄
demic to China [ J] . Acta Phytotaxonomica Sinicaꎬ 24: 157—
160
Meng Yꎬ Xie HYꎬ Nie ZL et al.ꎬ 2006. A karyomorphological study
on four species of Meconopsis Vig. ( Papaveraceae) from the
Hengduan Mountainsꎬ SW China [J] . Caryologiaꎬ 59: 1—6
Mory Bꎬ 1979. Beiträge zur Kenntnis der Sippenstruktur der Gattung
Glaucium Miller ( Papaveraceae ) [ J ] . Feddes Repertꎬ 89:
499—594
Nie ZLꎬ Wen Jꎬ Gu ZJ et al.ꎬ 2005. Polyploidy in the flora of the
Hengduan Mountains hotspotꎬ southwestern China [J] . Annals of
the Missouri Botanical Gardenꎬ 92: 275—306
Ruíz de Clavijo Eꎬ 1990. Números cromosomáticos de plantas occi ̄
dentalesꎬ 608-618 [J] . Anales del Jardín Botánico de Madridꎬ
47: 425—430
Ruiz  ̄Sanchez Eꎬ Rodriguez ̄Gomez Fꎬ Sosa Vꎬ 2012. Refugia and
geographic barriers of populations of the desert poppyꎬ Hunne ̄
mannia fumariifolia (Papaveraceae) [J] . Organisms Diversity &
Evolutionꎬ 12: 133—143
Runemark Hꎬ 2000. Mediterranean chromosome number reports 10
(1110-1188) [J] . Flora Mediterraneaꎬ 10: 386—402
Safonova INꎬ 1989. Sravnitelno ̄karyologicheskoe isuchenie podseme ̄
jstva Chelidonioideae semejstva Papaveraceae Juss [ J] . Plant
Karyologyꎬ 61—63
Safonova INꎬ 1991. Chromosome numbers in some species of the fami ̄
ly Papaveraceae [J]. Botanicˇeskii Žhurnal (Moscow & Leningrad)ꎬ
76: 904—905
Safonova INꎬ 1993. In: Takhtajan A ed.ꎬ Numeri Chromosomatum
Magnoliophytorum Florae URSSꎬ Moraceae-Zygophyllaceae [M].
Naukaꎬ Petropoli
Safonova INꎬ 1994. The karyotypical analysis of the genera Dicranos ̄
tigmaꎬ Hylomeconꎬ Macleayaꎬ Sanguinariaꎬ Stylophorum (Cheli ̄
donioideaeꎬ Papaveraceae) [J] . Botanicˇeskii Žhurnal (Moscow &
Leningrad)ꎬ 79: 70—76
Sha Wꎬ Wang LHꎬ Yang XJ et al.ꎬ 1995. Chromosome numbers of 20
species from northeast China [ J] . Journal of Wuhan Botanical
Researchꎬ 32: 180—182
Shatalova SAꎬ 2000. Chromosome numbers in vascular plants of the
Primorsky territory [J] . Botanicˇeskii Žhurnal (Moscow & Lenin ̄
grad)ꎬ 85: 152—156
Starodubtsev VNꎬ 1989. Chromosome numbers in species of the fami ̄
lies Aceraceaeꎬ Brassicaceaeꎬ Cyperaceaeꎬ Euphorbiaceaeꎬ Pa ̄
paveraceae and Ranunculaceae from the Soviet Far East [J] . Bo ̄
tanicˇeskii Žhurnal (Moscow & Leningrad)ꎬ 74: 1674—1675
Vent Wꎬ Mory Bꎬ 1973. Beitrage zur kenntnis der sippenstruktur der
gattungen Glacium Adans und Dicranostigma Hooker f. et Thom ̄
son (Papaveraceae) [J] . Gleditschiaꎬ 1: 33—41
Wu LHꎬ Wang SYꎬ 1997. A new species of Dicranostigma from
Henan and its karyotype [ J] . Bulletin of Botanica Researchꎬ
Harbinꎬ 17: 42—46
Wu ZYꎬ Chuang Hꎬ 1999. Dicranostigma [A] / / Wu ZY ed.ꎬ Flora
Reipublicae Popularis Sinicae [M]. Beijing: Science Pressꎬ 32:
61—64
Xiang CLꎬ Chen YPꎬ Funamoto T et al.ꎬ 2014. Cytology of thirteen
taxa in the genus Isodon (Lamiaceae: Nepetoideae) from China
[J] . Plant Diversity and Resourcesꎬ 36: 561—568
Xie HYꎬ Ash JEꎬ Linde CC et al.ꎬ 2014. Himalayan ̄Tibetan Plateau
Uplift Drives Divergence of Polyploid Poppies: Meconopsis Viguier
(Papaveraceae) [J] . PLOS Oneꎬ 9: e99177
Xu ZXꎬ Wei JHꎬ 1994. The chromosome numbers of some medicinal
taxa of Corydalis and their phylogenetic relationships [J] . China
Journal of Chinese Materia Medicaꎬ 19: 82—83
Yan GXꎬ Zhang SZꎬ Xue FH et al.ꎬ 2000. The chromosome numbers
and natural distribution of 38 forage plants in north China [ J] .
Grassland of Chinaꎬ 5: 1—5
Yang QEꎬ 2002. Cytology of the tribe Trollieae and of the tribe Cimic ̄
ifugeae in the Ranunculaceae: a comparative study [ J] . Acta
Phytotaxonomica Sinicaꎬ 40: 52—65
Zhang MLꎬ Su TYꎬ Magnus L et al.ꎬ 2008. Papaveraceae [ A] / /
Wu ZYꎬ Raven RHꎬ Hong DY eds.ꎬ Flora of China [M]. Bei ̄
jing: Science Pressꎻ St. Louis: Missouri Botanic Garden Pressꎬ
7: 261—428
Zhang YHꎬ Fu CXꎬ 1993. Chromosome numbers of 3 species and 1
variety in Corydalis [J] . Journal of Wuhan Botanical Researchꎬ
11: 383—384
Zhou CJꎬ Zhou SDꎬ Huang DQ et al.ꎬ 2012. Karyotypes of 25 popula ̄
tions of 15 species in Allium section Rhiziridium from China
[J] . Plant Diversity and Resourcesꎬ 34: 120—136
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