Chromosome counts and karyotypes in Chaetoseris and Stenoseris (Asteraceae-Cichorieae) from the Hengduan Mountains of SW China

Chaetoseris and Stenoseris are two morphologically close genera from the tribe Cichorieae of the sunflower family and they are endemic in alpine eastern Himalayas to the Hengduan Mountains of SW China.Mitotic chromosome numbers and karyotypes are reported for 12 populations representing eight species of Chaetoseris and two species of Stenoseris from the Hengduan Mountains region.Eight species are new and the other two provide confirmation of previous reference.All Chaetoseris and Stenoseris taxa are diploidy with 2n ＝ 16 and their basic number is tentatively suggested as x ＝ 8.Karyotypes of Chaetoseris and Stenoseris are similar to each other with 2A and 2B for the former and 2A for the latter.Cytological data of chromosomal numbers and karyotypes support a close relationship of the two genera.Currently no polyploids are found for these two genera and it seems that polyploidization has played a minor role in their evolutionary speciation in the Hengduan Mountains region.     

Karyotype and cytogeography of the genus Heracleum (Apiaceae) in the Hengduan Mountains

Abstract In the present study, the karyotypes of 34 populations belonging to 11 species and one variety of Heracleum from the Hengduan Mountains in China were examined. Chromosome numbers and the karyotypes of three species (H. souliei, H. kingdoni, and H. wenchuanense) are reported for the first time, as are the karyotypes of H. moellendorffii and H. henryi (tetraploid). Populations of H. candicans, H. franchetii, and H. kingdoni in the Hengduan Mountains were found to consist of a mixture of diploid and tetraploid plants. Except for four species of Heracleum, namely H. candicans, H. franchetii, H. henryi, and H. kingdoni, which have both diploid and tetraploid karyotypes, all other species of Heracleum are were found to be diploid. All karyotypes were found to belong to the 2A type of Stebbins, with the exception of H. candicans var. obtusifolium, which belongs to 2B, and H. hemsleyanum and H. franchetii (Mt. Dujuan, Daocheng, Sichuan, China), which belong to 1A. There was only a slight difference in the karyotype asymmetry index, which suggests a close kinship for species of Heracleum and that the entire phylogenetic development of Heracleum is relatively primitive. Species that exhibited advanced morphological features were also more advanced in karyotype structure, with the order of karyotype evolution being 1A→2A→2B. This phenomenon indicates that the species distributed in the Hengduan Mountains have not diverged completely and that the Hengduan Mountains are a relatively young and active area for the evolution of Heracleum. Polyploidization in Heracleum may be an important evolutionary mechanisms for some species, generating diversity. The biological attributes, distribution range, and the geological history of the genus have all played a part in accelerating the evolution through polyploidization or aneuploidization. It is known that as the distribution latitude of Heracleum decreases from north to south, the chromosome number, ploidy level, and asymmetry structure appear to increase. In the Hengduan Mountains, these tendencies are also evident. Finally, based on all the available cytogeographic data, we speculate that the more advanced tetraplont or aneuploid species of Heracleum in India may be derived from early diplont species that were distributed in the Caucasus region and Hengduan Mountains. The dispersal of Heracleum was from Eurasia to India, because this correlates with the emergence of the Himalayan Mountains through tectonic movement. Thus, the Hengduan Mountains are not only a center of diversity for Heracleum, but also a center of active speciation in modern times.     

Achene Wall Anatomy and Surface Sculpturing of Lactuca L. and Related Genera （Compositae： Lactuceae） with Notes on Their Systematic Significance

The achene wail anatomy and surface sculpturing of 14 species representing Lactuca L. and related genera were Investigated to evaluate inter-and Intrageneric relationships. The achene wall anatomy of the studied species can be divided Into two types： winged and ribbed. The winged type is present in Lactuca L., Pterocypsela Shlh, Clcerblta Wallr., Chaetoserls Shlh and Stenoseris Shlh, and is characterized by protruding wings and costae in transverse section. Winged type achene mesocarps are composed of parenchymatous cells and fiber ceils, and the distinct fibrous strands are confined to the costae or wings of the achenes. The ribbed type is present in Notoseris Shih and Paraprenanthes Chang ex Shih, and Is characterized by only having costae In transverse section. Fiber cells are continuous in the costae and intercostae, in surface sculpturing, the ornamentation and the shape of epidermal cells are different among these genera. The results Indicate that L. altalca and L. serrlola should be conspeclfic, and that L. dollchophylla is probably an Intermediate taxon between Lactuca and Chaetoserls. The results also support the separation of Pterocypseia, Paraprenanthes, Notoseris, Chaetoseris and Stenoserls from Lactuca and Clcerblta as independent entitles. Close affinities between Pterocypsela and Lactuca, Paraprenanthes and Notoserls, and Chaetoserls and Stenoserls are also proposed.     

横断山区两种菊科植物的核型研究

对横断山区两种菊科植物圆齿狗娃花和细叶亚菊进行了染色体数目报道和核型研究,其中圆齿狗娃花为首次报道,2n=18=2x=18m(2SAT);而细叶亚菊的为新的染色体数目,2n=72=8x=56m+16sm,与之前报道的36不同。研究结果发现菊科植物这两个属在横断山及其它地区同时存在二倍体与多倍体。     

Karyotype and cytogeography of the genus Heracleum （Apiaceae） in the Hengduan Mountains

In the present study, the karyotypes of 34 populations belonging to 11 species and one variety of Heracleum from the Hengduan Mountains in China were examined. Chromosome numbers and the karyotypes of three species （H. souliei, H. la＇ngdoni, and H. wenchuanense） are reported for the first time, as are the karyotypes of H. moellendorffii and H. henryi （tetraploid）. Populations of H. candicans, H. franchetii, and H. kingdoni in the Hengduan Mountains were found to consist of a mixture of diploid and tetraploid plants. Except for four species of Heracleum, namely H. candicans, H. franchetii, H. henryi, and H.kingdoni, which have both diploid and tetraploid karyotypes, all other species of Heracleum are were found to be diploid. All karyotypes were found to belong to the 2A type of Stebbins, with the exception ofH. candicans var. obtusifolium, which belongs to 2B, and H. hemsleyanum and H. franchetii （Mt. Dujuan, Daocheng, Sichuan, China）, which belong to 1A. There was only a slight difference in the karyotype asymmetry index, which suggests a close kinship for species of Heracleum and that the entire phylogenetic development of Heracleum is relatively primitive. Species that exhibited advanced morphological features were also more advanced in karyotype structure, with the order ofkaryotype evolution being 1A→2A→2B. This phenomenon indicates that the species distributed in the Hengduan Mountains have not diverged completely and that the Hengduan Mountains are a relatively young and active area for the evolution of Heracleum. Polyploidization in Heracleum may be an important evolutionary mechanisms for some species, generating diversity. The biological attributes, distribution range, and the geological history of the genus have all played a part in accelerating the evolution through polyploidization or aneuploidization. It is known that as the distribution latitude of Heracleum decreases from north to south, the chromosome number, ploidy level, and asymmetry structure appear to increase. In the Hengd     

Cytological study of Tibetia (Fabaceae) in the Hengduan Mountains region, China

The Hengduan Mountains comprise one of the world's most important hot spots of biodiversity. Tibetia (Ali) H.P. Tsui (Fabaceae), which has four or five species in two sections, is one of the genera endemic to the region. This paper describes for the first time the karyotype of three of those species. The chromosome counts of all three are 2n = 16. The karyotypes of the species examined contain chromosomes of variable karyotypic symmetry with centromeres at median and submedian positions that correlate with the morphological characteristics of the species. Karyotypic variation at the diploid level appears to be the predominant feature of chromosome evolution in the genus and may provide a clue to the study of evolutionary patterns of plants in this region. Received: June 12, 2001 / Accepted: September 12, 2001     

基于染色体计数和ITS序列初步探讨横断山区柴胡属植物(伞形科)的系统发育

横断山区是中国柴胡属Bupleurum植物的分布中心。本文对横断山区6个种2变种进行了染色体记数报道,其中4个种2变种是首次报道。对横断山区的10个种4个变种、中国北方（河北和黑龙江）的3个种的nrDNA ITS进行测序,同时从GenBank里面下载同属的来自非洲和地中海西部的16个nrDNA ITS序列数据,结合染色体数目变化结果,初步探讨了横断山区柴胡属植物的系统发育。结果表明横断山区可能是现代柴胡属植物的频度中心和多样分布中心之一。它们的祖先种可能是非洲北部的木本柴胡属植物B.fruticosum,或者是地中海西部的柴胡属植物,推测是通过中东和高加索扩散而形成的,其中与非洲南部特有种B.mundtii的亲缘关系也较近;染色体基数演化趋势是：8是较原始基数,6和7是次生基数,其染色体异基数变异和多倍化可能是物种形成、进化以及向外扩散的主要方式;在ITS系统发育树中,中国柴胡属植物染色体基数为8的种类聚为一支,染色体基数为6和7的种类聚为了一支,不支持舒璞等（1998）关于中国柴胡属的属下分类系统。结合已有的形态学、细胞学、孢粉学证据和ITS系统发育树,建议窄竹叶柴胡B.marginatum var.stenophyllum独立成种。     

Low incidence of polyploids and high uniformity of karyotypes displayed by Delphinium (Ranunculaceae) in the Hengduan Mountains region of south‐west China

The chromosome numbers and morphology in 92 populations belonging to 49 species and three varieties in the genus Delphinium L. (Ranunculaceae), mostly from the Hengduan Mountains region of south‐west China, were studied. Forty seven species and three varieties were diploid, with 2n = 16, one species was tetraploid, with 2n = 32, and one species had diploid and tetraploid cytotypes. Three species had B chromosomes, representing the first time the occurrence of B chromosomes has been reported in the genus. The karyotypes of all the diploid species were quite uniform, commonly bimodal, and usually consisted of one pair of large median‐centromeric (m), one pair of large submedian‐centromeric (sm), five pairs of medium‐sized subterminal‐centromeric (st), and one pair of smaller sm (rarely st) chromosomes. The low incidence of polyploids in Delphinium from the Hengduan Mountains region indicates that polyploidy has played a minor role in the speciation of this highly diversified genus in the region. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 158 , 172–188.     

Phylogenetic inference of the genus Bupleurum (Apiaceae) in Hengduan Mountains based on chromosome counts and nuclear ribosomal DNA ITS sequences

pleurum of Apiaceae in China. This paper reports chromosome numbers of six species and two varieties of Bupleurum, and for four species and two varieties their chromosome numbers are reported for the first time. The phylogeny of Bupleurum was investigated based on the ITS region of the nuclear ribosomal DNA (nrDNA) of 14 taxa from the Hengduan Mountains, 3 taxa from the North China (Hebei and Heilongjiang), and 16 taxa from Africa and the Mediterranean region. Varia-tions in chromosome numbers and the ITS sequences were used to infer phylogenetic relationships between Bupleurum species in Hengduan Mountains. The results showed that the Hengduan Mountains might represent one of the frequency and diversity centers for Bupleurum. The ancestors of Bupleurum species in the Hengduan Mountains may be related to the woody B. fruticosum in North Africa, or the species in the western Mediterranean region. It is postulated that the ancestral population migrated into Hengduan Mountains through the Middle East and the Caucasus. Furthermore, the neo-endemic B. mundtii in South Africa appeared to be a close relative of the species in the Hengduan Mountains. In the trend of basic chromosome number evolution, x = 8 should be regarded as the ancestral basic number, while x = 6, 7 as the derived ones. The Bupleurum species in the Hengduan Moun-tains have been undergoing changes in the basic chromosome numbers or the ploidy level. The ITS phylogenetic tree showed that the Chinese species were divided into two clades: one with the basic chromosome number x = 8, and the other with x = 6, 7. The results rejected the previous infrageneric classification of Bupleurum in China. We further suggested to raise B. marginatum var. stenophyllum to species rank based on the combined evidence from morphology, karyology, pollen morphology, and the ITS phylogenetic tree.     

横断山区被子植物染色体研究概况

横断山区作为全球生物多样性热点地区之一 ,染色体数目的研究对探讨植物区系起源和进化有重要的意义。本文对横断山区被子植物染色体数目报道方面的情况进行收集和整理并进行统计分析 ,结果发现共有 42科 1 4 3属 51 8种被子植物有染色体数目报道。     

Distribution of gamasid mites on small mammals in Yunnan Province, China

An investigation of gamasid mites on the body surface of small mammals was carried out in Yunnan Province of China from 1990 to 2004. The small mammal hosts were captured from 25 counties which represent five geographical subregions, namely Middle Subregion of Hengduan Mountains, Southern Subregion of Hengduan Mountains, Eastern Plateau Subregion of Yunnan, Western Plateau Subregion of Yunnan and Southern Moun- tainous Subregion of Yunnan. The captured 10 803 small mammal hosts belong to nine families, 29 genera and 52 species in four orders （Rodentia, Insectivora, Scandentia and Lagomorpha）. A total of 68 571 gamasid mites were collected from the body surface of the captured small mammal hosts and all the gamasid mites were identified to 10 families, 33 genera and 112 species. This paper lists all the mite species, together with their taxonomic position （genera and families） and their corresponding hosts. Much more mite species were found in the Middle Subregion of Hengduan Mountains than in other geographical subregions. The total individuals of mites and small mammals in the Middle Subregion of Hengduan Mountains are also the most plentiful in the five geographical subregions. Three dominant mite species and three dominant small mammal hosts were determined as the dominant species in the investigated areas of Yunnan Province. The dominant hosts are Rattus flavipectus （which accounts for 34.85% of the total individuals）, Apodemus chevrieri （13.43%） and Rattus norvegicus （10.40%） while the dominant gamasid mite species are Laelaps nuttalli （Hirst, 1915） （27.84%）, Laelaps echidninus （Berlese, 1887） （18.38%） and Laelaps guizhouensis （Gu et Wang, 1981） （14.79%）. The results showed the high species diversity of gamasid mites in Yunnan Province and the uneven distribution feature in different subregions.     

Species reassessment congruent with the phylogeographical study of the Biston falcata species group

The Himalaya, Hengduan, and Qinling mountains are very important mountain systems in China. Because they are geographically adjacent, species distributed in these three mountains usually have close relationships, but species/populations distributed in the first two mountain systems have higher genetic diversity and more complex phylogenetic structures than those found in the Qinling Mountains. In this study, we used molecular and morphological methods to explore the phylogeographical structure, species differentiation, and evolutionary history of the Biston falcata species group, which is widely distributed in the Himalaya, Hengduan, and Qinling mountains. Five species were identified; B. quercii is distributed in the Qinling Mountains and its adjacent areas, and the other four species are mainly distributed in the Himalaya and Hengduan Mountains. Four gradual species differentiation events occurred at approximately 3.07, 1.75, 1.50, and 1.30 Ma, corresponding to the stage with strong geological movements of the Qinghai‐Tibet Plateau. The low land between the Qinling and the northern Hengduan Mountains, the Sichuan Basin, the low land between the eastern and central Himalaya Mountains and the Mekong‐Salween Divide are four geographical barriers that resulted in multiple speciations. In addition, long‐range dispersal as another mechanism for speciation is non‐negligible. In spite of failing to confirm the ancestral area, our study provided direct evidence of the speciation induced by the geographical barriers and adaptation to the plateau.     

横断山区六种八居群鼠尾草属植物的核型分析

鼠尾草属(Salvia)是唇形科(Lamiaceae)最大的属,属下多种为民间常用草药,亦有供观赏的种类。为探究横断山区物种在细胞学水平的进化方式,讨论形态分类学与分子系统学之间的分类关系,该研究通过广泛收集染色体文献资料,采用植物常规压片法对采集自横断山地区6种8居群鼠尾草属植物进行核型分析,并构建了中国地区分布的鼠尾草属植物叶绿体系统发育树。统计结果表明:(1)全世界范围内报道了约23%的鼠尾草属植物染色体数据,其中分布在中国地区的鼠尾草属植物染色体报道率为32.10%,分布在横断山地区的鼠尾草属植物报道率为40.54%,(2)鼠尾草属植物染色体基数以x=8和x=11为主,分布在中国地区的鼠尾草属植物染色体基数均为x=8。实验结果表明:(1)西藏鼠尾草(S. wardii)核型数据为首次报道。(2)雪山鼠尾草(S. evansiana)首次在云南德钦地区发现二倍体居群。将细胞学数据结合叶绿体进化树开展染色体进化关联分析,论证多倍化可能不是鼠尾草属物种适应高海拔环境的主要机制,表明多倍体不是该属物种形成的主要进化途径而是以二倍体水平为主,推测染色体组的加倍可能是物种在形态学与分子系统学上分类关系不一致的原因之一。该研究丰富了横断山区鼠尾草属植物的染色体核型数据,结合区域分子系统树探讨染色体特征的进化关系,为今后深入研究该属物种的核型进化做出了探索,为开展祖先物种染色体基数推演分析补充了基础数据。     

Cytological studies on the Sino-Himalayan endemic Anisodus and four related genera from the tribe Hyoscyameae (Solanaceae) and their systematic and evolutionary implications

Cytological studies were carried out on eight species of five genera ( Anisodus , Atropanthe , Hyoscyamus , Mandragora and Przewalskia ) in the tribe Hyoscyameae (Solanaceae). First chromosome counts were reported in six species: Anisodus luridus , A. acutangulus , A. tanguticus , A. carniolicoides , Atropanthe sinensis and Mandragora caulescens , all with 2 n  = 8 x  = 48. Two records, for Hyoscyamus niger (2 n  = 2 x  = 34) and Przewalskia tangutica (2 n  = 4 x  = 44), were also confirmed. All species studied showed the proximal type of mitotic prophase chromosome condensation pattern. Three types of interphase nuclei were recognized: the round prochromosome type for Anisodus , Atropanthe and Mandragora , the rod prochromosome type for Przewalskia and the complex chromocentre type for Hyoscyamus . The cytological data supported the close relationship of all four species of Anisodus . Evidence from the interphase nuclei and chromosome base numbers supported the traditional classification of Hyoscyameae into two groups, i.e. Physochlaina praealta + Hyoscyamus (complex interphase type and x  = 7, 14, 17) and Przewalskia + Atropanthe + Anisodus + Scopolia + Atropa + Mandragora (prochromosome type and x  = 6, 11). Polyploidy is found in most species of the tribe in the Himalayan–Hengduan Mountains, as well as in the Mediterranean region, west–central Asia and eastern Asia. It seems that it probably occurred very early in Hyoscyameae evolution, before the uplift of the Himalayan–Hengduan Mountains. The Himalayan orogeny might have played a minor role in the polyploid evolution of plants in this tribe.  © 2005 The Linnean Society of London. Botanical Journal of the Linnean Society , 2005, 147 , 457–468.     

Cytogenetic studies of two land-planarian species from Brazil: Geoplana marginata and Issoca rezendei (Tricladida,Terricola)

The chromosome complement and the C-banded karyotypes of specimens of Geoplana marginata auct. and of Issoca rezendei (Schirch) were investigated. The diploid and fundamental numbers of the two species were identical (2n = 14; FN = 28). Their karyotypes were similar except for the morphology of chromosome pair 6. Their C-banding patterns differed in quantity and localization of the constitutive heterochromatin. The similarity in karyotypes of these species support the hypothesis, proposed earlier on morphological grounds, that the genera Geoplana Stimpson and Issoca Froehlich are closely related. G. marginata and I. rezendei are the first land planarian species of the Neotropical Region to have their karyotypes described.     

Cytological study of Tibetia (Fabaceae) in the Hengduan Mountains region,China

The Hengduan Mountains comprise one of the world's most important hot spots of biodiversity. Tibetia (Ali) H.P. Tsui (Fabaceae), which has four or five species in two sections, is one of the genera endemic to the region. This paper describes for the first time the karyotype of three of those species. The chromosome counts of all three are 2 n = 16. The karyotypes of the species examined contain chromosomes of variable karyotypic symmetry with centromeres at median and submedian positions that correlate with the morphological characteristics of the species. Karyotypic variation at the diploid level appears to be the predominant feature of chromosome evolution in the genus and may provide a clue to the study of evolutionary patterns of plants in this region.     

中国独活属的核型研究

本文首次报道了我国独活属10种(含1变种)的核型,其中8种的染色体数目亦为首次报道。研究表明它们的染色体数目均为2n＝2x＝22,为二倍体,其核型为中部着丝点和亚中部着丝点染色体组成,仅具1A和2A两种类型。有4种具随体染色体,在部分H．vicinum植株中发现有1条B染色体,在H．millefolium var．longilobum中同时存在2n＝22和2n＝24的非整倍体．不同种的染色体形态不同,为分类研究提供了细胞学依据。 本文在核型分析基础上讨论了该属分类群的划分、种间亲缘关系及演化,并结合我国毗邻地区该属细胞学资料和地理分布规律,提出我国西南部的横断山区是独活属的频度中心和多样性中心。     

五种齿突蟾在横断山南潜在地理分布预测

由于栖息地质量下降,近年来齿突蟾属物种种群数量急剧减少,明确齿突蟾属物种空间分布,是监测、管理、保护齿突蟾属物种的基础。横断山区可能是齿突蟾属的起源中心和分化中心,但齿突蟾属在横断山区的地理分布格局尚不明确。利用优化后Maxent模型,首次预测西藏齿突蟾Scutiger boulengeri、刺胸齿突蟾Scutiger mammatus、胸腺齿突蟾Scutiger glandulatus、圆疣齿突蟾Scutiger tuberculatus、贡山齿突蟾Scutiger gongshanensis 5种高海拔齿突蟾属物种在横断山南生物多样性保护优先区域的潜在地理分布,并分析其与环境因子的关系。结果显示,5种齿突蟾属物种在横断山南的潜在地理分布格局存在差异,西藏齿突蟾主要分布在横断山南的北部,圆疣齿突蟾主要分布在横断山南东北部的四川省境内,贡山齿突蟾主要分布在横断山南的西南部,刺胸齿突蟾和胸腺齿突蟾的潜在分布格局较为相似,在横断山南的中部、西北部地区都有较多分布,但胸腺齿突蟾潜在分布区更为碎片化。另外,横断山南北部地区的齿突蟾属丰富度明显高于南部地区。环境变量贡献率和刀切法结果显示温度因子和降水因子是决定横断山南齿突蟾属潜在分布的主要因素,最冷季降水量对西藏齿突蟾、贡山齿突蟾、圆疣齿突蟾潜在分布有重要影响,但它们对最冷季降水量的偏好存在差异。此外,研究也显示,通过评估潜在的Maxent参数组合,选择最佳的Maxent模型是有效且必要的。     

Chromosome analysis of five Brazilian species of poison frogs (Anura: Dendrobatidae)

Dendrobatid frogs have undergone an extensive systematic reorganization based on recent molecular findings. The present work describes karyotypes of the Brazilian species Adelphobates castaneoticus, A. quinquevittatus, Ameerega picta, A. galactonotus and Dendrobates tinctorius which were compared to each other and with previously described related species. All karyotypes consisted of 2n = 18 chromosomes, except for A. picta which had 2n = 24. The karyotypes of the Adelphobates and D. tinctorius species were highly similar to each other and to the other 2n = 18 previously studied species, revealing conserved karyotypic characteristics in both genera. In recent phylogenetic studies, all Adelphobates species were grouped in a clade separated from the Dendrobates species. Thus, we hypothesized that their common karyotypic traits may have a distinct origin by chromosome rearrangements and mutations. In A. picta, with 2n = 24, chromosome features of pairs from 1 to 8 are shared with other previously karyotyped species within this genus. Hence, the A. picta data reinforced that the C-banding pattern and the NOR location are species-specific traits in the genus Ameerega. Moreover, the Ameerega monophyletism proposed by previous phylogenetic studies indicates that the karyotypic differences among species in this genus result from a long divergence time.     

CHROMOSOMES,HYBRIDS AND PLOIDY OF SEDUM CREMNOPHILA AND ECHEVERIA LINGUIFOLIA (CRASSULACEAE)

Sedum cremnophila and Echeveria linguifolia have generally been placed in different genera on the basis of their flowers—largely because the petals are spreading in one and erect in the other—and the genera have been placed in different subfamilies. However, they are very similar vegetatively and in their unusual inflorescence, their karyotypes are similar (n = 33), and they readily hybridize to produce fertile F₁ hybrids. Study of hybrids of these two species with numerous others leads to the conclusion that each of the two is effectively diploid, with a genome consisting of 33 chromosomes that are all different and that do not pair with each other. Therefore, the good chromosome pairing and the fertility of the hybrid between them are the result of close structural and genetic homology between the corresponding chromosomes of the two species. Taxonomic revision to reflect their very close relationship is desirable. Some other species of Sedum and Echeveria also may need to be reclassified.