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     

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.     

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     

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。     

风毛菊属6种植物的核型分析

采用常规压片技术对分布于横断山区菊科(Compositae)风毛菊属(Saussurea DC.)的6种植物进行染色体数目和核型分析。结果表明:尖苞雪莲（S.polycolea var.acutisquama）核型公式为:2n=2x=32=20m+12sm,属2B型;球花雪莲核（S.globosa）型公式为:2n=2x=34=16m+18sm,属2B型;重齿风毛菊(S.katochaete)核型公式为:2n=2x=32=8m+18sm+6st,属3B型;柱茎风毛菊(S.columnaris)核型公式为:2n=2x=32=24m+8sm,属2B型;禾叶风毛菊(S.graminea)核型公式为:2n=2x=28=8m+18sm+2st,属3B型;长毛风毛菊(S.hieracioides)核型公式为:2n=2x=32=12m+16sm+4st,属2B型。6个种染色体中均未发现随体。其中尖苞雪莲和柱茎风毛菊染色体为首次报道。     

Variation in karyotypes of Dugesia japonica japonica (Turbellaria) from Ôsaka Prefecture,Central Japan

An analysis of the karyotypes of Dugesia japonica japonica Ichikawa et Kawakatsu, 1964, from 30 localities of seven river systems in Ôsaka Prefecture, Central Japan, revealed a total of 26 karyotypes of which 12 are new-found varieties. More than two karyotypes were found in many localities. The mixoaneuploidic triploid karyotype showed the widest distribution (21 localities), orthoploidic diploid karyotype and orthoploidic mixoploid of diploid and triploid karyotype were next (13 localities, respectively), and other karyotypes (triploidic aneuploid, orthoploidic triploid and mixoploid of triploid & tetraploid) were also found. Some correlation of karyotype with elevation was also detected.     

Karyotypes,nucleoli, and amphiplasty in hybrids between Hordeum vulgare L. and H. bulbosum L.

Chromosome measurements were carried out in Hordeum vulgare, H. bulbosum, and their diploid, triploid, and tetraploid hybrids. The chromosomes were classified by using relative values, and thus karyotypes were established. For comparison of these karyotypes both relative and absolute values were used. It was concluded that differential amphiplasty occurred, whereas neutral amphiplasty could not be demonstrated. In the hybrids the relative length of the parts of the chromosomes (long arm, short arm, satellite) was not changed in comparison with these lengths in the pure species. The karyotypes of both species had considerable similarities. From comparing the mean absolute genome lengths, it was, however, concluded that in the pure species, as well as in all hybrid types, the chromosomes of H. vulgare were longer than those of H. bulbosum. In the diploid and tetraploid hybrids the mean genome lengths were shorter than those in the pure species and the triploid hybrids. The differential amphiplasty was such that the secondary constriction of chromosome 6 of H. bulbosum, did not show up in the hybrids. This could be related to the suppression of nucleolar formation in the genome of H. bulbosum, because the maximum number of nucleoli in root tip cells equalled the number of satellite chromosomes. Finally it was found that the pattern of nucleolar fusion in diploid and triploid hybrids deviated from the expectation. The results were discussed in relation to chromosomal disturbances that occurred in the hybrid tissues and that resulted in elimination of chromosomes and other effects.     

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.     

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

Abstract 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.     

Phylogeny and biogeography of Chinese Heracleum (Apiaceae tribe Tordylieae) with comments on their fruit morphology

The Sino-Himalayan Mountain Region is a major center of diversity of Heracleum (Apiaceae, Tordylieae, Tordyliinae), yet the phylogenetic relationships among its rich diversity of endemic taxa are largely unknown. In this study, we (1) investigate the monophyly of Heracleum and ascertain the phylogenetic placements of those species of Heracleum from China that fall outside of the group; (2) estimate phylogenetic relationships within Heracleum sensu stricto, particularly among those species occurring in the Hengduan Mountain Region of southwestern China; and (3) reconstruct the biogeographic history of Heracleum sensu stricto. To achieve these objectives, Bayesian inference and maximum-parsimony analyses of the nuclear ribosomal DNA (nrDNA) internal transcribed spacer (ITS) region and five noncoding chloroplast DNA (cpDNA) loci (rps16-trnK, rps16 intron, trnQ-rps16, rpl32-trnL, and psbA-trnH), separately and in combination, were carried out. Fruit morphology of the Chinese Heracleum species was examined. Statistical dispersal-vicariance analysis (S-DIVA) was used to reconstruct the biogeographic history. In total, 114 accessions were considered, which included 88 species representing eight genera of subtribe Tordyliinae, 28 of the 29 species of Heracleum occurring in China, and 4 species of Angelica as outgroups. The genus Heracleum is monophyletic upon the exclusion of nine species of Heracleum from China and the inclusion of the Caucasian monotypic genera Mandenovia and Symphyoloma. These nine excluded species show affinities to Tetrataenium, Semenovia, and Angelica. The species of Heracleum endemic to southwestern China form a moderately to strongly supported clade; however, major discordance between the ITS- and cpDNA-derived phylogenies, lack of resolution in the trees, and polyphyly of several species preclude unequivocal hypotheses of phylogenetic relationship. The distinctly clavate vittae, shorter than the length of the mericarp, is a diagnostic feature for Heracleum sensu stricto. Heracleum originated in the West Palearctic, with three possible migration routes leading to its present-day distribution. Major topological conflicts between the ITS and cpDNA trees may be due to interspecific hybridization and/or incomplete lineage sorting. The taxonomic limits of many Chinese Heracleum species remain unclear, possibly because of recent radiation in the Hengduan Mountains.     

Karyotype studies in thirty‐two species of Lilium (Liliaceae) from China1

Fifty‐two populations representing thirty‐two taxa of Lilium L. from China are karyologically analyzed. The results showed that all populations have the same basic chromosome number x=12, and all species are diploid except Lilium tigrinum, which is triploid with 2n=36. In addition, one population of L. sulphureum is aneuploid with 2n=23. The karyotype evolution in the genus Lilium is mainly in terms of alterations of the fine structure of the chromosomes, not in ploidy or basic chromosome number variation. When combining previous studies and our results, we consider the major driving forces for evolution of Lilium to be an increase in karyotypic asymmetry, and unequal translocation and formation of secondary constrictions. The existence of intercalary satellites on the two largest pairs of chromosomes is correlated with the geographic distribution pattern of Lilium across the world. The Hengduan Mountains is a modern diversity and differentiation center of Lilium and it also could be the center of origin for this genus. In addition, some taxonomic conclusions were verified on the subgenus level. Among the species investigated, the chromosome number and karyotypes of 13 taxa were documented for the first time.     

A review of chromosomal variation in Dugesia japonica and D. ryukyuensis in the Far East

The chromosome numbers of Dugesia japonica Ichikawa et Kawakatsu, 1964, are n = 8, 2x = 16 and 3x = 24; those of Dugesia ryukyuensis Kawakatsu, 1976, are n = 7, 2x = 14 and 3x = 21. The karyotypes of both species include diploid, triploid and mixoploid; aneuploidic and mixoaneuploidic karyotypes may occur. In 785 specimens studied of D. japonica, the occurrence rates of specimens having each karyotype are substantially the same (29–37%). Diploid sexual specimens represented nearly 10% of the total and virtually no triploid or mixoploid sexual specimens were found. The diploid karyotype can be inherited by both sexual and asexual reproduction; the triploid and mixoploid karyotypes will be inherited only by asexual reproduction. In 51 specimens studied of D. ryukyuensis, the different karyotypes are diploid (ca 39%), triploid (ca 57%) and mixoploid (ca 4%). Diploid sexual specimens represented nearly 25% of the total; sexual specimens with tripooidic karyotypes made up nearly 27%. The diploid, triploid and mixoploid karyotypes were also found in juveniles hatched from cocoons. The diploid karytyype is inherited by both sexual and asexual reproductions; the other karyotypes may be inherited by parthenogenesis or self-fertilization (including pseudogamy) and asexual reproduction.     

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。     

Cytological studies on the genus Holcoglossum (Orchidaceae)

The karyotypes of ten species of Holcoglossum (Orchidaceae), a highly endangered and diversified genus from China, were investigated to study the infrageneric relationships, biogeography, and speciation patterns in the Hengduan Mountains. The karyotype formulae of the studied species are as follows: 2 n  = 38 = 20m + 18sm in H. subulifolium , 2 n  = 38 = 22m + 16sm in H. amesianum , 2 n  = 38 = 26m + 12sm (6 SAT) in H. lingulatum , 2 n  = 38 = 26m + 12sm in H. wangii , 2 n  = 38 = 10m + 28sm in H. kimballianum , 2 n  = 38 = 14m + 22sm + 2st in H. flavescens , 2 n  = 38 = 24m + 12sm + 2st in H. rupestre , 2 n  = 38 = 14m + 20sm + 4st in H. sinicum , 2 n  = 38 = 16m + 14sm + 8st in H. weixiense , and 2 n  = 76 in H. tsii . The karyotypes of two tropical species, H. amesianum and H. subulifolium , are the most primitive in the genus, whereas those of four temperate alpine species, H. sinicum , H. rupestre , H. weixiense , and H. flavescens , are more advanced. H. tsii is a tetraploid and H. rupestr e may be one of its ancestors. The low frequency of polyploidy in Holcoglossum in the Hengduan Mountains region supports the conclusion that chromosome stasis during rapid speciation is common there.  © 2007 The Linnean Society of London. Botanical Journal of the Linnean Society , 2007, 154 , 283–288.     

Karyosystematic studies on threeCrepis species (Asteraceae) endemic to Greece

This work examines the cytogeographical distribution, the morphological characters, and the karyotypes of threeCrepis species endemic to Greece (C. sibthorpiana, C. incana, andC. heldreichiana). C. sibthorpiana is diploid (2n = 2x = 8),C. incana is diploid (2n = 2x = 8) and tetraploid (2n = 4x = 16, 17), andC. heldreichiana is always dekaploid (2n = 10x = 40). The Giemsa positive bands, usually pairs of dots, are mainly centromeric inC. incana, while they are terminal inC. sibthorpiana (on the short arm of all chromosomes) and inC. heldreichiana (on both arms of all chromosomes). Intercalary C-bands are scarce and usually variable within karyotypes, individuals, and species. The most variable karyotype both in Feulgen and Giemsa preparations is that ofC. incana, in which also supernumerary chromosomes were observed, which are polysomic to standard set members. On the basis of morphological and karyological data the evolutionary relationships among the threeCrepis taxa are discussed.     

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.     

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。     

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。     

Karyotypes of some species of the genus Lessingianthus (Vernonieae,Asteraceae) and taxonomic implications

We determined the karyotypes of nine species of Lessingianthus, eight of which are here analyzed for the first time. The results include the first chromosome count for L. plantaginoides, which is tetraploid with 2n = 64. All species showed a high proportion of metacentric chromosomes combined with a lower proportion of submetacentric pairs. Only L. coriaceus had a subtelocentric chromosome pair. B chromosomes were observed in L. coriaceus and L. varroniifolius, which were subtelocentric. Differences among the karyotypes of the studied species were small, suggesting that karyotype diversity in the genus evolved by small changes in the structure of chromosomes. Karyotype features appear useful to distinguish Lessingianthus from the closely related genera Chrysolaena and Lepidaploa.