Cytogenetics, morphology and evolution of four subspecies of the Giant Sheep argali (Ovis ammon) of Asia

Dalai-lamae (Ovis ammon dalai-lamae), Gobi (O. a. darwini), Kara Tau (O. a. nigrimontana) and Tibetan (O. a. hodgsoni) argali share a 2n = 56 diploid chromosome number and a karyotype consisting of 2 pairs of biarmed and 25 pairs of acrocentric autosomes, a large acrocentric X and a minute Y chromosome. The Giemsa-banding patterns of the largest pair of biarmed chromosomes were identical to those of the largest biarmed chromosomes in all wild sheep and domestic sheep of the genus Ovis. The banding patterns of the second pair of biarmed chromosomes (metacentric) were identical to the third pair of biarmed chromosomes in Ovis with 2n = 54 and to the third largest pair of chromosomes in the 2n = 52 karyotype of Siberian snow sheep (O. nivicola). The G-banded karyotypes of dalai-lamae, darwini, hodgsoni and nigrimontana are consistent with all subspecies of argali (O. ammon), except that the Y chromosome is acrocentric instead of metacentric as typical of the argaliform wild sheep and Ovis. The Dalai-lamae and Tibetan argali specimens exhibit the light-colored, long-haired ruffs and body coloration typical of argalis from the Tibetan Plateau. The Gobi argali, from the extreme western Gobi, is similar to the dark phase argali.

Molecular systematics of pikas (genus Ochotona) inferred from mitochondrial DNA sequences

The phylogenetic relationships among worldwide species of genus Ochotona were investigated by sequencing mitochondrial cytochrome b and ND4 genes. Parsimony and neighbor-joining analyses of the sequence data yielded congruent results that strongly indicated three major clusters: the shrub-steppe group, the northern group, and the mountain group. The subgeneric classification of Ochotona species needs to be revised because each of the two subgenera in the present classification contains species from the mountain group. To solve this taxonomic problem so that each taxon is monophyletic, i.e., represents a natural clade, Ochotona could be divided into three subgenera, one for the shrub-steppe species, a second for the northern species, and a third for the mountain species. The inferred tree suggests that the differentiation of this genus in the Palearctic Region was closely related to the gradual uplifting of the Tibet (Qinghai-Xizang) Plateau, as hypothesized previously, and that vicariance might have played a major role in the differentiation of this genus on the Plateau, On the other hand, the North American species, O. princeps, is most likely a dispersal event, which might have happened during the Pliocene through the opening of the Bering Strait. The phylogenetic relationships within the shrub-steppe group are worth noting in that instead of a monophyletic shrub-dwelling group, shrub dwellers and steppe dwellers are intermingled with each other. Moreover, the sequence divergence within the sister tars of one steppe? dweller and one shrub dweller is very low. These findings support the hypothesis that pikes have entered the steppe environment several times and that morphological similarities within steppe dwellers were due to convergent evolution. (C) 2000 Academic Press.

Genetic diversity and taxonomic implication of Cordyceps sinensis as revealed by RAPD markers

Random amplified polymorphic DNA (RAPD) markers are used to investigate genetic variation and evolutionary relationships of 29 samples of Cordyceps sinensis from different geographical populations on the Qinghai-Tibet plateau. Out of 137 RAPD bands scored, 100 are polymorphic. A correlation is revealed between geographical distance and genetic distance. The molecular phylogenetic tree suggests that the 29 samples are divided into three notable clusters, corresponding to the geographical populations, i.e., the north population (NP), middle population (MP), and south population (SP). The NP consists of 7 northern samples from Menyuan, Maqu, and Luqu, the MP consists of 8 samples from Yushu and Chengduo, and the SP consists of 14 samples from Byma Snow Mountain, Renzhi Snow Moutain, Chongcaoxiwa, and Dacaodi. It is demonstrated that extensive genetic diversity is found among different geographical populations of C. sinensis. The genetic diversity pattern of C. sinensis may be caused by the founder effects. The taxonomic status of NP, MP, and SP populations should be that they are different subspecies rather than different species.

Molecular phylogeny of wood mice (Apodemus, Muridae) in East Asia

Sequences of the mitochondrial cytochrome b (1140 bp) and nuclear IRBP (1152 bp) genes were used to assess the evolutionary history of Apodemus, using the complete set of Asian species. Our results indicate that speciation in Asia involved three radiations, which supports an earlier study. The initial radiation yielded A. argenteus (Japanese endemic), A. gurkha (Nepalese endemic), and the ancestral lineage of the remaining Asian species. This lineage subsequently diverged into four groups: agrarius-chevrieri (agrarius group), draco-latronum-semotus (draco group), A. peninsulae, and A. speciosus (Japanese endemic). The final step consisted of divergence within two species groups as a consequence of the geography of the Yunnan-Guizhou plateau and Taiwan. The ecological ability of two Apodemus-species to inhabit one locality via niche partitioning likely drove the second radiation and shaped the basic geographical pattern seen today: A. argenteus and A. speciosus in Japan, A. agrarius and A. peninsulae in northern China, and the A. agrarius and A. draco groups in southern China. The three radiations are estimated to have occurred 7.5, 6.6, and 1.8-0.8 Mya respectively, using the IRBP clock, based on rat-mouse divergence 12 Mya. (C) 2003 The Linnean Society of London.

Molecular phylogeny and biogeography of Oriental voles: genus Eothenomys (Muridae, Mammalia)

Oriental voles of the genus Eothenomys are predominantly distributed along the Southeastern shoulder of the Qinghai-Tibetan Plateau. Based on phylogenetic analyses of the mitochondrial cytochrome b gene (1143 bp) obtained from 23 specimens (eight species)

Molecular phylogeny of Sinocyclocheilus (Cypriniformes : Cyprinidae) inferred from mitochondrial DNA sequences

More than 10 species within the freshwater fish genus Sinoncyclocbeilus adapt to caves and show different degrees of degeneration of eyes and pigmentation. Therefore, this genus can be useful for studying evolutionary developmental mechanisms, role of natural selection and adaptation in cave animals. To better understand these processes, it is indispensable to have background knowledge about phylogenetic relationships of surface and cave species within this genus. To investigate phylogenetic relationships among species within this genus, we determined nucleotide sequences of complete mitochondrial cytochrome b gene (1140 bp) and partial ND4 gene (1032 bp) of 31 recognized ingroup species and one outgroup species Barbodes laticeps. Phylogenetic trees were reconstructed using maximum parsimony. Bayesian, and maximum likelihood analyses. Our phylogenetic results showed that all species except for two surface species S. jii and S. macrolepis clustered as five major monophyletic clades (I, II, III, IV, and V) with strong supports. S. jii was the most basal species in all analyses, but the position of S. macrolepis was not resolved. The cave species were polyphyletic and occurred in these five major clades. Our results indicate that adaptation to cave environments has occurred multiple times during the evolutionary history of Sinocyclocheilus. The branching orders among the clades I, II, III, and IV were not resolved, and this might be due to early rapid radiation in Sinocyclocheilus. All species distributed in Yunnan except for S. rhinocerous and S. hyalinus formed a strongly supported monophyletic group (clade V), probably reflecting their common origins. This result suggested that the diversification of Sinocyclocheilus in Yunnan may correlate with the uplifting of Yunnan Plateau. © 2005 Published by Elsevier Inc.

Phylogenetic relationships in genus Niviventer (Rodentia : Muridae) in China inferred from complete mitochondrial cytochrome b gene

Chinese species of the genus Niviventer, predominantly distributed in the southeastern Tibetan Plateau and in Taiwan, are a diverse group and have not yet received a thorough molecular phylogenetic analysis. Here, we reconstructed the phylogenetic relatio

Natural selection on EPAS1 (HIF2 alpha) associated with low hemoglobin concentration in Tibetan highlanders

By impairing both function and survival, the severe reduction in oxygen availability associated with high-altitude environments is likely to act as an agent of natural selection. We used genomic and candidate gene approaches to search for evidence of such genetic selection. First, a genome-wide allelic differentiation scan (GWADS) comparing indigenous highlanders of the Tibetan Plateau (3,200 3,500 m) with closely related lowland Han revealed a genome-wide significant divergence across eight SNPs located near EPAS1. This gene encodes the transcription factor HIF2 alpha, which stimulates production of red blood cells and thus increases the concentration of hemoglobin in blood. Second, in a separate cohort of Tibetans residing at 4,200 m, we identified 31 EPAS1 SNPs in high linkage disequilibrium that correlated significantly with hemoglobin concentration. The sex-adjusted hemoglobin concentration was, on average, 0.8 g/dL lower in the major allele homozygotes compared with the heterozygotes. These findings were replicated in a third cohort of Tibetans residing at 4,300 m. The alleles associating with lower hemoglobin concentrations were correlated with the signal from the GWADS study and were observed at greatly elevated frequencies in the Tibetan cohorts compared with the Han. High hemoglobin concentrations are a cardinal feature of chronic mountain sickness offering one plausible mechanism for selection. Alternatively, as EPAS1 is pleiotropic in its effects, selection may have operated on some other aspect of the phenotype. Whichever of these explanations is correct, the evidence for genetic selection at the EPAS1 locus from the GWADS study is supported by the replicated studies associating function with the allelic variants.

Quantitative analysis of land mammal zoogeographical regions in China and adjacent regions

Quantitative analysis of land mammal zoogeographical regions in China and adjacent regions. Zoological Studies 43(1): 142-160. In this paper, our aim was to determine, by means of quantitative analysis, the distribution patterns of the land mammals in China and, adjacent regions using physiographical regions as operative geographical units (OGUs). Based, on the pre-sence or absence of 11 orders, 42 families, 197 genera, and 577 species of land mammals in their zoogeographical regions, which were used as OGUs, we studied the biotic boundary between the Oriental Region (OR) and the Palaearctic Region (PR), as well as subregion boundaries. The boundary's statistical significance was tested by G-test as described by McCoy et al. A significantly strong biotic boundary was found to separate the PR from the OR, and there is a weak biotic boundary in the PR, which divides it into 2 subregions. We concluded that the biotic boundary which separates the PR and OR is a strong boundary. We suggest that the Qinghai-Xizang Plateau should be regarded as a subregion of the PR, which can embody its characteristics of high elevations and a frigid climatic, which is called the Qing-Zang subregion of the PR (QZSP).

Taxonomic review of the genus Sinocrossocheilus Wu (Teleostei : Cyprinidae), with a description of four new species

All Sinocrossocheilus species, except S. microstomatus, are reviewed. Four new species, S. labiata, S. papillolabra, S. nigrovittata, and S. longibulla, are described. The genus Sinocrossocheilus differs from other genera of Cyprinidae by the last simple dorsal fin ray being unserrated and unossified, the last unbranched anal fin ray being unserrated and unossified, the 5-branched anal fin rays, the mouth gap being inferior, the rostral cap covering the lower jaw and connecting directly with the lower lip, a row of fleshy lobes on the lower jaw, and a cloudy black spot above the pectoral fin. Sinocrossocheilus labiata is small and has 22 predorsal scales; S. longibulla has a very large air bladder; S. papillolabra possesses a well-developed ventral fin and a wide band covered by fleshy papillae on the lower lip; and S. nigrovittata possesses black longitudinal stripes along the lateral line. Crossocheilus bamaensis and Crossocheilus liuchengensis are transferred to the genus Sinocrossocheilus. Sinocrossocheilus species are endemic to the central and eastern Yunnan-Guizhou Plateau of China, where river systems are anfractuous, including seasonal rivers, cave rivers, underground rivers, and streamlets between mountains. These separated rivers probably provide conditions for the allopatric speciation of the Sinocrossocheilus.

藏北色林错流域的水文特征

本文报道丁1997年和1998年夏季对藏北色林错等水体水文特征的考察情况结 果表明，色林错是一个大型深水湖泊，表层水温在下午4：30左右达到最高，清晨7：30左右时最 低浅水区域表层湖水在6月份的极端最高水温为2l 2℃．韫端最低水温为11 0℃，而H甲均水温 变幅仅5 6℃，其幅度大于在同一时段湖区内不同区域之间表层水温的变化色林错湖水的DH值 较高．为9 19 9 66；表层水的pH昼夜变化较小，仅O．15．最高值出现在凌晨．同时水体不同区域 及深度的pH值基本一致．其溶解氧变化在4．62—5 12mg／L，而且不同深度水层之间的变幅较小． 仅0 03—0 35mg／L；但其湖汉浆东如瑞溶解氧的昼夜变幅为4 58 5 59mg／I，；盐度的日变幅为 1 60％ 2 20％考察表明．藏北水体的盐分有一个从河流到湖泊、以及由流水向静水富集的趋 势；鱼类的分布与水体盐分的含母密切相关，即鱼类随着水体盐分的增加而消失，代之H嗜盐性卣 虫的出现及大量繁衍

Molecular phylogeny and biogeography of woolly flying squirrel (Rodentia : Sciuridae), inferred from mitochondrial cytochrome b gene sequences

To investigate the genetic diversity between the populations of woolly flying squirrels (Eupetaurus) from the eastern and western extremes of the Himalayas, partial mitochondrial cytochrome b gene sequences (390-810bp) that were determined from the museum specimens were analyzed using maximum parsimony (MP) and maximum likelihood (ML) methods. The molecular data reveal that the two specimens that were collected in northwestern Yunnan (China) are members of the genus Eupetaurus. Reconstructed phylogenetic relationships show that the populations of Eupetaurus in the eastern and western extremes of the Himalayas are two distinct species with significant genetic differences (12%) and diverged about 10.8 million years ago. Eupetaurus is significantly different from Petaurista and Pteromys. The level of estimated pairwise-sequence divergence observed between Eupetaurus and Petaurista or Pteromys is greater than that observed between Eupetaurus and Trogopterus, Belomys, Glaucomys, or Hylopetes. Considering the divergence time of the two Eupetaurus groups, the glaciations and the uplift of the Himalayas and Qinghai-Tibet plateau during the Pliocene-Pleistocene period might be the major factors affecting the present distribution of Eupetaurus along the Himalayas. (C) 2004 Elsevier Inc. All rights reserved.

Otolith microstructure of Oxygymnocypris stewartii (Cypriniformes, Cyprinidae, Schizothoracinae) in the Lhasa River in Tibet, China

Otolith microstructure of Oxygymnocypris stewartii collected from the Lhasa River was examined and described with regards to the early life history events. The monthly changes in the number of microincrements on the margin of the otolith were examined to validate the approximately daily periodicity of otolith increment formation. The microstructure of otoliths was used to detect changes in microincrement deposition patterns corresponding with events during early life. The annuli, microincrements and checks including the hatch check, yolk absorption check and several recurrent patterns in the otolith were described. Periodicity of the recurrent patterns was weekly, fortnightly and monthly. Through counting the number of the microincrements, it was confirmed that the primary growth of O. stewartii was in a period of 7 or 8 months from late March to October; it was estimated that O. stewartii might hatch between April and May.

Otolith characteristics and age determination of an endemic Ptychobarbus dipogon (Regan, 1905) (Cyprinidae: Schizothoracinae) in the Yarlung Tsangpo River, Tibet

We describe the microincrements, checks and annuli in the lapilli of the schizothoracine Ptychobarbus dipogon, an endemic species of the Tibetan plateau. We collected samples in the Yarlung Tsangpo River and its tributaries on a monthly basis (from April 2004 to August 2006). We describe the shape features of the three pairs of otoliths and document the full trajectory of lapillus development. We found that five to seven checks were clearly visible in the opaque zone of the first annulus. The pattern of 21-23 daily growth increments within each check might be explained as a lunar-induced deposition. We counted between 137 and 154 increments within the first annulus. Annuli appeared as a sequence of gradually declining increment widths, whereas false rings were characterized by abrupt checks. Our oldest estimates were 23(+)years for males and 44(+) for females. The time of annulus completion was clearly between March and April each year using monthly marginal increments analysis. We consider the factors responsible for daily increment formation as an endogenous circadian rhythm. Environmental information, such as strong sunlight and cold water temperatures in the Tibetan Plateau, could reinforce the endogenous daily cycle. Our results provided important data addressing the ecology and population dynamics of P. dipogon.

Phylogeography of Schizothorax o'connori (Cyprinidae: Schizothoracinae) in the Yarlung Tsangpo River, Tibet

Schizothorax o'connori is endemic to the Yarlung Tsangpo River on the Tibetan Plateau. We assessed the relative impacts of historical and contemporary factors in organizing genetic variation in S. o'connori populations using mitochondrial cytochrome b sequences. We analyzed 191 samples from 11 populations and identified 78 haplotypes. The phylogenetic analyses and analysis of molecular variance all supported the same conclusions of two well-differentiated east-west phylogroups, separated by the Tsangpo Great Gorge. The split between the two clades accounted for 58% of the genetic variance observed among the examined samples. Waterfalls as effective barriers played an important role in shaping the phylogeographical structure of this species. Analyses of migration rates revealed that upstream dispersal was limited crossing waterfalls. Our study revealed substantial spatial and temporal variation in the influence of landscape features on contemporary patterns of genetic structure in S. o'connori. Interglacial range expansions clearly left their mark on contemporary populations above the Tsangpo Great Gorge.