Biogeography and molecular phylogeny of the genus Schizothorax (Teleostei: Cyprinidae) in China inferred from cytochrome b sequences

Aim To test a vicariant speciation hypothesis derived from geological evidence of large‐scale changes in drainage patterns in the late Miocene that affected the drainages in the south‐eastern Tibetan Plateau. Location The Tibetan Plateau and adjacent areas. Methods The cytochrome b DNA sequences of 30 species of the genus Schizothorax from nine different river systems were analysed. These DNA sequences were analysed using parsimony, maximum likelihood and Bayesian methods. The approximately unbiased and Shimodaira–Hasegawa tests were applied to evaluate the statistical significance of the shortest trees relative to alternative hypotheses. Dates of divergences between lineages were estimated using the nonparametric rate smoothing method, and confidence intervals of dates were obtained by parametric bootstrapping. Results The phylogenetic relationships recovered from molecular data were inconsistent with traditional taxonomy, but apparently reflected geographical associations with rivers. Within the genus Schizothorax, we observed a divergence between the lineages from the Irrawaddy–Lhuit and Tsangpo–Parlung rivers, and tentatively dated this vicariant event back to the late Miocene (7.3–6.8 Ma). We also observed approximately simultaneous geographical splits within drainages of the south‐eastern Tibetan Plateau, the Irrawaddy, the Yangtze and the Mekong–Salween rivers in the late Miocene (7.1–6.2 Ma). Main conclusions Our molecular evidence tentatively highlights the importance of palaeoriver connections and the uplift of the Tibetan Plateau in understanding the evolution of the genus Schizothorax. Molecular estimates of divergence times allowed us to date these vicariant scenarios back to the late Miocene, which agrees with geological suggestions for the separation of these drainages caused by tectonic uplift in south‐eastern Tibet. Our results indicated the substantial role of vicariant‐based speciation in shaping the current distribution pattern of the genus Schizothorax.     

Leaf and infructescence fossils of Alnus (Betulaceae) from the late Eocene of the southeastern Qinghai–Tibetan Plateau

Plant fossils from the Qinghai–Tibetan Plateau (QTP), China are critical to understand not only the diversification history of plants there, but also the paleoenvironmental conditions. Alnus are deciduous trees, mainly distributed in temperate and subtropical regions of Eurasia and North America, and they are well known in the fossil records throughout the Cenozoic in the Northern Hemisphere. We collected numerous well‐preserved Alnus leaf and infructescence fossils from the Lawula Formation (～34.6 Ma with ⁴⁰Ar/³⁹Ar dating) at the present elevation of 3910 m a.s.l. in the southeastern QTP. Based on detailed morphological comparisons with existing and fossil species, these fossils show closest affinity to Alnus ferdinandi‐coburgii C. K. Schneid., and we refer to these fossils as A. cf. ferdinandi‐coburgii. These specimens comprise the oldest megafossil record of Alnus in the QTP, and provide solid evidence for the distribution of Alnus there as early as the late Eocene. Extant A. ferdinandi‐coburgii is distributed in areas with mean annual temperature values between 9.7 °C and 16.9 °C, and mean annual precipitation values ranging from 896.2 mm to 1161.2 mm; therefore, fossils of A. cf. ferdinandi‐coburgii suggest a much warmer and wetter climate during the late Eocene than today in the southeastern QTP. This finding is consistent with other evidence for continued uplift of the southeastern QTP after the late Eocene that might be due to the eastward extension of the QTP.     

云南剑川盆地上始新统双河组双壳类新材料及其地质意义*

青藏高原东南缘一直被认为是高原隆升造成物质逃逸的主要场所,很好地记录了反映高原隆升的沉积、构造和生物演化事件。剑川盆地是青藏高原东南缘的一个拉分盆地。作为剑川盆地最主要的沉积地层,双河组内仅有少量双壳类化石的报道,缺少详细的化石对比研究。本文系统描述双河组最新发现的双壳类共4属9种,分别为Unio,Acuticosta,Cuneopsis和Anodonta 4属,Acuticosta huoerguosica,Acuticosta sp.,Acuticosta subimmutata,Acuticostacf.chinensis,Acuticostacf.cuspidata,Acuticostacf.semiretiaria,Unio cf.subparallelus,Cuneopsis cf.subceltiformis和Anodonta cf.subpiscinalis 9种,均属于古异齿目珠蚌科。其中Acuticosta和Cuneopsis为现代东亚的特征属,Acuticosta chinensis和Cuneopsis subceltiformis为长江以南的现生种,而Acuticosta chinensis以往只在新疆有过化石记录。结合双河组的侵入脉体的年龄和介形虫标准化石的研究确定双河组的时代为晚始新世,故本次研究不仅为相关双壳类的产出时代可以延伸至晚始新世提供了有力的证据和新材料,而且反映出剑川盆地双河组在晚始新世为亚热带的淡水三角洲前缘-前三角洲的沉积环境。     

The Evolution and Paleobiogeography of Flying Squirrels (Sciuridae, Pteromyini) in Response to Global Environmental Change

Flying squirrels are strictly arboreal squirrels adopting a special gliding form of locomotion. This group of animals has a long history that has mirrored the vicissitude of forests. The discrepancy in the distribution between fossils and extant species indicates a mysterious evolution history requiring further exploration. This study compiles the worldwide fossils of Pteromyini to the species level in order to reproduce the spatiotemporal distribution pattern of flying squirrels and deduce the ancestral distribution according to dispersal-vicariance analysis of a phylogeny of the extant species. In addition, we reconstruct the paleoenvironmental background and find that flying squirrels probably originated in the Oligocene–Miocene transition from Europe and immediately dispersed to Asia and North America. Influenced by glaciation, CO₂ reduction, geologic movements and the Paratethys retreat, the Northern Hemisphere underwent climate deterioration and grassland expansion during the late Miocene, and thus the diversity of Pteromyini dramatically decreased. The uplift of the Tibet Plateau in addition to the strengthened Asian monsoons intensified the aridity in central Asia, but brought sufficient water to the densely forested regions of South and Southeast Asia. These forests are likely both refugia and diversification center for flying squirrels during glacial periods in the Quarternary. The subsequent connection and separation events among these heterogeneous habitats has probably been a driving force in the speciation of flying squirrels. Based on this work, we predict a bleak future for the flying squirrels, one which is closely associated with the fate of forests in Asia.     

Fruit fossils of Rosoideae (Rosaceae) from the late Pliocene of northwestern Yunnan,Southwest China

The subfamily Rosoideae Focke (Rosaceae) has a good fossil record in the Northern Hemisphere, but these fossil records are confined mainly to a few genera, whereas the majority, in particular those with herbaceous members, are still under‐represented. In this study, we describe new fruit fossils of Rosoideae, including Fragaria achenes and Rubus endocarps, from the late Pliocene of northwestern Yunnan, Southwest China. These fossils add new accounts to the fossil archive of Rosoideae and provide the first fossil record of Fragaria in East Asia. The new fossil findings provide a historical backdrop for the modern diversity and distribution of the subfamily in northwestern Yunnan, a topographically complex area accommodating a high diversity for many plant groups. Our Rubus fossils, in combination with other nearby coeval occurrences of the genus, suggest that Rubus was already establishing its modern diversity in northwestern Yunnan during the late Pliocene. This finding enriches our knowledge of the post‐Neogene diversification of flowering plants in northwestern Yunnan, which is thought to be largely driven by dramatic mountain uplifts and environmental complications associated with the southeastern extension of the Tibetan Plateau.     

Fossil fruits of Illigera (Hernandiaceae) from the Eocene of central Tibetan Plateau

Illigera (Hernandiaceae) is a liana genus distributed mainly in the tropical Asia and Africa. Previous fossil records suggested that Illigera was restricted in western North America during the Eocene. Recent paleobotanical investigation has unveiled a Paleogene flora that is totally different from today's vegetation in central Tibet. This provides novel insights into the paleoenvironmental change during the evolution of the Tibetan Plateau (TP). Here, we investigated 10 fruit impressions of Illigera from the early middle Eocene Niubao Formation in the Bangor Basin, central TP. The fossil winged fruits are characterized by their eroded fruit wings, well preserved fusiform locular areas with a median ridge bisecting the fruit, and short veins fanning radially outward. These features allow us to assign these fossils to Illigera eocenica, a species originally discovered in the Eocene of western North America. This is the second fossil occurrence of Illigera worldwide, and the first in Asia. Our finding suggests a warm and humid climate in the central TP during the early middle Eocene, and a close floristic link between Asia and North America during the Paleogene. We also propose a Northern Hemisphere origin and a Paleogene dispersal event from Northern Hemisphere to Africa for Illigera.     

Amber fossils reveal the Early Cenozoic dipterocarp rainforest in central Tibet

The palaeoenvironmental reconstruction of central Tibet is key to understanding the uplift history of the Tibetan Plateau, which had a profound influence on Cenozoic global climate and biotic change. Here we report an amber layer from the lower part of the Dingqing Formation (late Oligocene) in Lunpola of central Tibet, which is the first record of amber from Tibet. Herein we find that Lunpola amber is derived from dipterocarp trees, as determined by gas chromatography-mass spectrometry, which are restricted to and dominant in Asian rainforest nowadays. This amber forest represents the northernmost dipterocarp forest and is consistent with the hypothesis of out-of-India dispersal of Asian dipterocarps. The Lunpola amber most probably was derived from the lower part of the Niubao Formation (early–middle Eocene) and suggests a tropical/subtropical wet forest was present in central Tibet at least before the late Oligocene (probably early–middle Eocene).     

A unique record of Cercis from the late early Miocene of interior Asia and its significance for paleoenvironments and paleophytogeography

The climatic impacts of the Tibetan Plateau since the Neogene and the phytogeographic pattern changes of formerly widely-distributed forest communities on the plateau remain poorly constrained. Today, Cercis L. (Fabaceae) is a well-known arborescent genus typically distributed in subtropical to warm temperate zones of the Northern Hemisphere, and Paleogene fossil occurrences from Eurasia and North America show a long history of the genus in mid-low latitudes of the Northern Hemisphere. Here, we describe a fossil species, Cercis zekuensis sp. nov. based on well-preserved fruits from the early Miocene of the northeastern Tibetan Plateau. Detailed morphological comparison (e.g., ventral margin with a veinless wing) of extant and fossil members of Cercis and other genera confirmed validity of the present taxonomic identity. Based on the comparison with extant relatives and their climate preferences, this unexpected occurrence of thermophilic Cercis in northeastern Tibetan Plateau indicates this area had higher temperature and precipitation in the Miocene than today. Integrated with inferred (paleo-)temperature lapse rates, this indicates a low paleoelevation of less than 2.4 km. In contrast with the present-day alpine climate here (~3.7 km), such a low elevation facilitated a more favorable habitat with comparatively high biodiversity and warm temperate forests at that time, as were evidenced by co-occurring megafossils. Moreover, the present existence of Cercis implies the genus was widespread in interior Asia during the early Neogene and shows its modern disjunction or diversification between eastern and central Asia was possibly shaped by the late Cenozoic regional tectonic uplift and consequential environmental deterioration.     

Cyprininae phylogeny revealed independent origins of the Tibetan Plateau endemic polyploid cyprinids and their diversifications related to the Neogene uplift of the plateau

Origin and diversification of the Tibetan polyploid cyprinids (schizothoracins) may help us to explore relationships between diversification of the cyprinids and the Tibetan Plateau uplift. Cyprininae phylogeny was analyzed using mitochondrial and nuclear DNA sequences to trace origins of polyploidy and diversifications of schizothoracins. Ancestral states reconstruction for ploidy levels indicated that the Cyprininae was diploid origin and the schizothoracin clades tetraploid origins. There were two diversification rate shifts along with diversification of the cyprinine fishes in response to the Tibetan uplift. The unusual diversification shifts were located to branches subtending the clades of Tibetan polyploid cyprinids. Our analyses suggested that (i) phylogeny of Cyprininae recovered two independent origins of the Tibetan polyploidy schizothoracins; (ii) diversifications of the schizothoracins were closely related to the Neogene uplift of the Tibetan plateau in the following ways: the relatively ancient Late Oligocene-Middle Miocene adaptive radiation may be associated with the uplift of the southern Tibet and Himalaya; the Middle Miocene-Early Pleistocene lineage-specific diversification broadly coincident with major phase of the Neogene Tibetan uplift; and the most recent Pleistocene diversification shift in Schizothorax closely coincident with the successive Kunlun-Huanghe and Gonghe movements of the Tibetan uplift and the glaciation-induced climate oscillations on the plateau.     

Age of the latest marine sedimentation in the western Kunlun area constrained by planktic foraminifera

Surface uplift of the Tibetan Plateau in the Cenozoic has dramatically affected evolutions of climate, environment, and biota in East Asia. The onset time of plateau uplift can be roughly constrained by the termination of the latest marine sedimentation, which shows large spatial differences in the Tibetan Plateau. In the northwest Tibetan Plateau of the western Kunlun area, the Tielongtan Group represents the latest marine sedimentation. However, the age of its depositional cessation is still unknown. For the first time, we conduct taxonomic, paleoecologic and paleogeographic studies of planktic foraminifera in the limestones of the Tielongtan Group. Despite poor preservation and low abundance, ten planktic foraminiferal species are identified, including Globotruncana falsostuarti, G. cf. linneiana, G. cf. ventricosa, Planohedbergella prairiehillensis, P. cf. yaucoensis, Pseudotextularia nuttalli, Planoheterohelix globulosa, P. cf. praenuttalli, Muricohedbergella holmdelensis and Whiteinella brittnensis. The foraminiferal assemblages suggest an age of late Campanian to Maastrichtian, indicating a shallow, neritic marine environment. Combining available research data from other regions, we show that these planktic foraminifera were widely distributed in the Tarim basin, southern Tibet, and the Mediterranean area during the latest Cretaceous. Our result of planktic foraminifera in the Tielongtan Group implies that the surface uplift of the western Kunlun area did not begin until after ∼76–66 Ma.     

Cave Stedocys spitting spiders illuminate the history of the Himalayas and southeast Asia

Stedocys spitting spiders (Araneae: Scytodidae) inhabit subterranean environments and have poor dispersal abilities. The Cenozoic Indian–Eurasian collision affected the regional biota of this genus, which occurs in parts of Indochina. Phylogeographical pattern of Stedocys based on multigene DNA sequence datasets reveals how tectonic history drove four biological splits. The first split dates to the late Paleocene–Eocene and involves the Truong Son Mountain Range and Mekong River. The other splits associate with the Eocene–Oligocene transition, including the Tonkin (Beibu) Gulf, the Ma River, and the Red River. These events indicate four early uplifts of the Himalayas and Tibetan Plateau. Our results cannot reject the hypothesis that uplifting of the Himalayas and Tibetan Plateau region due to crustal thickening and the lateral extrusion of Indochina occurred synchronously during the Paleocene–Oligocene transition in reaction to the Indian–Eurasian collision. Species of Stedocys cluster into groups I and II. Their evolution involves one dispersal and four vicariance events, which formed the following five Indochinese clades: Hainan clade (I‐1); western Yunnan and central Laos clade (I‐2); central Vietnam clade (I‐3); northern Vietnam and southwestern China clade (I‐4); and Thailand clade (II‐1). The lateral extrusion of Indochina is the driver of these events. The drifting of Hainan Island to its present location owes to its southeastern movement from continental Vietnam and Guangxi, China around the Eocene–Oligocene boundary. This biogeographical pattern highlights the significant role geography plays in shaping evolutionary history in southeastern Asia. It also illuminates how the timing of geological events drives the distributions of species.     

A survey of the geographic distribution of Ophiocordyceps sinensis

Ophiocordyceps sinensis is one of the best known fungi in Traditional Chinese Medicine. Many efforts have been devoted to locating the production areas of this species resulting in various reports; however, its geographic distribution remains incompletely understood. Distribution of O. sinensis at the county level is clarified in this work based on both a literature search and fieldwork. More than 3600 publications related to O. sinensis were investigated, including scientific papers, books, and online information. Herbarium specimens of O. sinensis and field collections made by this research group during the years 2000-2010 were examined to verify the distribution sites. A total of 203 localities for O. sinensis have been found, of which 106 are considered as confirmed distribution sites, 65 as possible distribution sites, 29 as excluded distribution sites and three as suspicious distribution sites. The results show that O. sinensis is confined to the Tibetan Plateau and its surrounding regions, including Tibet, Gansu, Qinghai, Sichuan, and Yunnan provinces in China and in certain areas of the southern flank of the Himalayas, in the countries of Bhutan, India and Nepal, with 3,000 m as the lowest altitude for the distribution. The fungus is distributed from the southernmost site in Yulong Naxi Autonomous County in northwestern Yunnan Province to the northernmost site in the Qilian Mountains in Qilian County, Qinghai Province, and from the east edge of the Tibetan Plateau in Wudu County, Gansu Province to the westernmost site in Uttarakhand, India. The clarification of the geographic distribution of O. sinensis will lay the foundation for conservation and sustainable use of the species.     

Areas of endemism and patterns of diversity for aphids of the Qinghai-Tibetan Plateau and the Himalayas

Aim  The study aimed to identify areas of endemism for aphids in the Qinghai-Tibetan Plateau and the Himalayas (QTPH), and to test congruence between patterns of endemism and patterns of overall species richness identified in a previous study.
Location  The QTPH.
Methods  A distribution data base of 326 endemic aphids in the QTPH was compiled. The study area was divided into a grid of 2°× 2° operative geographical units. Parsimony analysis of endemicity (PAE) was used to identify areas of endemism, and the diversity patterns of endemic species were then mapped using GIS.
Results  We identified 326 endemic species belonging to 138 genera within Adelgidae and 14 subfamilies of Aphididae. Five areas of endemism were found using PAE analysis: the eastern Himalayas, the western Himalayas, north-western Yunnan, southern Tibet and the eastern QTPH. Maps of patterns of endemism identified four major centres for endemic aphids, namely the western Himalayas, the eastern Himalayas (or Sikkim-Assam Himalayas), north-western Hengduan Mountains and the mountains of southern Gansu Province, and three minor centres, southern Tibet, south-eastern Tibet and the eastern Qinghai Province in the north-eastern QTPH.
Main conclusions  Our study identifies major centres of aphid endemism. Furthermore, there is a noticeable congruence between patterns of endemism and patterns of species richness. The patterns of endemism were most likely influenced by the recent uplift of the QTPH.     

青南、藏北中侏罗世缅甸贝内部构造的研究及修订

通过缅甸贝内部构造及解剖学特征的研究和对比,对该属进行了修订和整理.发现过去置于该属中的90余种实际上包括了现在理解的9个以上属的内容.经与相近属内部和外部特征的比较,对缅甸贝的起源和演化做了初步研究.提出该属在晚 Bajocian 期起源于 Formosarhynchia,早 Bathonian 期得到了爆发性的发展和辐射,并在中、晚 Bathonian 期朝3个方向演化.通过对该属时空分布、共生生物、群落结构、伴生沉积岩及介壳稳定同位素和微量元素的综合分析,认为 Burmirhynchia-Holcothyris 群落主要生活于近岸浅水、含盐度偏低、水深可能小于 30m 的低能环境.     

Mitochondrial phylogeography and genetic diversity of Tibetan gazelle (Procapra picticaudata): implications for conservation

The Tibetan gazelle (Procapra picticaudata) is a threatened species and distributed on the Qinghai-Tibet Plateau of China (Qinghai Province, Tibet Autonomous Region and the adjacent Gansu Province, Sichuan Province, and Xinjiang Uigur Autonomous Region). Small peripheral populations of Tibetan gazelle were once found in northern Sikkim and Ladakh, but now these are close to extinction. To describe the evolutionary history and to assess the genetic diversity within this monotypic species and population structure among different geographic locations in China, we sequenced mitochondrial DNA from the control region (CR) and cytochrome (cyt) b gene for 46 individuals from 12 geographic localities in Qinghai, Tibet, Xinjiang, Gansu, and Sichuan. A total of 25 CR haplotypes and 16 cyt b haplotypes were identified from these gazelle samples. CR haplotype diversity (0.98+/-0.01) and nucleotide diversity (0.08+/-0.009) were both high. Phylogenetic trees indicate that the Tibetan gazelle in China can be divided into three main clades: Tibet, Sichuan (SCH) and Qinghai-Arjin Shan-Kekexili (QH-ARJ-KKXL). Analysis of molecular variance (AMOVA) and network analysis consistently support this geographic structure in both datasets. Significant differentiation between populations argues for the presence of management units (MUs). Such differentiation may reflect a geographic separation resulting from the uplift of the Qinghai-Tibet Plateau during the Late Pliocene and Pleistocene. Mismatch distribution analysis implies that Tibetan gazelle has undergone complex population changes. We suggest that the present population structure has resulted from habitat fragmentation during the recent glacial period on the Qinghai-Tibet Plateau and population expansion from glacial refugia after the glacial period. It is likely that the present populations of Tibetan gazelle exhibit a pattern reminiscent of several bottlenecks and expansions in the recent past.     

Molecular phylogenetic relationship of snow finch complex (genera Montifringilla, Pyrgilauda, and Onychostruthus) from the Tibetan plateau

The snow finch complex (Montifringilla, Pyrgilauda, and Onychostruthus) has its center of distribution on the Tibetan plateau, with six out of seven species in the genera occurring there. Phylogenetic relationships among these six species of three genera have been studied based on DNA sequence data obtained from the mitochondrial cytochrome b gene and the nuclear myoglobin gene. The results support monophyly of the snow finch complex group and three major evolutionary lineages are recognized. The first clade consists of ruficollis, blanfordi, and davidiana. These three taxa are sometimes placed in their own genus, Pyrgilauda, and the DNA data supports this. The three taxa nivalis, henrici, and adamsi have traditionally been placed in the genus Montifringilla, and they group together strongly in the present analysis. The results further suggest that nivalis and adamsi are more closely related to each other than are nivalis and henrici, despite that the latter two are often regarded as conspecific. The third distinct lineage within the snow finch complex consists of taczanowskii, which has been placed its own genus, Onychostruthus. This taxon has a basal position in the phylogenetic tree and is sister to all other snow finches. We estimated that taczanowskii split from the other taxa between 2 and 2.5 mya, i.e., about the time for the most recent uplift of the Tibetan plateau, "the Tibet movement", 3.6-1.7 mya. Cladogenesis within the Montifringilla and Pyrgilauda clades seems to be contemporary with the second phase of "Tibet movement" at 2.5 mya and the third phase at 1.7 mya and "Kunhuang movement" in 1.5-0.6 mya. The dramatic climatic and ecological changes following from the uplift of the Tibetan plateau, together with the cyclic contraction and expansion of suitable habitats during the Pleistocene, are probably the most important factors for the cladogenesis in snow finch complex.     

Geographical Distribution of Testate Amoebae in Tibet and Northwestern Yunnan and their Relationships with Climate

In Asia, especially in China, our knowledge of the distribution of testate amoebae is still limited. In this paper, the geographical distribution of testate amoebae in Tibetan Plateau and northwestern Yunnan Plateau, southwest China and their relationships with the climatic factors have been studied. We found testate amoebae shifted in the most dominant species and increased in species (or genus) richness from northwest to southeast. Further, the linear regression analyses revealed that both species richness and genus richness have higher positive correlations with the mean temperature of the warmest month and annual mean precipitation as contrasted with the mean altitude, which showed weak negative correlation. This indicates that the temperature and precipitation are more significant influences on the richness than the altitude. The cluster analysis based on the community structure, defined by Sørenson’s coefficient matrix, suggested four groups from the 10 physiographical regions. This geographical distribution pattern was also closely related with the climatic regionalization. The present climatic regionalization pattern of the study area originated from the uplift of Tibetan Plateau and mainly occurred in or after the late Pleistocene. Therefore, the geographical distribution of testate amoebae in our study area may have experienced complicated and drastic changes corresponding to the variation of the climate caused by the geological events.     

Into Tibet: An Early Pliocene Dispersal of Fossil Zokor (Rodentia: Spalacidae) from Mongolian Plateau to the Hinterland of Tibetan Plateau

This paper reports the fossil zokors (Myospalacinae) collected from the lower Pliocene (~4.4 Ma) of Zanda Basin, southwestern Tibet, which is the first record in the hinterland of Tibetan Plateau within the Himalayan Range. Materials include 29 isolated molars belonging to Prosiphneus eriksoni (Schlosser, 1924) by having characters including large size, highly fused roots, upper molars of orthomegodont type, m1 anterior cap small and centrally located, and first pair of m1 reentrants on opposing sides, high crowns, and high value of dentine tract parameters. Based on the cladistics analysis, all seven species of Prosiphneus and P. eriksoni of Zanda form a monophyletic clade. P. eriksoni from Zanda, on the other hand, is nearly the terminal taxon of this clade. The appearance of P. eriksoni in Zanda represents a significant dispersal in the early Pliocene from its center of origin in north China and Mongolian Plateau, possibly via the Hol Xil-Qiangtang hinterland in northern Tibet. The fast evolving zokors are highly adapted to open terrains at a time when regional climates had become increasingly drier in the desert zones north of Tibetan Plateau during the late Miocene to Pliocene. The occurrence of this zokor in Tibet thus suggests a rather open steppe environment. Based on fossils of large mammals, we have formulated an “out of Tibet” hypothesis that suggests earlier and more primitive large mammals from the Pliocene of Tibet giving rise to the Ice Age megafauna. However, fossil records for large mammals are still too poor to evaluate whether they have evolved from lineages endemic to the Tibetan Plateau or were immigrants from outside. The superior record of small mammals is in a better position to address this question. With relatively dense age intervals and numerous localities in much of northern Asia, fossil zokors provide the first example of an “into Tibet” scenario–earlier and more primitive taxa originated from outside of the Tibetan Plateau and the later the lineage became extinct in southwestern Tibet.     

藏族的高原适应——西藏藏族生物人类学研究回顾

藏族生活在具有世界屋脊之称的青藏高原, 特殊的生态环境和特殊的文化背景造就了藏族特殊的适应高原缺氧机制, 引起了国内外学者的广泛关注和浓厚的研究兴趣。本文根据国内外数据库的文献并结合我们的研究工作, 从高原适应的角度回顾了30多年藏族人类学研究。回顾显示, 藏族由于长期生活在高原缺氧的环境中, 不仅形态和机能发生了适应性变化, 而且体成分也表现出相应的变化, 体现了形态、机能和体成分的统一。这些变化是长期进化形成的, 与安第斯山人等有明显不同, 就是在同一高原生活的西藏、青海、四川、甘肃和云南的藏族乃至尼泊尔和印度藏族的体质也表现出地域差异, 这些差异的产生是多种因素所致, 两个关键性的基因是导致两大高原人口高原适应机制不同的最主要的原因。     

First fossil record of Cedrelospermum (Ulmaceae) from the Qinghai–Tibetan Plateau: Implications for morphological evolution and biogeography

Cedrelospermum Saporta is an extinct genus in the Ulmaceae with abundant fossil records in North America and Europe. However, so far, fossil records of this genus from Asia are sparse, which limits the interpretations of the morphological evolution and biogeographical history of the genus. Here we report well‐preserved fruits (Cedrelospermum tibeticum sp. nov.) and a leaf (Cedrelospermum sp.) of Cedrelospermum from the upper Oligocene Lunpola and Nyima basins in the Qinghai–Tibetan Plateau (QTP). This is the first fossil record of Cedrelospermum in the QTP, showing that this genus grew in this region during the late Oligocene. Cedrelospermum tibeticum fruits are double‐winged, morphologically similar to the Eocene and Oligocene double‐winged Cedrelospermum species from North America. This supports the hypothesis that Cedrelospermum migrated to Asia from North America by way of the Bering Land Bridge. Given that Cedrelospermum was a typical element of Northern Hemispheric flora in the Paleogene and Neogene, the presence of this genus indicates that the central region of the QTP was phytogeographically linked with other parts of the Northern Hemisphere during the late Oligocene. The morphological observations of C. tibeticum fruits and other double‐winged Cedrelospermum fruits suggest an evolutionary trend from obtuse to acute apex for the primary wing. Cedrelospermum tibeticum likely had warm and wet climatic requirements. This type of an environment possibly existed in the central QTP in the late Oligocene, thereby supporting the survival of C. tibeticum.