新生代全球变化与中国古植物区系的演变

新生代是地质历史最年轻的一个时代 ,当时全球板块运动和岩浆活动强烈而频繁 ,中生代开始的大陆漂移活动继续进行 ,并于早第三纪末开始喜马拉雅运动 ,古地中海逐渐封闭 ,经过一系列的板块碰撞和洋底扩张 ,晚第三纪时地壳构造的基本轮廓和古地理面貌逐渐接近现代。受地壳运动的控制 ,全球新生代气候波动明显 ,早第三纪总体比较温暖 ,晚第三纪气候发生明显分异 ,气温显著下降 ,第三纪末期开始有冰川活动 ,一直持续到更新世末期 ,期间发生多次冰期和间冰期 ,全新世气候逐渐转暖。受上述全球变化的影响 ,我国新生代植物区系纵向演变和横向分异明显 ,早第三纪植物区系主要是纬向分带 ,从古新世到渐新世 ,我国植物区系主要分为北方区和南方区 ;晚第三纪初 ,由于喜马拉雅运动使青藏高原及东部的横断山脉地区持续抬升 ,逐渐形成独立的高原植物区。总体来说 ,晚第三纪植物区系和植被类型逐渐接近现代。第四纪由于青藏高原继续强烈抬升 ,激发了东亚季风 ,使我国的植物区系经向分带更加明显 ,植物区系和植被类型与现代已基本一致 ,到了全新世晚期 ,植物区系的演变除了受全球变化的影响外 ,人类活动的作用也越来越明显     

From ‘third pole’ to north pole: a Himalayan origin for the arctic fox

The ‘third pole’ of the world is a fitting metaphor for the Himalayan–Tibetan Plateau, in allusion to its vast frozen terrain, rivalling the Arctic and Antarctic, at high altitude but low latitude. Living Tibetan and arctic mammals share adaptations to freezing temperatures such as long and thick winter fur in arctic muskox and Tibetan yak, and for carnivorans, a more predatory niche. Here, we report, to our knowledge, the first evolutionary link between an Early Pliocene (3.60–5.08 Myr ago) fox, Vulpes qiuzhudingi new species, from the Himalaya (Zanda Basin) and Kunlun Mountain (Kunlun Pass Basin) and the modern arctic fox Vulpes lagopus in the polar region. A highly hypercarnivorous dentition of the new fox bears a striking resemblance to that of V. lagopus and substantially predates the previous oldest records of the arctic fox by 3–4 Myr. The low latitude, high-altitude Tibetan Plateau is separated from the nearest modern arctic fox geographical range by at least 2000 km. The apparent connection between an ancestral high-elevation species and its modern polar descendant is consistent with our ‘Out-of-Tibet’ hypothesis postulating that high-altitude Tibet was a training ground for cold-environment adaptations well before the start of the Ice Age.     

Phylogeny of spiny frogs Nanorana (Anura: Dicroglossidae) supports a Tibetan origin of a Himalayan species group

Recent advances in the understanding of the evolution of the Asian continent challenge the long‐held belief of a faunal immigration into the Himalaya. Spiny frogs of the genus Nanorana are a characteristic faunal group of the Himalaya–Tibet orogen (HTO). We examine the phylogeny of these frogs to explore alternative biogeographic scenarios for their origin in the Greater Himalaya, namely, immigration, South Tibetan origin, strict vicariance. We sequenced 150 Nanorana samples from 62 localities for three mitochondrial (1,524 bp) and three nuclear markers (2,043 bp) and complemented the data with sequence data available from GenBank. We reconstructed a gene tree, phylogenetic networks, and ancestral areas. Based on the nuDNA, we also generated a time‐calibrated species tree. The results revealed two major clades (Nanorana and Quasipaa), which originated in the Lower Miocene from eastern China and subsequently spread into the HTO (Nanorana). Five well‐supported subclades are found within Nanorana: from the East, Central, and Northwest Himalaya, the Tibetan Plateau, and the southeastern Plateau margin. The latter subclade represents the most basal group (subgenus Chaparana), the Plateau group (Nanorana) represents the sister clade to all species of the Greater Himalaya (Paa). We found no evidence for an east–west range expansion of Paa along the Himalaya, nor clear support for a strict vicariance model. Diversification in each of the three Himalayan subclades has probably occurred in distinct areas. Specimens from the NW Himalaya are placed basally relative to the highly diverse Central Himalayan group, while the lineage from the Tibetan Plateau is placed within a more terminal clade. Our data indicate a Tibetan origin of Himalayan Nanorana and support a previous hypothesis, which implies that a significant part of the Himalayan biodiversity results from primary diversification of the species groups in South Tibet before this part of the HTO was uplifted to its recent heights.     

Ephemeral Pleistocene woodlands connect the dots for highland rattlesnakes of the Crotalus intermedius group

Aim To test how Pleistocene climatic changes affected diversification of the Crotalus intermedius species complex. Location Highlands of Mexico and the south‐western United States (Arizona). Methods We synthesize the matrilineal genealogy based on 2406 base pairs of mitochondrial DNA sequences, fossil‐calibrated molecular dating, reconstruction of ancestral geographic ranges, and climate‐based modelling of species distributions to evaluate the history of female dispersion. Results The presently fragmented distribution of the C. intermedius group is the result of both Neogene vicariance and Pleistocene pine–oak habitat fragmentation. Most lineages appear to have a Quaternary origin. The Sierra Madre del Sur and northern Sierra Madre Oriental are likely to have been colonized during this time. Species distribution models for the Last Glacial Maximum predict expansions of suitable habitat for taxa in the southern Sierra Madre Occidental and northern Sierra Madre Oriental. Main conclusions Lineage diversification in the C. intermedius group is a consequence of Pleistocene climate cycling. Distribution models for two sister taxa in the northern and southern Sierra Madre Occidental and northern Sierra Madre Oriental during the Last Glacial Maximum provide evidence for the expansion of pine–oak habitat across the Central Mexican Plateau. Downward displacement and subsequent expansions of highland vegetation across Mexico during cooler glacial cycles may have allowed dispersal between highlands, which resulted in contact between previously isolated taxa and the colonization of new habitats.     

榆科榉属的植物地理学

榉属是东亚—地中海间断分布局,分属于东亚植物区、地中海植物区及伊朗—土兰植物区。根据现存榉属植物的形态演化趋势、细胞学、孢粉学证据及其化石资料和古地质历史事件相结合分析,该属植物是第三纪温带森林的孑遗成分而且第三纪末期的喜马拉雅造山运动及第四纪的冰川作用是样属植物演化的主要动力。在欧洲,由于喜马拉雅造山运动及第四纪冰川作用的较强烈,使分布于此的榉属植物受到严厉的生存竞争,未能适应环境的物种遭受灭绝,少数进行适应性变异的物种在极端环境条件下生存了下来。而东亚地区由于自然环境相对较稳定,便成了榉属植物的避难所,因此也保留了榉属植物种较为古老的种类。     

Diversity and distribution of aphids in the Qinghai–Tibetan Plateau–Himalayas

Abstract.  1. Based on the geographic distribution database of museum specimens and published literature, the diversity and distribution of aphids in the Qinghai–Tibetan Plateau–Himalayas was studied using the GIS method.
2. A total of 646 aphid species/subspecies belonging to 195 genera in Adelgidae, Phylloxeridae, and 16 subfamilies of Aphididae were recorded from this region; among which Aphidinae shared the highest species proportion (343 species, 53.1%), followed by Greenidinae (63, 9.7%), Eriosomatinae (54, 8.3%), and Lachninae (48, 7.4%). A total of 326 species and 20 genera are endemic to this region.
3. The aphid fauna has diverse faunal components and shows affinity with other zoogeographic realms. It is also rich in alpine species.
4. The aphid fauna exhibits an asymmetric distribution pattern with few species found on the high surface of the plateau. Four distribution centres were mapped. These were (i) the associated mountainous areas of the Qinghai–Tibetan Plateau and the western Qinling Mountains and southern Gansu Province, (ii) the Hengduan Mountains Region, (iii) the eastern Himalayas, and (iv) the western Himalayas.
5. The aphid fauna of the region may have multiple origins, and has been influenced by geological and ecological factors, such as glaciers and diversified vegetation. Possible reasons for the species diversity and distribution patterns are discussed. The implications and priorities for conservation based, on these distribution patterns, are also discussed.     

藏北尼阿木底遗址发现的似阿舍利石器——兼论晚更新世人类向青藏高原的扩张

2013年夏,在藏北那曲地区申扎县羌塘高原旧石器调查中,从规模恢弘的尼阿木底遗址地表暴露的、数以万计的石制品中,分别采集到似阿舍利类型"手斧"9件和"薄刃斧"2件。本文对尼阿木底遗址所见的这类器物进行了细致的观察与分析,认为它们只是旧石器时代晚期勒瓦娄哇技术产品的石核,或偶尔为之的、与手斧和薄刃斧形似的石制品,并非真正意义上的手斧与薄刃斧。尼阿木底遗址不存在从选料、剥片、加工出成品,到使用、损坏和废弃等各个阶段的、明确的阿舍利类型石器工业产品生产体系的工艺链条,换言之,在尼阿木底生活的古人脑海中,并不存在一个加工阿舍利类型手斧和薄刃斧等工具的"概念模板"(Mental template)。联系到青藏高原其他地区以前所报道的同类材料,我们认为在青藏高原腹地目前所见的旧石器遗址中,还没有真正意义上的阿舍利石器工业类型的遗存。晚更新世末期时,来自于印巴次大陆方向的早期占领者,沿着喜马拉雅山脉、冈底斯山脉和昆仑山脉三条东西走向的巨大山系之间的通道,自高原西南方向开始向高原腹地扩张,其间阿舍利石器工业技术在非洲和欧亚大陆早已消弭,当时的青藏高原腹地并不具备重新产生该类石器工业技术的土壤。     

江西省鄱阳湖自然保护区昆虫区系初探

据对有较详细分布资料的220种昆虫的分析,鄱阳湖自然保护区昆虫属东洋界范畴,但已接近古北界南缘;昆虫种类形成了古北,东洋两大区系成分交叉重叠现象．亦是两大区系交叉过渡地带。在我国东部地区,与鄱阳湖共有种的分布规律为随着纬度的增高而递减。本文论述了鄱阳湖33种昆虫在我国东半部分布的南北限。     

横断山区高山绣线菊的谱系地理学研究

横断山区作为青藏高原东南部主要的一个冰期避难所,第四纪冰期气候的变化对该地区的植物地理分布和居群遗传结构都产生了重要的影响。为了揭示该地区物种分布的分子系统地理学结构,选取在该地区广泛分布的一种高山灌木-高山绣线菊的叶绿体trnL-trnF序列进行研究。采集了15个居群182个个体进行测序,共发现7个单倍型。总的遗传多样性较高(HT=0.809),但居群内遗传多样性较低(HS=0.236)。分子变异分析(AMOVA)结果表明分布区内高山绣线菊的遗传变异主要存在于居群间(84.48%),且居群间的遗传分化很高(GST=0.708,FST=0.84476,NST=0.863),有着显著的谱系地理学结构(NST>GST,P<0.01)和较低的居群间平均基因流(Nm=0.09)。单倍型的系统进化树和进化分支网络分析得到了相似的拓扑结构,7种单倍型都按照地理分布聚为三支:横断山区西部、横断山区东部以及两者的交接地带。本研究推测该物种在横断山区存在多个冰期避难所,而没有表现出大规模的种群集体扩张和迁移的现象。青藏高原隆升、第四纪气候的反复波动以及横断山区特殊的地理环境使得原来连续的居群片段化,并发生范围扩张,从而塑造了高山绣线菊的现代生物地理分布格局。     

青藏高原动物地理区的地位和东部界线问题

本文从历史时空的角度分析了青藏高原地区发生的鱼类区系的变化过程,和高原鱼类区系演化的相对独立性,认为这一鱼类区系的分化直接反映了高原隆升事件,并应当在动物地理区划上得到客观反映,即将青藏高原作为一个独立的区划单元。由于青藏高原的隆升和造成古北区、东洋区分化的第三纪末、第四纪初的全球性气候变冷在时间上相近、波及的范围都巨大,共同促成了欧亚大陆古北区、东洋区和青藏高原区特有类群的分化,作为对这些地史事件反映的青藏高原区、古北区和东洋区,在动物地理区划上应该具有相同的地位．另外,依据高原鱼类的分布范围和青藏高原对非高原鱼类的阻碍作用,讨论了青藏高原的界限及其划分问题。     

Combining the least cost path method with population genetic data and species distribution models to identify landscape connectivity during the late Quaternary in Himalayan hemlock

Himalayan hemlock (Tsuga dumosa) experienced a recolonization event during the Quaternary period; however, the specific dispersal routes are remain unknown. Recently, the least cost path (LCP) calculation coupled with population genetic data and species distribution models has been applied to reveal the landscape connectivity. In this study, we utilized the categorical LCP method, combining species distribution of three periods (the last interglacial, the last glacial maximum, and the current period) and locality with shared chloroplast, mitochondrial, and nuclear haplotypes, to identify the possible dispersal routes of T. dumosa in the late Quaternary. Then, both a coalescent estimate of migration rates among regional groups and establishment of genetic divergence pattern were conducted. After those analyses, we found that the species generally migrated along the southern slope of Himalaya across time periods and genomic makers, and higher degree of dispersal was in the present and mtDNA haplotype. Furthermore, the direction of range shifts and strong level of gene flow also imply the existence of Himalayan dispersal path, and low area of genetic divergence pattern suggests that there are not any obvious barriers against the dispersal pathway. Above all, we inferred that a dispersal route along the Himalaya Mountains could exist, which is an important supplement for the evolutionary history of T. dumosa. Finally, we believed that this integrative genetic and geospatial method would bring new implications for the evolutionary process and conservation priority of species in the Tibetan Plateau.     

When east met west: the sub-fossil footprints of the west European hedgehog and the northern white-breasted hedgehog during the Late Quaternary in Europe

The distinct distribution of the west European hedgehog Erinaceus europaeus and the northern white-breasted hedgehog Erinaceus roumanicus and their separate refugial origins after the Pleistocene is a well-known example in the zoogeography of the Holarctic. Among the Late Quaternary faunal assemblages, the west European hedgehog is recorded at 269 sites whereas the northern white-breasted hedgehog is recorded only at 52 sites in Europe. The distribution patterns of the temporal and spatial Glacial records of the west European hedgehog show a general trend: a strong restriction to glacial refugia (the Iberian and Italian Peninsulas) during the Weichselian Glacial until the end of the Last Glacial Maximum, and a colonization of southern France during the early Late Glacial between 14 000 and 125 00 ¹⁴C years BP (15 000–12 800 cal. BC). Whereas the British Isles could have already been colonised by the end of the Pre-Boreal, in the rest of Central Europe E. europaeus was clearly distributed there in the Boreal for the first time. The west European hedgehog is an absolute Holocene faunal element in Central Europe. It appears in most parts of Central Europe during the Early Holocene, when the west European hedgehog met its eastern relative, which probably was similarly sensitive. After meeting each other, the distribution limit of both Erinaceus species in Central Europe seems to have been relatively constant in its geographic extent. Because of the clear climatic correlation, E. europaeus should be considered as an indicator species for temperate climatic conditions of the Holocene fauna. This should be considered during the reconstruction of climatic conditions with the help of the analysis of quaternary faunal material.     

横断山区昆虫区系初探

1981—1984年,中国科学院青藏高原科学考察队在我国西南部的横断山区进行了大规模的多学科综合考察,收集昆虫标本17万多号.横断山区特殊的自然地理条件孕育着独特的昆虫区系.种类繁庶、成分复杂,特有种,尤其高山特有种相当丰富,物种分化显著,并具有地域上的狭布性,构成本区昆虫区系特征.作者根据古北、东洋、高山特有三种主要成分的分布,试提出古北、东洋两大区系在本区的分异界线.典型东洋区系成分一般限于海拔2,800—3,000m以下地区,约与亚热带常绿阔叶林分布上线一致.     

江西武夷山自然保护区蝶类区系结构及垂直分布

据有较详细分布资料的227种蝶类昆虫分析,武夷山蝶类属东洋界范畴;垂直分布研究结果表明,武夷山蝶类从山麓至山顶,随着海拔升高,东洋界种类递减,古北界成分逐渐增多,尤以海拔1100m以上更加明显;在海拔1800m以上东洋界种被古北及古北跨区系种所替代。本文对武夷山蝶类水平分布的南北限和垂直分布的上限及它们的相互关系作了探讨。     

Evolutionary history of Apocheima cinerarius (Lepidoptera: Geometridae), a female flightless moth in northern China

The alterations in the phylogeographical structures of insects in response to the uplift of the Qinghai–Tibet Plateau and the Quaternary glaciations in eastern Asia, particularly in northern China, remain largely unknown. In this study, we selected Apocheima cinerarius, a moth with flightless females, using molecular data (complete mitochondrial genomes and nuclear data) and ecological niche modelling (ENM) to investigate the effects of paleoclimatic changes on the evolutionary history of insects in the area of northern China. The phylogenetic tree of complete mitochondrial genomes indicated that there were two lineages, the western and eastern lineages. The nuclear gene analyses also detected unique haplotypes in each lineage. Time of the most recent common ancestor (TMRCA) of the two lineages was approximately in Early–Middle Pleistocene. Bayesian skyline plots revealed that the western lineage underwent a population expansion event after the Last Glacial Maximum, whereas the eastern lineage underwent expansion between the Last Interglacial and the Last Glacial Maximum. Our results suggest that A. cinerarius expanded eastward from western sites until the moth was distributed across the entire region of northern China. Then, A. cinerarius underwent contraction into isolated glacial refugia followed by subsequent expansion driven by Pleistocene climate changes, which established a narrow sympatric area. Our results indicate that the Quaternary environmental fluctuations had profound influences on the diversification and demography of an insect in northern China, and the same species in north‐western China and north‐eastern China have different demographic histories.     

原羚属分类地位研究兼论中国羚羊的分类

中国羚羊类动物在世界羚羊类动物中有着特殊的地位,其中普氏原羚、藏原羚和藏羚是中国特有的珍稀物种,普氏原羚是极度濒危动物.开展中国羚羊类动物的分类和进化研究是当务之急.本研究利用形态特征、头骨测量及线粒体DNA的12S rRNA(440 bp)和16S rRNA(590 bp)对原羚属的分类地位及中国其他几种羚羊分类地位进行了探讨,发现对质量性状和数量性状数据的聚类分析的结果差异大,而且质量性状的聚类结果比较符合传统的分类系统.分子分类也得到相似结果,但在属下的分析存在较大差异.7种牛科动物的12S rRNA和16S rRNA序列差异分别为0.2%～7.1%和0.3%～8.4%.系统进化分析表明原羚属3种羚羊形成单系群,且蒙古瞪羚与普氏原羚的关系较藏原羚与普氏原羚的关系近.赛加羚应归宿于羚羊亚科,而不是与羊亚科的藏羚形成独立的赛加羚亚科.原羚属的进化可能与青藏高原的隆起有关.     

江西武夷山蝶类昆虫区系概貌

本文分析了江西武夷山区的蝶类昆虫的种类、分布、数量结构和区系组成。其１１２种蝶类隶属１１个区系或混合系,区系成分以东洋系为主体,但古北区系成分占有相当的比重,这一结构表明,江西武夷山处于南北种类昆虫交流汇合的重心地段,其位置接近于东洋区系的北缘。     

青藏高原及其周边地区高山植物谱系地理学研究进展

青藏高原及其周边地区是世界上高山植物最丰富的地区,通过谱系地理学研究可以探讨高山植物演化历史与高原隆升和第四纪冰期的关系。根据对已经报道的36种高山植物的谱系地理分析,其谱系地理模式主要表现为:一、冰期退却到高原边缘的避难所,冰后期回迁到高原面;二、地理隔离造成冰期存在多处避难所(含微型避难所),冰后期发生局域性扩张。青藏高原在晚第三纪的快速隆升促进了物种的分化和成种,而第四纪冰期更是加剧了物种的快速分化,高原隆升和第四纪周期性气候波动是形成青藏高原高山植物现代谱系地理格局的主要原因。横断山脉地区作为第四纪冰期高山植物的主要避难所,在进化生物学和保护生物学方面具有重要启示。最后在物种选择、采样策略、基因片段选择和研究方法等4个方面提出青藏高原地区谱系地理学的研究方向。     

横断山区蚜虫区系的特异性与历史渊源

文章系统分析了横断山区蚜虫区系的特异性,主要表现在:区系成分的复杂性、古北与东洋成分的交融性、区系组成的特有性和物种分布的间断性.进一步结合地质历史事件对横断山区蚜虫区系的历史渊源及其特异性演化进行了初步讨论.根据蚜虫类分布特点,建议古北区和东洋区在横断山区的分界线南移至丽江一线;横断山区蚜虫和其他地区的间隔分布可能是由地质历史上板块运动和青藏高原的隆升及第四纪冰期-间冰期的循环往复造成的;横断山区蚜虫区系是一个复合区系,同其他动物地理区划如新北区、澳洲区、非洲区等的蚜虫区系都有密切的历史渊源,青藏高原和横断山区的隆起及第四纪冰期-间冰期的循环往复又使得本区蚜虫区系有着独特的演化历程,从而造成了本区蚜虫区系的特有性.     

Comparative phylogeography and evolutionary history of schizothoracine fishes in the Changtang Plateau and their implications for the lake level and Pleistocene climate fluctuations

The water level oscillation of endorheic lakes and extent change of glaciers associated with the Asian monsoon are known as prominent representatives of climatic and environmental events in the Tibetan Plateau during the Quaternary. However, details process in spatial and temporal changes are still debated. We use the schizothoracines as a palaeoclimatic proxy to test two hypotheses concerning the evolution of Quaternary glaciations and lakes of the Changtang Plateau: (1) the Tibetan glaciations generally tended to decrease since the middle Pleistocene; (2) the lakes expansion was driven by summer monsoon rainfall. Based on a wide range‐wide sampling throughout in the Changtang Plateau and its adjacent drainages, we constructed phylogeny and demographic histories of schizothoracines in the Changtang Plateau. Our results showed that the populations of the exorheic rivers and lakes in southern Tibet possessed higher genetic variability, earlier coalescent and expansion times than those of the endorheic lakes in the Changtang Plateau. Population expansions are highly consistent with phases of strong summer monsoon and high lake level during interglacial stages. The maximum growth rate intervals showed three pulses from 64.7 to 54.8, 39.6 to 31.0, and 14.9 to 2.4 kya respectively. The significant positive correlations were found between regional precipitation and genetic diversity, as well as coalescence time of populations in the endorheic lakes. We suggested that the demographic history of the schizothoracines reflects the spatial and temporal changes in climate and lake level, in particular, in regional precipitation gradients associated with changes of the South Asian monsoon, and supports the climatic hypothesis of a general diminishing tend in Tibetan glaciations in the Tibetan Plateau since the middle Pleistocene.