中国大陆第四纪冰期潜在植物避难所研究进展

研究第四纪冰期潜在植物避难所,对了解当今植物分布格局和未来的演变具有重要意义,是全球变化生态学研究的热点问题。根据已发表的文献,本文综述了我国大陆第四纪冰期冰川的发生规模、范围和对植物分布的影响,探讨了冰期潜在植物避难所形成的特点,包括特殊地理位置的生物多样性和避难所残遗种分布等问题。本文还讨论了孢粉分析和分子生物技术的方法在确定冰期植物避难所方面的应用,以及我国科学家在该领域重要研究结果。同时,本文还介绍了国外在该领域的研究进展和我们存在的问题,以期促进我国第四纪冰期植物避难所研究工作的深入开展。     

基于AFLP分析的伯乐树(Bretschneidera sinensis)谱系地理学研究

伯乐树（Bretschneidera sinensis Hemsl.）是主要分布于中国的濒危物种。采用AFLP分子标记对分布于中国11个省的24个伯乐树居群192个个体进行谱系地理学研究。结果显示,伯乐树有相对较高的遗传多样性水平,基因多样性指数（He）和Shannon指数（Ⅰ）分别为0.2728和0.4070。伯乐树居群间的遗传分化远大于居群内遗传分化,遗传分化系数G_ST=0.7138,基因流N_m=0.2005。通过聚类分析、STRUCTURE分析和BAPS分析发现,24个伯乐树居群可形成4大地理居群组和进化谱系;云贵高原东部地区居群遗传多样性较高,可能是伯乐树在中国的扩散中心和冰期避难所,伯乐树在冰期后由此向外进行居群扩散;南岭地区各居群遗传多样性水平普遍高于其他地区,与邻近地区各居群的亲缘关系较近,可能为伯乐树演化历史上的另一个冰期避难所。     

四照花物种分布格局模拟及冰期避难所推测

气候是影响物种多样性的重要自然因素,同时也影响生物的生长发育及其分布。该文以广泛分布于中国的观花植物四照花为研究对象,通过植物标本库查询和野外实地调查确定四照花在中国的分布情况,利用MaxEnt模型和ArcGIS软件模拟四照花的历史分布格局、当前分布格局及未来的分布趋势,计算不同时期物种分布面积的变化并分析物种分布变化产生的原因。研究结果显示:四照花当前的分布主要受降水量变异系数、最冷月份最低温、最冷季节平均温度和温度季节变化4个生物气候因子影响。此外,基于四照花在末次盛冰期、全新世中期和当前的分布格局的变化推测:冰期来临时,中西部的大巴山地区和东部天目山地区是四照花潜在分布的核心区,为四照花的冰期避难所。     

基于MaxEnt模型和GIS的青藏高原紫花针茅分布格局模拟

气候变化对物种分布的影响是生物地理学研究的热点问题.本研究以广泛分布在青藏高原高寒草甸区域的建群种紫花针茅为研究对象,通过植物标本库查询和实地调查了解紫花针茅在青藏高原的分布情况,利用Max Ent模型模拟紫花针茅在青藏高原历史、当前及未来的分布趋势,分析紫花针茅各历史时期的分布格局,探讨了物种分布变化的产生原因.结果表明:青藏高寒草地的针茅属物种具有较高的物种多样性,代表性植物紫花针茅的主要分布区域为青藏高原腹地及沿喜马拉雅地区;紫花针茅的分布强烈地受最暖季降水量、最湿季降水量和年均降水量的控制.依据紫花针茅在末次冰期的分布格局以及青藏高原的地理地质特点,研究认为:冰期来临时,羌塘以南藏北核心区和喜马拉雅西部阿里地区是紫花针茅潜在分布的核心区,此处提供了比其他地区更适合紫花针茅生存的栖息地,这些区域成为紫花针茅的避难所,当前的紫花针茅都是从这些避难所迁移并扩展开来.生物避难所的存在有助于理解青藏高原高寒植被起源和分化的相关问题.     

鸟类分子系统地理学研究进展

分子系统地理学是当代生物地理学的重要分支,是以分子生物学方法重建种内和种上水平的系统发育关系,阐释进化历史,并通过分析近缘生物类群的系统发育关系与其空间和时间分布格局间的相关性构建生物区系历史的研究,是分子生物学与生物地理学结合的产物。中性进化学说和溯祖理论分析的建立,以及线粒体DNA和微卫星标记等分子遗传标记的应用,为分子系统地理学研究的开展提供了理论和实践基础。近年来,分子系统地理方法在鸟类学研究中的应用揭示了许多不同于传统认知的发现,为准确而深入的了解鸟类分子系统地理格局的差异和不同类群的起源中心提供了新颖的证据。目前的研究多从隔离分化说和扩散说的角度对鸟类分子系统地理格局的成因进行分析,而迁徙行为不同对鸟类系统地理格局的影响为成因的解释提供了新的角度。结合区域特点的比较分子系统地理研究,在更广泛的地域和更多类群中开展研究是我国鸟类分子系统地理研究的方向。此外,展望了第二代测序技术对分子生态生物地理研究具有的潜在促进作用。     

松鼠分子生态学研究进展

松鼠由于受到非法猎捕、栖息地破坏及欧洲部分地区的北美灰松鼠生态入侵,导致种群数量锐减,现已被世界自然保护联盟(IUCN)列为近危种,我国吉林省已将其列入省重点保护野生动物名录.分子生物学研究方法的快速发展,尤其是基于mtDNA片段开展的相关研究,以及已筛选出并能应用于松鼠研究的微卫星位点的应用,使松鼠分子生态学研究不断深入.本文对松鼠的分子系统发育、遗传多样性和分子系统地理学等分子生态学内容进行了综述,并提出松鼠分子生态学未来研究的展望:进一步探讨松鼠与日本松鼠的系统分化关系;松鼠连续种群、隔离种群和集合种群的遗传多样性比较分析;利用核基因其他标记分析松鼠分子系统地理学问题;探讨亚洲是否存在第四纪冰期避难所.     

松鼠分子生态学研究进展

松鼠由于受到非法猎捕、栖息地破坏及欧洲部分地区的北美灰松鼠生态入侵,导致种群数量锐减,现已被世界自然保护联盟(IUCN)列为近危种,我国吉林省已将其列入省重点保护野生动物名录.分子生物学研究方法的快速发展,尤其是基于mtDNA片段开展的相关研究,以及已筛选出并能应用于松鼠研究的微卫星位点的应用,使松鼠分子生态学研究不断深入.本文对松鼠的分子系统发育、遗传多样性和分子系统地理学等分子生态学内容进行了综述,并提出松鼠分子生态学未来研究的展望:进一步探讨松鼠与日本松鼠的系统分化关系;松鼠连续种群、隔离种群和集合种群的遗传多样性比较分析;利用核基因其他标记分析松鼠分子系统地理学问题;探讨亚洲是否存在第四纪冰期避难所.     

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。     

东北亚地区鼩鼱科动物分子生态学研究进展

在东北亚地区（中国的东北地区和内蒙古东北部、日本、朝鲜半岛、蒙古国、俄罗斯的远东地区）具有较为丰富的鼩鼱科类群。分子生物学方法的快速发展,使东北亚地区鼩鼱科动物分子生态学研究不断深入。对鼩鼱科动物的分子系统发育、遗传多样性和分子系统地理学等分子生态学内容进行了综述。提出鼩鼱科动物分子生态学研究未来的发展：1）东北亚地区第四纪冰期避难所的研究;2）同域分布的鼩鼱科动物比较系统地理学研究;3）中国东北地区鼩鼱科动物在东北亚分布区的系统地理学地位;4）新型分子标记和分析方法的发展。     

药食同源植物薤白的谱系地理学研究

对我国14个地区的药食同源植物薤白（Allium macrostemon）的叶绿体基因片段（psbA-trnH、rps16和trnL-F）与核基因片段（ITS）进行测序分析,揭示薤白的遗传变异分布式样、单倍型地理分布格局,并推断其在第四纪冰期的避难所。结果表明：薤白叶绿体基因（cpDNA）遗传多样性低于核基因（nrDNA）遗传多样性（cpDNA：H_T=0.868;nrDNA：H_T=0.890）。cpDNA和nrDNA的分子变异分析（AMOVA）结果显示：薤白遗传变异主要发生在居群间（cpDNA：92.84%;nrDNA：98.40%）,存在遗传分化（cpDNA：N_st=0.918,G_st=0.866,F_st=0.928;nrDNA：N_st=0.984,G_st=0.855,F_st=0.984）,且N_st均大于G_st,表明该物种具有明显的谱系地理结构。在薤白居群中,共检测到11个叶绿体单倍型和14个nrDNA基因型;单倍型网络图及地理分布图表明,叶绿体单倍型H3、核DNA基因型H1频率最高,位于网络结构图的中心位置,可能为古老单倍型。此外,冰期避难所假说认为遗传多样性高、拥有古老单倍型和较多特有单倍型的区域可能是该物种的冰期避难所,因此推测薤白在第四纪冰期时可能在大盘山、天水和通化地区存在多个冰期避难所。这些分析可为类似草本植物的进化提供参考,丰富对东亚草本植物分子系统与生物地理学的认识。     

Evolutionary history of Lissotriton helveticus: multilocus assessment of ancestral vs. recent colonization of the Iberian Peninsula

The Pleistocene was characterized by climatic changes that greatly altered the distribution of organisms. Population extinctions, bottlenecks, isolation, range expansions and contractions were often associated with glaciations, leaving signatures in the spatial patterns of genetic diversity across species. Lissotriton helveticus belongs to a Pan-European lineage of newts that were strongly affected by glaciations and represent an excellent model to analyse the effect of generalized climatic changes in phylogeographic patterns. We studied the genetic diversity of the species using data from two mitochondrial and three nuclear genes analyzed in a Bayesian phylogenetic framework to investigate the historical processes shaping spatial patterns of genetic diversity. Mitochondrial haplotypes cluster in four different groups present in the Iberian Peninsula and of Pleistocene origin, probably by allopatric fragmentation. Nuclear genes present no obvious geographic structure patterns, suggesting gene flow and generalized incomplete lineage sorting. Populations north of the Pyrenees are closely related to those from northeastern Iberia, suggesting recent range expansion from this region. Historical demographic analyses indicate a demographic expansion starting about 100,000years ago and more recent population declines. Compared to other Lissotriton species, L. helveticus includes only relatively young genetic lineages, suggesting a Central European pre-Pleistocene distribution followed by complete extirpation of the species during glaciations in that area. Historical demographic trends in the Iberian Peninsula are reversed with respect to the more Mediterranean species Lissotriton boscai, indicating different responses of both species to climate changes. Diversity patterns among Lissotriton species seem to be defined by four main factors: ancestral distributions, colonization capabilities, interactions with other species and effective population sizes. Differences in these factors define two types of species, referred to as "R" (refugia) and "S" (sanctuaries) that explain part of the diversity in patterns of genetic diversity created by glaciations in Western Europe.     

Historical biogeography: the new synthesis

Since Darwin first noted the impact of past ice ages on the distribution of organisms, biogeographers have debated whether Pleistocene glaciations shaped evolutionary patterns. A new synthesis of population genetics and palaeoclimatology promises unprecedented insights into Pleistocene evolutionary history.     

Phylogeographic history of grey wolves in Europe

Background  

While it is generally accepted that patterns of intra-specific genetic differentiation are substantially affected by glacial history, population genetic processes occurring during Pleistocene glaciations are still poorly understood. In this study, we address the question of the genetic consequences of Pleistocene glaciations for European grey wolves. Combining our data with data from published studies, we analysed phylogenetic relationships and geographic distribution of mitochondrial DNA haplotypes for 947 contemporary European wolves. We also compared the contemporary wolf sequences with published sequences of 24 ancient European wolves.     

Phylogeography and palaeodistribution modelling of Nassauvia subgenus Strongyloma (Asteraceae): exploring phylogeographical scenarios in the Patagonian steppe

The Patagonian steppe is an immense, cold, arid region, yet phylogeographically understudied. Nassauvia subgen. Strongyloma is a characteristic element of the steppe, exhibiting a continuum of morphological variation. This taxon provides a relevant phylogeographical model not only to understand how past environmental changes shaped the genetic structure of its populations, but also to explore phylogeographical scenarios at the large geographical scale of the Patagonian steppe. Here, we (1) assess demographic processes and historical events that shaped current geographic patterns of haplotypic diversity; (2) analyze hypotheses of isolation in refugia, fragmentation of populations, and/or colonization of available areas during Pleistocene glaciations; and (3) model extant and palaeoclimatic distributions to support inferred phylogeographical patterns. Chloroplast intergenic spacers, rpl32–trnL and trnQ–5′rps16, were sequenced for 372 individuals from 63 populations. Nested clade analysis, analyses of molecular variance, and neutrality tests were performed to assess genetic structure and range expansion. The present potential distribution was modelled and projected onto a last glacial maximum (LGM) model. Of 41 haplotypes observed, ten were shared among populations associated with different morphological variants. Populations with highest haplotype diversity and private haplotypes were found in central‐western and south‐eastern Patagonia, consistent with long‐term persistence in refugia during Pleistocene. Palaeomodelling suggested a shift toward the palaeoseashore during LGM; new available areas over the exposed Atlantic submarine platform were colonized during glaciations with postglacial retraction of populations. A scenario of fragmentation and posterior range expansion may explain the observed patterns in the center of the steppe, which is supported by palaeomodelling. Northern Patagonian populations were isolated from southern populations by the Chubut and the Deseado river basins during glaciations. Pleistocene glaciations indirectly impacted the distribution, demography, and diversification of subgen. Strongyloma through decreased winter temperatures and water availability in different areas of its range.     

Disentangling causes of disjunction on the South Island of New Zealand: the Alpine fault hypothesis of vicariance revisited

Many elements of the flora and fauna of New Zealand's South Island show disjunct distributions with conspecific populations or closely-related species that occur in the north-west and south separated by a central gap. Three events have been implicated to account for this pattern: Pleistocene glaciations, Pliocene mountain building, or displacement along the Alpine fault, the border of the Pacific and Australian plates stretching diagonally across the South Island from south-west to north-east that formed during the Miocene. Disjunct distributions of species level taxa are probably too young to be due to Alpine fault vicariance. It has therefore been suggested that the biogeographical impact of the Alpine fault, if any, should be apparent on deeper phylogenetic levels. We tested this hypothesis by reconstructing the phylogenetic relationships of the hydrobiid gastropods of New Zealand based on mitochondrial DNA fragments of cytochrome oxidase subunit I (CO I ) and 16S rDNA. The creno- and stygobiont species of this family are typically poor dispersers. Therefore, ancient patterns of distribution may be conserved. The phylogenetic reconstructions were in accordance with the Alpine fault hypothesis uniting genera occurring on either side of the fault. Divergence estimates based on a molecular clock of CO I indicated splits predating the Pliocene uplift of the Alps.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 361–374.     

Molecular evidence for Pleistocene refugia at the eastern edge of the Tibetan Plateau

The role of the Quaternary ice ages in forming the contemporary genetic structure of populations has been well studied in a number of global regions. However, due to the different nature of glaciations and complex topography, their role in shaping eastern Eurasian genetic diversity, particular in areas surrounding the Tibetan Plateau have remained largely unstudied. We aimed to address this question by examining the genetic structure of an alpine forest-associated taxon, the blood pheasant (Ithaginis cruentus) to infer its phylogeographic history. We detected three phylogenetic lineages and four current population groups. By comparing molecular and palaeovegetation data, we found that major glaciations during the Pleistocene have had a major impact upon the current genetic diversity of this species. Coalescent simulations indicate that the populations retreated to different refugia during some glacial periods in the Pleistocene, but persisted through the last glacial maximum (LGM). The most significant recent population expansion was found to have occurred before the LGM, during which palaeoclimatic data indicate that the climate was both warmer and wetter than today. In contrast, during the LGM populations may have adopted an altitudinal shift strategy in order to track changes in alpine glaciers, exemplifying a general response for montane species in the region where alpine glaciations were not large enough to cause qualitative changes in vegetation. Although analysis based on a plumage related gene showed that divergent selection may have contributed to current patterns of intra-specific diversity, demographic isolation is inferred to have played a more dominant role.     

High levels of population subdivision in a morphologically conserved Mediterranean toad (Alytes cisternasii) result from recent,multiple refugia: evidence from mtDNA,microsatellites and nuclear genealogies

Pleistocene glaciations often resulted in differentiation of taxa in southern European peninsulas, producing the high levels of endemism characteristic of these regions (e.g. the Iberian Peninsula). Despite their small ranges, endemic species often exhibit high levels of intraspecific differentiation as a result of a complex evolutionary history dominated by successive cycles of fragmentation, expansion and subsequent admixture of populations. Most evidence so far has come from the study of species with an Atlantic distribution in northwestern Iberia, and taxa restricted to Mediterranean‐type habitats remain poorly studied. The Iberian Midwife toad (Alytes cisternasii) is a morphologically conserved species endemic to southwestern and central Iberia and a typical inhabitant of Mediterranean habitats. Applying highly variable genetic markers from both mitochondrial and nuclear genomes to samples collected across the species’ range, we found evidence of high population subdivision within A. cisternasii. Mitochondrial haplotypes and microsatellites show geographically concordant patterns of genetic diversity, suggesting population fragmentation into several refugia during Pleistocene glaciations followed by subsequent events of geographical and demographic expansions with secondary contact. In addition, the absence of variation at the nuclear β‐fibint7 and Ppp3caint4 gene fragments suggests that populations of A. cisternasii have been recurrently affected by episodes of extinction and recolonization, and that documented patterns of population subdivision are the outcome of recent and multiple refugia. We discuss the evolutionary history of the species with particular interest in the increasing relevance of Mediterranean refugia for the survival of genetically differentiated populations during the Pleistocene glaciations as revealed by studies in co‐distributed taxa.     

Impact of ice ages on circumpolar molecular diversity: insights from an ecological key species

We address the impact of the ice age cycles on intraspecific cpDNA diversity, for the first time on the full circumboreal-circumarctic scale. The bird-dispersed bog bilberry (or arctic blueberry, Vaccinium uliginosum) is a key component of northern ecosystems and is here used to assess diversity in previously glaciated vs. unglaciated areas and the importance of Beringia as a refugium and source for interglacial expansion. Eighteen chloroplast DNA haplotypes were observed in and among 122 populations, grouping into three main lineages which probably diverged before, and thus were affected more or less independently by, all major glaciations. The boreal 'Amphi-Atlantic lineage' included one haplotype occurring throughout northern Europe and one occurring in eastern North America, suggesting expansion from at least two bottlenecked, glacial refugium populations. The boreal 'Beringian lineage' included seven haplotypes restricted to Beringia and the Pacific coast of USA. The 'Arctic-Alpine lineage' included nine haplotypes, one of them fully circumpolar. This lineage was unexpectedly diverse, also in previously glaciated areas, suggesting that it thrived on the vast tundras during the ice ages and recolonized deglaciated terrain over long distances. Its largest area of persistence during glaciations was probably situated in the north, stretching from Beringia and far into Eurasia, and it probably also survived the last glaciation in southern mountain ranges. Although Beringia apparently was important for the initial divergence and expansion of V. uliginosum as well as for continuous survival of both the Beringian and Arctic-Alpine lineages during all ice ages, this region played a minor role as a source for later interglacial expansions.     

Land planarians (Platyhelminthes) as a model organism for fine-scale phylogeographic studies: understanding patterns of biodiversity in the Brazilian Atlantic Forest hotspot

The Brazilian Atlantic Forest is one of the richest biodiversity hotspots of the world. Paleoclimatic models have predicted two large stability regions in its northern and central parts, whereas southern regions might have suffered strong instability during Pleistocene glaciations. Molecular phylogeographic and endemism studies show, nevertheless, contradictory results: although some results validate these predictions, other data suggest that paleoclimatic models fail to predict stable rainforest areas in the south. Most studies, however, have surveyed species with relatively high dispersal rates whereas taxa with lower dispersion capabilities should be better predictors of habitat stability. Here, we have used two land planarian species as model organisms to analyse the patterns and levels of nucleotide diversity on a locality within the Southern Atlantic Forest. We find that both species harbour high levels of genetic variability without exhibiting the molecular footprint of recent colonization or population expansions, suggesting a long-term stability scenario. The results reflect, therefore, that paleoclimatic models may fail to detect refugia in the Southern Atlantic Forest, and that model organisms with low dispersal capability can improve the resolution of these models.     

High genetic variation in marginal fragmented populations at extreme climatic conditions of the Patagonian Cypress Austrocedrus chilensis

Knowledge about current patterns of genetic structure of populations together with the evolutionary history of a species helps to understand and predict the adaptation of populations to future climate change. We assayed variation at nuclear microsatellite markers among peripheral vs. continuous populations of the temperate South American species Austrocedrus chilensis, to investigate the role of historical vs. demographical forces in shaping population genetic structure. This species occurs in continuous populations in the west and central distribution range, but becomes highly fragmented at the eastern limit, which comprised ice-free areas during Quaternary glaciations and has extreme climatic conditions at present times. Bayesian analysis methods identified two contrasting patterns of genetic structure; (I) populations from humid, mesic and peri-glacial regions formed a single deme with relatively low genetic differentiation and high admixture levels whereas (II) a highly heterogeneous genetic structure with low level of admixture was found in the steppe, towards the east and northeast limit of the distribution range. In the steppe, population fragmentation, restricted gene flow and isolation-by-distance were also inferred. In addition, several small steppe populations showed high genetic diversity and divergent gene pools, suggesting that they constitute ancient refuges from pre-Holocene glaciations with just a subgroup of them contributing significantly to post-glacial spread. These results are discussed in relation to patterns of genetic variation found for other temperate species and the contribution of the particular southern Andes topography and climate to post-glacial spread.