生物冰期避难所与冰期后的重新扩散

冰期（尤其是更新世冰期）对当今生物的空间分布格局和遗传结构产生了深远影响,研究生物冰期避难所对于了解不同生物区系间的关系、物种形成以及生物多样性保护具有十分重要的意义。生物冰期避难所的确定员韧是根据特有种的分布、化石、抱粉等证据推测而来,分子遗传标记为冰期生物避难所以及冰期后重新扩散路线的研究提供了有力的工具。本文以北美和欧洲为例,介绍了分子遗传标记在生物避难所以及冰期后再扩散路线研究中的应用和一些结论。我国存在许多东西走向的大山,减缓了冰期时低温对生物的影响,为许多生物提供了避难场所,但我国有关研究开展得很少。员后,对我国该领域提出了应优先开展的研究方向。     

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。     

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.     

中国东北温带针阔混交林植物物种的谱系地理研究进展

中国东北温带针阔混交林是暖温带植被与寒温带植被的交错区,是研究第四纪气候变化对遗传格局影响的关键区域。古植被重建研究显示,在末次冰盛期时,针阔混交林向南退缩至25°–30°N之间。然而,谱系地理研究则表明针阔混交林植物在末次冰盛期时可在35°N以北的一个或多个避难所中原地存活。长白山和朝鲜半岛是针阔混交林植物最重要的两个避难所,在其他区域还可能存在多个避难所,最北可达小兴安岭和俄罗斯远东地区。在避难所种群间冰期或冰期后发生扩张的过程中,形成了复杂的遗传多样性分布模式。单个避难所向北扩张的种群历史并不一定导致种群内遗传多样性随纬度升高而显著降低。在多个避难所模式下,种群内遗传多样性一般呈均匀分布。以往研究揭示了第四纪,尤其是末次冰盛期对针阔混交林植物进化历史的影响,但仍主要是乔木成分,且集中于单一物种。因此,东北和南方地区近缘种或姊妹种的适应性进化比较及针阔混交林的群落构建机制检验是未来研究的两个发展趋势。     

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。     

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。     

横断山地区海仙报春的谱系地理学研究

由于横断山地区特殊的地理效应,第四纪冰期气候的反复变化对该地区植物的地理分布和居群遗传结构产生重要影响。为了揭示该地区分布的物种的谱系地理结构,检测了该地区特有的、生于湿地环境的植物海仙报春（Primula poissonii）的叶绿体trnL—F和trnT—L两个序列变异。共对13个居群167个个体的两个叶绿体片段进行测序,共发现11个单倍型。三种单倍型被共享,其他单倍型都只存在于单个居群内。分子变异分析（AMOVA）结果表明分布区内海仙报春的遗传变异主要存在于居群间（99．08％）,且居群间的遗传分化很高（GST=0．916,FST=0．99077,NST=0．973）,有着极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=0．08）。结合嵌套分支分析（NCA）,本文的研究结果推测该物种在冰期可能存在多个避难所,并没有表现出大多数温带植物在第四纪冰期所经历过的种群集体扩张和迁移的现象,异域片段化被认为是该物种现有单倍型分布格局形成的主要原因。海仙报春的谱系地理结构是气候的反复波动和横断山特殊的地理环境互相作用的结果。     

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。     

基于叶绿体DNA trnT-trnF序列研究祁连圆柏的谱系地理学

由于青藏高原的地貌效应,第四纪冰期气候的反复变化应对现今该地区生物的地理分布及其居群遗传结构产生重大影响。本文对这一地区特有分布物种祁连圆柏Juniperus przewalskii Kom.整个分布区内20居群392个个体的trnT-trnF序列变化进行了研究;共发现3种单倍型(haplotype),构成两种地理区域:高原台面上的居群主要固定Hap A,而Hap A、Hap B和Hap C在高原边缘居群均有分布。所有居群总的遗传多样性HT = 0.511,GST= 0.847。在低海拔的高原边缘,Hap A、Hap B和Hap C高频率固定在不同的居群中,表明可能存在多个不同的避难所,居群反复缩小和扩张的瓶颈效应造成了遗传多样性的丢失。而边缘的一个居群含有两种相似单倍型频率则可能是冰期后迁移融合而成或者该居群在冰期经受的瓶颈作用更弱。高原台面东部间断分布的居群只固定Hap A,表明它们可能经历了冰期后共同的回迁过程和由此产生的奠基者效应。我们的研究结果表明祁连圆柏在冰期可能存在多个避难所,瓶颈效应和奠基者效应造成了这些居群现在的遗传多样性分布式样。     

“横断山脉——喜马拉雅山脉”,菊科蒿属植物区系地理一特殊的分区

本文论述了"横断山脉——喜马拉雅山脉"森林植物亚区是菊科蒿属(Artemisia Linn.)植物区系地理一特殊的分区。该分区含蒿属植物116种、21变种,作者参考了孢粉及地质资料后提出该亚区既是第四纪冰期蒿属植物的"避难所",又是该属植物"次生的起源中心"与现代分布区的"第二个密集中心"。现该亚区保存有少数第三纪蒿属古特有种与次生的原始种,同时还含众多冰后期分化、衍生出的区域性特有种,后者有64种13变种之多。文中将该亚区分4地区成分分述,附各地区成分蒿属植物的代表种,同时还与周围地区的区系成分作比较。     

Species persistence in northerly glacial refugia of Europe: a matter of chance or biogeographical traits?

Aim The southern European peninsulas (Iberian, Italian and Balkan) are considered to have been refugia for many European species of plants and animals during the climatic extremes of the Pleistocene ice ages. A number of recent studies (fossil and genetic), however, have provided evidence for full‐glacial survival of some species beyond these peninsulas. Here we explore the biogeographical traits of these species, and ask whether they possessed certain characteristics that enabled them to persist in more northerly refugia. Location Europe. Methods Fossil and genetic evidence for refugial localities of species that survived in Europe during the last full‐glacial was obtained from the literature (totalling 90 species: 34 woody plants and 56 vertebrates). Forty‐seven of these species (23 woody plants and 24 vertebrates) had fossil evidence, whereas the remaining 43 species (11 woody plants and 32 vertebrates) had only genetic evidence. All species were scored according to their present geographical distribution, habitat preference and life‐history traits. The species were classified on the basis of these traits using hierarchical cluster analysis. Analysis of similarities was used to examine differences in vertebrate and woody plant species groups that survived only in southerly refugia and those that also persisted in more northerly locations. Non‐metric multi‐dimensional scaling was used to examine patterns observed between and within groups. Results Results from our analysis of species with fossil and genetic evidence for survival in refugia reveal that species that survived only in southerly refugia were large‐seeded trees or thermophilous vertebrates. In contrast, species that had a full‐glacial distribution, including more northerly locations, were wind‐dispersed, habitat‐generalist trees with the ability to reproduce vegetatively, and habitat‐generalist mammals with present‐day northerly distributions. Main conclusions Analysis of the geographical distribution, habitat preference and life‐history traits of the species studied suggests that underlying biogeographical traits may have determined their response to Pleistocene glaciation. The traits most commonly found in present populations with a northerly distribution in Europe enabled the same species to exist much farther north than the southern European peninsulas during the full‐glacial. It is possible that many of these species are now in restricted populations, within the ‘warm‐stage’ refugia of the current interglacial. The northerly full‐glacial survival of a number of woody plants and vertebrate species has significant implications for understanding migration rates of these species in response to climate change. It also has important implications for understanding current patterns of genetic diversity of European species. We suggest that both fossil and genetic evidence should be used to identify and prioritize for conservation of refugial localities in southern and northern Europe.     

Environmental confirmation of multiple ice age refugia for Pacific cod, Gadus macrocephalus

The concept of species surviving through quaternary climatic extremes by retreating to glacial refugia, and then evolving genetically during re-population movements of the following interglacial, has been in the literature for over 40 years. Recently, advances in genetic analysis have enabled this concept to be validated and theories regarding population expansions and contractions to be built. For the major Northern Hemisphere species of cod, Gadus morhua (Atlantic cod) and Gadus macrocephalus (Pacific cod), genetic analysis has suggested retreat to separate refugia on both sides of their respective ocean basins during the last glacial period. Ecosystem niche modelling has previously confirmed that environmental conditions during the last glacial were compatible with the existence of these separate refugia for Atlantic cod. Here it is shown that such modelling also confirms a reduced core glacial distribution for G. macrocephalus, but probable refugia on either side of the Pacific. Existing mitochondrial DNA analyses suggest two separate glacial populations in the northwest Pacific, which modelling confirms, with predicted separate marine refugia in the land-locked Sea of Japan basin and the Sea of Okhotsk. Existing mitochondrial DNA for the northeast Pacific populations is less conclusive regarding whether there were one or two separate refugia off this coast, and their location. Using environmental niche models this study shows the glacial NE Pacific environment could support two marine refugia, one centred in the Aleutians/Gulf of Alaska and the other off British Columbia. The intervening Cordilleran Ice Sheet, and the glacially sub-aerial and ice-free Queen Charlotte Islands shelf, is hypothesised to have constrained exchange between glacial stocks either side of the Islands. The postulated southern marine refugium is off-shore from an established terrestrial refugium, suggesting greater dependence of species and ecosystems during environmental change. An earth system approach to evolutionary change could enhance understanding of past and future ecosystems.     

中国亚热带地区阔叶林植物的谱系地理历史

中国亚热带地区因丰富的植物物种多样性备受生物地理学研究关注,丰富的多样性与中新世以来的地质气候变化密切相关。谱系地理学已成为探讨植物分布模式和遗传格局受地质气候变化影响的主要手段。总结了该地区阔叶林植物对中新世以来地质气候变化的响应模式和种群分化的历史成因。在中新世和上新世时,由于全球变冷、青藏高原抬升和亚洲内陆干旱,阔叶林植物被迫向南退缩,形成不同的谱系。同时,亚洲季风的增强为遗传多样性的增加提供了良好的环境。在更新世冰期和间冰期时,大部分落叶阔叶林和常绿阔叶林植物在多个避难所间存在长期隔离,不同的避难所种群各自经历局部地区的收缩和扩张。长期的隔离使不同谱系间进一步分化,形成高水平的遗传多样性和遗传分化。少部分植物在冰期时向南退缩,并在间冰期时明显向北扩张。最后,就分化时间的准确估计和探究遗传格局背后的机制两方面展望了未来的研究趋势。     

Postglacial colonization of Europe by the barbastelle bat: agreement between molecular data and past predictive modelling

The barbastelle (Barbastella barbastellus) is a rare forest bat with a wide distribution in Europe. Here, we combine results from the analysis of two mtDNA fragments with species distribution modelling to determine glacial refugia and postglacial colonization routes. We also investigated whether niche conservatism occurs in this species. Glacial refugia were identified in the three southern European peninsulas: Iberia, Italy and the Balkans. These latter two refugia played a major role in the postglacial colonization process, with their populations expanding to England and central Europe, respectively. Palaeo‐distribution models predicted that suitable climatic conditions existed in the inferred refugia during the last glacial maximum (LGM). Nevertheless, the overlap between the current and the LGM distributions was almost inexistent in Italy and in the Balkans, meaning that B. barbastellus populations were forced to shift range between glacial and interglacial periods, a process that probably caused some local extinctions. In contrast, Iberian populations showed a ‘refugia within refugium’ pattern, with two unconnected areas containing stable populations (populations that subsisted during both glacial and interglacial phases). Moreover, the match between LGM models and the refugial areas determined by molecular analysis supported the hypothesis of niche conservatism in B. barbastellus. We argue that geographic patterns of genetic structuring, altogether with the modelling results, indicate the existence of four management units for conservation: Morocco, Iberia, Italy and UK, and Balkans and central Europe. In addition, all countries sampled possessed unique gene pools, thus stressing the need for the conservation of local populations.     

Phylogeographic insights of the lowland species Cheirolophus sempervirens in the southwestern Iberian Peninsula

The southwestern Iberian Peninsula is an important biogeographic region, showing high biodiversity levels and hosting several putative glacial refugia for European flora. Here, we study the genetic diversity and structure of the Mediterranean, thermophilous plant Cheirolophus sempervirens (Asteraceae) across its whole distribution range in SW Iberia, as a tool to disentangle some of the general biogeographic patterns shaping this southern refugia hotspot. Null genetic diversity was observed in the cpDNA sequencing screening. Nonetheless, AFLP data revealed high levels of among-population genetic differentiation correlated to their geographic location. Our results suggest longer species persistence in southern Iberian refugia during glacial periods and subsequent founder effects northwards due to colonizations in warmer stages (i.e., the southern richness to northern purity pattern). Additionally, our phylogeographic analyses indicate the presence of two separate genetic lineages within Ch. sempervirens, supporting the hypothesis of multiple minor refugia for SW Iberia in agreement with the refugia within refugia model.     

Palaeoclimatic models help to understand current distribution of Caucasian forest species

Spatial and temporal constraints on dispersal explain the absence of species from areas with potentially suitable conditions. Previous studies have shown that post‐glacial recolonization has shaped the current ranges of many species, yet it is not completely clear to what extent interspecific differences in range size depend on different dispersal rates. The inferred boundaries of glacial refugia are difficult to validate, and may bias spatial distribution models (SDMs) that consider post‐glacial dispersal constraints. We predicted the current distribution of 12 Caucasian forest plants and animals, factoring in the effective geographical distance from inferred glacial refugia as an additional predictor. To infer glacial refugia, we tested the transferability of the current SDMs based on the distribution of climatic variables, and projected the most transferable ones onto two climate scenarios simulated for the Last Glacial Maximum (LGM). We then calculated least‐cost distances from the inferred refugia, using elevation as a friction surface, and recalculated the current SDMs incorporating the distances as an additional variable. We compared the predictive powers of the initial with the final SDMs. The palaeoclimatic simulation that best matched the distribution of species was assumed to represent the closest fit to the true palaeoclimate. SDMs incorporating refugial distance performed significantly better for all but one studied species, and the Model for Interdisciplinary Research on Climate (MIROC) climatic simulation provided a more convincing pattern of the LGM climate than the Community Climate System Model (CCSM) simulation. Our results suggest that the projection of suitable habitat models onto past climatic conditions may yield realistic boundaries of glacial refugia, and that the current distribution of forest species in the study region is strongly associated with locations of former refugia. We inferred six major forest refugia throughout western Asia: (1) Colchis; (2) western Anatolia; (3) western Taurus; (4) the upper reaches of the Tigris River; (5) the Levant; and (6) the southern Caspian basin. The boundaries of the modelled refugia were substantially broader than the refugia boundaries inferred solely from pollen records. Thus, our method could be used to: (1) improve models of current species distributions by considering the dispersal histories of the species; and (2) validate alternative reconstructions of palaeoclimate with current distribution data. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 231–248.