Climate oscillation during the Quaternary associated with landscape heterogeneity promoted allopatric lineage divergence of a temperate tree Kalopanax septemlobus (Araliaceae) in East Asia

We investigated the biogeographic history of Kalopanax septemlobus, one of the most widespread temperate tree species in East Asia, using a combined phylogeographic and palaeodistribution modelling approach. Range-wide genetic differentiation at nuclear microsatellites (G'(ST) = 0.709; 2205 samples genotyped at five loci) and chloroplast DNA (G(ST) = 0.697; 576 samples sequenced for 2055 bp at three fragments) was high. A major phylogeographic break in Central China corresponded with those of other temperate species and the spatial delineation of the two temperate forest subkingdoms of East Asia, consistent with the forests having been isolated within both East and West China for multiple glacial-interglacial cycles. Evidence for multiple glacial refugia was found in most of its current range in China, South Japan and the southernmost part of the Korean Peninsula. In contrast, lineage admixture and absence of private alleles and haplotypes in Hokkaido and the northern Korean Peninsula support a postglacial origin of northernmost populations. Although palaeodistribution modelling predicted suitable climate across a land-bridge extending from South Japan to East China during the Last Glacial Maximum, the genetic differentiation of regional populations indicated a limited role of the exposed sea floor as a dispersal corridor at that time. Overall, this study provides evidence that differential impacts of Quaternary climate oscillation associated with landscape heterogeneity have shaped the genetic structure of a wide-ranging temperate tree in East Asia.     

山东植物区系的演变和来源

1　现代植物区系山东省位于我国东部、黄河下游 ,北濒渤海 ,东临黄海 ,地理范围介于北纬 34°2 3′～38°2 4′,东经 1 1 4°4 8′～ 1 2 2°4 3′之间。全省总面积为 1 5.72万 km2 ,占全国总面积的 1 .6%。属暖温带季风气候 ,沿海比较湿润 ,地带性植被主要是暖温带落叶阔叶林和松、栎类针阔叶混交林。山东省在中国植物区系的分区地位隶属于泛北极植物区、中国 -日本森林植物亚区、华北植物地区 [1 ]。据最近研究统计 ,现有野生维管植物 1 47科、61 4属 ,约 1 547种 (包括变种 ,下同 )。其中蕨类植物 2 4科 39属 1 0 5种 ,裸子植物 3科 3属 …     

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。     

The Quaternary evolutionary history,potential distribution dynamics,and conservation implications for a Qinghai–Tibet Plateau endemic herbaceous perennial,Anisodus tanguticus (Solanaceae)

Various hypotheses have been proposed about the Quaternary evolutionary history of plant species on the Qinghai–Tibet Plateau (QTP), yet only a handful of studies have considered both population genetics and ecological niche context. In this study, we proposed and compared climate refugia hypotheses based on the phylogeographic pattern of Anisodus tanguticus (three plastid DNA fragments and nuclear internal transcribed spacer regions from 32 populations) and present and past species distribution models (SDMs). We detected six plastid haplotypes in two well‐differentiated lineages. Although all haplotypes could be found in its western (sampling) area, only haplotypes from one lineage occurred in its eastern area. Meanwhile, most genetic variations existed between populations (F_ST = 0.822). The SDMs during the last glacial maximum and last interglacial periods showed range fragmentation in the western area and significant range contraction in the eastern area, respectively, in comparison with current potential distribution. This species may have undergone intraspecific divergence during the early Quaternary, which may have been caused by survival in different refugia during the earliest known glacial in the QTP, rather than geological isolation due to orogenesis events. Subsequently, climate oscillations during the Quaternary resulted in a dynamic distribution range for this species as well as the distribution pattern of its plastid haplotypes and nuclear genotypes. The interglacial periods may have had a greater effect on A. tanguticus than the glacial periods. Most importantly, neither genetic data nor SDM alone can fully reveal the climate refugia history of this species. We also discuss the conservation implications for this important Tibetan folk medicine plant in light of these findings and SDMs under future climate models. Together, our results underline the necessity to combine phylogeographic and SDM approaches in future investigations of the Quaternary evolutionary history of species in topographically complex areas, such as the QTP.     

Phylogeographic insights into cryptic glacial refugia

The glacial episodes of the Quaternary (2.6 million years ago-present) were a major factor in shaping the present-day distributions of extant flora and fauna, with expansions and contractions of the ice sheets rendering large areas uninhabitable for most species. Fossil records suggest that many species survived glacial maxima by retreating to refugia, usually at lower latitudes. Recently, phylogeographic studies have given support to the existence of previously unknown, or cryptic, refugia. Here we summarise many of these insights into the glacial histories of species in cryptic refugia gained through phylogeographic approaches. Understanding such refugia might be important as the Earth heads into another period of climate change, in terms of predicting the effects on species distribution and survival.     

Molecular phylogeography and intraspecific divergence of Spiraea alpina (Rosaceae) distributed in the Qinghai-Tibetan Plateau and adjacent regions inferred from nrDNA

The Qinghai-Tibetan plateau (QTP) uplift had a decisive effect on climatic and eco-environmental evolution in East Asia during the Quaternary. In the current study phylogeographic structure and diversification history of Spiraea alpina across the QTP were investigated for the first time based on nuclear internal transcribed spacer. The nuclear internal transcribed spacers (ITS_1a–ITS₄) were generated for a total of 284 individuals distributed within 31 natural populations. A clear phylogeographic structure was found for S. alpina. The results showed that this species colonized in three different glacial refugia during the Quaternary extensive glaciation and expanded during the Interglacial period. Analysis of molecular variance (AMOVA) showed 74.13% genetic diversity among populations and 25.87% genetic variation within populations with distinct phylogeographic structure (F_ST = 0.741*). The estimated divergence time revealed that the main lineages of S. alpina diversified during the Quaternary 1.2–0.6 million years ago. The study concluded that severe climatic oscillations during Quaternary and the uplift of QTP had a profound effect on intraspecific divergence of S. alpina.     

The Tertiary Vegetation and Flora and Floristic Regions in China

The development or adaptation radiation of angiosperms has been closely related to the paleoclimate and paleoenvironment. Several environmental factors, influencing each other, have affected the evolution or development of angiosperms, which falls into different stages. The lines between these stages are not consistent with the traditional geological ages. Floristic regionalization might be better based on the stages of the development. Angiosperms have become important or dominant elements in the flora since the middle and late period of the Late Cretaceous. The development of angiosperms may be divided into four stages: (1) Initial stage: The angiosperms were not yet well established during the Early Cretaceous. The number of species and individuals was limited, the leaf small, mostly with entire margine, irregular venation and poor differentiation of venation; (2) Flourished stage: The angiosperms increased dramatically and became abundant. The leaf was larger with more regular venation and the differentiation of venation was remarkable. The percentage of angiosperms was from 40%-60% in the flora, and eventually arose, becoming predominant, with important families well presented. This was probably from the Late Cretaceous to the Early Tertiary; (3) Herbs-Flourished stage: Some of woody plants were extinct, while herbs greatly increased, because of the changes in climate, sea level and mountain uplift during the Neogene period; (4)Quaternary stage: The mountain glacier in China, influenced by global glacial-interglacial alternation due to the climatic fluctuation, advanced and regressed during the Quaternary. The climatic fluctuation apparently affected the distribution of plants. But the components of the flora were similar to or slightly different from the features of the present one. The aim of the present paper is to discuss the floristic regions at the second and third stages. The Late Cretaceous-Early Tertiary stage: A. The Late Cretaceous-Paleogene flora of northern China 1. Warm-temperate to subtropical deciduous broad-leaved and conifer forests in North and Northeastern China 2. Dry subtropical flora of Northwestern and Central China B. Southern China Late Cretaceous-Paleogene flora 3. Deciduous and evergreen forests with coniferous elements in the subtropical coastal region of east China 4. Evergreen forests of South China 5. Evergreen forests of lower mountains in the Tethys The Neogene stage-Herbs-flourished stage: 1. Temperate forests and grasslands to semidesert-desert floras of northwestern China; 2. Warm temperate deciduous forests of north and northeastern China; 3. Warm temperate to subtropical deciduous and evergreen forests of central and east China; 4. Evergreen forests of the subtropical and tropics mangrove vegetation of south China; 5. Subtropical evergreen and deciduous forests on the Yunnan and Xizang plateau.     

长江中游(以湖北湖南为主)的植物生物多样性及其保护对策

在对植物调查研究的基础上,对长江中游(湖北、湖南为主)的植物区系、植被、生物多样性保护及保护对策进行了系统的论述。根据两省土著种子植物名录统计,本区共有202科1476属7037种(包括种下等级),其中裸子植物7科30属64种;被子植物196科1445属6973种。以鄂湘为代表的华中区分布类型复杂、物种丰富、起源古老,而且特有的科、属、种多,特有度高。北温带木本植物属高度集中,体现了长江中游具有古第三纪．泛北极植物区系的代表性。拥有众多的东亚特有属、东亚．北美间断分布属和中国特有属,它们既是本地植物区系的特色,也代表了中国植物区系的核心。本地中山以亮叶水青冈(Fagus lucida)为主的阔叶林系第三纪．泛北中生落叶阔叶林的后裔。在植被区划上,本区属于常绿阔叶林区域,包括北亚热带常绿阔叶．落叶阔叶混交林地带及中亚热带常绿阔叶林两个植被地带。后者分为北部典型常绿阔叶林亚地带,南部含华南植物区系成分的常绿阔叶林亚地带。本区主要的自然植被类型(以原生类型为主)有171个类型(相当于群系formations)。从生物多样性保护的三个层次上(物种、群落、景观)对保护现状、特点、保护方针和策略等进行了探讨并提出了建议。     

Molecular evidence for glacial refugia of mountain plants in the European Alps

Many mountain ranges have been strongly glaciated during the Quaternary ice ages, and the locations of glacial refugia of mountain plants have been debated for a long time. A series of detailed molecular studies, investigating intraspecific genetic variation of mountain plants in the European Alps, now allows for a first synopsis. A comparison of the phylogeographic patterns with geological and palaeoenvironmental data demonstrates that glacial refugia were located along the southwestern, southern, eastern and northern border of the Alps. Additional glacial refugia were present in central Alpine areas, where high-elevation plants survived the last glaciation on ice-free mountain tops. The observed intraspecific phylogeographies suggest general patterns of glacial survival, which conform to well-known centres of Alpine species diversity and endemism. This implies that evolutionary or biogeographic processes induced by climatic fluctuations act on gene and species diversity in a similar way.     

Multiple glacial refugia for cool‐temperate deciduous trees in northern East Asia: the Mongolian oak as a case study

In East Asia, temperate forests are predicted to have retracted southward to c. 30° N during the last glacial maximum (LGM) based on fossil pollen data, whereas phylogeographic studies have often suggested glacial in situ survival of cool‐temperate deciduous trees in their modern northern ranges. Here we report a study of the genetic diversity and structure of 29 natural Mongolian oak (Quercus mongolica) populations using 19 nuclear simple sequence repeat (nSSR) loci and four chloroplast DNA fragments. Bayesian clustering analysis with nSSRs revealed five groups, which were inferred by approximate Bayesian computation (ABC) to have diverged in multiple refugia through multiple glacial–interglacial cycles. Analysis of chloroplast DNA variation revealed four lineages that were largely but incompletely geographically disjunct. Ecological niche modelling (ENMs) indicated a southward range shift of the oak's distribution at the LGM, although high suitability scores were also evident in the Changbai Mts. (Northeast China), the Korean Peninsula, areas surrounding the Bohai Sea, and along the coast of the Russian Far East. In addition, endemic chloroplast DNA haplotypes and nuclear lineages occurred in high‐latitude northern areas where the ENM predicted no suitable habitat. The combined evidence from nuclear and chloroplast DNA, and the results of the ENM clearly demonstrate that multiple northern refugia, including cryptic ones, were maintained across the current distributional range of the Mongolian oak during the LGM or earlier glacial periods. Though spatially limited, postglacial expansions from these refugia have led to a pattern of decreased genetic diversity with increasing latitude.     

Allopatric divergence and phylogeographic structure of the plateau zokor (Eospalax baileyi), a fossorial rodent endemic to the Qinghai–Tibetan Plateau

Aim Most species of temperate regions are believed to have shifted to lower latitudes or elevations during the glacial periods of the Quaternary. In this study we test whether this phylogeographic assumption is also true for the plateau zokor (Eospalax baileyi), a fossorial rodent endemic to the climate-sensitive Qinghai–Tibetan Plateau (QTP), which ranges in elevation from 2600 to 4600 m. Location The QTP of western China. Methods Phylogeographic analyses were conducted based on the mitochondrial cytochrome b gene sequences of 193 individuals from 20 populations over the entire range of the species. Results A total of 54 haplotypes identified in the present study clustered into four geographically correlated clades located in the interior of the QTP (clade A) and at the plateau edge (B, C and D). Molecular calibrations suggest that the interior plateau (A) and plateau-edge (B–D) clades diverged at 1.2 Ma and that the three plateau-edge clades diverged between 0.85 and 0.80 Ma. These estimates are concordant with diastrophism and glaciation events in the QTP. Coalescent tests rejected both the hypothesis that all current populations originated from a single refugium at a low elevation during the Last Glacial Maximum (LGM) and the hypothesis that the two lineages diverged during the LGM. The tests instead supported the hypothesis that there were four refugia during the LGM, and that the four clades diverged prior to the late Pleistocene. Main conclusions Our results suggest that Quaternary diastrophisms and glaciations repeatedly promoted allopatric divergence of the plateau zokor into geographical clades, and that these regional clades subsequently persisted at high elevations, rather than migrating to the low-elevation plateau edge during subsequent glacial ages.     

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.     

The middle Miocene palynoflora and palaeoenvironments of Eskihisar (Yatağan basin,south‐western Anatolia): a combined LM and SEM investigation

We investigated a palynological section from middle Miocene sediments at Eskihisar (south‐western Anatolia) to establish biogeographic links of the palynoflora and to infer the palaeoenvironment. Four algal palynomorphs, nine spore taxa, eight gymnosperms, three monocots and 67 dicot pollen types were encountered and investigated using the ‘single grain method’ that combines light microscopy and scanning electron microscopy. Two pollen zones reflect different phases of basin development. Zonal vegetation remained fairly stable across the section and reflects heterogeneous environments including broad‐leaved deciduous forest, subtropical forest and sclerophyllous and semi‐evergreen oak forest. Conifers were accessory elements in the broad‐leaved deciduous forest communities and replaced these at higher elevations. Some herbaceous taxa (Plumbaginaceae) indicate scattered occurrences of sandy and/or rocky soils. Biogeographic affinities are general Northern Hemisphere, North American and East Asian, as also suggested by the macrofossil record. Only two taxa provide potential biogeographic links with the African flora. This suggests that biome shifts of plant taxa between African subtropical/tropical biomes and Anatolian (western Eurasian) temperate forests and shrublands may have been rare in the middle Miocene.     

Evidence of a highly complex phylogeographic structure on a specialist river bird species, the dipper (Cinclus cinclus)

This study details the phylogeographic pattern of the white-throated dipper (Cinclus cinclus), a Palearctic, temperate, passerine bird that is exclusively associated with flowing water. Our results reveal a complex phylogeographic structure with at least five distinct lineages for the Western Palearctic region. As for many species of the Western Palearctic fauna and flora, this genetic structure is probably linked to the isolation of populations in different southern refuges during glacial periods. Furthermore, the isolation of populations in Scandinavia and/or Eastern regions, but also in Morocco and probably in Corsica, was accentuated by ecological and biogeographic barriers during Quaternary interglacial periods. During glacial periods, Italy, Sicily and the Balkano-Carpathian region acted as major refuge zones for the dipper. At the end of the last ice age, Western Europe was repopulated by dippers from an Italian refuge, while Eastern Europe was recolonised by Balkano-Carpathian birds. A large contact zone between these two lineages was evidenced and extends from Luxembourg to Hungary. Finally, our results indicate the need to clarify the taxonomic status of the dipper, especially concerning the European subspecies whose validity appears uncertain.     

东亚暖温带落叶阔叶林的起源

以对北京百花山种子植物区系起源的分析为例,探讨了暖温带落叶阔叶林的起源,认为:世界暖温带落叶阔叶林在植物区系起源上是统一的,东亚暖温带落叶阔叶林是基本类型,中国西南的亚热带山地是它们的发源地。     

云南中山湿性常绿阔叶林起源的探讨

云南中山湿性常绿阔叶林广泛分布于海拔1800~3400 m的山地,这类常绿阔叶林在生态外貌上具有亚热带常绿阔叶林特征,但其分布生境却是暖温带-温带气候,且植物区系组成与热带植物区系有古老的渊源。该文以研究得比较深入的哀牢山中山湿性常绿阔叶林为例,对其植物区系与其气候条件的不协调进行剖析。哀牢山的中山湿性常绿阔叶林记录了种子植物110科386属821种。在植物区系科的地理成分上,无论是含种数多的科,还是所有科的分布区类型,都是以热带分布科为主,体现了它的远古热带起源背景。在该种子植物区系中,热带分布属占总属数的47.75%,热带分布种占总种数的33.45%,在近代演化上仍体现出明显的热带亲缘特征。按照哀牢山中山湿性常绿阔叶林分布地区的暖温带-温带气候条件,它相当于中国东部的暖温带落叶阔叶林地区。在中国东部地区类似气候条件的地带性植被的植物区系中,热带分布属通常只占总属数的25%~27%,云南中山湿性常绿阔叶林植物区系的地理成分与所在地的温带气候明显不协调。对这种现象的解释,笔者认为结合哀牢山地区曾经是低海拔夷平面,在第四纪以来才迅速抬升形成的地质历史,可能这里曾经在第四纪以前的热带-亚热带性质的植物区系和常绿阔叶林随着地形的迅速抬升,逐渐适应了因海拔上升而形成的温带气候,成为在云南高海拔山地存在的所谓中山湿性常绿阔叶林,其植物区系和植被的热带-亚热带性质及常绿阔叶生态外貌并未因气候改变而发生改变。     

海南岛第三纪沉积环境与古植物群落的多样性及其变迁

海南岛第三纪沉积类型及古植物群落复杂多样,早第三纪主要为河湖泊,湖泊相和湖沼相等沉积,晚期发生海侵,开始出现海陆交相沉积;古植物群落主要有：温带山地针叶林和落叶阔叶林,亚热带常绿和落叶阔叶林,晚晚开始出现红树林,总体属温暖湿珠亚热带气候,晚第三纪海侵加大,沉积为型为滨海相,滨海－浅海相,浅海相及深水广海相等海相沉积,在滨海地区红树林面积大大增加,盆地中心平原地上展为低地热带雨林,总体属炎热潮湿的热带气候,但在海南岛西部和西北部仍发育有山地雪松和云杉林,上新世时植被类型逐渐接近现今当地的面貌,深入研究本区第三纪古环境与古植物群落的变迁对海南岛现代生物多样性形成机制的研究具有重要意义。     

Persistence in peripheral refugia promotes phenotypic divergence and speciation in a rainforest frog

It is well established from the fossil record and phylogeographic analyses that late Quaternary climate fluctuations led to substantial changes in species' distribution, but whether and how these fluctuations resulted in phenotypic divergence and speciation is less clear. This question can be addressed through detailed analysis of traits relevant to ecology and mating within and among intraspecific lineages that persisted in separate refugia. In a biogeographic system (the Australian Wet Tropics [AWT]) with a well-established history of refugial isolation during Pleistocene glacial periods, we tested whether climate-mediated changes in distribution drove genetic and phenotypic divergence in the rainforest frog Cophixalus ornatus. We combined paleomodeling and multilocus genetics to demonstrate long-term persistence within, and isolation among, one central and two peripheral refugia. In contrast to other AWT vertebrates, the three major lineages differ in ecologically relevant morphology and in mating call, reflecting divergent selection and/or genetic drift in the peripheral isolates. Divergence in mating call and contact zone analyses suggest that the lineages now represent distinct species. The results show that climate shifts can promote genetic and phenotypic divergence and, potentially, speciation and direct attention toward incorporating adaptive traits into phylogeographic studies to better resolve the mechanisms of speciation.     

罗霄山脉种子植物区系的特有现象与残遗现象

罗霄山脉种子植物区系非常丰富, 是连接华东、华南、华中植物区系的重要通道, 区系组成具有明显的古老性和过渡性特征。本文针对该区域种子植物的特有现象和残遗现象, 分析了其组成特点及形成原因。结果表明: (1)罗霄山脉共有中国特有科3科, 特有属55属, 特有种1,624种, 罗霄山脉区域特有种43种7变种; 特有属中以古特有属为主, 并以温带成分占优势(达55.91%), 体现出明显的古老性和孑遗性。(2)罗霄山脉共有孑遗属165属, 以热带亚洲分布、东亚-北美间断分布、东亚特有及中国特有成分为主, 包括木本属132属。(3)受中新世气候波动以及第四纪冰期的影响, 许多北方热带植物群的常绿成分、北极-第三纪的落叶成分, 在罗霄山脉得以保存下来成为古特有和孑遗成分。特有现象、残遗现象的分析结果表明罗霄山脉是一个重要的生物避难所, 其对中国东部植物区系的保存和重新扩散具有重要意义。     

Molecular evidence for glacial expansion and interglacial retreat during Quaternary climatic changes in a montane temperate pine (Pinus kwangtungensis Chun ex Tsiang) in southern China

It has been suggested that the eastern Asian temperate flora have responded to Quaternary climatic changes in a different way compared with temperate plants in Europe and Northern America. However, knowledge about their phylogeographic structure and evolutionary history is still limited. In this study we investigated mitochondrial DNA variation in 17 populations of Pinus kwangtungensis, a five-needled pine inhabiting in isolated mountains of southern China and northern Vietnam. A total of ten mitochondrial haplotypes (mitotypes) were characterized by the polymorphisms of two fragments (cox-1-2 and nad7 intron1), and total genetic diversity was high (h _T = 0.847). The construction of phylogenetic relationships of the ten mitotypes detected three major, well-distinct clades, largely corresponding to four population groups identified by SAMOVA analysis. SAMOVA also indicated that most genetic variance should be attributed to among-group differentiation (F _CT = 0.868), consistent with the substantial genetic structure found within P. kwangtungensis (G _ST = 0.751). The genetic distances of P. kwangtungensis weakly but significantly correlated with geographical distances (R = 0.228, P = 0.03), revealing a pattern of isolation-by-distance. Demographic analysis did not detect any bottleneck events in the recent history of P. kwangtungensis. These results clearly suggested that there were three major refugia for this montane temperate conifer during warm stages (along with other minor refugia), and that the species would have tracked Quaternary climatic changes by expanding to nearby lowlands in the glacial periods, and by retreating to mountaintops (the refugia) during interglacial warmer stages, experiencing fragmentation and isolation among refugia.