Quaternary refugia are associated with higher speciation rates in mammalian faunas of the Western Palaearctic

Quaternary climate changes have contributed to shape the biogeographic distribution of extant species. The combination of climatic niche conservatism and glacial‐interglacial cycles forced many species to retract their range limits for surviving under the advance of Pleistocene ice‐sheets. Refugia offered geographical opportunities for species to retreat, persist and, later on, begin recolonization processes under favourable environmental conditions. Here we explore the hypothesis that refugia have been not only crucial for the survival of multiple species but also acted as speciation centres for Western Palaearctic mammals. We define ‘recurrent massive refugia’ as those geographical regions that have historically accumulated the highest levels of co‐occurring species for several Quaternary cycles. Our assemblage‐level analyses identify the existence of refugia within the Iberian Peninsula and the Atlantic French margin that were recurrently selected by most mammals. The topographic heterogeneity, climatic stability and microhabitat availability of these refugial areas may have offered suitable habitat conditions for multiple species during different climatic events over time. Using a Bayesian analysis of macroevolutionary mixtures we detected that the higher level of divergence and accumulative evolutionary changes in mammals of the Western Palaearctic are found in refugia. The continuous retractions and expansions of species’ ranges during the Pleistocene promoted temporal changes in the composition and richness of communities in this biogeographic region. The reorganization of ecological composition driven by cyclical climatic events may have favoured the emergence of biotic interactions and ecological responses conducive to novel selective pressures. Our findings suggest, first, that multiple climatic changes in the form of glacial‐interglacial transitions during the Quaternary have left a detectable imprint on the observed geographical patterns of species richness in mammalian faunas of the Western Palaearctic, and second, highlight the importance of refugia for the preservation of species (‘museums’) and as centers of speciation and endemism (‘cradles’) as well.     

Multilocus phylogeography of the European ground squirrel: cryptic interglacial refugia of continental climate in Europe

The theory of classical and cryptic Pleistocene refugia is based mainly on historical changes in temperature, and the refugia are usually defined within a latitudinal gradient. However, the gradient of oceanic–continental climate (i.e. longitudinal) was also significantly variable during glacial cycles with important biotic consequences. Range‐wide phylogeography of the European ground squirrel (EGS) was used to interpret the evolutionary and palaeogeographical history of the species in Europe and to shed light on its glacial–interglacial dynamic. The EGS is a steppe‐inhabiting species and the westernmost member of the genus in the Palaearctic region. We have analysed 915 specimens throughout the present natural range by employing mitochondrial DNA sequences (cytochrome b gene) and 12 nuclear microsatellite markers. The reconstructed phylogeography divides the species into two main geographical groups, with deep substructuring within both groups. Bulgaria is the centre of the ancestral area, and it also has the highest genetic diversity within the species. The northernmost group of the EGS survived in the southern part of Pannonia throughout several glacial–interglacial cycles. Animals from this population probably repeatedly colonized areas further to the north and west during the glacial periods, while in the interglacial periods, the EGS distribution contracted back to this Pannonian refugium. The EGS thus represents a species with a glacial expansion/interglacial contraction palaeogeographical dynamics, and the Pannonian and southeastern Balkanian steppes are supported as cryptic refugia of continental climate during Pleistocene interglacials.     

Genetic consequences of interglacial isolation in a steppe bird

In response to climate changes that have occurred during Pleistocene glacial cycles, taxa associated to steppe vegetation might have followed a pattern of historical evolution in which isolation and fragmentation of populations occurred during the short interglacials and expansion events occurred during the long glacial periods, in contrast to the pattern described for temperate species. Here, we use molecular genetic data to evaluate this idea in a steppe bird with Palaearctic distribution, the little bustard (Tetrax tetrax). Overall, extremely low genetic diversity and differentiation was observed among eight little bustard populations distributed in Spain and France. Mismatch distribution analyses showed that most little bustard populations expanded during cooling periods previous to, and just after, the last interglacial period (127,000-111,000 years before present), when steppe habitats were widespread across Europe. Coalescent-based methods suggested that glacial expansions have resulted in substantial admixture in Western Europe due to the existence of different interglacial refugia. Our results are consistent with a model of evolution and genetic consequences of Pleistocene cycles with low between-population genetic differentiation as a result of short-term isolation periods during interglacials and long-term exchange during glacial periods.     

Phylogeography of Potamon ibericum (Brachyura: Potamidae) identifies Quaternary glacial refugia within the Caucasus biodiversity hot spot

Refugia are critical for the maintenance of biodiversity during the periods of Quaternary climatic oscillations. The long‐term persistence of refugial populations in a large continuous refugium has resulted in a homogenous pattern of genetic structure among populations, while highly structured evolutionary lineages characterize the restriction of refugial populations to smaller subrefugia. These mechanisms have resulted in the identification of hot spots of biodiversity within putative glacial refugia. We studied phylogeography of Potamon ibericum (Brachyura: Potamidae) in the drainages of the western Caucasus biodiversity hot spot (i.e., Colchis and the Caucasus) to infer spatial genetic structure and potential refugia for a freshwater crab in this region. These areas have traditionally considered as a refugium due to the presence of Tertiary relict species. We integrated population genetic data and historical demographic analysis from cytochrome oxidase subunit I sequences and paleoclimatic data from species distribution modeling (SDM). The results revealed the lack of phylogeographic structure and provided evidence for demographic expansion. The SDM presented a rather homogenous and large refugium that extended from northeast Turkey to Colchis during the last glacial period. In contrast to these findings, previous phylogeographic study on P. ibericum of the eastern Caucasus biodiversity hot spot (i.e., Hyrcania) identified multiple independent refugia. By combining these results, we explain the significance of this important western Palearctic hot spot of biological diversity in shaping the geographic distribution of intraspecific genetic diversity in a freshwater taxon.     

Alpine biogeography of Parnassian butterflies during Quaternary climate cycles in North America

Growth of alpine glaciers during the Pleistocene had profound effects on montane landscapes in North America and the organisms now inhabiting alpine ecosystems. Biogeography of this region has often been viewed as a system of sky islands despite the fact that species richness patterns deviate from a strict island biogeographic model. One explanation is that alpine species are not in equilibrium because of late Quaternary geographic range shifts. Genetic data can provide evidence of nonequilibrium dynamics and the distributional shifts that occur during glaciation events in alpine landscapes. Using mitochondrial and nuclear sequence data, we examine the evolutionary history of butterflies in the Parnassius phoebus complex. We test explicit, alternative models of the biogeographic history of Parnassius smintheus and Parnassius behrii , including an equilibrium island model, ancestral radiation and fragmentation, an expanding alpine archipelago and an alpine archipelago refuge model. Our results support the alpine archipelago refuge model, in which alpine butterflies undergo population contraction during glacial climates followed by population expansion during interglacial phases. While butterflies can disperse between distant mountain ranges during glacial periods, gene flow is rare. We find evidence of recent connectivity between California and Colorado, population expansion events following deglaciation ∼20 000 years B.P., and small population sizes during the last glacial period. An analysis of lineage splitting suggests that morphological differences in P. smintheus and P. behrii are the result of late Pleistocene divergence (∼48 000 years B.P.) with limited gene flow. Our results demonstrate that spatially complex and nonequilibrium population dynamics influence alpine diversity patterns.     

Stable persistence of relict populations involved evolutionary shifts of reproductive characters in the genus Tanakaea (Saxifragaceae)

Tertiary relicts often show evolutionary stasis in morphology and ecology and have been hypothesized to retain stable population sizes in refugia. However, recent studies have reported that some relicts evolutionarily shifted their physiology, ecology, and morphology and experienced various patterns of demography. To understand the historical survival of relict plants, a multidimensional study investigating the evolution of ecological and morphological traits as well as population demographic history is needed. The genus Tanakaea (Saxifragaceae) comprises two species in China and Japan. These species share most vegetative characteristics and are sometimes treated as a single species. The distribution pattern is relictual, as the populations are confined to small areas in mesic warm temperate forests less influenced by Quaternary glacial climates. Focusing on the relictual plant group, this study tested the hypotheses of evolutionary stasis and population stability in long-term refugia. Genetic analyses using plastome sequences and genome-wide single-nucleotide polymorphisms revealed divergence of the two species approximately 6.8 million years ago and strong genetic differentiation of the regional populations. Demographic analysis revealed that almost all populations retained stable population sizes during glacial–interglacial climate changes, supporting the traditional view. However, morphological assessments revealed a simultaneous shift in breeding systems (from hermaphrodite to dioecy/non-clonal to clonal reproduction) in Japanese species and intraspecific differentiation of leaf traits. Therefore, the relict species do not show evolutionary stasis in every aspect. Changes in reproductive characteristics may have contributed to their long-term in situ survival.     

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.     

Phylogeography of the Japanese pipistrelle bat,Pipistrellus abramus,in China: the impact of ancient and recent events on population genetic structure

The influence of Pleistocene climatic oscillations on shaping the genetic structure of Asian biota is poorly known. The Japanese pipistrelle bat occurs over a wide range in eastern Asia, from Siberia to Japan. To test the relative impact of ancient and more recent events on genetic structure in this species, we combined mitochondrial (cytochrome b) and microsatellite markers to reconstruct its phylogeographic and demographic history on continental China and its offshore islands, Hainan Island and the Zhoushan Archipelago. Our mitochondrial DNA tree recovered two divergent geographical clades, indicating multiple glacial refugia in the region. The first clade was mainly confined to Hainan Island, indicating that gene flow between this population and the continent has been restricted, despite being repeatedly connected to the mainland during repeated glacial episodes. By contrast, haplotypes sampled on the Zhoushan Archipelago were mixed with those from the mainland, suggesting a recent shared history of expansion. Although microsatellite allele frequencies showed clear discontinuities across the sampling range, supporting the current isolation of both Hainan Island and the Zhoushan Archipelago, we also found clear evidence of more recent back colonization, probably via post‐glacial expansion or, in the latter case, occasional long distance dispersal. The results obtained highlight the importance of using multiple sets of markers for teasing apart the roles of ancient and more recent events on population genetic structure. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 582–594.     

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 effects of the Late Quaternary glacial–interglacial cycles on Anatolian ground squirrels: range expansion during the glacial periods?

The present study aimed to understand how Anatolian ground squirrels, Spermophilus xanthoprymnus (Bennett, 1835), have responded to global climate changes through the Late Quaternary glacial–interglacial cycles. Accordingly, ecological niche modelling was used, together with molecular phylogeography. Using species occurrence data compiled from field observations and relevant sources and the maximum entropy machine learning algorithm in MAXENT, an ecological niche model was developed to predict the potential geographical distribution of S. xanthoprymnus under reconstructed past (the Last Interglacial, approximately 130 000–116 000 years ago and the Last Glacial Maximum, 21 000 years ago) and present (1950–2000) bioclimatic conditions. In addition, using cytochrome b mitochondrial DNA sequences deposited in GenBank and the Bayesian skyline plot in BEAST, demographic events (population fluctuations) were further assessed over the history of Anatolian ground squirrels. Combined ecological niche modelling and molecular phylogeography revealed that S. xanthoprymnus, itself also a temperate (mid‐latitude) species, has responded to global climate changes through the Late Quaternary glacial–interglacial cycles in a fashion converse to that of most temperate (mid‐latitude) species: its range expanded rather than contracted during the glacial periods and contracted rather than expanded during the interglacial periods. In other words, Anatolian ground squirrels have been in refugia during the interglacial periods, suggesting that the classical paradigm of glacial range contraction and interglacial range expansion for temperate species may not be as general as previously assumed. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 19–32.     

Glacial refugia and the phylogeography of Steller's sea lion (Eumatopias jubatus) in the North Pacific

Mitochondrial DNA sequence data were used to examine the phylogeographic history of Steller's sea lions (Eumetopias jubatus) in relation to the presence of Plio-Pleistocene insular refugia. Cytochrome b and control region sequences from 336 Steller's sea lions reveal phylogenetic lineages associated with continental refugia south of the ice sheets in North America and Eurasia. Phylogenetic analysis suggests the genetic structure of E. jubatus is the result of Pleistocene glacial geology, which caused the elimination and subsequent reappearance of suitable rookery habitat during glacial and interglacial periods. The cyclic nature of geological change produced a series of independent population expansions, contractions and isolations that had analogous results on Steller's sea lions and other marine and terrestrial species. Our data show evidence of four glacial refugia in which populations of Steller's sea lions diverged. These events occurred from approximately 60,000 to 180,000 years BP and thus preceded the last glacial maximum.     

Mountainous genus Anterastes (Orthoptera,Tettigoniidae): autochthonous survival across several glacial ages via vertical range shifts

Although the high‐latitude range margins in Europe and North America are intensively studied, attention is gradually turned towards the taxa/populations inhabiting glacial refugia. Here, we evaluate the genealogical history of the cold‐adapted Anatolio‐Balkan genus Anterastes especially to test the possible effects of intrarefugial vertical range shifts during climatic oscillations of the Quaternary. Using concatenated data from sequences of COI+16S and ITS1–5.8S–ITS2, intrageneric relationships and the time of speciation events were estimated. Thirteen different demographic analyses were performed using a data set produced from sequences of 16S. Different phylogenetic analyses recovered similar lineages with high resolution. The molecular chronogram estimated speciation events in a period ranging from 5.60 to 1.22 Myr. Demographic analyses applied to 13 populations and five lineages suggested constant population size. Genetic diversity is significantly reduced in a few populations, while not in others. Fixation indices suggested extremely diverged populations. In the light of these data, the following main conclusions were raised: (i) although glacial refugia are the biodiversity hotspots, species level radiation of the cold‐adapted lineages is mainly prior to the Mid‐Pleistocene transition; (ii) heterogeneous topography provides refugial habitats and allows populations to survive through vertical range shifts during climatic fluctuations; (iii) prolonged isolation of refugial populations do not always result in reduced intrapopulation diversity, but in high level of genetic differentiation; (iv) the cold‐adapted lineages with low dispersal ability might have not colonised the area out of Anatolian refugium during interglacial periods; and (v) populations of invertebrates may have restricted ranges, but this does not mean that they have small effective population size.     

Has past climate change affected cold‐specialized species differentially through space and time?

Quaternary climate change has been strongly linked to distributional shifts and recent species diversification. Montane species, in particular, have experienced enhanced isolation and rapid genetic divergence during glacial fluctuations, and these processes have resulted in a disproportionate number of neo‐endemic species forming in high‐elevation habitats. In temperate montane environments, a general model of alpine population history is well supported, where cold‐specialized species track favourable climate conditions downslope during glacial episodes and upslope during warmer interglacial periods, which leads to a climate‐driven population or species diversification pump. However, it remains unclear how geography mediates distributional changes and whether certain episodes of glacial history have differentially impacted rates of diversification. We address these questions by examining phylogenomic data in a North American clade of flightless, cold‐specialized insects, the ice crawlers (Insecta: Grylloblattodea: Grylloblattidae: Grylloblatta). These low‐vagility organisms have the potential to reveal highly localized refugia and patterns of spatial recolonization, as well as a longer history of in situ diversification. Using continuous phylogeographic analysis of species groups, we show that all species tend to retreat to nearby low‐elevation habitats across western North America during episodes of glaciation, but species at high latitude exhibit larger distributional shifts. Lineage diversification was examined over the course of the Neogene and Quaternary periods, with statistical analysis supporting a direct association between climate variation and diversification rate. Major increases in lineage diversification appear to be correlated with warm and dry periods, rather than with extreme glacial events. Finally, we identify substantial cryptic diversity among ice crawlers, leading to high endemism across their range. This diversity provides new insights into highly localized glacial refugia for cold‐specialized species across western North America.     

Population Dynamics Following the Last Glacial Maximum in Two Sympatric Lizards in Northern China

Phylogeographic studies of Eremias lizards （Lacertidae） in East Asia have been limited, and the impact of major climatic events on their population dynamics remains poorly known. This study aimed to investigate population histories and refugia during the Last Glacial Maximum of two sympatric Eremias lizards （E. argus and E. brenchleyi） inhabiting northern China. We sequenced partial mitochondrial DNA from the ND4 gene for 128 individuals of E. argus from nine localities, and 46 individuals of E. brenchleyi from five localities. Forty-four ND4 haplotypes were determined from E. argus samples, and 33 from E. brenchleyi samples. Population expansion events began about 0.0044 Ma in E. argus, and 0.031 Ma in E. brenchleyi. The demographic history of E. brenchleyi indicates a long-lasting population decline since the most recent common ancestor, while that of E. argus indicates a continuous population growth. Among-population structure was significant in both species, and there were multiple refugia across their range. Intermittent gene flow occurred among expanded populations across multiple refugia during warmer phases of the glacial period, and this may explain why the effective population size has remained relatively stable in E. brenchleyi and grown in E. argus.     

DNA barcoding reveals 24 distinct lineages as cryptic bird species candidates in and around the Japanese Archipelago

DNA barcoding using a partial region (648 bp) of the cytochrome c oxidase I (COI) gene is a powerful tool for species identification and has revealed many cryptic species in various animal taxa. In birds, cryptic species are likely to occur in insular regions like the Japanese Archipelago due to the prevention of gene flow by sea barriers. Using COI sequences of 234 of the 251 Japanese‐breeding bird species, we established a DNA barcoding library for species identification and estimated the number of cryptic species candidates. A total of 226 species (96.6%) had unique COI sequences with large genetic divergence among the closest species based on neighbour‐joining clusters, genetic distance criterion and diagnostic substitutions. Eleven cryptic species candidates were detected, with distinct intraspecific deep genetic divergences, nine lineages of which were geographically separated by islands and straits within the Japanese Archipelago. To identify Japan‐specific cryptic species from trans‐Paleartic birds, we investigated the genetic structure of 142 shared species over an extended region covering Japan and Eurasia; 19 of these species formed two or more clades with high bootstrap values. Excluding six duplicated species from the total of 11 species within the Japanese Archipelago and 19 trans‐Paleartic species, we identified 24 species that were cryptic species candidates within and surrounding the Japanese Archipelago. Repeated sea level changes during the glacial and interglacial periods may be responsible for the deep genetic divergences of Japanese birds in this insular region, which has led to inconsistencies in traditional taxonomies based on morphology.     

Comparative phylogeography of two coastal polychaete tubeworms in the Northeast Atlantic supports shared history and vicariant events

The historic processes which have led to the present-day patterns of genetic structure in the marine coastal fauna of the Northeast Atlantic are still poorly understood. While tectonic uplifts and changes in sea level may have caused large-scale vicariance, warmer conditions during glacial maxima may have allowed pockets of diversity to persist to a much wider extent than in the Northwestern Atlantic. The large-scale geographic distribution of deeply divergent lineages of the coastal polychaete tubeworms Pectinaria koreni (two clades) and Owenia fusiformis (three clades) were compared using a fragment of the mitochondrial cytochrome oxidase I gene (mtCOI). All lineages were present along the biogeographic transition zone on the north coast of Brittany (France) and we found evidence pointing towards congruence in the timing of cladogenic events between Pectinaria sp. (P. auricoma/P. belgica and P. koreni) and Owenia sp., suggesting a shared history of vicariant events. More conserved 16SrRNA sequences obtained from four species of Pectinariidae together with mtCOI sequences of P. koreni seem consistent with an initial establishment of pectinariids in the north, and a southward colonization of the Northeast Atlantic. Phylogeographic patterns in O. fusiformis were also consistent with a north/south pattern of lineage splitting and congruent levels of divergence were detected between lineages of both species. We observed signatures of both persistence in small northern glacial refugia, and of northwards range expansion from regions situated closer to the Mediterranean. However, whether the recolonization of the Northeast Atlantic by both species actually reflects separate interglacial periods is unclear with regards to the lack of molecular clock calibration in coastal polychaete species.     

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.     

Quaternary refugia in southwestern Iran: insights from two sympatric moth species (Insecta, Lepidoptera)

Alternating glacial and interglacial periods led to range shifts (contractions and expansions), persistence in distinct glacial refugia and extinction events in various temperate organisms. Today, the integrative analysis of molecular markers and spatial distribution models conducted for multiple taxa allows the detection of phylogeographical patterns, thus reconstructing major biogeographical events in their shared evolutionary history. In this study, the effects of past climate change on the evolutionary history of two sympatric moth species (Gnopharmia colchidaria s.l. and G. kasrunensis) and their host plants (Prunus scoparia and P. fenzliana) were inferred for the largely neglected biodiversity hot spot Iran. We complementarily analyzed the population structure of both moth species (187 specimens, based on COI) in congruence with batched species distribution models (SDMs) for all four taxa and for the times of the Last Glacial Maximum (21 ky BP), 6 ky BP and today. Coincidence of SDMs and the distribution of haplotype lineages indicated a shared refugium for the southwestern Zagros Mountains and potential species-specific refugial areas in the southern Caucasus and the Kope-Dagh Mountains. Both moth species experienced past population expansion.     

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.     

Decades-long phylogeographic issues: complex historical processes and ecological factors on genetic structure of alpine plants in the Japanese Archipelago

Mountain regions are important places for biodiversity, where organisms could persist throughout prolonged periods and accumulate genetic divergence as well as promote speciation. Roles of mountains for biodiversity have been exclusively discussed in regions that have specifically diverse species or covered with ice-sheets during the Pleistocene glacial periods, whereas the importance of mountainous regions in East Asia has been less disputed. High mountains in the Japanese Archipelago, located at the eastern edge of the Eurasia continent, have one of southernmost populations of alpine and arctic-alpine plants that are also distributed in the northern Pacific and/or the circumarctic regions. Phylogeographic studies on the Japanese alpine plants have excluded their possible ephemeral occurrence during the current warm period, and rather, suggest persistence of alpine plants throughout several cycles of climate changes in the Pleistocene on high mountains in central Honshu, the main island of the Japanese Archipelago. In this review, I look through decade long phylogeographic studies and show complicated patterns of range dynamics of Japanese alpine plants. In addition, I note recent findings of genetic relationships of Japanese populations of alpine and/or arctic-alpine plants with those in northern regions and their possible ecological divergence in the Japanese Archipelago. Taken together, I provide several issues for understanding historical processes that established distribution of alpine plants following climate changes as well as their diversification and propose importance of Japanese populations of alpine plants on biodiversity in alpine communities across broader range, especially in the northern Pacific region.