The importance of marginal population hotspots of cold‐adapted species for research on climate change and conservation

Areas hosting hotspots of low‐latitude marginal populations of cold‐adapted plant species could be key areas for understanding geographical attributes that result in refugia during climatic shifts as well as the conservation of genetic diversity in the face of climate change. Low‐latitude populations of cold‐adapted plants are important because they may harbour the combination of alleles that foster persistence in a warmer climate. Consequently, identification of areas where arctic‐alpine, circumpolar and circumboreal species reach the low‐latitude ends of their distribution will present a unique opportunity to uncover processes that shaped current biogeographical patterns, as well as prepare for future scenarios. Here, we identify 35 main marginal population hotspots (19 and 16 areas in North America and Europe, respectively) of 183 plant taxa. These hotspots represent areas where southern marginal populations of cold‐adapted species co‐occur. The identification of hotspots was based on geographic overlap of southernmost locations of the target species, in a 50 × 50 km grid. With a threshold of two species in a single grid cell or in two contiguous cells, the analysis revealed that hotspots are in most cases located in the southern portion of major mountain chains. However, hotspots also occur in lowland areas at high latitudes (Fennoscandia, Alaska, Hudson Bay) which do not necessarily correspond to known cold‐ or warm‐stage refugia (e.g. Alps). Rockies and Sierra Nevada both in California and Spain, Apennines, and the southern Scandes, maintain their hotspot status even with more stringent cut‐off thresholds (>3 and >5 species per cell group). From a conservation point of view, our analysis reveals that only a small portion of the hotspots are currently included within protected areas. We discuss the importance of marginal population hotspots to future research on climate change and, finally, outline how conservation strategies can capitalize on the knowledge gained from studying climate change effects on cold‐adapted plants.     

The genetic pattern of population threat and loss: a case study of butterflies

Recent decreases in biodiversity in Europe are commonly thought to be due to land use and climate change. However, the genetic diversity of populations is also seen as one essential factor for their fitness. Genetic diversity in species across the continent of Europe has been recognized as being in part a consequence of ice age isolation in southern refugia and postglacial colonization northwards, and these phylogeographical patterns may themselves affect the adaptability of populations. Recent work on butterfly species with different refugia, colonization paths and genetic structures allows this idea to be examined. The 'chalk-hill blue' pattern is one of decreasing genetic diversity from south to north, whereas the 'woodland ringlet' pattern shows greater genetic diversity in eastern than in western lineages. Comparison of population demographic trends in species with these biogeographical patterns reveals higher rates of decrease with lower genetic diversity. This indicates reduced adaptability due to genetic impoverishment as a result of glacial and postglacial range changes. Analysis of phylogeographical pattern may be a useful guide to interpreting demographic trends and in conservation planning.     

Unravelling biodiversity,evolution and threats to conservation in the Sahara‐Sahel

Deserts and arid regions are generally perceived as bare and rather homogeneous areas of low diversity. The Sahara is the largest warm desert in the world and together with the arid Sahel displays high topographical and climatic heterogeneity, and has experienced recent and strong climatic oscillations that have greatly shifted biodiversity distribution and community composition. The large size, remoteness and long‐term political instability of the Sahara‐Sahel, have limited knowledge on its biodiversity. However, over the last decade, there have been an increasing number of published scientific studies based on modern geomatic and molecular tools, and broad sampling of taxa of these regions. This review tracks trends in knowledge about biodiversity patterns, processes and threats across the Sahara‐Sahel, and anticipates needs for biodiversity research and conservation. Recent studies are changing completely the perception of regional biodiversity patterns. Instead of relatively low species diversity with distribution covering most of the region, studies now suggest a high rate of endemism and larger number of species, with much narrower and fragmented ranges, frequently limited to micro‐hotspots of biodiversity. Molecular‐based studies are also unravelling cryptic diversity associated with mountains, which together with recent distribution atlases, allows identifying integrative biogeographic patterns in biodiversity distribution. Mapping of multivariate environmental variation (at 1 km × 1 km resolution) of the region illustrates main biogeographical features of the Sahara‐Sahel and supports recently hypothesised dispersal corridors and refugia. Micro‐scale water‐features present mostly in mountains have been associated with local biodiversity hotspots. However, the distribution of available data on vertebrates highlights current knowledge gaps that still apply to a large proportion of the Sahara‐Sahel. Current research is providing insights into key evolutionary and ecological processes, including causes and timing of radiation and divergence for multiple taxa, and associating the onset of the Sahara with diversification processes for low‐mobility vertebrates. Examples of phylogeographic patterns are showing the importance of allopatric speciation in the Sahara‐Sahel, and this review presents a synthetic overview of the most commonly hypothesised diversification mechanisms. Studies are also stressing that biodiversity is threatened by increasing human activities in the region, including overhunting and natural resources prospection, and in the future by predicted global warming. A representation of areas of conflict, landmines, and natural resources extraction illustrates how human activities and regional insecurity are hampering biodiversity research and conservation. Although there are still numerous knowledge gaps for the optimised conservation of biodiversity in the region, a set of research priorities is provided to identify the framework data needed to support regional conservation planning.     

Evolutionary melting pots: a biodiversity hotspot shaped by ring diversifications around the Black Sea in the Eastern tree frog (Hyla orientalis)

Hotspots of intraspecific genetic diversity, which are of primary importance for the conservation of species, have been associated with glacial refugia, that is areas where species survived the Quaternary climatic oscillations. However, the proximate mechanisms generating these hotspots remain an open issue. Hotspots may reflect the long‐term persistence of large refugial populations; alternatively, they may result from allopatric differentiation between small and isolated populations, that later admixed. Here, we test these two scenarios in a widely distributed species of tree frog, Hyla orientalis, which inhabits Asia Minor and southeastern Europe. We apply a fine‐scale phylogeographic survey, combining fast‐evolving mitochondrial and nuclear markers, with a dense sampling throughout the range, as well as ecological niche modelling, to understand what shaped the genetic variation of this species. We documented an important diversity centre around the Black Sea, composed of multiple allopatric and/or parapatric diversifications, likely driven by a combination of Pleistocene climatic fluctuations and complex regional topography. Remarkably, this diversification forms a ring around the Black Sea, from the Caucasus through Anatolia and eastern Europe, with terminal forms coming into contact and partially admixing in Crimea. Our results support the view that glacial refugia generate rather than host genetic diversity and can also function as evolutionary melting pots of biodiversity. Moreover, we report a new case of ring diversification, triggered by a large, yet cohesive dispersal barrier, a very rare situation in nature. Finally, we emphasize the Black Sea region as an important centre of intraspecific diversity in the Palearctic with implications for conservation.     

How well are biodiversity drivers reflected in protected areas? A representativeness assessment of the geohistorical gradients that shaped endemic flora in Japan

Protected areas function as a lifeboat that can preserve the origins and maintenance of biodiversity. We assessed the representativeness of biodiversity in existing protected areas in Japan using a distribution dataset and phylogenetic tree for 5565 Japanese vascular plant species. We first examined the overlap of species distribution with the existing protected areas and identified the minimum set representing all plant species. Second, we evaluated the relative importance of environmental variables in explaining the spatial arrangement of protected areas using a random forest model. Finally, we clarified how potential drivers of plant diversity were sufficiently captured within the protected areas network. Although the protected areas captured the majority of species, nearly half of the minimum set areas were selected from outside the existing protected areas. The locations of existing protected areas are mainly associated with geographical and socio-economic factors rather than key biodiversity features (including evolutionary distinctiveness). Moreover, critical biodiversity drivers, which include current climate, paleoclimatic stability, and geographical isolation, were biasedly emulated within the existing protected areas. These findings demonstrate that current conservation planning fails to represent the ecological and evolutionary processes relevant to species sorting, dispersal limitation, and allopatric speciation. In particular, under-representativeness of historically stable habitats that function as evolutionary hotspots or refugia in response to climate changes may pose a threat to the long-term persistence of Japan’s endemic biota. This study provides a fundamental basis for developing prioritization measures to retain species assembly processes and in situ diversification along current climatic and geohistorical gradients.     

Patterns of freshwater biodiversity in Europe: lessons from the spring snail genus Bythinella

Aim We analyse patterns of biodiversity in the spring snail genus Bythinella, a group of highly isolated and stenotopic freshwater species. We aim to test: (1) whether there are European areas of increased diversity (i.e. ‘hotspots’), (2) whether the potential hotspots inferred show qualitative differences in biodiversity characteristics such as endemicity, distinctiveness of taxa, age of lineages or degree of fragmentation, and (3) whether these hotspots match the Pleistocene refugia of Bythinella spp. Location Europe, Asia Minor. Methods The analyses are based on genetic data from 717 Bythinella specimens sampled at 194 sites. We used haplotypes as operational units in all analyses. To test hypothesis 1, mean pairwise genetic distances between Bythinella populations within each 1° × 1° geographical grid cell sampled in Europe were calculated. Within individual mountain ranges, grid cells with high diversity were grouped with neighbouring ones and hotspots were identified based on pre‐defined criteria. Then, to test hypothesis 2, different biodiversity indices of these regions were calculated and compared. Finally, to test hypothesis 3, the spatial distribution of the identified hotspots was compared with the known Pleistocene refugia of Bythinella spp. Results Five areas showed increased levels of genetic diversity: the Massif Central/Pyrenees, the western and eastern Alps, and the western and eastern Carpathians. These regions showed qualitative differences in biodiversity, with the eastern Carpathians holding the highest number of (endemic) haplotypes, the oldest and most distinct lineages and the highest degree of fragmentation. Only three of the five detected hotspots matched previously identified Pleistocene refugia for Bythinella spp. Main conclusions The genetic diversity of Bythinella spp. is not randomly distributed throughout Europe. Some of the hotspots we identify coincide with those found in other freshwater taxa; others have not previously been reported. Thus, spring organisms may reflect a unique evolutionary history that is distinct from lentic and lotic taxa. Our findings may be useful for conservation purposes even though the species‐level taxonomy of the genus is still under discussion.     

Use of cross-taxon congruence for hotspot identification at a regional scale

One of the most debated problems in conservation biology is the use of indicator (surrogate) taxa to predict spatial patterns in other taxa. Cross-taxon congruence in species richness patterns is of paramount importance at regional scales to disclose areas of high conservation value that are significant in a broader biogeographical context but yet placed in the finer, more practical, political context of decision making. We analysed spatial patterns of diversity in six arthropod taxa from the Turkish fauna as a regional case study relevant to global conservation of the Mediterranean basin. Although we found high congruence in cross-taxon comparisons of species richness (0.241相似文献   

Do hotspots fall within protected areas? A Geographic Approach to Planning analysis of regional freshwater biodiversity

相似文献   

Genetically diverse but with surprisingly little geographical structure: the complex history of the widespread herb Carex nigra (Cyperaceae)

Aim Based on extensive range‐wide sampling, we address the phylogeographical history of one of the most widespread and taxonomically complex sedges in Europe, Carex nigra s. lat. We compare the genetic structure of the recently colonized northern areas (front edge) and the long‐standing southern areas (rear edge), and assess the potential genetic basis of suggested taxonomic divisions at the rank of species and below. Location Amphi‐Atlantic, central and northern Europe, circum‐Mediterranean mountain ranges, central Siberia, Himalayas. Methods A total of 469 individuals sampled from 83 populations, covering most of the species’ range, were analysed with amplified fragment length polymorphism (AFLP) and chloroplast DNA (cpDNA) markers. Bayesian clustering, principal coordinates analysis, and estimates of diversity and differentiation were used for the analysis of AFLP data. CpDNA data were analysed with statistical parsimony networks and maximum parsimony and Bayesian inference of phylogenetic trees. Results Overall genetic diversity was high, but differentiation among populations was limited. Major glacial refugia were inferred in the Mediterranean Basin and in western Russia; in addition, there may have been minor refugia in the North Atlantic region. In the southern part of the range, we found high levels, but geographically quite poorly structured genetic diversity, whereas the levels of genetic diversity varied among different areas in the north. North American populations were genetically very similar to the European populations. Main conclusions The data are consistent with extensive gene flow, which has obscured the recent history of the taxon. The limited differentiation in the south probably results from the mixing of lineages expanding from several local refugia. Northward post‐glacial colonization resulted in a leading‐edge pattern of low diversity in the Netherlands, Belgium, Scotland and Iceland, whereas the observed high diversity levels in Fennoscandia suggest broad‐fronted colonization from the south as well as from the east. The patterns found in the American populations are consistent with post‐glacial colonization, possibly even with anthropogenic introduction from Europe. Our data also suggest that the tussock‐forming populations of C. nigra, often referred to as a distinct species (Carex juncella), represent an ecotype that has originated repeatedly from different populations with creeping rhizomes.     

Population genetic structure and diversity of the Apennine endemic stream frog, Rana italica--insights on the Pleistocene evolutionary history of the Italian peninsular biota

For most species in the Western Palaearctic region, southern Mediterranean peninsulas have been identified as major Quaternary refugia and hotspots of intraspecific diversity, and thus, as areas of particular relevance for the conservation of the evolutionary potential. We analysed the patterns of geographical variation among 26 populations of the Italian stream frog, using both nuclear (allozymes) and mitochondrial (partial cytochrome b sequences) markers. Phylogenetic, phylogeographical and population genetic analyses suggested that the species survived the last glacial–interglacial cycles in two distinct refugia, one restricted to the tip of the Calabrian peninsula, at the extreme south of the species' range, the other spanning from central Calabria to central Apennines and showing evidences for further population subdivision therein. Historical demographic tests suggested a significant population expansion from the latter, which most likely began around the last pleniglacial. This expansion would have led to the rapid colonization of the northern Apennines to the north, and to a secondary contact and population admixture with the population from the southern refugium in southern central Calabria. A comparison of the evolutionary history inferred for the Italian stream frog with the data emerging for other codistributed species suggests: (i) the generality of a multiple-refugia scenario for the Italian peninsula, (ii) the possible occurrence of at least one suture zone in southern Italy, and (iii) that for most species, this Pleistocene refugium is not only a hotspot, but also a melting pot of intraspecific genetic diversity. Finally, the conservation implications of these results are also briefly highlighted.     

Questioning the effectiveness of the Natura 2000 Special Areas of Conservation strategy: the case of Crete

Aim This study examines the effectiveness of the selected ‘network’ of Natura 2000 Special Areas of Conservation (SACs) at a regional scale in Greece, in terms of its representativeness of plant biodiversity. Location The island of Crete is used as a case study because it is considered to be one of the 10 hotspots for biodiversity in the Mediterranean Basin. Methods Hotspot analysis and complementarity algorithms are used to define priority areas for conservation and calculate their spatial overlap with the Natura 2000 SACs in Crete. Results The various categories of hotspots contain subsamples of plant categories, used for their definition. Spatial overlap among different categories of hotspots, areas of complementary diversity and Natura 2000 SCAs is low. Main conclusions The results show that the Natura 2000 SACs ‘network’ in Crete seems insufficient to ensure satisfactory representation of the regional plant biodiversity elements.     

Phylogeography and biogeography of the lower Central American Neotropics: diversification between two continents and between two seas

Lower Central America (LCA) provides a geologically complex and dynamic, richly biodiverse model for studying the recent assembly and diversification of a Neotropical biota. Here, we review the growing literature of LCA phylogeography studies and their contribution to understanding the origins, assembly, and diversification of the LCA biota against the backdrop of regional geologic and climatic history, and previous biogeographical inquiry. Studies to date reveal that phylogeographical signal within taxa of differing distributions reflects a diversity of patterns and processes rivalling the complexities of LCA landscapes themselves. Even so, phylogeography is providing novel insights into regional diversification (e.g. cryptic lineage divergences), and general evolutionary patterns are emerging. Congruent multi‐taxon phylogeographic breaks are found across the Nicaraguan depression, Chorotega volcanic front, western and central Panama, and the Darién isthmus, indicating that a potentially shared history of responses to regional‐scale (e.g. geological) processes has shaped the genetic diversity of LCA communities. By contrast, other species show unique demographic histories in response to overriding historical events, including no phylogeographic structure at all. These low‐structure or incongruent patterns provide some evidence for a role of local, ecological factors (e.g. long‐distance dispersal and gene flow in plants and bats) in shaping LCA communities. Temporally, comparative phylogeographical structuring reflects Pliocene–Pleistocene dispersal and vicariance events consistent with the timeline of emergence of the LCA isthmus and its major physiographic features, e.g. cordilleras. We emphasise the need to improve biogeographic inferences in LCA through in‐depth comparative phylogeography projects capitalising on the latest statistical phylogeographical methods. While meeting the challenges of reconstructing the biogeographical history of this complex region, phylogeographers should also take up the critical service to society of applying their work to the conservation of its fascinating biodiversity.     

Do biodiversity hotspots match with rodent conservation hotspots?

Biodiversity hotspots are used widely to designate priority regions for conservation efforts. It is unknown, however, whether the current network of hotspots adequately represents globally threatened taxonomic diversity for whole plant and animal groups. We used a mammalian group traditionally neglected in terms of conservation efforts, the rodents, in order to test whether biodiversity hotspots match the current distribution of threatened taxa (genera and species). Significantly higher numbers of threatened rodent genera and species fell within biodiversity hotspots; nonetheless over 25% of the total threatened genera and species did not occur in any biodiversity hotspot. This was particularly true for the Australian region, where 100% of the threatened genera and species fell outside biodiversity hotspots, with many threatened taxa found in Papua-New Guinea. We suggest to officially including Papua New Guinea among biodiversity hotspots for rodents, and also the steppic/semidesert areas of central Asia.     

Historical climate modelling predicts patterns of current biodiversity in the Brazilian Atlantic forest

Aim We aim to propose validated, spatially explicit hypotheses for the late Quaternary distribution of the Brazilian Atlantic forest, and thereby provide a framework for integrating analyses of species and genetic diversity in the region. Location The Atlantic forest, stretching along the Brazilian coast. Methods We model the spatial range of the forest under three climatic scenarios (current climate, 6000 and 21,000 years ago) with BIOCLIM and MAXENT. Historically stable areas or refugia are identified as the set of grid cells for which forest presence is inferred in all models and time projections. To validate inferred refugia, we test whether our models are matched by the current distribution of the forest and by fossil pollen data. We then investigate whether the location of inferred forest refugia is consistent with current patterns of species endemism and existing phylogeographical data. Results Forest models agree with pollen records and predict a large area of historical forest stability in the central corridor (Bahia), as well as a smaller refuge (Pernambuco) along the Brazilian coast, matching current centres of endemism in multiple taxa and mtDNA diversity patterns in a subset of the species examined. Less historical stability is predicted in coastal areas south of the Doce river, which agrees with most phylogeographical studies in that region. Yet some widely distributed taxa show high endemism in the southern Atlantic forest. This may be due to limitations of the modelling approach, differences in ecology and dispersal capability, historical processes not contemplated by the current study or inadequacy of the available test data sets. Main conclusions Palaeoclimatic models predict the presence of historical forest refugia in the Atlantic rain forest and suggest spatial variation in persistence of forests through the Pleistocene, predicting patterns of biodiversity in several local taxa. The results point to the need for further studies to document genetic and species endemism in the relatively poorly known and highly impacted areas of Atlantic rain forests of north‐eastern Brazil.     

The Importance of Rotational Crops for Biodiversity Conservation in Mediterranean Areas

Nowadays we are seeing the largest biodiversity loss since the extinction of the dinosaurs. To conserve biodiversity it is essential to plan protected areas using a prioritization approach, which takes into account the current biodiversity value of the sites. Considering that in the Mediterranean Basin the agro-ecosystems are one of the most important parts of the landscape, the conservation of crops is essential to biodiversity conservation. In the framework of agro-ecosystem conservation, farmland birds play an important role because of their representativeness, and because of their steady decline in the last Century in Western Europe. The main aim of this research was to define if crop dominated landscapes could be useful for biodiversity conservation in a Mediterranean area in which the landscape was modified by humans in the last thousand years and was affected by the important biogeographical phenomenon of peninsula effect. To assess this, we identify the hotspots and the coldspots of bird diversity in southern Italy both during the winter and in the breeding season. In particular we used a scoring method, defining a biodiversity value for each cell of a 1-km grid superimposed on the study area, using data collected by fieldwork following a stratified random sampling design. This value was analysed by a multiple linear regression analysis and was predicted in the whole study area. Then we defined the hotspots and the coldspots of the study area as 15% of the cells with higher and lower value of biodiversity, respectively. Finally, we used GAP analysis to compare hotspot distribution with the current network of protected areas. This study showed that the winter hotspots of bird diversity were associated with marshes and water bodies, shrublands, and irrigated crops, whilst the breeding hotspots were associated with more natural areas (e.g. transitional wood/shrubs), such as open areas (natural grasslands, pastures and not irrigated crops). Moreover, the results underlined the negative effects of permanent crops, such as vineyards, olive groves, and orchards, in particular during the winter season. This research highlights the importance of farmland areas mainly for wintering species and the importance of open areas for breeding species in the Mediterranean Basin. This may be true even when the species’ spatial distribution could be affected by biogeography. An important result showed that the hotspots for breeding species cannot be used as a surrogate for the wintering species, which were often not considered in the planning of protected areas.     

Identifying refugia from climate change using coupled ecological and genetic data in a transitional Mediterranean‐temperate tree species

Populations occurring in areas of overlap between the current and future distribution of a species are particularly important because they can represent “refugia from climate change”. We coupled ecological and range‐wide genetic variation data to detect such areas and to evaluate the impacts of habitat suitability changes on the genetic diversity of the transitional Mediterranean‐temperate tree Fraxinus angustifolia. We sampled and genotyped 38 natural populations comprising 1006 individuals from across Europe. We found the highest genetic diversity in western and northern Mediterranean populations, as well as a significant west to east decline in genetic diversity. Areas of potential refugia that correspond to approximately 70% of the suitable habitat may support the persistence of more than 90% of the total number of alleles in the future. Moreover, based on correlations between Bayesian genetic assignment and climate, climate change may favour the westward spread of the Black Sea gene pool in the long term. Overall, our results suggest that the northerly core areas of the current distribution contain the most important part of the genetic variation for this species and may serve as in situ macrorefugia from ongoing climate change. However, rear‐edge populations of the southern Mediterranean may be exposed to a potential loss of unique genetic diversity owing to habitat suitability changes unless populations can persist in microrefugia that have facilitated such persistence in the past.     

Multilocus approach reveals an incipient differentiation process in the Stone-curlew, <Emphasis Type="Italic">Burhinus oedicnemus</Emphasis> around the Mediterranean basin

The Stone-curlew Burhinus oedicnemus, a steppe bird species, is mainly distributed in the Mediterranean and Macaronesian regions, which are considered hotspots of biodiversity with priority for animal and plant species richness conservation. In this study, we investigated the genetic diversity of the Stone-curlew in the Mediterranean basin and in the Canary Islands by applying a multilocus approach. We analysed mitochondrial and nuclear markers in order to evaluate the genetic structure and the congruence between morphological subspecies and geographic samples. We found a significant level of genetic differentiation between Mediterranean and Canary Island populations with all markers. Both in the Mediterranean basin and in the Canary Islands, we found a significant level of genetic diversity with nuclear markers only. We identified seven population groups, including insular populations. The four subspecies described for the Western Palaearctic were confirmed with some changes in distribution range. In spite of habitat fragmentation and negative population trend, the Stone-curlew showed a significant level of genetic diversity and gene flow among continental populations. However, islands constitute important reservoirs of genetic diversity and a potential for the evolution of the species.     

Biogeography of freshwater fishes of the Balkan Peninsula

Delineating biogeographical regions is a critical step towards the establishment and evaluation of conservation priorities. In the present study, we analysed the distribution patterns of the freshwater fish of an understudied European biodiversity hotspot, the Balkan Peninsula. Based on the most extensive available database of native freshwater fish species distributions, we performed a hierarchical clustering analysis to identify the major biogeographical regions of the Balkan Peninsula. We also highlighted the ‘hottest hotspots’ of freshwater fish diversity across the delimited biogeographical regions by describing the patterns of species richness, endemic and vulnerable species; indicator species were also determined. The bioregionalisation scheme consisted of eight groups of drainage basins that correspond to distinct regions of the Balkan Peninsula. Overall, the delineated biogeographical regions varied in terms of species richness, endemism, vulnerability (i.e. extinction threats) and indicator species composition. From a conservation perspective, this study emphasises the prioritisation of areas characterised by high levels of irreplaceability (endemism) and vulnerability (i.e. the Attikobeotia region, Ionian Sea and Prespa Lakes) and stresses the necessity of implementing a network of protected freshwater areas across the Balkan Peninsula.     

Molecular phylogeography of Dryas integrifolia: glacial refugia and postglacial recolonization

Chloroplast DNA variation in the Arctic plant species Dryas integrifolia (Rosaceae) was analysed in relation to both the present-day geographical distribution of populations and to Pleistocene fossil records of this species. The phylogeographical structure was weak but the analysis of haplotype diversity revealed several groups of haplotypes having present-day geographical ranges that overlap locations postulated from geographical and fossil evidence to have been glacial refugia. Based on this information we infer that two important refugial sources of Arctic recolonization by this species were Beringia and the High Arctic. Two other putative refugia, located southeast of the ice sheet and along coastal regions of the eastern Arctic may have served as sources for recolonization of smaller portions of the Arctic. The genetic substructure in the species is mostly due to variation among populations regardless of the ecogeographical region in which they are found. Spatial autocorrelation at the regional scale was also detected. High levels of diversity both within populations and ecogeographical regions are probably indicative of population establishment from several sources possibly combined with recent gene flow.     

Increased genetic structuring of isolated Salamandra salamandra populations (Caudata: Salamandridae) at the margins of the Carpathian Mountains

The traditional southern Pleistocene refugia hypothesis in Europe has lately been challenged for several animal and plant species. The Carpathian Basin, especially at the marginal regions, is one of the recently recognized biodiversity hotspots in Europe. Marginal populations are prone to have lower genetic diversity and higher genetic differentiation than central populations. Here, we examined one mitochondrial DNA fragment (D‐loop) and nine nuclear (microsatellite) loci to describe the genetic diversity and phylogeographical pattern of fire salamander (Salamandra salamandra) populations in the Carpathian Basin with focusing on the southern margins of the Western Carpathians, where isolated populations of this species are present. Analyses of microsatellites indicated reduced genetic diversity for most of the isolated populations. Based on the mitochondrial DNA, only two haplotypes were found, whereas the analyses with the nuclear markers revealed a more recent genetic split between Western (Alpine) and Eastern (Carpathian) populations, and separated the Apuseni Mountains population (part of the Western Carpathians). Using approximate Bayesian computation analyses, we identified the most probable colonization scenario for the isolated North Hungarian Carpathian Basin populations. The split between isolated salamander populations from the central populations in the Carpathian Mountains dates back to the beginning of the Late Pleistocene, while the split between most of the Hungarian populations can be associated with the Last Glacial Maximum. We found evidence for long‐time isolation between the marginal Carpathian Basin and central populations. Our results also show that S. salamandra survived glacial periods in the temperate forests of north‐east Pannonia (North Hungarian Mountains), confirming that the Carpathian Basin served as important northerly refugia during the Pleistocene climatic oscillations.