Beringian Refugium Research Articles

Landscape Genomics Provides Evidence of Ecotypic Adaptation and a Barrier to Gene Flow at Treeline for the Arctic Foundation Species Eriophorum vaginatum.

Global climate change has resulted in geographic range shifts of flora and fauna at a global scale. Extreme environments, like the Arctic, are seeing some of the most pronounced changes. This region covers 14% of the Earth’s land area, and while many arctic species are widespread, understanding ecotypic variation at the genomic level will be important for elucidating how range shifts will affect ecological processes. Tussock cottongrass (Eriophorum vaginatum L.) is a foundation species of the moist acidic tundra, whose potential decline due to competition from shrubs may affect ecosystem stability in the Arctic. We used double-digest Restriction Site-Associated DNA sequencing to identify genomic variation in 273 individuals of E. vaginatum from 17 sites along a latitudinal gradient in north central Alaska. These sites have been part of 30 + years of ecological research and are inclusive of a region that was part of the Beringian refugium. The data analyses included genomic population structure, demographic models, and genotype by environment association. Genome-wide SNP investigation revealed environmentally associated variation and population structure across the sampled range of E. vaginatum, including a genetic break between populations north and south of treeline. This structure is likely the result of subrefugial isolation, contemporary isolation by resistance, and adaptation. Forty-five candidate loci were identified with genotype-environment association (GEA) analyses, with most identified genes related to abiotic stress. Our results support a hypothesis of limited gene flow based on spatial and environmental factors for E. vaginatum, which in combination with life history traits could limit range expansion of southern ecotypes northward as the tundra warms. This has implications for lower competitive attributes of northern plants of this foundation species likely resulting in changes in ecosystem productivity.

Continent-wide population genomic structure and phylogeography of North America's most destructive conifer defoliator, the spruce budworm (Choristoneura fumiferana).

The spruce budworm, Choristoneura fumiferana, is presumed to be panmictic across vast regions of North America. We examined the extent of panmixia by genotyping 3,650 single nucleotide polymorphism (SNP) loci in 1975 individuals from 128 collections across the continent. We found three spatially structured subpopulations: Western (Alaska, Yukon), Central (southeastern Yukon to the Manitoba–Ontario border), and Eastern (Manitoba–Ontario border to the Atlantic). Additionally, the most diagnostic genetic differentiation between the Central and Eastern subpopulations was chromosomally restricted to a single block of SNPs that may constitute an island of differentiation within the species. Geographic differentiation in the spruce budworm parallels that of its principal larval host, white spruce (Picea glauca), providing evidence that spruce budworm and spruce trees survived in the Beringian refugium through the Last Glacial Maximum and that at least two isolated spruce budworm populations diverged with spruce/fir south of the ice sheets. Gene flow in the spruce budworm may also be affected by mountains in western North America, habitat isolation in West Virginia, regional adaptations, factors related to dispersal, and proximity of other species in the spruce budworm species complex. The central and eastern geographic regions contain individuals that assign to Eastern and Central subpopulations, respectively, indicating that these barriers are not complete. Our discovery of previously undetected geographic and genomic structure in the spruce budworm suggests that further population modelling of this ecologically important insect should consider regional differentiation, potentially co‐adapted blocks of genes, and gene flow between subpopulations.

The phylogeography of coho salmon based on mtDNA control region variation: Asian populations descend from the Beringian Refugium

The phylogeography of coho salmon based on mtDNA control region variation: Asian populations descend from the Beringian Refugium

Morphological Variability among Spawning Populations of Bering CiscoCoregonus laurettae

Bering Cisco Coregonus laurettae is an anadromous coregonine species known almost exclusively from northwestern North America and with only three documented spawning populations, all in Alaska. Previous studies of Bering Cisco phenotypic variation examined individuals collected primarily in coastal rearing habitats where population affiliation was not known. Here we compare meristic counts and morphometric ratios of pre-spawning adults among the three known populations: one each in the Yukon, Kuskokwim, and Susitna rivers in Alaska. We also compare meristic data with those previously reported for this species. Populations in the Yukon and Kuskokwim rivers were very similar, while the population in the Susitna River was significantly divergent for certain meristic counts and morphometric ratios. Our findings are consistent with recent genetic analyses that found the Susitna River population to be the most divergent of the three populations. While the Yukon and Kuskokwim river populations survived the Wisconsinan Ice Age in the Beringian Refugium, the Susitna River population colonized the drainage following ice retreat sometime in the last few thousand years. The population's divergence from the source population in the Yukon or Kuskokwim river could be due to a founder effect or adaptation to different environmental conditions.

Environmental selection during the last ice age on the mother-to-infant transmission of vitamin D and fatty acids through breast milk

Because of the ubiquitous adaptability of our material culture, some human populations have occupied extreme environments that intensified selection on existing genomic variation. By 32,000 years ago, people were living in Arctic Beringia, and during the Last Glacial Maximum (LGM; 28,000-18,000 y ago), they likely persisted in the Beringian refugium. Such high latitudes provide only very low levels of UV radiation, and can thereby lead to dangerously low levels of biosynthesized vitamin D. The physiological effects of vitamin D deficiency range from reduced dietary absorption of calcium to a compromised immune system and modified adipose tissue function. The ectodysplasin A receptor (EDAR) gene has a range of pleiotropic effects, including sweat gland density, incisor shoveling, and mammary gland ductal branching. The frequency of the human-specific EDAR V370A allele appears to be uniquely elevated in North and East Asian and New World populations due to a bout of positive selection likely to have occurred circa 20,000 y ago. The dental pleiotropic effects of this allele suggest an even higher occurrence among indigenous people in the Western Hemisphere before European colonization. We hypothesize that selection on EDAR V370A occurred in the Beringian refugium because it increases mammary ductal branching, and thereby may amplify the transfer of critical nutrients in vitamin D-deficient conditions to infants via mothers' milk. This hypothesized selective context for EDAR V370A was likely intertwined with selection on the fatty acid desaturase (FADS) gene cluster because it is known to modulate lipid profiles transmitted to milk from a vitamin D-rich diet high in omega-3 fatty acids.

Phylogeography of the Chydorus sphaericus Group (Cladocera: Chydoridae) in the Northern Palearctic.

The biodiversity and the biogeography are still poorly understood for freshwater invertebrates. The crustacean Chydorus sphaericus-brevilabris complex (Cladocera: Chydoridae) is composed of species that are important components of Holarctic freshwater food webs. Recent morphological and genetic study of the complex has indicated a substantial species diversity in the northern hemisphere. However, we know little of the geographic boundaries of these novel lineages. Moreover, a large section of the Palearctic remains unexamined at the genetic level. Here we attempt to address the biodiversity knowledge gap for the Chydorus sphaericus group in the central Palearctic and assess its diversity and biogeographic boundaries. We sequenced nuclear (ITS-2) and mitochondrial (COI) gene regions of Chydorus specimens across the Palearctic and compared them with already available Holarctic sequences. We detected six main clades in the C. sphaericus group in the Palearctic, of which two of the groups are novel. Three of the more divergent clades are geographically widespread. The central portion of Eurasia (the Yenisey River basin) appears to be a narrow zone of secondary contact for phylogroups that expanded from European and Beringian refugia. As such, the previously unsampled central Palearctic represents an important region for understanding the evolutionary consequences of Pleistocene climatic oscillations on the Chydorus sphaericus group.

Genome-wide set of SNPs reveals evidence for two glacial refugia and admixture from postglacial recolonization in an alpine ungulate.

Past glaciation events have played a major role in shaping the genetic diversity and distribution of wild sheep in North America. The advancement of glaciers can isolate populations in ice-free refugia, where they can survive until the recession of ice sheets. The major Beringian refugium is thought to have held thinhorn sheep (Ovis dalli) populations during times of glacial advance. While isolation in the major refugium can account for much of the genetic and morphological diversity seen in extant thinhorn sheep populations, mounting evidence suggests the persistence of populations in smaller minor refugia. We investigated the refugial origins of thinhorn sheep using ~10000 SNPs obtained via a cross-species application of the domestic sheep ovine HD BeadChip to genotype 52 thinhorn sheep and five bighorn sheep (O. canadensis) samples. Phylogenetic inference revealed a distinct lineage of thinhorn sheep inhabiting British Columbia, which is consistent with the survival of a group of thinhorn sheep in a minor refugium separate from the Beringian refugium. Isolation in separate glacial refugia probably mediated the evolution of the two thinhorn sheep subspecies, the white Dall's sheep (O. d. dalli), which persisted in Beringia, and the dark Stone's sheep (O. d. stonei), which utilized the minor refugium. We also found the first genetic evidence for admixture between sheep from different glacial refugia in south-central Yukon as a consequence of post glacial expansion and recolonization. These results show that glaciation events can have a major role in the evolution of species inhabiting previously glaciated habitats and the need to look beyond established refugia when examining the evolutionary history of such species.

First record of the Holarctic least shrew (Sorex minutissimus) and associated helminths from Canada: new light on northern Pleistocene refugia

We report the first Canadian record of the Holarctic least shrew (Sorex minutissimus Zimmermann, 1780) and associated helminth worms, collected along the Dempster Highway in central Yukon in 2014. We identify the specimen based on morphological characters, characterize the habitat, report other mammals and helminth species associated with this specimen, and use mitochondrial DNA sequences to place the specimen within a phylogenetic context and address Pleistocene refugial hypotheses. Although long considered an Eurasian endemic, the diminutive least shrew was first reported from Alaska in 1994. Our new record for Canada indicates that the species may occur at least as far east as the MacKenzie River and DNA variation suggests this species persisted only in the Beringian refugium in North America during the Last Glacial Maximum. The discovery of a new mammal and associated parasites for Canada points to the urgent need for more detailed information on high-latitude biotas in North America, data that are best obtained through museum-based field surveys, particularly for small, cryptic species.

Beringian sub-refugia revealed in blackfish (Dallia): implications for understanding the effects of Pleistocene glaciations on Beringian taxa and other Arctic aquatic fauna.

BackgroundPleistocene climatic instability had profound and diverse effects on the distribution and abundance of Arctic organisms revealed by variation in phylogeographic patterns documented in extant Arctic populations. To better understand the effects of geography and paleoclimate on Beringian freshwater fishes, we examined genetic variability in the genus Dallia (blackfish: Esociformes: Esocidae). The genus Dallia groups between one and three nominal species of small, cold- and hypoxia-tolerant freshwater fishes restricted entirely in distribution to Beringia from the Yukon River basin near Fairbanks, Alaska westward including the Kuskokwim River basin and low-lying areas of Western Alaska to the Amguema River on the north side of the Chukotka Peninsula and Mechigmen Bay on the south side of the Chukotka Peninsula. The genus has a non-continuous distribution divided by the Bering Strait and the Brooks Range. We examined the distribution of genetic variation across this range to determine the number and location of potential sub-refugia within the greater Beringian refugium as well as the roles of the Bering land bridge, Brooks Range, and large rivers within Beringia in shaping the current distribution of populations of Dallia. Our analyses were based on DNA sequence data from two nuclear gene introns (S7 and RAG1) and two mitochondrial genome fragments from nineteen sampling locations. These data were examined under genetic clustering and coalescent frameworks to identify sub-refugia within the greater Beringia refugium and to infer the demographic history of different populations of Dallia.ResultsWe identified up to five distinct genetic clusters of Dallia. Four of these genetic clusters are present in Alaska: (1) Arctic Coastal Plain genetic cluster found north of the Brooks Range, (2) interior Alaska genetic cluster placed in upstream locations in the Kuskokwim and Yukon river basins, (3) a genetic cluster found on the Seward Peninsula, and (4) a coastal Alaska genetic cluster encompassing downstream Kuskokwim River and Yukon River basin sample locations and samples from Southwest Alaska not in either of these drainages. The Chukotka samples are assigned to their own genetic cluster (5) similar to the coastal Alaska genetic cluster. The clustering and ordination analyses implemented in Discriminant Analysis of Principal Components (DAPC) and STRUCTURE showed mostly concordant groupings and a high degree of differentiation among groups. The groups of sampling locations identified as genetic clusters correspond to geographic areas divided by likely biogeographic barriers including the Brooks Range and the Bering Strait. Estimates of sequence diversity (θ) are highest in the Yukon River and Kuskokwim River drainages near the Bering Sea. We also infer asymmetric migration rates between genetic clusters. The isolation of Dallia on the Arctic Coastal Plain of Alaska is associated with very low estimated migration rates between the coastal Alaska genetic cluster and the Arctic Coastal Plain genetic cluster.ConclusionsOur results support a scenario with multiple aquatic sub-refugia in Beringia during the Pleistocene and the preservation of that structure in extant populations of Dallia. An inferred historical presence of Dallia across the Bering land bridge explains the similarities in the genetic composition of Dallia in West Beringia and western coastal Alaska. In contrast, historic and contemporary isolation across the Brooks Range shaped the distinctiveness of present day Arctic Coastal Plain Dallia. Overall this study uncovered a high degree of genetic structuring among populations of Dallia supporting the idea of multiple Beringian sub-refugia during the Pleistocene and which appears to be maintained to the present due to the strictly freshwater nature and low dispersal ability of this genus.Electronic supplementary materialThe online version of this article (doi:10.1186/s12862-015-0413-2) contains supplementary material, which is available to authorized users.

Geographical origins, migration patterns and refugia of Sibbaldia procumbens, an arctic–alpine plant with a fragmented range

AbstractAimMany plants, especially at high latitudes, have both widespread and highly discontinuous geographical distributions. To increase understanding of how such patterns originate, we examine genetic patterns in the arctic–alpine plant Sibbaldia procumbens. We evaluate the contributions of refugia and the role of long‐distance dispersal in shaping the current range of this species.LocationNorthern Hemisphere, especially North America.MethodsWe sampled Sibbaldia from 176 localities, including 168 for S. pro‐cumbens. We analysed sequence variation in three plastid DNA non‐coding regions (the atpI–atpH and trnL–trnF intergenic spacers and the trnL intron), performed Bayesian phylogenetic analyses and statistical parsimony analyses on the combined sequences, and analysed the geographical patterns of haplotype distribution and genetic diversity using data from all populations.ResultsSibbaldia procumbens probably originated in the mountains of South and East Asia. We identified highly distinct clades in Europe and North America, which overlapped on oceanic islands of the North Atlantic indicating long‐distance dispersal capability. The North American clade included two lineages, one in California and the other widely distributed across the continent and North Atlantic. Haplotype diversity in the latter lineage was markedly higher to the south, suggesting mid–late Pleistocene southward displacement of North American populations with subsequent migration northwards into previously glaciated regions. In Europe, disjunct geographical regions generally harboured distinct haplotypes.Main conclusionsMultiple Pleistocene refugia for S. procumbens occurred in both North America and Europe. North American refugia existed in California and in the southern Rocky Mountains, but in contrast with most widespread arctic–alpine species we found no evidence for a Beringian refugium. Cryptic refugia may have existed within the Cordilleran Ice Sheet. Episodes of range expansion and contraction and long‐distance dispersal have all contributed to the genetic structure and widespread but fragmented distribution of this species.

Phylogeography of a Holarctic rodent (Myodes rutilus): testing high‐latitude biogeographical hypotheses and the dynamics of range shifts

AbstractAimWe used the Holarctic northern red‐backed vole (Myodes rutilus) as a model organism to improve our understanding of how dynamic, northern high‐latitude environments have affected the genetic diversity, demography and distribution of boreal organisms. We tested spatial and temporal hypotheses derived from previous mitochondrial studies, comparative phylogeography, palaeoecology and the fossil record regarding diversification of M. rutilus in the Palaearctic and Beringia.LocationHigh‐latitude biomes across the Holarctic.MethodsWe used a multilocus phylogeographical approach combined with species distribution models to characterize the biogeographical and demographic history of M. rutilus. Our molecular assessment included widespread sampling (more than 100 localities), species tree reconstruction and population genetic analyses.ResultsThree well‐differentiated mitochondrial lineages correspond to geographical regions, but nuclear genes were less structured. Multilocus divergence estimates indicated that diversification of M. rutilus was driven by events occurring before c. 100 ka. Population expansion in all three clades occurred prior to the Last Glacial Maximum (LGM) and presumably led to secondary contact. Species distribution modelling predicted a broad LGM distribution consistent with population and range expansion during this period.Main conclusionsThe biogeographical history of M. rutilus differs from other boreal forest‐associated species. Well‐differentiated clades and the existence of secondary contact zones indicate prolonged isolation and persistence in Eurasian and Beringian refugia. Dynamic demographic and distributional changes emphasize the impact of pre‐LGM glacial–interglacial cycles on contemporary geographical structure. The Bering Strait was not a significant factor in the diversification of northern red‐backed voles.

Phylogeography of northern Dolly VardenSalvelinus malma malmabased on analysis of mitochondrial DNA

The northern Dolly Varden, Salvelinus malma malma, is a typical representative of arctic fauna distributed in northeastern Asia and northwestern North America. Because its spawning habitats were affected by Pleistocene glacial advances over most of its natural range, S. m. malma is among the most interesting objects of phylogeographic and microevolutionary studies. We reconstructed the genealogy of mtDNA haplotypes from 27 Alaskan and Asian populations to study the influence of glacial and geological vicariance events on the contemporary population genetic structure, phylogeographic subdivision and distribution of the northern Dolly Varden. Analysis of restriction site states in three PCR-amplified mtDNA regions (ND1/ND2, ND5/ND6, Cytb/D-loop; 47% of the mitochondrial genome) resolved 75 haplotypes in 436 fish. Similar patterns of subspecific variation apparent from hierarchical diversity and nested clade analyses of mtDNA haplotypes identify weak spatial differentiation and low levels of divergence. Our results suggest that (1) demographic history has been influenced by historical range expansions and recent isolation by distance, (2) present populations from Asia and North America were colonized from one main Beringian Refugium, and (3) that this taxon's ancestral population probably experienced a bottleneck in the Beringian Refugium during the late Pleistocene (Wisconsin) glacial period.

Phylogeography of northern Dolly Varden Salvelinus malma (Salmoniformes: Salmonidae) from Asia and North America: An analysis based on the mitochondrial DNA genealogy

Spawning in habitats affected by Pleistocene glacial advances over most of its natural range, northern Dolly Varden Salvelinus malma malma typifies Arctic fauna distributed in northeastern Asia and northwestern North America. We reconstructed a genealogy of mtDNA haplotypes from 27 Alaskan and Asian populations to study the influence of historical events on the phylogeography and contemporary population genetic structure. Analysis of molecular variance partitioned most of the mtDNA variability to the intrapopulation component (72.5%) with much reduced differences between populations (21.1%) and regions (6.4%). Similar patterns of variation apparent from hierarchical diversity and nested clade phylogeographical analysis (NCPA) of mtDNA haplotypes identify weak spatial differentiation and low levels of divergence. These findings suggest (1) that demographic history has been influenced by historical range expansions and recent isolation by distance, (2) that present populations from Asia and North America were colonized from one main Beringian Refugium, and (3) that this taxon’s ancestral population probably experienced a bottleneck in the Beringian Refugium during the late Pleistocene (Wisconsin) glacial period. The genealogical and NCPA analyses, and mismatch distribution of S. m. malma mtDNA haplotypes do not confirm the assumptions about presence of the two refugia on the territories of the Beringian Land, in which allopatric S. m. malma ancestral populations evolved, and independent origin of the Sea of Okhotsk populations.

Genetic consequences of glacial survival: the late Quaternary history of balsam poplar (Populus balsamifera L.) in North America

AbstractAim Beringia, the unglaciated region encompassing the former Bering land bridge, as well as the land between the Lena and Mackenzie rivers, is recognized as an important refugium for arctic plants during the last ice age. Compelling palaeobotanical evidence also supports the presence of small populations of boreal trees within Beringia during the Last Glacial Maximum. The occurrence of balsam poplar (Populus balsamifera) in Beringia provides a unique opportunity to assess the implications of persistence in a refugium on present‐day genetic diversity for this boreal tree species.Location North America.Methods We sequenced three variable non‐coding regions of the chloroplast genome (cpDNA) from 40 widely distributed populations of balsam poplar across its North American range. We assessed patterns of genetic diversity, geographic structure and historical demography between glaciated and unglaciated regions of the balsam poplar’s range. We also utilized a coalescent model to test for divergence between regions.Results Levels of genetic diversity were consistently greater for populations at the southern margin (θW = 0.00122) than in the central (θW = 0.00086) or northern (θW = 0.00034) regions of the current distribution of balsam poplar, and diversity decreased with increasing latitude (R2 = 0.49, P < 0.01). We detected low, but significant, structure (FCT = 0.05, P = 0.05), among regions of P. balsamifera’s distribution. The cpDNA genealogy was shallow, however, showing an absence of highly differentiated chloroplast haplotypes. Coalescent analyses supported a model of divergence between the southern ice margin and the northern unglaciated region of balsam poplar’s distribution, but analyses of other regional comparisons did not converge.Main conclusions The palaeobotanical record supports the presence of a Beringian refugium for balsam poplar, but we were unable to definitively identify the presence of known refugial populations based on genetic data alone. Balsam poplar populations from Beringia are not a significant reservoir of cpDNA diversity today. Unique alleles that may have been present in the small, isolated populations that survived within Beringia were probably lost through genetic drift or swamped by post‐glacial, northward migration from populations south of the ice sheets.

Constraining the Late Wisconsinan retreat of the Laurentide ice sheet from western Canada using luminescence ages from postglacial aeolian dunes

Optically stimulated luminescence (OSL) dating of quartz extracts from postglacial aeolian dunes from central Alberta in western Canada points to a landscape that was free of ice as early as 15 ka. Data from profiles where multiple ages have been obtained indicate an increase in depositional age with depth, suggesting that older aeolian sands underlie the dated sequences. The OSL ages present plausible minimum age constraints for the retreat of the Laurentide Ice Sheet (LIS) towards the end of the Late Wisconsinan glaciation. Previous reconstructions of the LIS recession have relied on radiocarbon chronologies, despite the scarcity of contemporaneous radiocarbon-bearing material for large parts of western Canada. While the OSL chronology may be deemed concordant with ice sheet margin retreat models determined using radiocarbon data, there appears to be a systematic lag in the radiocarbon ages which may reflect that aeolian activity is initiated prior to the proliferation of organic material. The OSL chronology reported in this study does not preclude the emergence of a wide deglacial corridor between the LIS and the Cordilleran Ice Sheet by 15 ka or earlier. The possibility of such a passage would resuscitate the notion of an ice-free corridor that appeared early enough to afford the first peoples of the Americas a navigable inland migratory passage from Beringia to south of the North American ice sheets. More broadly, the corridor would also have allowed genetic exchanges between the Beringian refugium and the American middle and low latitudes.

Multilocus phylogeography and population structure of common eiders breeding in North America and Scandinavia

Aim Glacial refugia during the Pleistocene had major impacts on the levels and spatial apportionment of genetic diversity of species in northern latitude ecosystems. We characterized patterns of population subdivision, and tested hypotheses associated with locations of potential Pleistocene refugia and the relative contribution of these refugia to the post-glacial colonization of North America and Scandinavia by common eiders (Somateria mollissima). Specifically, we evaluated localities hypothesized as ice-free areas or glacial refugia for other Arctic vertebrates, including Beringia, the High Arctic Canadian Archipelago, Newfoundland Bank, Spitsbergen Bank and north-west Norway. Location Alaska, Canada, Norway and Sweden. Methods Molecular data from 12 microsatellite loci, the mitochondrial DNA (mtDNA) control region, and two nuclear introns were collected and analysed for 15 populations of common eiders (n = 716) breeding throughout North America and Scandinavia. Population genetic structure, historical population fluctuations and gene flow were inferred using F-statistics, analyses of molecular variance, and multilocus coalescent analyses. Results Significant inter-population variation in allelic and haplotypic frequencies were observed (nuclear DNA FST = 0.004‐0.290; mtDNA FST = 0.051‐0.927). Whereas spatial differentiation in nuclear genes was concordant with subspecific designations, geographic proximity was more predictive of inter-population variance in mitochondrial DNA haplotype frequency. Inferences of historical population demography were consistent with restriction of common eiders to four geographic areas during the Last Glacial Maximum: Belcher Islands, Newfoundland Bank, northern Alaska and Svalbard. Three of these areas coincide with previously identified glacial refugia: Newfoundland Bank, Beringia and Spitsbergen Bank. Gene-flow and clustering analyses indicated that the Beringian refugium contributed little to common eider post-glacial colonization of North America, whereas Canadian, Scandinavian and southern Alaskan post-glacial colonization is likely to have occurred in a stepwise fashion from the same glacial refugium. Main conclusions Concordance of proposed glacial refugia used by common eiders and other Arctic species indicates that Arctic and subarctic refugia were important reservoirs of genetic diversity during the Pleistocene. Furthermore, suture zones identified at MacKenzie River, western Alaska/Aleutians and Scandinavia coincide with those identified for other Arctic vertebrates, suggesting that these regions were strong geographic barriers limiting dispersal from Pleistocene refugia.

Pleistocene Bison Skeleton (Bison bison cf. occidentalis) from Clayhurst Crossing, British Colombia

Remains of a large old male bison skeleton from a gravel pit near Clayhurst Crossing are tentatively referred to the extinct western bison (Bison bison cf. occidentalis). It is the most complete late Pleistocene "medium-horned" bison so far reported from Canada, and is therefore described in detail. Four accelerator mass spectrometry (AMS) radiocarbon dates on bone from the specimen provide a weighted average age of 10,500±90 yr BP, supporting an earlier suggestion that river terraces in the mid-Peace area began developing before 10,000 years ago. Evidently Bison bison antiquus occurred in southern Alberta about 11,000 years ago, and both it and Bison bison occidentalis occupied the Peace River area shortly before 10,000 years ago. It is hypothesised that by at least 9700 BP, B. b. occidentalis had spread southern Alberta. These data suggest that biotic interchange between the former Beringian refugium and the southern interior Plains of North America occurred prior to 10,500 years ago.

Holarctic phylogeography of the root vole (Microtus oeconomus): implications for late Quaternary biogeography of high latitudes

A species-wide phylogeographical study of the root vole (Microtus oeconomus) was performed using the whole 1140 base pair mitochondrial (mt) cytochrome b gene. We examined 83 specimens from 52 localities resulting in 65 unique haplotypes. Our results demonstrate that the root vole is divided into four main mtDNA phylogenetic lineages that seem to have largely allopatric distributions. Net divergence estimates (2.0-3.5%) between phylogroups, as well as relatively high nucleotide diversity estimates within phylogroups, indicate that the distinct phylogeographical structure was initiated by historical events that predated the latest glaciation. European root voles are divided into a Northern and a Central mtDNA phylogroup. The mtDNA data in concert with fossil records imply that root voles remained north of the classical refugial areas in southern Europe during the last glacial period. The currently fragmented populations in central Europe belong to a single mtDNA phylogroup. The Central Asian and the North European lineages are separated by the Ural Mountains, a phylogeographical split also found in collared lemmings (Dicrostonyx) and the common vole (M. arvalis). The Beringian lineage occurs from eastern Russia through Alaska to northwestern Canada. This distribution is congruent with the traditional boundaries of the Beringian refugium and with phylogeographical work on other organisms. In conclusion, similarities between the phylogeographical patterns in the root vole and other rodents, such as Arctic and subarctic lemmings, as well as more temperate vole species, indicate that late Quaternary geological and climatic events played a strong role in structuring northern biotic communities.

Phylogeography of lemmings (Lemmus): no evidence for postglacial colonization of Arctic from the Beringian refugium.

Beringia is considered as an important glacial refugium that served as the main source for colonization of formerly glaciated Arctic regions. To obtain high resolution views of Arctic refugial history, we examined mitochondrial cytochrome b phylogeography in the northern genus of rodents, Lemmus (true lemmings), sampled across its circumpolar distribution. Strong phylogeographical structure suggests vicariant separation over several glacial-interglacial periods and does not provide evidence supporting the importance of Beringia for extensive colonization of formerly glaciated regions. Rather than a source of postglacial colonization, Beringia represents an area of intraspecific endemism previously undetected by biogeographical analysis. Existing phylogeographical structure suggests that vicariant separation by glacial barriers was an important factor generating genetic divergence and, thus, increasing genetic diversity in lemmings on continental and circumpolar scales. However, there is little evidence for the direct effect of the last glaciation on the level of genetic variation and allele genealogy in lemmings on a regional geographical scale. This finding implies that the population genetic models of postglacial colonization suggested for temperate taxa might have limited applicability for Arctic species.

High Genetic Heterogeneity in Chum Salmon in Western Alaska, the Contact Zone between Northern and Southern Lineages

Genetic relationships among 64 spawning populations of chum salmon Oncorhynchus keta in western Alaska were studied using allele frequency data from 40 protein-encoding loci. Two major lineages of chum salmon inhabiting Alaska were detected using clustering and multidimensional scaling analyses of Cavalli-Sforza and Edwards' chord distances. Populations of the northwest Alaska lineage occur in the largely unglaciated areas of Alaska north of the Alaska Peninsula (Beringia, the Beringian Refugium), and the Alaska Peninsula–Gulf of Alaska lineage occurs in the glaciated and unglaciated areas of the Alaska Peninsula, Kodiak Island, and southcentral Alaska. The two lineages come into contact in the 150-km area separating Herendeen Bay and Port Heiden on the northern Alaska Peninsula; this area may represent a major zoogeographic contact zone. Genetic data also suggest the lineages come in contact in upper Cook Inlet; the population representing the Susitna River drainage, which drains into Cook Inlet and the Gulf of Alaska, shows affinity to the northwest Alaska lineage. Genetic variability was higher in the Alaska Peninsula–Gulf of Alaska lineage than in the northwest Alaska lineage. A comparison of allele frequency data collected in this study with data available for Pacific Rim populations suggests that populations of the Alaska Peninsula–Gulf of Alaska lineage were derived from Cascadia (the Pacific Refugium) and belong to a larger southern lineage, which includes populations from southeast Alaska, British Columbia, and the Pacific Northwest. In contrast, populations from northwest Alaska appear to be derived from a northern lineage with affinities to Asian populations.