The brown bear has proved a useful model for studying Late Quaternary mammalian phylogeography. However, information is lacking from northern continental Eurasia, which constitutes a large part of the species' current distribution. We analysed mitochondrial DNA sequences (totalling 1943 bp) from 205 bears from northeast Europe and Russia in order to characterize the maternal phylogeography of bears in this region. We also estimated the formation times of the sampled brown bear lineages and those of its extinct relative, the cave bear. Four closely related haplogroups belonging to a single mitochondrial subclade were identified in northern continental Eurasia. Several haplotypes were found throughout the whole study area, while one haplogroup was restricted to Kamchatka. The haplotype network, estimated divergence times and various statistical tests indicated that bears in northern continental Eurasia recently underwent a sudden expansion, preceded by a severe bottleneck. This brown bear population was therefore most likely founded by a small number of bears that were restricted to a single refuge area during the last glacial maximum. This pattern has been described previously for other mammal species and as such may represent one general model for the phylogeography of Eurasian mammals. Bayesian divergence time estimates are presented for different brown and cave bear clades. Moreover, our results demonstrate the extent of substitution rate variation occurring throughout the phylogenetic tree, highlighting the need for appropriate calibration when estimating divergence times.

We estimated the phylogenetic relationships of brown bear maternal haplotypes from countries of northeastern Europe (Estonia, Finland and European Russia), using sequences of mitochondrial DNA (mtDNA) control region of 231 bears. Twenty-five mtDNA haplotypes were identified. The brown bear population in northeastern Europe can be divided into three haplogroups: one with bears from all three countries, one with bears from Finland and Russia, and the third composed almost exclusively of bears from European Russia. Four haplotypes from Finland and European Russia matched exactly with haplotypes from Slovakia, suggesting the significance of the current territory of Slovakia in ancient demographic processes of brown bears. Based on the results of this study and those from the recent literature, we hypothesize that the West Carpathian Mountains have served either as one of the northernmost refuge areas or as an important movement corridor for brown bears of the Eastern lineage towards northern Europe during or after the last ice age. Bayesian analyses were performed to investigate the temporal framework of brown bear lineages in Europe. The molecular clock was calibrated using Beringian brown bear sequences derived from radiocarbon-dated ancient samples, and the estimated mutation rate was 29.8% (13.3%-47.6%) per million years. The whole European population and Western and Eastern lineages formed about 175,000, 70,000 and 25,000 years before present, respectively. Our approach to estimating the time frame of brown bear evolution demonstrates the importance of using an appropriate mutation rate, and this has implications for other studies of Pleistocene populations.

Aim Using sequences of complete mitochondrial genomes, our aims were: (1) to investigate the matrilineal phylogeographical structure, migration patterns and lineage coalescence times in a large, continuous population of brown bears (Ursus arctos); and (2) to develop a novel spatial genetic method to identify migration corridors and barriers. Location North‐western Eurasia: from eastern European Russia to the Baltic Sea. Methods We sequenced the complete mitochondrial genomes of 95 brown bears. The phylogeographical resolution of complete genomes was compared to that derived from subsets of the genome, including the most commonly used shorter sequences. We conducted network and Bayesian phylogeographical analyses and developed a novel, spatially explicit, individual‐based approach (called DResD) for identifying migration corridors and barriers. Results Analysis of mitogenome sequences revealed five haplogroups, specific to particular geographical areas, exhibiting far greater resolving power than shorter sequences. Estimated coalescence times for the haplogroups ranged from 7.7 to 15.2 ka, suggesting that their divergence took place after the last glaciation. We found several migration trends, including a large westward migration from eastern European Russia towards Finland. We also found evidence of a potential barrier and a migration corridor in the south‐west of the study area. Main conclusions The use of complete mitochondrial genomes from a brown bear population in north‐western Eurasia allowed us to identify phylogeographical structure, signatures of demographic history and spatial processes that had not previously been detected using shorter sequences. These findings have implications for studies on other species and populations, especially those exhibiting low mtDNA diversity. The relatively recent divergence estimates for haplogroups highlight the significance not only of the last glaciation but also of climatic fluctuations during the post‐glacial period for the divergence of mammal populations in Europe. Our spatial genetic method represents a new tool for the analysis of genetic data in a geographical context and is applicable to any data that yield genetic distance matrices, including microsatellites, amplified fragment length polymorphisms (AFLPs) and single‐nucleotide polymorphisms (SNPs).

Knowledge of population structure and genetic diversity and the spatio-temporal demographic processes affecting populations is crucial for effective wildlife preservation, yet these factors are still poorly understood for organisms with large continuous ranges. Available population genetic data reveal that widespread mammals have for the most part only been carefully studied at the local population scale, which is insufficient for understanding population processes at larger scales. Here, we provide data on population structure, genetic diversity and gene flow in a brown bear population inhabiting the large territory of northwestern Eurasia. Analysis of 17 microsatellite loci indicated significant population substructure, consisting of four genetic groups. While three genetic clusters were confined to small geographical areas-located in Estonia, southern Finland and Leningrad oblast, Russia-the fourth cluster spanned a very large area broadly falling between northern Finland and the Arkhangelsk and Kirov oblasts of Russia. Thus, the data indicate a complex pattern where a fraction of the population exhibits large-scale gene flow that is unparalleled by other wild mammals studied to date, while the remainder of the population appears to have been structured by a combination of demographic history and landscape barriers. These results based on nuclear data are generally in good agreement with evidence previously derived using mitochondrial markers, and taken together, these markers provide complementary information about female-specific and population-level processes. Moreover, this study conveys information about spatial processes occurring over multiple generations that cannot be readily gained using other approaches, e.g. telemetry.

Aim Climatic changes during the Late Pleistocene had major impacts on populations of plant and animal species. Brown bears and other large mammals are likely to have experienced analogous ecological pressures and phylogeographical processes. Here, we address several unresolved issues regarding the Late Pleistocene demography of brown bears: (1) the putative locations of refugia; (2) the direction of migrations across Eurasia and into North America; and (3) parallels with the demographic histories of other wild mammals and modern humans. Location Eurasia and North America. Methods We sequenced 110 complete mitochondrial genomes from Eurasian brown bears and combined these with published sequences from 138 brown bears and 33 polar bears. We used a Bayesian approach to obtain a joint estimate of the phylogeny and evolutionary divergence times. The inferred mutation rate was compared with estimates obtained using two additional methods. Results Bayesian phylogenetic analysis identified seven clades of brown bears, with most individuals belonging to a very large Holarctic clade. Bears from the widespread clade 3a1, which has a distribution from Europe across Asia to Alaska, shared a common ancestor about 45,000 years ago. Main conclusions We suggest that the Altai‐Sayan region and Beringia were important Late Pleistocene refuge areas for brown bears and propose large‐scale migration scenarios for bears in Eurasia and to North America. We also argue that brown bears and modern humans experienced a demographic standstill in Beringia before colonizing North America.