Alexei V. Abramov

Mitogenetic structure of brown bears (Ursus arctos L.) in northeastern Europe and a new time frame for the formation of European brown bear lineages

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.

Sudden expansion of a single brown bear maternal lineage across northern continental Eurasia after the last ice age: a general demographic model for mammals?

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.

Phylogeography of the Russian flying squirrel (Pteromys volans): implication of refugia theory in arboreal small mammal of Eurasia.

A phylogeographical study of the Russian (Siberian) flying squirrel (Pteromys volans) was carried out using the complete mitochondrial (mt) cytochrome b gene sequences with special reference to the refugia theory for the arboreal traits of this species. We examined 31 specimens from 24 localities, resulting in 28 haplotypes. One breeding specimen with a unique haplotype from Hokkaido, Japan was included in the phylogenetic analysis. There were three mtDNA lineages: Hokkaido, Far Eastern, and northern Eurasia. Divergence data among lineages demonstrated that the Hokkaido group separated from the other groups during the Holsteinian interglacial. The phylogeographical pattern of P. volans is different from that previously reported for terrestrial rodents associated with treeless habitats. Unlike grasslands, forests decreased during glaciation and moved southward because of the cold and arid environmental conditions. The glacial refugia of P. volans would have been associated with forest dynamics in the Pleistocene.

Intrageneric diversity of the cytochrome b gene and phylogeny of Eurasian species of the genus Mustela (Mustelidae, Carnivora).

Abstract To illuminate molecular phylogenetic relationships among Eurasian species of the genus Mustela (Mustelidae, Carnivora), we determined nucleotide sequences of the complete mitochondrial cytochrome b gene region (1,140 base pairs). Molecular phylogenetic trees, constructed using the neighbor-joining and the maximum likelihood methods, showed the common topology of species relationships to each other. The American mink M. vison first branched off and was positioned very remotely from the other species of Mustela. Excluding M. vison, the ermine M. erminea first split from the rest of the species. Two small body-sized weasels, the least weasel M. nivalis and the mountain weasel M. altaica, comprised one cluster (named “the small weasel group”). The other species formed another cluster, where the remarkably close relationships among the domestic ferret M. furo, the European polecat M. putorius, and the steppe polecat M. eversmanni were noticed with 87–94% bootstrap values (named “the ferret group”), supporting the history that the ferret was domesticated from M. putorius and/or M. eversmanni. The European mink M. lutreola was the closest to the ferret group. The genetic distance between the Siberian weasel M. sibirica and the Japanese weasel M. itatsi corresponded to differences of interspecific level, while the two species were relatively close to M. lutreola and the ferret group. These results provide invaluable insight for understanding the evolution of Mustela as well as for investigating the hybridization status between native and introduced species for conservation.

Low Genetic Diversity in Japanese Populations of the Eurasian Badger Meles meles (Mustelidae, Carnivora) Revealed by Mitochondrial Cytochrome b Gene Sequences

Abstract To assess the level of genetic variations of the Eurasian badger Meles meles in Japan, the entire sequences (1,140 base pairs) of the mitochondrial cytochrome b gene were phylogenetically examined. Most of substitutions between haplotypes were transitions resulting in synonymous mutations. A phylo-genetic tree reconstructed by sequence differences clearly showed that Japanese populations of Meles meles were differentiated from continental populations (from the Baikal area and eastern Europe) of M. meles. By contrast, genetic distances among Japanese populations were much smaller, and their geographic structures did not reflect geographic distances between sampling localities. The results indicate that polymorphisms of the ancestral populations still remain via loss of haplotypes by population size changes. In addition, M. meles could have occupied the present habitats in Japanese main islands (Honshu, Shikoku, and Kyushu) in a short period, possibly after the last glacial age.

MORPHOLOGICAL VARIABILITY AND EVOLUTION OF THE BACULUM (OS PENIS) IN MUSTELIDAE (CARNIVORA)

Abstract We examined morphological variability and evolution of the baculum (os penis) across the Mustelidae through allometric analyses and character mapping. Fifty-four species and 26 genera (including 2 fossil forms) were examined with numerous caniform out-group species. Allometric analyses showed that bacular length is relatively constant across mustelids and caniforms; only a tendency to a slightly shortened baculum in mephitines was observed. Character mapping revealed the ancestral mustelid baculum to be an elongated rod-shaped bone that lacks a urethral groove and possesses a simple, nondistinct distal tip. This form is largely retained in mephitines and, to a lesser degree, in lutrines. From the ancestral condition, it is possible to derive forms with a more complicated head that has projections and openings (e.g., melines, Eira barbara, Galictis, Gulo gulo, Martes) or spoon-shaped and cup-shaped processes (e.g., Ictonyx, Mellivora capensis). Another evolutionary trajectory involves the distal tip of the baculum becoming hook-shaped and the urethral groove well developed (e.g., Mustela, Vormela peregusna). Although the structure of the baculum distinguishes closely related species, many features are derived independently in more distantly related forms. Therefore, bacular structure provides restricted phylogenetic information and should be analyzed in concert with other data sources (e.g., morphology of the basicranial region).

Molecular phylogeny and taxonomy of the genus Mustela (Mustelidae, Carnivora), inferred from mitochondrial DNA sequences: New perspectives on phylogenetic status of the back-striped weasel and American mink

ABSTRACT To further understand the phylogenetic relationships among the mustelid genus Mustela, we newly determined nucleotide sequences of the mitochondrial 12S rRNA gene from 11 Eurasian species of Mustela, including the domestic ferret and the American mink. Phylogenetic relationships inferred from the 12S rRNA sequences were similar to those based on previously reported mitochondrial cytochrome b data. Combined analyses of the two genes demonstrated that species of Mustela were divided into two primary clades, named “the small weasel group” and “the large weasel group”, and others. The Japanese weasel (Mustela itatsi) formerly classified as a subspecies of the Siberian weasel (M. sibirica), was genetically well-differentiated from M. sibirica, and the two species clustered with each other. The European mink (M. lutreola) was closely related to “the ferret group” (M. furo, M. putorius, and M. eversmanii). Both the American mink of North America and the back-striped weasel (M. strigidorsa) of Southeast Asia were more closely related to each other than to other species of Mustela, indicating that M. strigidorsa originated from an independent lineage that differs from other Eurasian weasels. Based on biochemical, cytogenetic, and molecular differences as well as morphological evidence, it is proposed that the American mink be elevated to a distinct mustelid genus, Neovison.

The Laptev Sea walrus Odobenus rosmarus laptevi: an enigma revisited

The walrus (Odobenus rosmarus) is in some current systematic schemes divided into three subspecies: O. r. rosmarus in the North Atlantic, O. r. divergens in the North Pacific and O. r. laptevi in the Laptev Sea. These three subspecies have been described as differing in body size, but the taxonomic status of O. r. laptevi is disputed. The current study applies molecular and morphometric methods to assess the taxonomic status of O. r. laptevi and to analyse the systematic and phylogeographic relationships between the three purported walrus subspecies. Tusk length and tusk circumference were measured from the few skulls available of O. r. laptevi, and the obtained values were within the ranges reported for Pacific walruses. Thus, morphologically, subspecies status for O. r. laptevi is not supported according to the Amadon–Mayr ‘75% rule’. Phylogenetic analyses and haplotype networks based on mitochondrial nucleotide sequence data of NADH dehydrogenase 1, 16S rRNA, cytochrome oxidase I and the d‐loop of the control region of the historic O. r. laptevi bone material and contemporary O. r. rosmarus and O. r. divergens showed that the Laptev Sea walrus groups with individuals from the North Pacific. Thus, the mitochondrial sequence data do not support the recognition of three walrus subspecies as reciprocally monophyletic evolutionary units with independent evolutionary histories. Only O. r. rosmarus and O. r. divergens meet this criterion with the present sampling. Accordingly, we recommend that Odobenus r. laptevi be abandoned and the Laptev walrus instead be recognized as the westernmost population of the Pacific walrus, Odobenus r. divergens. However, further research is recommended to assess whether the Laptev walrus could be considered as a significant unit in terms of conservation and management, since it is unique in several ecological parameters.

Comparative phylogeography between the ermine Mustela erminea and the least weasel M. nivalis of Palaearctic and Nearctic regions, based on analysis of mitochondrial DNA control region sequences.

Abstract Phylogeography of the ermine Mustela erminea and the least weasel M. nivalis from Palae-arctic and Nearctic regions were investigated based on mitochondrial DNA control region sequences. Mustela erminea exhibited a very low level of genetic variation, and geographic structures among populations were unclear. This may indicate that M. erminea recently reoccupied a wide territory in Eurasia following the last glacial retreat. In comparison with M. erminea, genetic variations within and among populations of M. nivalis were much greater. Molecular phylogenetic relationships showed that two lineages of M. nivalis occurred in the Holarctic region: one spread from the Eurasian region to North America, and the other occurred in south-eastern Europe, the Caucasus and Central Asia. The results suggest either mitochondrial DNA introgression among populations of south-eastern Europe, the Caucasus and Central Asia, or ancestral polymorphisms remaining in those populations. Contrastive phylogeographic patterns between the two mustelid species could reflect differences of their migration histories in Eurasia after the last glacial age.

Phylogeographic Sympatry and Isolation of the Eurasian Badgers (Meles, Mustelidae, Carnivora): Implications for an Alternative Analysis using Maternally as Well as Paternally Inherited Genes

In the present study, to further understand the phylogenetic relationships among the Eurasian badgers (Meles, Mustelidae, Carnivora), which are distributed widely in the Palearctic, partial sequences of the mitochondrial DNA (mtDNA) control region (539–545 base-pairs) as a maternal genetic marker, and the sex-determining region on the Y-chromosome gene (SRY: 1052–1058 base-pairs), as a paternal genetic marker, were examined. The present study revealed ten SRY haplotypes from 47 males of 112 individuals of the Eurasian Continent and Japan. In addition, 39 mtDNA haplotypes were identified from those animals. From the phylogeography of both the uniparentally inherited genes, four lineages were recognized as Japanese, eastern Eurasian, Caucasian, and western Eurasian. The distribution patterns of the mtDNA lineages showed the existence of a sympatric zone between the eastern and western Eurasian lineages around the Volga River in western Russia. Furthermore, the present study suggested that in the Japanese badgers, the larger genetic differentiation of the Shikoku population was attributable to geographic history in the Japanese islands.