Charrs of the genus Salvelinus (including Salvethymus) represent a monophyletic group of salmo nid fishes that diverged from the common ancestor without subdivision into subgenera. The phylogenesis of the genus is characterized by four cycles of mitochondrial genome divergence. The first one, belonging to the Late Miocene-the border between Miocene and Pliocene (6 to 4 million years ago)-was associated with the consecutive divergence of the S. fontinalis, S. namaycush, S. levanidovi, and S. leucomaenis basal branches. Two divergence events, including separation of the ancestral lineage of Western Pacific group of S. m. krascheninnikovi and the following segregation of the common ancestor into two mitochondrial phyla, happened within the period of 3 to 2 million years ago. The next cycle is attributed to the time interval of about 1 million years ago and includes the divergence of both phyla. In one phylum, a relatively quick isolation of Arctic and Eastern Pacific phylogroups, along with the divergence of the latter phylogroup into S. confluentus and S. m. lordi lineages, took place. At the same time, the second phylum diverged into the S. m. malma and S. alpinus phylogenetic groups. At the final stage (Middle to Late Pleistocene), differentiation of the taxa within the phylogenetic groups took place.

-Phylogenetic analysis of charrs of the genus Salvelinus was performed using PCR-RFLP analysis of mitochondrial DNA (mtDNA). Three fragments of mtDNA (ND1/ND2, ND5/ND6 and Cyt b/D-loop) were amplified by PCR and examined for restriction site variation using 13 restriction endonucleases. Analysis of 313 sites of over 7672 bp of charr mtDNA was conducted. Six phylogenetic lineages of the mtDNA were revealed, corresponding to six separate charr taxa: Salvelinus leucomaenis Pallas; Salvelinus levanidovi Chereshnev, Skopetz & Gudkov; Salvelinus taranetzi Kaganovsky; Salvelinus malma krascheninnikovi Taranetz; Salvelinus malma malma Walbaum and Salvelinus alpinus alpinus. The Levanidov (S. levanidovi) and white-spotted (S. leucomaenis) charrs represented the most divergent lineages. These are probably closest to a common ancestor of the genus. Salvelinus taranetzi was the next divergent lineage. Salvelinus malma krascheninnikovi was placed between S. leucomaenis and S. taranetzi. Salvelinus malma malma and S. a. alpinus were sister taxa, and were on the last branch to diverge. The divergence between S. m. malma and S. a. alpinus inferred from mtDNA nucleotide sequences was 1.1%; between S. m. malma and S. taranetzi 2.8%; between S. m. malma and S. m. krascheninnikovi 4%; between S. taranetzi, S. m. krascheninnikovi and S. a. alpinus 2.8%. Salvelinus levanidovi and S. leucomaenis were almost equally remote from the other charrs by some 9% and 7%, respectively. The extent of genetic differences between the northern S. m. malma populations and the European S. a. alpinus populations indicates that they diverged recently. The existence of many putative charr species with different divergence times in northeastern Asia suggests that this region may be one of the centres of speciation of the genus Salvelinus.

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The geographic distribution pattern of mitochondrial DNA (control region) sequence polymorphisms from 73 populations of a salmonid fish, Dolly Varden Salvelinus malma, over most of its range in the North Pacific rim, was examined to assess how its spatial population genetic structure has been molded. The observed 68 haplotypes were grouped into three main lineages, which correspond to western, central, and eastern regions in the North Pacific. The two outlier-haplotype groups gave close agreement with DNA types from two congeneric species, white-spotted charr S. leucomaenis and Arctic charr S. alpinus, respectively. These results suggest that the present-day genetic structure of S. malma reflects historical patterns of isolation and re-colonization, and also historical hybridization with co-distributed species. We also placed the haplotypes of S. malma within our study areas into a pre-existing evolutionary relationship of S. alpinus and S. malma throughout the Northern Hemisphere. Phylogenetic analysis revealed that the Western Lineage S. malma was basal to all other lineages of S. malma and S. alpinus. Our data serve as a biogeographic hypothesis for salmonid fishes that the Sea of Japan and/or Sea of Okhotsk regions represents a place of origin for S. malma and S. alpinus groups currently distributed in circumpolar regions.

Lake Kronotskoye (the Kronotsky Biosphere State Reserve, south-eastern Kamchatka Peninsula) contains three closely related resident morphotypes charrs, which are considered to be either independent species (white charr Salvelinus albus, longhead charr Salvelinus kronocius, Schmidt's charr Salvelinus schmidti) or a united lacustrine-riverine charrs, represented by several phenotypes. Salvelinus malma malma is isolated from the lake charr populations by an upstream migration barrier in the Kronotskaya River, which flows out of the lake. We examined the level of genetic variability and divergence of mtDNA both between charrs from Kronotsky Lake and between resident lacustrine charrs and the hypothetical ancestor species S. m. malma. The RFLP-PCR analysis was used to examine six regions (ND1/ ND2, ND3/ND4L/ND4, ND5/ND6, COI/COII/A8, A8/A6/COIII/ND3, and Cytb/D-loop), comprising *80% of the mtDNA. Significantly different levels of diversity were found among the populations of lacustrine charrs. S. albus and S. schmidti had the highest indices of mtDNA diversity among the investigated populations from the different habitats. Heterogeneity tests revealed highly significant differentiation among lake populations and among riverine (Kronotskaya River) and lake (Lake Kronotskoye) populations of charrs, indicating their reproductive isolation. Hierarchical analysis of molecular variance revealed the following regularities of diversity distribution: the high proportion of interpopulation variation (93.25%) and low but statistically significant subdivision between charr populations (6.75%, P \ 0.001). Results of the present study suggest that the populations of S. albus, S. kronocius, S. schmidti belong to the S. m. malma phylogenetic group. The divergence value of mtDNA of resident charrs (0.0357-0.0010%) does not exceed the intraspecific variability of S. m. malma. The analysis of the mtDNA haplotypes genealogy of charrs showed that the low values of nucleotide divergence reflect a short period from the beginning of divergence from the ancestral lineages and are due to ancestral polymorphism, as well as to haplotype exchange between the diverged phylogenetic groups as a result of introgressive hybridization.Electronic supplementary material The online version of this article (

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