Nikolai E. Dokuchaev

Intraspecific phylogeny and geographical variation of six species of northeastern Asiatic Sorex shrews based on the mitochondrial cytochrome b sequences

Intraspecific phylogeny and genetic variation were investigated based on nucleotide sequences of the mitochondrial cytochrome b gene in six soricine shrew species, Sorex unguiculatus, S. caecutiens, S. shinto, S. gracillimus, S. minutissimus and S. hosonoi, collected primarily from northeastern Asia. Maximum likelihood trees and a phylogenetic network were generated to estimate intraspecific phylogenies. S. minutissimus showed high congruence between phylogenetic position and geographical origin and S. gracillimus showed low congruence. In contrast, there was no congruence between phylogeny and geography in S. unguiculatus and the S. caecutiens from Sakhalin‐Eurasia. Positive correlation between genetic and geographical distances was found in S. minutissimus and S. gracillimus, but not in the other species (or regional populations). The results of the phylogenetic and genetic analyses suggest that S. minutissimus and S. gracillimus have occupied their present ranges for a longer time than the other species if we assume a stepping‐stone model of population structure. In addition, there was no contradiction between the present investigations and the hypotheses of multiple immigration by S. gracillimus and a single immigration by S. unguiculatus into Hokkaido Island. It is proposed that these six northeastern Asian species experienced different historical processes of range expansion and dispersal despite the fact that some of them currently show similar patterns of distribution.

Beringia: Intercontinental exchange and diversification of high latitude mammals and their parasites during the Pliocene and Quaternary

ABSTRACT Beringia is the region spanning eastern Asia and northwestern North America that remained ice-free during the full glacial events of the Pleistocene. Numerous questions persist regarding the importance of this region in the evolution of northern faunas. Beringia has been implicated as both a high latitude refugium and as the crossroads (Bering Land Bridge) of the northern continents for boreal mammals. The Beringian Coevolution Project (BCP) is an international collaboration that has provided material to assess the pattern and timing of faunal exchange across the crossroads of the northern continents and the potential impact of past climatic events on differentiation. Mammals and associated parasite specimens have been collected and preserved from more than 200 field sites in eastern Russia, Alaska and northwestern Canada since 1999. Previously, fossils and taxonomic comparisons between Asia and North America mammals have shed light on these events. Molecular phylogenetics based on BCP specimens is now being used to trace the history of faunal exchange and diversification. We have found substantial phylogeographic structure in the Arctic and in Beringia in mustelid carnivores, arvicoline rodents, arctic hares and soricine shrews, including spatially concordant clades and contact zones across taxa that correspond to the edges of Beringia. Among the tapeworms of these mammalian hosts, new perspectives on diversity have also been developed. Arostrilepis horrida (Hymenolepididae) was considered to represent a single widespread and morphologically variable species occurring in a diversity of voles and lemmings in eastern and western Beringia and more broadly across the Holarctic region. The BCP has demonstrated a complex of at least 10 species that are poorly differentiated morphologically. The diversity of Paranoplocephala spp. and Anolocephaloides spp. (Anoplocephalidae) in Beringia included relatively few widespread and morphologically variable species in arvicolines. BCP collections have changed this perspective, allowing the recognition of a series of highly endemic species of Paranoplocephala that demonstrate very narrow host specificity, and additional species complexes among arvicolines. Thus, extensive, previously unrecognized, diversity for tapeworms of 2 major families characterizes the Beringian fauna. By elucidating evolutionary relationships and phylogeographic variation among populations, species and assemblages, refined views of the sequence and timing of biotic expansion, geographic colonization and impact of episodic climate change have been developed for Beringia. Ultimately, Beringia was a determining factor in the structure and biogeography of terrestrial faunas across the Nearctic and Neotropical regions during the Pliocene and Quaternary.

Genetic diversity of Echinococcus spp. in Russia

In Russia, both alveolar and cystic echinococcoses are endemic. This study aimed to identify the aetiological agents of the diseases and to investigate the distribution of each Echinococcus species in Russia. A total of 75 Echinococcus specimens were collected from 14 host species from 2010 to 2012. Based on the mitochondrial DNA sequences, they were identified as Echinococcus granulosus sensu stricto (s.s.), E. canadensis and E. multilocularis. E. granulosus s.s. was confirmed in the European Russia and the Altai region. Three genotypes, G6, G8 and G10 of E. canadensis were detected in Yakutia. G6 was also found in the Altai region. Four genotypes of E. multilocularis were confirmed; the Asian genotype in the western Siberia and the European Russia, the Mongolian genotype in an island of Baikal Lake and the Altai Republic, the European genotype from a captive monkey in Moscow Zoo and the North American genotype in Yakutia. The present distributional record will become a basis of public health to control echinococcoses in Russia. The rich genetic diversity demonstrates the importance of Russia in investigating the evolutionary history of the genus Echinococcus.

A spatial aspect on mitochondrial DNA genealogy in Apodemus peninsulae from East Asia.

Apodemus peninsulae is a field mouse that inhabits the broad-leafed forests of temperate Eurasia. We examined the mitochondrial cytochrome b gene in 57 individuals of A. peninsulae from northeastern Asia, including Siberia, Primorye, Magadan region, Sakhalin, Hokkaido, and the Korean Peninsula. The genealogy of the mitochondrial DNA (mtDNA) in A. peninsulae was shown to have substantial geographic affinity, suggesting geographic architecture of northeastern Asia, including the islands of Sakhalin and Hokkaido, played important roles on the cladogenesis. Taking into account the presence of region-specific anciently divergent mtDNA types, three parts of the regions of Primorye, Siberia, and the Korean Peninsula can be denoted as refugia for A. peninsulae during the substantial period of the Quaternary glacial ages. Among the geographic regions examined, Primorye is likely to be the most influential one, from which the mtDNA is thought to have migrated to the neighboring regions of Sakhalin, Hokkaido, the Magadan region, and Siberia during the evolution of this species.

High-latitude diversification within Eurasian least shrews and Alaska tiny shrews (Soricidae)

Abstract A novel shrew was discovered recently in Alaska and described based on morphological characters as Sorex yukonicus. This species is closely allied to Sorex minutissimus, a widespread shrew ranging through Eurasia. Together their distribution spans Beringia, a large Pleistocene nonglaciated area that connected Asia and North America. Beringia was repeatedly divided due to raised sea levels during Pleistocene interglacials and subsequently reconnected during glacials. We tested predictions related to the influence of large-scale geologic events on genetic variability through a phylogeographic analysis of both species of shrew using evidence from 3 independent genetic loci. We found low genetic divergence between S. minutissimus and S. yukonicus across continents. However, major phylogeographic breaks were found for Eurasian and Maritime Northeast Asia populations. Neither species is reciprocally monophyletic for any of the loci examined. Coalescence times for all pairwise population comparisons within both species fall within the Wisconsinan–Weichselian glacial (<130 thousand years ago), and significant population expansion estimates date to the Holocene suggesting that divergence between these taxa is minimal and may not warrant recognition of 2 distinct species. Phylogeographic relationships and sequence divergence estimates place populations of North American S. yukonicus and Siberian S. minutissimus as most closely related, and together they are sister to European S. minutissimus. We conclude that populations east and west of the Bering Strait represent a single Holarctic species, S. minutissimus. Temporal changes in range based on ecological niche predictions and a comparative assessment of other codistributed taxa provide a preliminary view of potential Last Glacial Maximum refugia in northern Asia.

GEOGRAPHIC AND HOST RANGE OF THE NEMATODE SOBOLIPHYME BATURINI ACROSS BERINGIA

The nematode Soboliphyme baturini Petrov, 1930, was found to represent a single species with a relatively broad geographic range across Beringia and northwestern North America on the basis of the assessment of molecular sequence data for adult and juvenile parasites. Refuted are hypotheses suggesting that several cryptic species could be partitioned either among an array of mustelid definitive hosts or across the vast region that links North America and Eurasia. Host specificity for this species is examined on the basis of a comprehensive list for definitive hosts, derived from new field surveys and existing literature for S. baturini. Only 5 mustelids (Gulo gulo, Martes americana, M. caurina, M. zibellina, and Neovison vison) appear to have significant roles in the life history, persistence, and transmission of this nematode. Soboliphyme baturini readily switches among M. americana, M. caurina, Mustela erminea, or N. vison at any particular locality throughout its geographic range in North America, although Martes spp. could represent the source for nematodes in a broader array of mustelids. Molecular analyses (243 base pairs of mitochondrial gene nicotinamide dehydrogenase ;obND4;cb) suggest that hypotheses for host specificity across an array of mustelid definitive hosts are not supported. The life cycle of S. baturini is explored through a review of diet literature for 2 marten species, M. americana and M. caurina, and other mustelids across the Holarctic. Shrews (Soricomorpha: Soricidae) comprise >8% of prey for these species of Martes, suggesting their putative role as paratenic hosts. Juvenile nematodes found in the diaphragms of soricids are genetically identical to adult S. baturini found in the stomachs of mustelids at the same locations in both Asia and North America, corroborating a role in transmission for species of Sorex.

Biogeographical history of northeastern Asiatic soricine shrews (insectivora, mammalia)

A hypothetical biogeographical history of northeastern Asiatic soricine shrews in the late Quaternary was developed by integrating their present distributions, fossil records, a hypothetical phylogeny, and geological investigations. First, a biological area cladogram of the northeastern Asiatic region was constructed by applying the vicariance hypothesis to the phylogeny of thecaecutiens/shinto group, a monophyletic group proposed by Ohdachi et al. (1997). Comparing the biological area cladogram with a geological hypothesis by Ohshima (1990,1991,1992), we hypothesized a geographical history of northeastern Asia. Species were then located on the dendrogram of the geographical history, referring to the present distributions, fossil records, and phylogeny of shrews. According to our hypothesis, higher species diversity of the northern region of northeastern Asia (Hokkaido, Sakhalin, and Eastern Siberia) was achieved by several series of colonizations and habitat expansion. On the other hand, the shrew communities of the southern region (Honshu, Sado, Shikoku, and Kyushu) were created by extinction and isolation followed by speciation.

Nearctic Shrews, Sorex spp., as Paratenic Hosts of Soboliphyme baturini (Nematoda: Soboliphymidae)

Abstract Third-stage larvae (L3) of Soboliphyme baturini were discovered for the first time in shrews, Sorex cinereus and Sorex tundrensis from Alaska and the Nearctic. Shrews were found to be infected with L3 at Suloia Lake, southeastern Alaska, Togiak National Wildlife Refuge, southwestern Alaska, and at the Yukon-Charley Rivers National Preserve near the Yukon-Alaska border. Larvae in shrews from Alaska were morphologically indistinguishable from those known in both insectivores and arvicoline rodents from Russia. The occurrence of S. baturini in Alaskan insectivores further establishes shrews as important hosts in the transmission of S. baturini among mustelids and other carnivores and indicates for the first time the basis for a paratenic cycle in the Nearctic.

Reappraisal of Hydatigera taeniaeformis (Batsch, 1786) (Cestoda: Taeniidae) sensu lato with description of Hydatigera kamiyai n. sp.

The common cat tapeworm Hydatigera taeniaeformis is a complex of three morphologically cryptic entities, which can be differentiated genetically. To clarify the biogeography and the host spectrum of the cryptic lineages, 150 specimens of H. taeniaeformis in various definitive and intermediate hosts from Eurasia, Africa and Australia were identified with DNA barcoding using partial mitochondrial cytochrome c oxidase subunit 1 gene sequences and compared with previously published data. Additional phylogenetic analyses of selected isolates were performed using nuclear DNA and mitochondrial genome sequences. Based on molecular data and morphological analysis, Hydatigera kamiyai n. sp. Iwaki is proposed for a cryptic lineage, which is predominantly northern Eurasian and uses mainly arvicoline rodents (voles) and mice of the genus Apodemus as intermediate hosts. Hydatigera taeniaeformis sensu stricto (s.s.) is restricted to murine rodents (rats and mice) as intermediate hosts. It probably originates from Asia but has spread worldwide. Despite remarkable genetic divergence between H. taeniaeformis s.s. and H. kamiyai, interspecific morphological differences are evident only in dimensions of rostellar hooks. The third cryptic lineage is closely related to H. kamiyai, but its taxonomic status remains unresolved due to limited morphological, molecular, biogeographical and ecological data. This Hydatigera sp. is confined to the Mediterranean and its intermediate hosts are unknown. Further studies are needed to classify Hydatigera sp. either as a distinct species or a variant of H. kamiyai. According to previously published limited data, all three entities occur in the Americas, probably due to human-mediated introductions.

Intraspecific Phylogeny and Nucleotide Diversity of the Least Shrews, the Sorex minutissimus-S. yukonicus Complex, Based on Nucleotide Sequences of the Mitochondrial Cytochrome b Gene and the Control Region

Abstract. Phylogenetic analysis was conducted for various populations of the Sorex minutissimus-S. yukonicus complex based on mitochondrial gene (cytochrome b and/or the control region) sequences. Sorex minutissimus was divided into some monophyletic groups in Eurasia; it was divided into 2 main groups, eastern and western Eurasian clades, based on combined data of the cytochrome b and the control region. Monophyly of shrews from Hokkaido-Sakhalin, Primorye, Mongolia-Transbaikalia, southeastern Finland was strongly supported respectively in most analyses. Sorex yukonicus was phylogenetically close to S. minutissimus in eastern Siberia. Some shrews from western and central Siberia were included in the clade of southeastern Finland. Also, most shrews from central-northern Finland and Norway made a clade close to but different from the southeastern Finland clade. This finding suggests that Fennoscandian shrews might consist of individuals which were recolonised from various refugia after the Last Glacial Maximum. Nucleotide diversity of shrews from Hokkaido and Alaska was low. Three regional groups in Kamchatka-Sakha, Sakhalin, and Mongolia-Transbaikalia tended to have medium nucleotide diversity. In contrast, shrews from Cisbaikalia-western Siberia and Fennoscandia had high nucleotide diversity. The S. minutissimus-S. hosonoi group appears to have experienceed a quit different biogeographic history from two shrews with similar ranges, the S. caecutiens-S. hosonoi group and S. tundrensis.