Chromosomal Evolution of the Common Shrew Sorex araneusL. from the Southern Urals and Siberia in the Postglacial Period

This paper summarizes a series of studies on chromosomal geography of the common shrew Sorex araneusL. in Siberia and the Southern Urals. Chromosomal races inhabiting the Southern Urals and the Western Siberian Plain sequentially replace each other in the latitudinal direction. In this region, karyotypes of each two adjacent races differ from each other by a single whole-arm reciprocal translocation. In the Eastern Siberian and Altai branches, the neighboring races differ mainly in the number or set of metacentric chromosomes. Analysis of the race distribution in the common shrew in the context of paleoecology of the glacial and postglacial period allowed us to reconstruct the sequence of events leading to the establishment of the present-day structure of the species S. araneus.     

Hybrid zones between chromosome races of the common shrew in West Siberia

In West Siberia, the whole species range of the common shrew (Sorex araneus L.) is shared by two parapatric chromosome races — Novosibirsk and Tomsk. These races form a hybrid zone with each other. In addition, on the western margin of the range there is a hybrid zone between the Novosibirsk race and the Uralian race Serov, and in the east, the Tomsk race forms a hybrid zone with the East Siberian race Strelka hybrid. The structures of the three hybrid zones are very different and depend on the karyotypic state of the races in contact. A comparative analysis of the hybrid zones between the chromosome races of the common shrew in West Siberia is discussed together with the role of these zones in maintaining variability in natural populations.     

Population structure of common juniper in the Cis-Urals and Southern Urals

The population structure of common juniper (Juniperus communis L.) growing in the Cis-Urals and Southern Urals has been studied using 17 morphological traits of generative and vegetative organs. A multivariate analysis of ten coenopopulations has recognized three phenotypically different local populations: Cis-Ural, forest Cis-Ural forest-steppe, and Southern Ural mountain populations. The Cis-Ural forest population is strictly associated with lowland pine forests of the northwestern part of the Bashkir Cis-Urals. The Cis-Ural forest-steppe population is located in the northwestern part of the Bashkir Cis-Urals and the southeastern part of the Udmurt Cis-Urals. The Southern Ural mountain population is located in the central part of the Southern Urals and is associated mainly with mountain pine and dark coniferous forests. The last population is divided into forest and forest-edge subpopulations; the first one is represented by typical undergrowth locations, whereas the second is associated with open steppelike slopes and forest edges. In general, based on morphological traits of generative organs, the revealed local subpopulations hold an intermediate position between the Eastern European and Siberian populations of common juniper. Based on the morphological traits of vegetative organs, Cis-Ural populations are considered related to the populations of the European part of Russia, whereas the mountain Southern Ural population resemble Siberian populations. Concerning morphological traits of generative organs, the intrapopulation phenotypic diversity of common juniper is higher for mountain habitats; in the case of vegetative organs, the maximum diversity is observed for lowland habitats. The character of phenotypic differentiation determines the need to conserve the gene pool of common juniper of the Cis-Urals and southern Urals on a population basis.     

Molecular variability in the commom shrew Sorex araneus L. from European Russia and Siberia inferred from the length polymorphism of DNA regions flanked by short interspersed elements (Inter-SINE PCR) and the relationships between the Moscow and Seliger chromosome races

Genetic exchange among chromosomal races of the common shrew Sorex araneus and the problem of reproductive barriers have been extensively studied by means of such molecular markers as mtDNA, microsatellites, and allozymes. In the present study, the interpopulation and interracial polymorphism in the common shrew was derived, using fingerprints generated by amplified DNA regions flanked by short interspersed repeats (SINEs)-interSINE PCR (IS-PCR). We used primers, complementary to consensus sequences of two short retroposons: mammalian element MIR and the SOR element from the genome of Sorex araneus. Genetic differentiation among eleven populations of the common shrew from eight chromosome races was estimated. The NP and MJ analyses, as well as multidimensional scaling showed that all samples examined grouped into two main clusters, corresponding to European Russia and Siberia. The bootstrap support of the European Russia cluster in the NJ and MP analyses was respectively 76 and 61%. The bootstrap index for the Siberian cluster was 100% in both analyses; the Tomsk race, included into this cluster, was separated with the bootstrap support of NJ/MP 92/95%.     

Holarctic phylogeography of the tundra shrew (Sorex tundrensis) based on mitochondrial genes

A range‐wide phylogeographic study of the tundra shrew (Sorex tundrensis) was performed using cytochrome b and cytochrome oxidase I (COI) mitochondrial genes. The results based on 121 specimens from 42 localities demonstrate that the tundra shrew is divided into five main mitochondrial DNA phylogenetic lineages with largely parapatric distribution. In addition to a single Nearctic clade (Alaska) four Palearctic clades are identified: Western (Northen Urals, Kazakhstan, South‐West Siberia), Eastern (from East Transbaikalia and the Middle Amur to Chukotka), South Central (Central Siberia, the Altai, the Dzhungarian Alatau) and North Central (Northern Siberia, Central Yakutia). Date estimates obtained by use of a molecular clock corrected for potential rate decay suggest Late Pleistocene age for the most recent common ancestor of all contemporary tundra shrew populations. Relatively high genetic divergence between phylogroups (0.95–1.6%) indicates that the observed phylogeographic structure was initiated by historical events that predated the Last Glacial Maximum. We assume that, being more cold‐ and arid‐tolerant, tundra shrew underwent expansion during an early cold phase of the Last Glacial and spread through its recent range earlier than most of other Siberian red‐toothed shrews. Comparative phylogeographic analysis of Siberian shrews and rodents suggests that evolutionary histories of species associated with azonal or open habitats show important differences compared to forest species. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101 , 721–746.     

Molecular variability in the common shrew Sorex araneus L. from european russia and siberia inferred from the length polymorphism of DNA regions flanked by short interspersed elements (inter-SINE PCR) and the relationships between the moscow and seliger chromosome races

Genetic exchange among chromosomal races of the common shrew Sorex araneus and the problem of reproductive barriers have been extensively studied by means of such molecular markers as mtDNA, microsatellites, and allozymes. In the present study, the interpopulation and interracial polymorphism in the common shrew was derived, using fingerprints generated by amplified DNA regions flanked by short interspersed repeats (SINEs)—interSINE PCR (IS-PCR). We used primers, complementary to consensus sequences of two short retroposons: mammalian element MIR and the SOR element from the genome of Sorex araneus. Genetic differentiation among eleven populations of the common shrew from eight chromosome races was estimated. The NJ and MP analyses, as well as multidimensional scaling showed that all samples examined grouped into two main clusters, corresponding to European Russia and Siberia. The bootstrap support of the European Russia cluster in the NJ and MP analyses was respectively 76 and 61%. The bootstrap index for the Siberian cluster was 100% in both analyses; the Tomsk race, included into this cluster, was separated with the bootstrap support of NJ/MP 92/95%.     

The microsatellite polymorphism and gene flow in the contact zone of four common shrew (Sorex araneus L., Mammalia) chromosome races

The variation of microsatellite loci in 130 individuals of four common shrew chromosome races (Moscow, Western Dvina, Seliger, and St. Petersburg) contacting on the Valdai Hills was studied. A low level of genetic differences between the chromosome races, which differ at three-five fixed diagnostic metacentric chromosomes, was found. The genetic differentiation within the races is more considerable as compared with that between the races. A high deficiency in heterozygotes was recorded; presumably, this is connected with regular variation in the population sizes. It is assumed that the fixation of centric chromosome fusions was supported by selection (drive) in the evolution of the common shrew against the background of a neutral evolution of the microsatellite loci.     

Genetic variation and population differentiation in Siberian fir Abies sibirica lebed. Inferred from allozyme markers

Genetic variation in 24 populations of Siberian fir Abies sibirica Lebed. from the Urals, West Siberia, East Siberia, South Siberia, and the Baikal region were examined using allozyme markers. Three out of fifteen allozyme loci proved to be polymorphic. Heterozygosity He was 6.6-9.6%, which is substantially lower than that in other widely spread boreal conifers. Our results suggest that the Siberian fir populations are subdivided into four geographic groups: (1) the Baikal Lake group, (2) the Sayan and the Altai group, (3) the Middle and Southern Urals group, and (4) Subpolar and Northern Urals group. This pattern of geographic differentiation may be explained by the preservation of the Siberian fir during the last glacial maximum (18 000-22000 years B.P.) in isolated refugia with subsequent recolonization of the present area. FST in the populations examined was 10.16%, which is comparable to the estimate for Larix sibirica (7.9%), a conifer species having a similar range and pattern of geographic population differentiation.     

Density-dependent processes determine the distribution of chromosomal races of the common shrew <Emphasis Type="Italic">Sorex araneus</Emphasis> (Lipotyphla,Mammalia)

The common shrew is subdivided into 74 chromosomal races, widely distributed in the postglacial area from the Britain Islands to Lake Baikal. Based on 1969 karyotypes from 216 localities, we present for the first time a map of ranges of 25 chromosomal races (except the Altai race) currently known in Russia. We revealed two centers of high karyotypic diversity: the western (near Baltic Sea) and the eastern (near Baikal Lake). The studied races were categorized as small-, medium-, and large-range races, and small-range races concentrated around those two centers of karyotypic diversity. We did not find any significant association between race range size and ecological zone, latitude, or the ambient temperature. Physical barriers, such as Ural Mountain or rivers, do not limit race distribution. The width of rivers that divide a range of a single race or ranges of two different races does not differ. We supposed that the occupation of an area by a race could limit the invasion of a different race from an adjacent area and expansion of its range, thus contributing to race parapatric distribution alone without additional effects of physical barriers. Based on karyotype similarity and geographic localization, we combined races into four “karyotypic chains,” in which the races can be derived from one another consequently by a single chromosomal translocation. The present distribution of the common shrew races in Russia supports the idea that it has resulted from recolonization from refugia governed by the density-dependent processes.     

Lack of mitochondrial DNA divergence between chromosome races of the common shrew, Sorex araneus, in Sweden. Implications for interpreting chromosomal evolution and colonization history

The common shrew, Sorex araneus, has one of the most variable karyotypes among mammals, displaying numerous chromosome races throughout its distribution. The six chromosome races present in Sweden can be categorized in two different karyotypic groups, the west and north European karyotypic groups (western and northern). Three races belonging to the western group are considered to have arisen through whole arm reciprocal translocations (WARTs). Race formation through this process requires a bottleneck event. In the present study we sequenced a part of the mitochondrial DNA (mtDNA) genome to investigate molecular differences between the chromosome races in Sweden. We found no mtDNA differentiation between the mainland chromosome races or the karyotypic groups. Genetic variation is as large between populations within a race as between populations among the races or karyotypic groups, suggesting that the karyotypic groups might have originated in a common glacial refugium. The noticeable exception is the Oland race, which shows higher mtDNA diversity compared to the other Swedish races, indicating a divergent origin difficult to explain. Mitochondrial DNA variation in Sweden suggests that most haplotypes arose in situ and that the populations has undergone a rapid size expansion. Altogether, the mtDNA data are in agreement with the WART hypothesis, which still holds as the most plausible variant of karyotype evolution for three of the chromosome races of the common shrew in Sweden.     

Algae as indicators of a biogeographical zonation in the early carboniferous of the Urals,Siberia and Northeast Russia

Summary Algal assemblages of the Urals and Siberia belong to two biogeographical belts: The equatorial Mediterranean and the Siberian belt. They differ in the composition of their agla floras. The end of the Famennian and the beginning of the tournaisian were indicated by smaller differences due to short-term connections of the basins which lie in different climatic belts. The development of the microfloras of the Urals and Siberias during the late Tournaisian exhibits two different trends. These differences were intensified during the Visean and reached their climax in the Serpukhovian. In general, redalgae, especially Stacheinaceae, were predominant in the late Carboniferous basins of the Urals. The Visean and Serpukhovian algae of Siberia comprising usually more than 20 genera (e.g.Calcifolium, Fasciella, Ungdarella) were widely distributed in the Urals as well as in other provinces of the Mediterranean belt. The biogeographical zonation based on the study of algal floras confirms the existence of an Eurasian realm in the Mediterranean belt and several Siberian provinces as well as an intermediate Taimyr-Chukotkian area, which had already been postulated from earlier foraminiferal studies. All of these provinces exhibit distinct features in the development of their algal associations, e.g. the Kolyma-Omolon province is characterized by its predominance of green algae, while in the Altai-Sayan and the Verkhoyansk provinces red algae prevail. These characteristic trends can be observed over rather long stretches of time.     

Genetic and morphological variation in a partially isolated population of Caucasian shrew sorex Satunini (Mammalia)

Morphological and genetic variation at microsatellite loci of Caucasian shrew Sorex satunini Ogn. is examined and compared with that of the common shrew S. araneus L. Genetic distance at microsatellite loci between the common shrew and Caucasian shrew proved to be threefold higher than between chromosome races of the common shrew. The Caucasian shrew manifested low polymorphism in studies of both microsatellites and morphometric mandibular traits. The heterozygote deficit was also typical. These properties may be a consequence of partial isolation of the population and gene drift.     

Genetic variation and population differentiation in Siberian fir Abies sibirica ledeb. inferred from allozyme markers

Genetic variation in 24 populations of Siberian fir Abies sibirica Ledeb. from the Urals, West Siberia, East Siberia, South Siberia, and the Baikal region were examined using allozyme markers. Three out of fifteen allozyme loci proved to be polymorphic. Heterozygosity H _e was 6.6–9.6%, which is substantially lower than that in other widely spread boreal conifers. Our results suggest that the Siberian fir populations are subdivided into four geographic groups: (1) the Baikal Lake group, (2) the Sayan and the Altai group, (3) the Middle and Southern Urals group, and (4) Subpolar and Northern Urals group. This pattern of geographic differentiation may be explained by the preservation of the Siberian fir during the last glacial maximum (18 000–22 000 years B.P.) in isolated refugia with subsequent recolonization of the present area. F _ST in the populations examined was 10.16%, which is comparable to the estimate for Larix sibirica (7.9%), a conifer species having a similar range and pattern of geographic population differentiation.     

Genetic and morphological variation in a partially isolated population of Caucasian shrew <Emphasis Type="Italic">Sorex satunini</Emphasis> (Mammalia)

Morphological and genetic variation at microsatellite loci of Caucasian shrew Sorex satunini Ogn. is examined and compared with that of the common shrew S. araneus L. Genetic distance at microsatellite loci between the common shrew and Caucasian shrew proved to be threefold higher than between chromosome races of the common shrew. The Caucasian shrew manifested low polymorphism in studies of both microsatellites and morphometric mandibular traits. The heterozygote deficit was also typical. These properties may be a consequence of partial isolation of the population and gene drift.     

Natural hybridization between extremely divergent chromosomal races of the common shrew (Sorex araneus, Soricidae, Soricomorpha): hybrid zone in European Russia

The Moscow and Seliger chromosomal races of the common shrew differ by Robertsonian fusions and possibly whole‐arm reciprocal translocations (WARTs) such that their F₁ hybrids produce a chain‐of‐eleven configuration at meiosis I and are expected to suffer substantial infertility. Of numerous hybrid zones that have been described in the common shrew, those between the Moscow and Seliger races involve the greatest chromosomal difference. We collected 211 individuals from this zone to generate a total dataset of 298 individuals from 187 unique global positioning system (GPS) locations within the vicinity of interracial contact. We used a geographic information system (GIS) to map the location of the hybrid zone, which follows a direct route between two lakes, as would be anticipated from tension zone theory. Even within the central area of the hybrid zone, there is a much higher frequency of pure race individuals than hybrid, making this a clear example of a bimodal zone in the sense of Jiggins & Mallet (2000) . The zone runs through good habitat for common shrews, but nevertheless it is very narrow (standard cline widths: 3–4 km), as would be anticipated from low hybrid fitness. There is clear potential for an interruption to gene flow and build‐up of reproductive isolation. As found in some other hybrid zones, there is a high frequency of novel genetic variants, in this case, new chromosomal rearrangements. Here, we report a de novo Robertsonian fission and a de novo reciprocal translocation, both for the first time in the common shrew. There is an extraordinarily high frequency of de novo mutations recorded in F₁ hybrids in the zone and we discuss how chromosomal instability may be associated with such hybrids. The occurrence of a de novo Robertsonian fission is of considerable significance because it provides missing evidence that fissions are the basis of the novel acrocentric forms found and apparently selected for in certain common shrew hybrid zones.     

Western Siberia rivers and the boundaries between chromosomal races of the common shrew (<Emphasis Type="Italic">Sorex araneus</Emphasis> L. Lipotyphla,Mammalia)

In Western Siberia, most boundaries between common shrew chromosomal races have been found to pass along the banks of rivers, mainly those flowing in the meridional direction. The races Serov and Novosibirsk co-inhabited the right bank of the Irtysh. The easternmost point of the Novosibirsk race has been found on the middle Yenisei River, while the race Tomsk in this area was only on the right (eastern) bank.     

Narrow hybrid zone between Moscow and Western Dvina chromosomal races and specific features of population isolation in common shrew Sorex araneus (mammalia)

The contact zone between Moscow and Western Dvina chromosomal races of common shrew Sorex araneus L. at the south of the Valdai Hights was traced over a distance of 20 km. Within this, close to parapatric, contact zone of chromosomal races the width of sympatry zone was about 500 m (the narrowest among currently known hybrid zones), and the proportion of hybrids was 24.3%. It was shown that in bimodal hybrid zones between chromosomal races of common shrew the width of sympatry zones varied from 0.5 to 13 km. This width does not correlate with the cytogenetic features of the hybrids, and seems to be determined by competitive relations between the races. The hybrid proportion is determined by the type of hybrid heterozygosity, and decreased in the race sympatry zone from 33-40 to 21.5-25.2%. The decrease of the hybrid proportion can be associated with the abnormal fertility of either the first generation, or the backcross hybrids.     

Distribution of two chromosome races of the common shrew (<Emphasis Type="Italic">Sorex araneus</Emphasis> L.) in the hybrid zone: Can a change of the dispersal mode maintain independent gene frequencies?

Combination of different dispersal modes may itself, without external obstacles, lead to the appearance of subdivided populations and maintain the existence of independent population systems. The common shrew, a mammal convenient for studying different levels of intraspecific differentiation, was the object of the study. Empirical data have been used for simulation taking into account the change of dispersal modes in the population area. The obtained results agree with empirical data on the distribution of races and hybrids in the hybrid zone of chromosome races Moscow and Seliger. Change of the dispersal mode may maintain independent population dynamics and, in the case of chromosome races, prevent the migration of parental individuals into the territory of the other race.     

Peculiarities and Determinants of Regeneration of Siberian Larch on the Upper Limit of Its Growth in the Urals

Quantitative and qualitative characteristics of cones and seeds and the emergence and survival of seedlings in the Siberian larch (Larix sibirica Ledeb.) have been studied in different parts of the ecotone at the upper boundary of arboreal vegetation (treeline ecotone) on Serebryanskii Kamen’ Mountain (Northern Urals) and on the hills (with a landmark 312 m above sea level) surrounding Chernaya Mountain (Polar Urals) from 2005 to 2011. We have found a decrease in the parameters of cones, number of seeds in the cones, their viability in laboratory with an increase in altitude, and differences in the number of seedlings between parts of the treeline ecotone. It is shown that the formation of Siberian larch generations on the Northern Urals occurs only after moist years. It is noted that, in the Polar Urals, Siberian larch produces a big seed crop every 2–3 years. It is proven that the seed production and mortality of seedlings in Siberian larch are influenced not only by air temperature and soil moisture, but also by wind load, snow depth, and soil temperature rapidly changing along the slope.     

Racial and population variability of phenotypic (cranial) characters in the common shrew Sorex araneus L., 1758

Variability of the cranial properties of chromosomal races Serov, Manturovo and Pechora of the common shrew were studied. A consistent increase of scull size in the Serov race with moving from the plain to highlands and a skull size decrease from low to high latitudes was detected. Interpopulation variability among different races was shown to be comparable with interracial variability or to exceed it. This suggests microevolution at the level of local populations.