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.     

Morphometric difference between the Novosibirsk and Tomsk chromosome races of<Emphasis Type="Italic">Sorex araneus</Emphasis> in a zone of parapatry

A hybrid zone between the Novosibirsk and Tomsk chromosome races of the common shrewSorex araneus Linnaeus, 1758 was found near Novosibirsk city (West Siberia, Russia) in an area unimpeded by geographic barriers. In this zone, the shrews of both races and their hybrids were trapped and karyotyped and 22 features of their cranial and postcranial skeleton were measured. Canonical discriminant analysis revealed 3 distinct groups of individuals, which corresponded to the 3 karyotypic categories involved in the analysis. The first discriminant function reflected the differences in the size of skeletal elements. The Novosibirsk shrews and the hybrids were significantly smaller than the Tomsk shrews. The second discriminant function was interpreted as a parameter of skeletal proportionality. The hybrids were significantly less proportional than the parental races. This study revealed one of the clearest examples of morphological differentiation between chromosome races of the common shrew.     

Intraspecies phylogenetic connections in Sorex araneus L.: a southern Baltic subgroup of chromosome races

Results of chromosome G-banding were used to identify two new karyotypic races of the common shrew (Sorex araneus L.) in the European part of Russia. The chromosomal diagnosis of race St. Petersburg included five metacentrics (hk, ip, jl, mq, and nr) and two acrocentrics (g and o) (2Na = 20); the diagnosis of race West Dvina included six metacentrics (gm, hk, ip, jl, no, and qr) (2Na = 18). The phylogenetic significance of the chromosomal markers gm, hk, and ip is considered and the possibility of reticulate evolution in the species S. araneus is discussed.     

Multilevel subdivision in the <Emphasis Type="Italic">araneus</Emphasis> species group of the genus <Emphasis Type="Italic">Sorex</Emphasis>: 1. Chromosomal differentiation

This review summarizes the available data on multilevel differentiation in the araneus species group of the genus Sorex (Eulipotyphla, Mammalia). Communication 1 gives basic information about the biology and morphological and karyotypic differentiation of some species belonging to this genus, as well as about the subdivision of one of the species, the common shrew (Sorex araneus), into intraspecific chromosomal races. The list of the 75 races known so far is included (26 of them dwell in Russia).     

Craniometric differences between karyotypic races of the common shrew Sorex araneus (Mammalia) as a result of limited hybridization

The contact points of four karyotypic races (St. Petersburg, Moscow, Seliger and West Dvina) of the common shrew Sorex araneus L. were studied at the Valdai Hills (European Russia) in an area unimpeded by geographic barriers. The populations of the races are separated by narrow hybrid zones that represent the most complex heterozygous hybrid karyotypes. At these points of contact, the morphometric differentiation of karyotype races was examined in 12 cranial measurements in 190 shrews of a known karyotype. A comparison of the mean values in studied samples of immature shrews revealed statistically significant differences and the correlation of some measurements which describe the level of musculus temporalis. It has been proposed that morphometric differences in the karyotypic races were preserved and accumulated because of a 50% reduction of the frequencies of hybrids. The deviation from the Hardy-Weinberg ration in the frequencies of the genotype and haploid sets of chromosomes in the hybrid zones can be attributed to a number of fatalities of hybrid embryos or the nonrandom mating of karyotypic races. The ethological isolation might arise in the evolution of some karyotypic races from the reduced fitness of the hybrids.     

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.     

Intraspecific Phylogenetic Relationships in Sorex araneus L.: The Southern Baltic Subgroup of Chromosome Races

Results of chromosome G-banding were used to identify two new karyotypic races of the common shrew (Sorex araneus L.) in the European part of Russia. The chromosomal diagnosis of race St. Petersburg included five metacentrics (hk, ip, jl, mq, and nr) and two acrocentrics (g and o) (2Na = 20); the diagnosis of race West Dvina included six metacentrics (gm, hk, ip, jl, no, and qr) (2Na = 18). The phylogenetic significance of the chromosomal markers gm, hk, and ip is considered and the possibility of reticulate evolution in the species S. araneus is discussed.     

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%.     

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.     

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%.     

A new chromosomal race of the common shrew <Emphasis Type="Italic">Sorex araneus</Emphasis> (Lipotyphla,Mammalia) found in Russia

Karyotyping of the common shrews Sorex araneus from the poorly studied region of the Northwest of Russia revealed the belonging of individuals to two chromosomal races, Lemi and St. Petersburg. The Lemi race, previously known only from Finland, was discovered for the first time on the territory of Russia and thus replenished the list of S. araneus chromosomal races that are distributed within the Russian part of the species range (27 races out of 74 known in the whole species range at the moment). The studied individuals showed a high level of chromosomal polymorphism, caused by Rb translocations.     

Phenotypic Variation across Chromosomal Hybrid Zones of the Common Shrew (Sorex araneus) Indicates Reduced Gene Flow

Sorex araneus, the Common shrew, is a species with more than 70 karyotypic races, many of which form parapatric hybrid zones, making it a model for studying chromosomal speciation. Hybrids between races have reduced fitness, but microsatellite markers have demonstrated considerable gene flow between them, calling into question whether the chromosomal barriers actually do contribute to genetic divergence. We studied phenotypic clines across two hybrid zones with especially complex heterozygotes. Hybrids between the Novosibirsk and Tomsk races produce chains of nine and three chromosomes at meiosis, and hybrids between the Moscow and Seliger races produce chains of eleven. Our goal was to determine whether phenotypes show evidence of reduced gene flow at hybrid zones. We used maximum likelihood to fit tanh cline models to geometric shape data and found that phenotypic clines in skulls and mandibles across these zones had similar centers and widths as chromosomal clines. The amount of phenotypic differentiation across the zones is greater than expected if it were dissipating due to unrestricted gene flow given the amount of time since contact, but it is less than expected to have accumulated from drift during allopatric separation in glacial refugia. Only if heritability is very low, N_e very high, and the time spent in allopatry very short, will the differences we observe be large enough to match the expectation of drift. Our results therefore suggest that phenotypic differentiation has been lost through gene flow since post-glacial secondary contact, but not as quickly as would be expected if there was free gene flow across the hybrid zones. The chromosomal tension zones are confirmed to be partial barriers that prevent differentiated races from becoming phenotypically homogenous.     

A hybrid zone between the Kirillov and Petchora chromosomal races of the common shrew (Sorex araneus L., 1758) in northeastern European Russia: a preliminary description

Karyotypes of the Petchora and Kirillov chromosomal races of the common shrew differ by six Robertsonian metacentrics with monobrachial homology, such that interracial F₁ hybrids produce a ring-of-six configuration at meiosis I and are expected to suffer infertility. Mapping of 52 karyotyped individuals by using a unique global positioning system (GPS) revealed that the Kirillov-Petchora hybrid zone is positioned close to the river Mezen, which separated these races, and so may limit the migration of shrews across the contact zone. Although the population density of shrews was found to be markedly different with respect to habitats, the zone runs through a mosaic of habitats that are similar for both the Petchora and the Kirillov sides. This is one of the narrowest chromosomal hybrid zones among those studied in Sorex araneus with a standard cline width of about 1 km. The center of the cline is located on a bank occupied by the Petchora race at a distance of 0.4 km away from a riverine barrier. Interestingly, both the Kirillov race and hybrid individuals were found on a small island in the middle of a river fully flooded each spring. The frequencies of karyotypic variants allow us to consider the zone as an example of a bimodal zone. New Robertsonian and de novo whole-arm reciprocal translocations (WART) chromosomal variants found in the zone could be regarded as evidences of current evolutionary process in chromosomal hybrid zones.     

Chromosomal evolution of the Common Shrew Sorex araneus L.from the Southern Ural and Siberia in the postglacial period]

This paper summarizes a series of studies on chromosomal geography of the common shrew Sorex araneus L. 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 branch, 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 paleophysiology of the glacial period allowed us to reconstruct the sequence of events leading to the establishment of the present-day structure of the species.     

Interracial and population variability of phenotypic (cranial) characters in the common shrew <Emphasis Type="Italic">Sorex araneus</Emphasis> L., 1758

Variability of the cranial properties of chromosomal races Serov, Manturovo and Petchora 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.     

Narrow hybrid zone between Moscow and Western Dvina chromosomal races and specific features of population isolation in common shrew <Emphasis Type="Italic">Sorex araneus</Emphasis> (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.     

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.     

Craniometric study of the common shrew (Sorex araneus L. 1758) from different localities and chromosomal races across Germany and Europe

A total of 440 skulls of common shrews, Sorex araneus, from Germany and Europe (Croatia, Hungary, Austria, and Norway) were studied. The material represented six chromosomal races (Ulm, Laska, Drnholec, Mooswald, Jütland, and Abisko) assignable to the Western European and Northern European karyotypic groups. The race of a few samples was not determined. Twenty-one linear measurements were taken on skulls and mandibles and used in this study. Pearson correlations and multiple linear regressions were used to see the relationship of the cranial variables to altitude, latitude, the chromosomal race, and the geographic location. The results from the tested samples differed; the most negative correlations to latitude were found in the samples assigned to the Western European karyotypic group (WEK), the least negative ones in the samples of the Ulm race. These results indicate the converse of Bergmann's rule. But taking into consideration all the samples of the different karyotypic groups across Europe, the correlations to latitude included positive ones, which would indicate that Sorex follows Bergmann's rule in some of the variables. The studied material of different karyotypic groups could not be differentiated in discriminant analyses. The separation of the studied races within the WEK alone was slightly better, but about 30 % of ungrouped cases remained. Only the separation of the regional samples within one chromosomal race revealed better results but was still very different between the races. This indicates that within the races, regional differences might be strong enough for a separation of the samples and that within a karyotypic group, and even more so across karyotypic groups, regional differences conceal racial differences.     

No apparent reduction of gene flow in a hybrid zone between the West and North European karyotypic groups of the common shrew, Sorex araneus

The common shrew, Sorex araneus, exhibits an unusually high level of karyotypic variation. Populations with identical or similar karyotypes are defined as chromosome races, which are, in turn, grouped into larger evolutionary units, karyotypic groups. Using six microsatellite markers, we investigated the genetic structure of a hybrid zone between the Sidensjö and Abisko chromosome races, representatives of two distinct karyotypic groups believed to have been separated during the last glacial maximum, the West European karyotypic group (western group) and the North European karyotypic group (northern group), respectively. Significant F_ST values among populations suggest some weak genetic structure. All hierarchical levels show similar levels of genetic differentiation, equivalent to levels of genetic structure in several intraracial studies of common shrew populations from central Europe. Notably, genetic differentiation was of the same order of magnitude between and within karyotypic groups. Although the genetic differentiation was weak, the correlation between genetic and geographical distance was positive and significant, suggesting that the genetic variation observed between populations is a function of geographical distance rather than racial origin. Hence, considerable chromosomal differences do not seem to prevent extensive gene flow.     

Genetic and karyotypic structure in the shrews of the Sorex araneus group: are they independent?

The species of the common shrew (Sorex araneus) group are morphologically very similar but exhibit high levels of karyotypic variation. Here we used genetic variation at 10 microsatellite markers in a data set of 212 individuals mostly sampled in the western Alps and composed of five karyotypic taxa (Sorex coronatus, Sorex antinorii and the S. araneus chromosome races Cordon, Bretolet and Vaud) to investigate the concordance between genetic and karyotypic structure. Bayesian analysis confirmed the taxonomic status of the three sampled species since individuals consistently grouped according to their taxonomical status. However, introgression can still be detected between S. antinorii and the race Cordon of S. araneus. This observation is consistent with the expected low karyotypic complexity of hybrids between these two taxa. Geographically based cryptic substructure was discovered within S. antinorii, a pattern consistent with the different postglaciation recolonization routes of this species. Additionally, we detected two genetic groups within S. araneus notwithstanding the presence of three chromosome races. This pattern can be explained by the probable hybrid status of the Bretolet race but also suggests a relatively low impact of chromosomal differences on genetic structure compared to historical factors. Finally, we propose that the current data set (available at http://www.unil.ch/dee/page7010_en.html#1) could be used as a reference by those wanting to identify Sorex individuals sampled in the western Alps.