Cytogenetic control of a hybrid zone between two <Emphasis Type="Italic">Sorex araneus</Emphasis> chromosome races before breeding season

Two chromosome races of common shrew, Moscow and Seliger, differ in the arm combination in 11 diagnostic chromosomes (Robertsonian metacentrics/acrocentrics). Homozygotes of both pure races, simple Robertsonian heterozygotes of Seliger race, and complex heterozygotes (F₁ hybrids) were detected in the found earlier hybrid zone of these races, in the spring before the breeding season. The g/o heterozygote was first discovered in race Seliger, whose chromosome formula typically contains acrocentrics g and o. The m/q heterozygote was recorded for the second time. Meiosis was studied in 16 males representing five detected karyotypic categories. No abnormal in pairing of homologs in either sex trivalent common for the species (XY1Y2) or autosome trivalents (g/o and m/q) was detected at diakinesis-metaphase I. Two hybrids displayed a theoretically expected and unimpaired meiotic configuration in a form of a very long chain comprising 11 monobrachial homologs (g/gm/mq/qp/pr/rk/ki/ih/hn/no/o). The results are discussed in terms of hypotheses on fertility of complex heterozygotes and limited gene flow in hybrid zone.     

Meiotic studies of male common shrews (Sorex araneus L.) from a hybrid zone between chromosome races

Thirty-three adult male common shrews (Sorex araneus L.) were collected from a hybrid zone between two chromosomal races that differed in Robertsonian metacentrics. Anaphase I nondisjunction frequencies were estimated on the basis of metaphase II counts. RIV and CV complex heterozygotes (four-element rings and five-element chains at meiosis I, respectively) had substantially higher nondisjunction rates than homozygotes and simple Robertsonian heterozygotes. However, at least in the case of RIV-forming hybrids, increased nondisjunction frequency did not result from malsegregation of the heterozygous complex. Extra elements found in hyperploid spreads were most frequently acrocentrics, that could not originate from a fully metacentric multivalent. Complex heterozygotes were also characterized by higher frequencies of univalents observed at diakinesis I. However, univalents did not originate from complex configurations, which were regularly formed with usually one chiasma per chromosome arm. Hence, we suppose that the presence of multivalents in the cell affects pairing and segregation of other elements at meiosis I.     

The complex hybrid zone between the Abisko and Sidensjö chromosome races of Sorex araneus in Sweden

Six chromosome races of the common shrew occur in Sweden, each with its characteristic arm combination of metacentric chromosomes. G-banded karyotypes were analysed from 201 common shrews in 14 localities of the northern hybrid zone in Sweden. Analyses from another 64 shrews from seven localities outside the hybrid zone w ere included for comparison. The shrews were classified with respect to karyotype into any of five categories: (1) Abisko race, (2) Sidensjö race, (3) hybrids between the parental races, (4) pseudohybrids (a type of hybrid), and (5) AT with all race-specific chromosomes (h, i, n, o, p, r) present as telocentrics. Hybrids occurred at a frequency close to Hardy-Weinberg expectation in the centre of the hybrid zone. Chromosome polymorphism of Robertsonian type was common and 43 different karyotypes were found among the specimens studied. The polymorphism involved six metacentric pairs in the Abisko and three in the Sidensjö race. The frequency of the Sidensjö race-specific metacentric hi decreased and the frequency of the Abisko race-specific hn increased from south-west to north-east along a transect across the hybrid zone. The number of race-specific telocentrics reached a peak 13 km north-east of the hi-hn cline centre. The estimated standardized cline width for chromosomes hi and hn was 16.0 km. The extension of the Sidensjö race is comparatively narrow [c. 50 km in the region of the investigation), and it is regarded to be a 'hybrid race' between the Uppsala race, which colonized Sweden from the south-west, and the Abisko race which arrived from the north-east after the most recent glaciation. The origin of the Sidensjö race is thus less than 10 000 years old, because earlier this area was covered by glacial ice.     

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.     

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.     

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

Chromosomal races of the common shrew differ in sets of metacentric chromosomes and on contact may produce hybrids with extraordinarily complex configurations at meiosis I that are associated with reduced fertility. There is an expectation that these may be some of the most extreme tension zones available for study and therefore are of interest as potential sites for reproductive isolation. Here, we analyse one of these zones, between the Novosibirsk race (characterized by metacentrics go, hn, ik, jl, mp and qr) and the Tomsk race (metacentrics gk, hi, jl and mn and acrocentrics o, p, q and r), which form hybrids with a chain-of-nine (CIX) and a chain-of-three (CIII) configuration at meiosis I. At the Novosibirsk-Tomsk hybrid zone, the CIX chromosomes form clines of 8.53 km standardized width on average, whereas the cline for the CIII chromosomes was 52.83 km wide. The difference in these cline widths fits with the difference in meiotic errors expected with the CIX and CIII configuration, and we produce estimates of selection against hybrids with these types of configurations, which we relate to dispersal and age of the hybrid zone. The hybrid zone is located at the isocline at 200 m altitude above sea level; this relationship between the races and altitude is suggested at both coarse and fine scales. This indicates adaptive differences between the races that may in turn have been promoted by the chromosome differences. Thus, the extreme chromosomal divergence between the Novosibirsk and Tomsk may be associated with genic differentiation, but it is still striking that, despite the large chromosomal differences, reproductive isolation between the Novosibirsk and Tomsk races has not occurred.     

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.     

Identification of a novel WART-like chromosome rearrangement in complex heterozygotes in an interracial hybrid zone of the common shrew Sorex araneus L

Karyotypes uncharacteristic of pure races or hybrids were identified in the interracial hybrid zones of the common shrew Sorex araneus L. that were recently discovered in the European part of Russia. This suggests the actual existence in natural populations of WART-like rearrangements (whole-arm reciprocal translocations) along with Robertsonian fusions of acrocentrics. Demonstration of new and still rare chromosome variants is the aim of this communication.     

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.     

Hierarchical analyses of genetic differentiation in a hybrid zone of Sorex araneus (Insectivora: Soricidae)

Microsatellites are used to unravel the fine-scale genetic structure of a hybrid zone between chromosome races Valais and Cordon of the common shrew ( Sorex araneus ) located in the French Alps. A total of 269 individuals collected between 1992 and 1995 was typed for seven microsatellite loci. A modified version of the classical multiple correspondence analysis is carried out. This analysis clearly shows the dichotomy between the two races. Several approaches are used to study genetic structuring. Gene flow is clearly reduced between these chromosome races and is estimated at one migrant every two generations using R -statistics and one migrant per generation using F -statistics. Hierarchical F - and R -statistics are compared and their efficiency to detect inter- and intraracial patterns of divergence is discussed. Within-race genetic structuring is significant, but remains weak. F _ST displays similar values on both sides of the hybrid zone, although no environmental barriers are found on the Cordon side, whereas the Valais side is divided by several mountain rivers. We introduce the exact G -test to microsatellite data which proved to be a powerful test to detect genetic differentiation within as well as among races. The genetic background of karyotypic hybrids was compared with the genetic background of pure parental forms using a CRT–MCA. Our results indicate that, without knowledge of the karyotypes, we would not have been able to distinguish these hybrids from karyotypically pure samples.     

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.     

Identification of a novel WART-like chromosome rearrangement in complex heterozygotes in an interracial hybrid zone of the common shrew Sorex araneus L.

Karyotypes uncharacteristic of pure races or hybrids were identified in the interracial hybrid zones of the common shrew Sorex araneus L. that were recently discovered in the European part of Russia. This suggests the actual existence in natural populations of WART-like rearrangements (whole-arm reciprocal trans-ocations) along with Robertsonian fusions of acrocentrics. Demonstration of new and still rare chromosome variants is the aim of this communication.     

Variability of a cytochrome b region in different chromosome races and populations of the common shrew Sorex araneus L., 1758

The nucleotide sequence (572 bp) of the cytochrome b gene of the common shrew Sorex araneus was analyzed. In total, 92 animals of five chromosome races from 12 localities were studied. The median haplotype network has a pronounced star-like structure. The central haplotype common for all samples, except for the southern island sample of the race Sok, accounts for about 36%. The main characteristics of molecular variation in our work correspond to those obtained in other studies. We revealed the lack of a correlation between the genetic and geographic distances and also population structurization of the species. On the basis of variation of the haplotypes in the samples, a scenario of colonization of the postglacial territories by females of one or several close matrilines with subsequent rapid subdivision of the population into independent populations is discussed.     

A hybrid zone with coincident clines for autosomal and sex-specific markers in the Sorex araneus group

In hybrid zones, endogenous counter-selection of hybrids is usually first expressed as reduced fertility or viability in hybrids of the heterogametic sex, a mechanism known as Haldane's rule. This phenomenon often leads to a differential of gene flow between sex-linked markers. Here, we address the possibility of a differential gene flow for Y chromosome, mtDNA and autosomal markers across the hybrid zone between the genetically and chromosomally well-differentiated species Sorex antinorii and Sorex araneus race Vaud. Intermarker comparison clearly revealed coincidental centre and very abrupt clines for all three types of markers. The overall level of genetic differentiation between the two species must be strong enough to hinder asymmetric introgression. Cyto-nuclear mismatches were also observed in the centre of hybrid zone. The significantly lower number of mismatches observed in males than in females possibly results from Y chromosome-mtDNA interactions. Results are compared with those previously reported in another hybrid zone between S. antinori and S. araneus race Cordon.     

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.     

Morphometric differentiation between the Manturovo and Serov chromosome races of the common shrewSorex araneus

Morphometric differentiation between the Manturovo and Serov chromosome races of the common shrewSorex araneus Linnaeus, 1758, in Northeastern European Russia was studied using 27 measurements of the skull in 953 specimens. Discriminant and cluster analyses showed that shrews belonging to different chromosome races were well differentiated. No regular dependence between morphological changes of the skull and longitude was observed and there was no association between geographic and morphological distance. Multiple regression analysis revealed that 24.7% of the total morphological variance could be explained by seven geoclimatic variables. We suggest that karyotypic divergence may play a significant role in differentiating skull morphology in the Manturovo and Serov races of the common shrew. We also suppose that selection may affect the skull morphology of different chromosome races in this species.     

Contact zones between chromosome races of the common shrew Sorex araneus L. (insectivora, mammalia) at the margins of the Veps stage of the Valdai ice sheet

Cytogenetic analysis of the common shrew populations in the Valdai Hills revealed the presence of four distinct chromosome races (Western Dvina, Seliger, St. Petersburg, and Moscow). The geographic distribution of these races relative to the borders of the Late Valdai glaciation area is described and discussed. Five contact zones between chromosomal races were found, located directly on the ice-marginal formations of the Veps stage. Two races, Seliger and St. Petersburg, are located exclusively in the former ice sheet area. Attention is drawn to the fact that many of the races located in the last glacial area from the Valdai Hills through Scandinavia are characterized by endemic chromosomes (gi, gp, mo, or, etc.). It is concluded that the situation in the former glaciation area could favor the preservation of tundra or forest vegetation refuges and populations of some small mammal species.     

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.     

Distribution of two karyotypic races of Sorex araneus (Insectivora, Soricidae) in Switzerland and the post-glacial recolonization of the Valais: First results

Two chromosome races of Sorex araneus were found in southern and western Switzerland. It is suggested they were issued by introgression of different sets of metacentrics, coming respectively from north-east and south, into a primitive “acrocentric” population which recolonized the country from the west after the last glaciation.