Gene-geographical variation and genetic differentiation in red-backed voles of the genus Clethrionomys (Rodentia,Cricetidae) in the Okhotskii region

Thirteen enzyme systems and three nonenzyme proteins were electrophoretically analyzed in red-backed voles of the genus Clethrionomys. In total, 25 loci were interpreted. Gene-geographic variation was studied and indices of genetic variability and differentiation were determined. By the distribution of electrophoretic variants of hemoglobin, C. rutilus was shown to be divided into two geographical groups (northern and southern). A low level of genetic differentiation was revealed in the island isolates of C. rutilus and C. rufocanus. Separation of C. rufocanus, C. rex, and C. sicotanensis into a superspecies complex was confirmed. A study of differential G- and C-banding on C. rutilus and C. rufocanus chromosomes did not reveal intraspecific variation of autosomes. In these species, karyotypes of voles from Kamchatka Peninsula were studied for the first time. They appeared to be morphologically similar to the karyotypes continental voles by both autosomes and sex chromosomes.     

Chromosomal variation in social voles: a Robertsonian fusion in Günther’s vole

The study reports on chromosomes in several populations of social voles from south-eastern Europe and the Middle East. The standard karyotypes of individuals of Microtus hartingi and Microtus guentheri originating from both south-eastern Europe and Asia Minor comprised 54 mostly acrocentric chromosomes. However, variation between populations was found in the amount and distribution of C-heterochromatin in certain autosomes and the sex chromosomes. Furthermore, a specific pattern of argyrophilic nucleolar organizer region distribution was recorded in different geographic populations. In a population from Asia Minor, a heterozygous centric fusion of two autosomes was found. The G-banded karyotypes of M. guentheri and Microtus socialis were compared, and tandem fusions of autosomes were suggested as possible mechanism of the divergence. The karyotypes of the nine currently recognized species of social voles are reviewed, and implications of chromosomal data for systematics are evaluated.     

The diet of <Emphasis Type="Italic">Mustela erminea</Emphasis> L. in the Magadan oblast

The study describes the diet of Mustela erminea L. in the Magadan oblast. The major component is Clethrionomys voles in the oblast, C. rufocanus in the Sea of Okhotsk area and C. rutilus in the basin of the Kolyma River. In winter the percentage of mammals declines by about 20%. The stoat compensates for the lack of voles by scavenging fish collected from the beds of ephemeral lakes and streams. Birds, insects, and plant foods are minor components of the stoat’s diet.     

Spatial distribution of red-backed voles of the genus <Emphasis Type="Italic">Myodes</Emphasis> in the Khanka Plain,Primorskii Krai

The authors present new data on spatial-temporal distribution of the northern red-backed vole (Myodes rutilus Pallas) and the large-toothed redback vole (M. rufocanus Sundervall) for the main habitat types in the Khanka Plain, Primorskii Krai. Data on seasonal and long-term abundance for 2003-2005 are reported. The study was stimulated by contradictory data on the distribution of the Myodes voles in the Khanka Plain.     

A Chromosomal Investigation of the West European Species of <Emphasis Type="BoldItalic">Corixa</Emphasis> Geoffroy (Heteroptera: Corixidae)

C-banded karyotypes are provided for the five west European species of Corixa Geoffroy. All have 2n = 22 autosomes + sex chromosomes which are XY (♂), XX (♀). Some specimens of C. punctata have a small B-chromosome, while some C. panzeri have a fairly large one. The karyotypes of C. punctata, C. dentipes, C. affinis and C. panzeri are all distinctive, but the karyotypes of C. punctata and C. iberica appear indistinguishable. The Scottish material of C. punctata and C. iberica used in this study shows morphological intergradation between the two, suggesting that they hybridise where their ranges overlap. Meiosis shows very fine even pachytene bands in C. punctata, and that the autosomes form true bivalents during diplotene in C. affinis and C. panzeri, while the sex chromosomes appear closely associated. At first metaphase the sex chromosomes may appear either closely associated or may have separated. There is no evidence of the presence of m-chromosomes.     

Chromosomal evolution in the genus Clethrionomys

The karyotypes of Clethrionomys rufocanus and C. glareolus are presented using G- and C-banding techniques. There is intraspecific polymorphism in Clethrionomys rufocanus for a pericentric inversion. The main difference between the karyotypes of the two species is considered to result from a reciprocal translocation.     

Karyology and karyotype analysis of diploid freshwater planarian populations of the Dugesia gonocephala group (Platyhelminthes,Tricladida) found in Sardinia

Karyology and karyotype analysis were carried out on freshwater planarian populations of the Dugesia gonocephala group. The strains studied were all diploid with chromosomic number 2n = 16; n = 8. They came from 12 sites mainly localized on the west of the island of Sardinia. Three karyotypes indicated with the letters A, B and C were found in which eight homomorphic pairs of chromosomes were easily identified. In karyotype A all chromosomes are metacentric. Ten populations of the twelve examined showed this karyotype which appears to be the most common. In karyotype B the seventh pair of chromosomes is submetacentric. This karyotype is quite common having been previously found in another eight Sardinian localities. Karyotype C differs from the others in having submetacentric third and seventh pairs of the chromosome complement. It was found in only one locality. The differences observed between these three karyotypes could be interpreted either as sign of differentiation at species level, or as an intraspecific variation due to chromosome mutations (pericentric inversions). This revised version was published online in July 2006 with corrections to the Cover Date.     

Chromosomes of Syrphidae

This report describes the karyotypes of fourteen species of the tribe Chrysotoxini (Diptera, Syrphidae) with the single genus Chrysotoxum. The species are listed in the Table on p. 240. All have 2n=10 chromosomes, usually consisting of short apparently telocentric sex chromosomes plus two subtelocentric and two submetacentric pairs of autosomes.     

Increased differentiation and reduced gene flow in sex chromosomes relative to autosomes between lineages of the brown creeper Certhia americana

The properties of sex chromosomes, including patterns of inheritance, reduced levels of recombination, and hemizygosity in one of the sexes may result in the faster fixation of new mutations via drift and natural selection. Due to these patterns and processes, the two rules of speciation to describe the genetics of postzygotic isolation, Haldane's rule and the large‐X effect, both explicitly include quicker evolution on sex chromosomes relative to autosomes. Because sex‐linked mutations may be the first to become fixed in the speciation process, and appear to be due to stronger genetic drift (in birds), we may identify pronounced genetic differentiation in sex chromosomes in taxa experiencing recent speciation and diverging mainly via genetic drift. Here, we use nine sex‐linked and 21 autosomal genetic markers to investigate differential divergence and introgression between marker types in Certhia americana. We identified increased levels of genetic differentiation and reduced levels of gene flow on sex chromosomes relative to autosomes. This pattern is similar to those observed in other recently‐divergent avian species, providing another case study of the earlier role of sex chromosomes in divergence, relative to autosomes. Additionally, we identify three markers that may be under selection between Certhia americana lineages.     

Densities of microtime rodents along a pollution gradient from a copper-nickel smelter

Population densities of microtine rodents were studied along an air pollution gradient in the Kola Peninsula, Russia, by long-term and short-term trapping. The study area is affected by high sulphur dioxide and heavy metal emissions from the Severonikel copper-nickel smelter in Monchegorsk. The density of Clethrionomys rufocanus, the most abundant vole species in the area, was lowest close to the smelter and increased with distance up to the farthest, less polluted trapping sites. Clethrionomys glareolus, C. rutilus and Lemmus lemmus were absent from the most severely damaged area and were also scarce at the moderately polluted area 28 km south of the smelter. Although the population of C. glareolus has previously been cyclic at the moderately polluted area, we were unable to demonstrate any regular cycle. The most likely explanation for the low number of microtine rodents in the damaged and moderately polluted areas in a decrease in the quantity of important food plants: epiphytic lichens for C. glareolus and possibly C. rutilus, mosses for L. lemmus and seed plants, especially Vaccinium myrtillus, for C. rufocanus. Close to the smelter, direct toxic effects of heavy metals may also reduce population densities. The results show that pollutants may change the relative proportions of microtine species.     

Transfer of mitochondrial DNA from the northern red-backed vole (Clethrionomys rutilus) to the bank vole (C. glareolus)

Summary Using a silver staining method to detect DNA fragments produced by restriction enzymes, it was possible to compare mitochondrial DNAs (mtDNAs) from 85 individuals of the bank vole (Clethrionomys glareolus) trapped at 25 localities in Fennoscandia. There are two distinctly different mtDNA lineages, one occurring in southern and central Fennoscandia and the other in the northern parts. A fragment comparison method shows about 12.7% nucleotide sequence divergence between these two lineages. This major difference between animals of the same species could theoretically be explained by intraspecific lineage survivorship independent of species hybridization, or by introduction of an atypical mtDNA via hybridization with a closely related species. Analysis of mtDNAs from the two otherClethrionomys species present in Fennoscandia (C. rutilus andC. rufocanus) shows that the mtDNA of northernC. glareolus is very similar to that ofC. rutilus and that the mtDNA lineages of these two species cluster together in a phenogram, with small genetic distances among them. By contrast, electrophoresis of proteins encoded by 17 nuclear loci reveals fixed allelic differences between these two species at 8 loci. Hence the presence of two distinctly different mtDNA lineages withinC. glareolus may be a consequence of a limited episode of hybridization betweenC. glareolus andC. rutilus, probably during the postglacial recolonization of Fennoscandia 8000–13,000 years ago.     

Sexual maturation and age-related dynamics of corticosterone in Clethrionomys rutilus and Cl. rufocanus voles (Rodentia, Cricetidae) under experimental conditions

Differences in the mechanisms regulating reproduction between the northern red-backed vole (Clethrionomys rutilus) and large-toothed red-backed vole (Cl. rufocanus) have been revealed under vivarium conditions. In Cl. rufocanus, the date of birth is the most important ??signal?? factor for sexual maturation of male young of the year, while the effect of population density is significant only for males of the spring generation. Males of the summer generation fail to mature in the same year and are completely excluded from reproduction, which may be accounted for by a rise in the level of corticosterone (measured in fecal samples). Such a mechanism for regulating the numbers of reproductive individuals is absent in Cl. rutilus voles. Throughout the breeding season, males of this species respond to high population density by a decrease in the rate of sexual maturation. No definite relationship between sexual maturation and the level of adrenocortical activity has been revealed in Cl. rutilus.     

Genetic variation and phylogeography of the bank vole (<Emphasis Type="Italic">Clethrionomys glareolus</Emphasis>, Arvicolinae,Rodentia) in Russia with special reference to the introgression of the mtDNA of a closely related species,red-backed vole (<Emphasis Type="Italic">Cl. rutilus</Emphasis>)

Totally, 294 bank voles (Clethrionomys glareolus) and 18 red-backed voles (Cl. rutilus) from 62 sites of European Russia were studied. Incomplete sequences (967 bp) of the mitochondrial cytochrome b gene were determined for 93 Cl. glareolus individuals from 56 sites and 18 Cl. rutilus individuals from the same habitats. Analysis of the cytochrome b gene variation has demonstrated that practically the entire European part of Russia, Ural, and a considerable part of Western Europe are inhabited by bank voles of the same phylogroup, displaying an extremely low genetic differentiation. Our data suggest that Cl. glareolus very rapidly colonized over the presently occupied territory in the post-Pleistocene period from no more than two (central European and western European) refugia from ancestral populations with a small effective size. PCR typing of the mitochondrial cytochrome b gene allowed us to assess the scale of mtDNA introgression from a closely related species, Cl. rutilus, and to outline the geographical zone of this introgression. Comparison with the red-backed vole haplotypes in the habitats shared by both species favors the hypothesis of an ancient hybridization event (mid-Holocene) and a subsequent introgression. These results suggest that the hybridization took place in the southern and middle Pre-Ural region.     

RAPD Fingerprinting of Common Bream Abramis brama L., Roach Rutilus rutilus L., and Their F1 Hybrids (A. brama × R. rutilus and R. rutilus × A. brama)

The polymerase chain reaction with arbitrary primers (RAPD–PCR) was used to study and to evaluate the genetic variation in the hybrid progeny of two Cyprinidae species, common bream Abramis brama L. and roach Rutilus rutilusL. Genetic polymorphism was studied in 20 fishes (young of the current year) obtained in four individual crosses: R. rutilus × R. rutilus (RR), A. brama × A. brama (AA), R. rutilus × A. brama (RA), and A. brama × R. rutilus (AR). Amplification spectra obtained with eight primers contained 288 fragments, 97.6% of which proved to be polymorphic. The proportion of polymorphic fragments was 75.0% in the RR progeny, 58.1% in the AA progeny, 84.9% in the AR progeny, and 77.8% in the RA progeny. Classification analysis in the space of principal components was performed with the first four components, which together accounted for 64% of the total variance of the character under study. The individual contributions of components I, II, III, and IV were 26.8, 16.8, 11.5, and 8.9%, respectively. Fishes of the two pure species and the hybrid progeny (direct and reverse hybrids together) were clearly differentiated in the space of principal components I and II. The best differentiation of the four samples (RR, AA, RA, and AR) was observed in the space of principal components II and IV. Possible causes of high genetic variation in interspecific hybrids are discussed.     

The role of chromosome rearrangements in the evolution of mole voles of the genus Ellobius (Rodentia,Mammalia)

Modern mole voles of the genus Ellobius are characterized by species-specific features of autosomes and sex chromosomes. Owing to the use of the Zoo-FISH method, the nomenclature of chromosomes was refined and nonhomologous Robertsonian translocations indistinguishable by G-staining were identified for Ellobius tancrei, which is a species with a wide chromosome variation of the Robertsonian type. The electron-microscopic analysis of synaptonemal complexes in F₁ hybrids of forms with 2n = 50 and 2n = 48 revealed the formation of a closed SC-pentavalent composed of three metacentrics with monobrachial homology and two acrocentrics. Segregation of chromosomes of such complex systems is impeded by disturbances in the nucleus architecture leading to the formation of unbalanced gametes and to a dramatic reduction in fertility of hybrids. Our data support the hypothesis that the formation of monobrachial homologous metacentric chromosomes can be considered as a way of chromosomal speciation.     

Can landscape properties predict occurrence of grey-sided voles?

There has been a long-term decline in spring and fall numbers of Clethrionomys rufocanus in boreal Sweden in 1971–2005. Previous studies on permanent sampling plots in the centre of 2.5 × 2.5 km landscapes suggested that habitat fragmentation (sensu destruction) could have contributed to the decline. Therefore, we tested these findings in a field study and compared trapping results on the central sampling plots of landscapes with a low degree of fragmentation (LDF) and of “hot spot” type with trapping results in managed forest landscapes with a high degree of fragmentation (HDF). We predicted that C. rufocanus would be more common on the LDF plots. We used our permanent plots supplemented with a new sample of plots, mainly of the rare LDF type, inside or just outside the long-term study area. Very few voles were trapped on both plot types, and no difference was found. However, a subsequent pilot study with trapping in a national park with large areas of pristine, unfragmented forest yielded more voles than in the managed, more fragmented, areas. Consequently, the initial field study data and some other recent data were also re-analysed from a “local patch quality” perspective. This alternative approach revealed the positive importance of large focal patches of forest >60 years old and their content of old-growth (pine) forest (>100 years). Interestingly, at the landscape level, the frequency distribution of patches of forest >60 years old, old-growth (>100 years), and especially of old-growth pine forest (>100 years), relative to the properties of plots with C. rufocanus, suggested that there are few forest patches left that are suitable for C. rufocanus. Our current results suggest that habitat fragmentation cannot be excluded as a contributing cause to the long-term decline of C. rufocanus in boreal Sweden.     

Genetic diversity of flat-headed vole (<Emphasis Type="Italic">Alticola strelzowi</Emphasis> (Kastschenko, 1899)) inferred from cytochrome <Emphasis Type="Italic">b</Emphasis> variation

Variation of the cytochrome b gene fragment was examined in 27 flat-headed voles Alticola strelzowi from different parts of the species range. A total of 15 haplotypes were described, while the species is characterized by low levels of genetic differentiation and polymorphism. The haplotypes form three haplogroups, one of which corresponded to the subspecies A. s. strelzowi, and the other two, to A. s. desertorum. Based on different indices, the level of genetic polymorphism in the later subspecies was considered to be higher than in the first one. Phylogeographic analysis suggested post-glacial dispersal of flat-headed voles from a single refugium located in Western Altai. Using different techniques, relatively recent colonization of the Central Altai territory was demonstrated (subspecies A. s. strelzowi), which determined low level of genetic variation in this territory.     

Variability of mitochondrial DNA cytochrome <Emphasis Type="Italic">b</Emphasis> gene in the red vole <Emphasis Type="Italic">Clethrionomys rutilus</Emphasis> Pallas, 1779, population in the flood-plain middle stream of the Kolyma River

The intrapopulation variability of a cytochrome b gene fragment and the corresponding amino acid sequence was studied in the red vole Clethrionomys rutilus Pallas, 1779 from the flood-plain of the Kolyma River. Wide polymorphic variability of these properties was observed. Differences in the cytochrome b gene sequence were determined between the red voles of the studied population and the species collected in neighborhoods of Novosibirsk and Omsk. The revealed results point to the urgency of the cytochrome b gene nucleotide sequence and the variants of the respective amino acid sequence as genetic markers of originality of different red vole populations.     

Giant sex chromosomes retained within the Portuguese lineage of the field vole (<Emphasis Type="Italic">Microtus agrestis</Emphasis>)

The field vole (Microtus agrestis) is characterised by extremely large blocks of heterochromatin on both the X and Y chromosome. Some other Microtus also have blocks of heterochromatin on their sex chromosomes but not as extensive and always of independent origin from the heterochromatic expansion found in M. agrestis. Coupled with evidence of geographic variation in large heterochromatic blocks within other species (e.g. in the western hedgehog Erinaceus europaeus), it might be expected that field voles would show substantial variation in size and disposition of the sex chromosome heterochromatin. In fact, only minor variation has been described up to now. Those studies conducted previously were largely on field voles from central and northern Europe. Here, we describe the karyotype of field voles from Portugal, of interest because recent molecular studies have shown field voles from western Iberia to be a separate evolutionary unit that might be considered a cryptic species, distinct from populations further to the east. The two Portuguese field voles (one female, one male) that we examined also had essentially the same karyotype as seen in other field voles, including the giant sex chromosomes, but with small differences in the structure of the Y chromosome from that described previously. The finding that field voles throughout Europe show relatively little variation in their giant sex chromosomes is consistent with molecular data which suggest a recent origin for this complex of species/near-species.     

Parental care in captive red-backed vole (Clethrionomys rutilus)

Parental behavior in ten pairs of male and female red-backed voles, Clethrionomus rutilus, was studied under laboratory conditions. Significant sex differences were found in total time spent in the nest, time spent by adults alone in the nest, grooming time, and nest-building activity; in half of the males under study, the level of paternal care was extremely low. These findings indicate different paternal contributions of female and male red-backed voles into the rearing of young, in agreement with the spatial-ethological structure of populations of this species. Parental behavior of C. rutilus is discussed in comparison with that of other species of the subfamily Microtinae.