Genetic Divergence and Similarity of Introduced Populations of European Beaver (Castor fiber L., 1758) from Kirov and Novosibirsk Oblasts of Russia

Potential factors were analyzed that affected the formation of gene pools of two introduced beaver populations founded in the 1940s–1950s by releasing beavers from Voronezh oblast and Belarus into rivers of the Kirov and Novosibirsk oblasts. The populations from these two regions were shown to differ in four allozyme loci (Ahd-2, Gpi, Es-5, and Dia-2). Within Kirov oblast, the samples from three tributaries of the Vyatka River differed in three other loci (Ck-1, Gp-9, and Trf). It is suggested that these features of the allele distribution at these loci are explained by the history of the introduced populations, hybridization between beavers from Voronezh oblast and Belarus, and founder effect during the subsequent artificial and spontaneous migration. Generally, genetic differences between the populations in all studied loci were very small: D = 0.02 between populations from different regions and D = 0.01 between populations from the tributaries of third-order rivers of the Kirov oblast. At the same time, disequilibrium was detected for ten polymorphic loci in the combined sample from the populations of Kirov oblast, which indicates the existence of interpopulation heterogeneity at the level of local populations of fourth- and fifth-order rivers. The unexpected finding of a genetic similarity between one studied individual of Tuvinian beaver subspecies (Castor fiber tuvinicus) and C. f. orientoeuropeas was recorded.     

Population-genetic structure of beaver (Castor fiber L., 1758) communities and estimation of effective reproductive size Ne of an elementary population

The absence of panmixia at all hierarchical levels of the European beaver communities down to individual families implies a complex organization of the population-genetic structures of the species, in particular, a large intergroup component of gene diversity in the populations. Testing this assumption by analysis of 39 allozyme loci in the communities of reintroduced beaver from the Vyatka river basin (Kirov oblast) has shown that only the beaver colonies exhibit high intergroup gene diversity (Gst = 0.32) whereas this parameter is much lower when estimated among beaver groups from individual Vyatka River tributaries and among localities of one of the tributaries (0.07 and 0.11, respectively). The data suggesting genetic heterogeneity among individual settles within colonies have been obtained. The factors affecting the structure of the beaver communities of the lower hierarchical ranks are considered: the common origin, founder effect, selection, gene drift, assortative mating, and social and behavior features of the species. The conclusion is drawn that the founder effect could be the primary factor of population differentiation only at the time of their formation. The heterogeneity among colonies and among settles is maintained largely by isolation of colonies from one another. The strong interspecific competition for food resources, which is behaviorally implemented in the species at the level of minimal structural units (individual settles) creates a profound and unique population-genetic subdivision of the species. These results substantiate the suggestion that an elementary population (micropopulation) of European beaver is a colony, i.e., a set of related settles of different types. Based on ecological and genetic parameters, the effective reproductive size Ne of the minimum beaver population was estimated to be equal to three animals. This extremely low value of effective reproductive population size largely explains the high tolerance of European beaver to inbreeding and striking viability of the species, which from the early 19th century has been for more than hundred years on the brink of survival in the condition which would made any other mammalian species vanish from the Earth.     

The gene pool of the Belgorod oblast population: Changes in population genetic relationships during the past 50 years

The model of the Belgorod oblast population has been used to demonstrate different effects of administrative reforms on microevolution in human populations. For the populations that formerly belonged to Kursk oblast, changes in the regional administrative structure (after which some of them remained in Kursk oblast and others were included into Belgorod oblast) have lead to an increase in the genetic distances between them. However, other populations (formerly belonging to Voronezh oblast) have become genetically closer to one another, although these populations now belong to different administrative regions (Belgorod and Voronezh oblasts).     

Recolonizing lost habitat—how European beavers (<Emphasis Type="Italic">Castor fiber</Emphasis>) return to south-western Germany

For the last decades, the European beaver (Castor fiber) has been recolonizing its original habitats. Reintroductions of beavers from different relict populations into southern Germany have resulted in several admixed populations, which are spreading out along various river systems. The eastern part of the German state of Baden-Württemberg is a melting pot of colonization waves originating from various introduced populations. The aim of this study was to exemplify origins and dispersal behaviour of beavers in this region using genetic fingerprint methods. Sequence analysis of hypervariable region 1 (HV1) of the mitochondrial control region and fragment length analyses at 11 microsatellite loci resulted in genetic profiles for 84 samples. The study region is being populated from three different local origins of beavers: the north of the Main-Tauber district, the Neckar River near the city of Mosbach and the Danube tributaries in the east. Main-Tauber samples were most diverse, including microsatellite alleles and HV1 haplotypes specific to C. f. albicus (from the German Elbe relict population). In view of the geographical proximity of this region to a release site of C. f. albicus in the Spessart area, this finding strongly suggests gene flow between beaver populations of different provenience. Two remaining local origins at the Neckar and Danube tributaries are closely related to each other, thus possibly descending from the same original (admixed) population. This study is intended to serve as a starting point for follow-up fine-scale research into dispersal behaviour of European beavers currently recolonizing their original habitats.     

Population-Genetic Structure of Beaver (Castor fiber L., 1758) Communities and Estimation of Effective Reproductive Size N e of an Elementary Population

The absence of panmixia at all hierarchical levels of the European beaver communities down to individual families implies a complex organization of the population-genetic structures of the species, in particular, a large intergroup component of gene diversity in the populations. Testing this assumption by analysis of 39 allozyme loci in the communities of reintroduced beaver from the Vyatka river basin (Kirov oblast) has shown that only the beaver colonies exhibit high intergroup gene diversity (G _st = 0.32) whereas this parameter is much lower when estimated among beaver groups from individual Vyatka River tributaries and among localities of one of the tributaries (0.07 and 0.11, respectively). The data suggesting genetic heterogeneity among individual settles within colonies have been obtained. The factors affecting the structure of the beaver communities of the lower hierarchical ranks are considered: the common origin, founder effect, selection, gene drift, assortative mating, and social and behavior features of the species. The conclusion is drawn that the founder effect could be the primary factor of population differentiation only at the time of their formation. The heterogeneity among colonies and among settles is maintained largely by isolation of colonies from one another. The strong interspecific competition for food resources, which is behaviorally implemented in the species at the level of minimal structural units (individual settles) creates a profound and unique population-genetic subdivision of the species. These results substantiate the suggestion that an elementary population (micropopulation) of European beaver is a colony, i.e., a set of related settles of different types. Based on ecological and genetic parameters, the effective reproductive size N _e of the minimum beaver population was estimated to be equal to three animals. This extremely low value of effective reproductive population size largely explains the high tolerance of European beaver to inbreeding and striking viability of the species, which from the early 19th century has been for more than hundred years on the brink of survival in the condition which would made any other mammalian species vanish from the Earth.     

Zooplankton in the zones of confluence of unregulated rivers

The hydrophysical characteristics and indices of zooplankton are given for the confluence zones of small and medium tributaries and channels of the Don and Khoper rivers within the territory of Voronezh oblast. Three types of confluence areas of tributaries and recipient rivers are described. In these areas the water masses mix and habitats differing in physical parameters of water and structures of aquatic animal communities are formed.     

Dynamics of food resources for beavers in settlements colonized and abandoned several times

The state of food resources for Eurasian beavers (Castor fiber Linnaeus, 1758) after several cycles of habitat colonization and abandonment has been analyzed in three beaver settlements on small rivers of Novgorod oblast. It has been shown that tree stands surrounding the settlements differ in composition and degree of damage but are similar in that they contain no forage for beavers: aspen is absent, and birch trees are large and require great efforts to cut them down and utilize. The undergrowth and understory also contain no aspen, while abundant willows and birch are represented by young, thin-stemmed plants unevenly distributed over the area. Tree species prevailing in abandoned settlements are not favored by beavers, implying that the animals must have expended extra energy for foraging. Supposedly, populations of beavers on small rivers in the southern taiga subzone, in habitats modified by their activities, will maintain their abundance far below the maximum possible level, depending on the rate of forage plant regrowth against the background of deteriorating habitat quality.     

Admixture of two phylogeographic lineages of the Eurasian beaver in Poland

The Eurasian beaver (Castor fiber) represents an uncommon example of an endangered species in which the restoration programs proved a spectacular success and led to enormous spatial and demographic expansion. Documented reintroduction of beavers in Poland has been conducted using animals of the eastern European origin, most likely derived from the eastern mtDNA lineage. However demographic and spatial expansion of beavers from Germany, which represent the western lineage, may have led to admixture of these two genetically distinct entities in Poland. We detected significant genetic differentiation between the populations from W and NE Poland both in mitochondrial DNA control region and microsatellites, but also substantial admixture including apparent first-generation migrants between regions. Our results indicate that beavers from the western mtDNA lineage have contributed considerably to the Polish population, particularly in W Poland. As there have been no adequately documented translocations of beavers from the western European populations to Poland, the observed situation appears to result from natural migration or range expansion from the west. In contrast to previous findings we detected a substantial diversity in mtDNA control region, which indicates that either the variation in relict populations has been underestimated, or that additional relict beaver populations survived at the end of the 19th century in Poland and Germany as indicated by considerable similarity of ancient and extant mtDNA haplotypes. The implications of our findings for beaver conservation and management are discussed.     

Ancient mitochondrial DNA and the genetic history of Eurasian beaver (Castor fiber) in Europe

After centuries of human hunting, the Eurasian beaver Castor fiber had disappeared from most of its original range by the end of the 19th century. The surviving relict populations are characterized by both low genetic diversity and strong phylogeographical structure. However, it remains unclear whether these attributes are the result of a human‐induced, late Holocene bottleneck or already existed prior to this reduction in range. To investigate genetic diversity in Eurasian beaver populations during the Holocene, we obtained mitochondrial control region DNA sequences from 48 ancient beaver samples and added 152 modern sequences from GenBank. Phylogeographical analyses of the data indicate a differentiation of European beaver populations into three mitochondrial clades. The two main clades occur in western and eastern Europe, respectively, with an early Holocene contact zone in eastern Europe near a present‐day contact zone. A divergent and previously unknown clade of beavers from the Danube Basin survived until at least 6000 years ago, but went extinct during the transition to modern times. Finally, we identify a recent decline in effective population size of Eurasian beavers, with a stronger bottleneck signal in the western than in the eastern clade. Our results suggest that the low genetic diversity and the strong phylogeographical structure in recent beavers are artefacts of human hunting‐associated population reductions. While beaver populations have been growing rapidly since the late 19th century, genetic diversity within modern beaver populations remains considerably reduced compared to what was present prior to the period of human hunting and habitat reduction.     

The Genetic Legacy of Multiple Beaver Reintroductions in Central Europe

The comeback of the Eurasian beaver (Castor fiber) throughout western and central Europe is considered a major conservation success. Traditionally, several subspecies are recognised by morphology and mitochondrial haplotype, each linked to a relict population. During various reintroduction programs in the 20th century, beavers from multiple source localities were released and now form viable populations. These programs differed in their reintroduction strategies, i.e., using pure subspecies vs. mixed source populations. This inhomogeneity in management actions generated ongoing debates regarding the origin of present beaver populations and appropriate management plans for the future. By sequencing of the mitochondrial control region and microsatellite genotyping of 235 beaver individuals from five selected regions in Germany, Switzerland, Luxembourg, and Belgium we show that beavers from at least four source origins currently form admixed, genetically diverse populations that spread across the study region. While regional occurrences of invasive North American beavers (n = 20) were found, all but one C. fiber bore the mitochondrial haplotype of the autochthonous western Evolutionary Significant Unit (ESU). Considering this, as well as the viability of admixed populations and the fact that the fusion of different lineages is already progressing in all studied regions, we argue that admixture between different beaver source populations should be generally accepted.     

The influence of Eurasian beaver Castor fiber activity on fish assemblages in small steppe rivers in Russia

In recent decades, the Eurasian beaver (Castor fiber) has once again become the keystone species in small river ecosystems in Russia. In many places, beaver activity has resulted in a significant change in lotic habitats, affecting the diversity, density, and biomass of aquatic organisms, including fish. While many studies have considered the ecosystem impacts of beavers, relatively few have focused on understanding the influence of beaver activity on steppe rivers. We conducted the first quantitative study of beaver impacts on fish assemblages in beaver-influenced and beaver-free sites on two small steppe rivers in the Don River basin in Russia. The presence of beavers altered the habitats in small steppe rivers and affected the diversity, density, and biomass of fish. A comparison of the number of species, density, and biomass of fish in six types of river habitats showed that these parameters were lower in beaver ponds than at riverine sites without beaver activity. Three fish species primarily preferred a single habitat type. Barbatula barbatula was found in riffles, Misgurnus fossilis in old beaver ponds, and Eudontomyzon mariae in abandoned beaver ponds. Beavers impacted fish distribution and density by changing dissolved oxygen, pH, and water current velocity. Overall, our results showed that the presence of beavers led to a temporary homogenization of fish habitats at a local scale in the valleys of small steppe rivers because beavers occupied these rivers only for a short period. However, habitat heterogeneity may increase if the beaver population stays stable or expands in the future.

     

Polymorphism of Y-chromosomal microsatellites in Russian populations from the northern and southern Russia as exemplified by the populations of Kursk and Arkhangel'sk Oblast

Allelic polymorphisms at five Y-chromosomal microsatellite loci (DYS19, DYS390, DYS391, DYS392, and DYS393) were typed in 87 individuals from male population samples from two geographically isolated regions (Arkhangelsk oblast and Kursk oblast) of the European part of Russia. The populations examined demonstrated substantial differences in the distribution of the DYS392 (P = 0.005) and DYS393 (P = 0.003) alleles. Estimates of genetic relationships between these populations and some other European populations (including Eastern-Slavic) showed that irrespectively of the measure of genetic distance chosen, Arkhangelsk population was closer to the populations belonging to the Finno-Ugric linguistic group (Saami and Estonians) and to the Estonian geographical neighbors, Latvians, while Kursk population was the member of a cluster formed by Eastern-Slavic populations (Russians of Novgorod oblast, Ukrainians, and Belarussians). Phylogenetic analysis of the most frequent haplotypes indicated that these differences between Kursk and Arkhangelsk populations were associated with high prevalence in the latter of major haplotypes characteristic primarily of the Finno-Ugric populations.     

Developing forensic reference database by 18 autosomal STR for DNA identification in Republic of Belarus

For the Republic of Belarus, development of a forensic reference database on the basis of 18 autosomal microsatellites (STR) using a population dataset (N = 1040), “familial” genotypic dataset (N = 2550) obtained from expertise performance of paternity testing, and a dataset of genotypes from a criminal registration database (N = 8756) is described. Population samples studied consist of 80% ethnic Belarusians and 20% individuals of other nationality or of mixed origin (by questionnaire data). Genotypes of 12346 inhabitants of the Republic of Belarus from 118 regional samples studied by 18 autosomal microsatellites are included in the sample: 16 tetranucleotide STR (D2S1338, TPOX, D3S1358, CSF1PO, D5S818, D8S1179, D7S820, THO1, vWA, D13S317, D16S539, D18S51, D19S433, D21S11, F13B, and FGA) and two pentanucleotide STR (Penta D and Penta E). The samples studied are in Hardy–Weinberg equilibrium according to distribution of genotypes by 18 STR. Significant differences were not detected between discrete populations or between samples from various historical ethnographic regions of the Republic of Belarus (Western and Eastern Polesie, Podneprovye, Ponemanye, Poozerye, and Center), which indicates the absence of prominent genetic differentiation. Statistically significant differences between the studied genotypic datasets also were not detected, which made it possible to combine the datasets and consider the total sample as a unified forensic reference database for 18 “criminalistic” STR loci. Differences between reference database of the Republic of Belarus and Russians and Ukrainians by the distribution of the range of autosomal STR also were not detected, corresponding to a close genetic relationship of the three Eastern Slavic nations mediated by common origin and intense mutual migrations. Significant differences by separate STR loci between the reference database of Republic of Belarus and populations of Southern and Western Slavs were observed. The necessity of using original reference database for support of forensic expertise practice in the Republic of Belarus was demonstrated.     

Haematology and Serum Biochemistry Parameters and Variations in the Eurasian Beaver (Castor fiber)

Haematology parameters (N = 24) and serum biochemistry parameters (N = 35) were determined for wild Eurasian beavers (Castor fiber), between 6 months – 12 years old. Of the population tested in this study, N = 18 Eurasian beavers were from Norway and N = 17 originating from Bavaria but now living extensively in a reserve in England. All blood samples were collected from beavers via the ventral tail vein. All beavers were chemically restrained using inhalant isoflurane in 100% oxygen prior to blood sampling. Results were determined for haematological and serum biochemical parameters for the species and were compared between the two different populations with differences in means estimated and significant differences being noted. Standard blood parameters for the Eurasian beaver were determined and their ranges characterised using percentiles. Whilst the majority of blood parameters between the two populations showed no significant variation, haemoglobin, packed cell volume, mean cell haemoglobin and white blood cell counts showed significantly greater values (p<0.01) in the Bavarian origin population than the Norwegian; neutrophil counts, alpha 2 globulins, cholesterol, sodium: potassium ratios and phosphorus levels showed significantly (p<0.05) greater values in Bavarian versus Norwegian; and potassium, bile acids, gamma globulins, urea, creatinine and total calcium values levels showed significantly (p<0.05) greater values in Norwegian versus Bavarian relict populations. No significant differences were noted between male and female beavers or between sexually immature (<3 years old) and sexually mature (≥3 years old) beavers in the animals sampled. With Eurasian beaver reintroduction encouraged by legislation throughout Europe, knowledge of baseline blood values for the species and any variations therein is essential when assessing their health and welfare and the success or failure of any reintroduction program. This is the first study to produce base-line blood values and their variations for the Eurasian beaver.     

Sequence diversity of the MHC DRB gene in the Eurasian beaver (Castor fiber)

Major histocompatibility complex (MHC) genes, coding molecules which play an important role in immune response, are the most polymorphic genes known in vertebrates. However, MHC polymorphism in some species is limited. MHC monomorphism at several MHC class I and II loci was previously reported for two neighbouring northern European populations of the Eurasian beaver (Castor fiber) and reduced selection for polymorphism has been hypothesized. Here, we analysed a partial sequence of the second exon of the MHC II DRB locus from seven relict European and Asian beaver populations. We detected 10 unique alleles among 76 beavers analysed. Only a western Siberian population was polymorphic, with four alleles detected in 10 individuals. Each of the remaining populations was fixed for a different allele. Sequences showed considerable divergence, suggesting the long persistence of allelic lineages. A significant excess of nonsynonymous substitutions was detected at the antigen binding sites, indicating that sequence evolution of beaver DRB was driven by positive selection. Current MHC monomorphism in the majority of populations may be the result of the superimposition of the recent bottleneck on pre-existing genetic structure resulting from population subdivision and differential pathogen pressure.     

Selective effect of the duration of the critical temperature period on some allozyme loci of Atlantic salmon Salmo salar L. (Salmonidae)

Genetic structure of juvenile fish from the populations of the Atlantic salmon Salmo salar inhabiting the rivers of Murmansk oblast, Arkhangelsk oblast, and Karelia, as well as of juveniles from hatcheries was examined at five allozymic loci: aspartate aminotransferase (AAT-4*), isocitrate dehydrogenase (IDHP-3*), iditol dehydrogenase (IDDH-2*), esterase D (ESTD*), and malic enzyme (MEP-2*). High genetic differentiation of both natural and "hatchery" juvenile fish was revealed. It was demonstrated that the gene pool of juveniles at three of the five loci tested was to a considerable degree formed by natural selection. In this case, the role of limiting factor was played by the duration of critical temperature regime in the rivers. The differentiation factors for juveniles from nature were not established, however, their clusterization pattern pointed to a possible role of natural selection in this process.     

Recovery and status of native and introduced beavers Castor fiber and Castor canadensis in France and neighbouring countries

• 1 The Eurasian beaver Castor fiber suffered a drastic reduction in both geographical range and population size, due to human persecution, until the end of the 19th century. After the adoption of protection measures, natural expansion and reintroductions led to the recovery of this species over much of its European range.
• 2 We review historical events that led to the recovery of beavers in France, and summarize the status of beavers in various river systems. Beaver establishment in France is a story of overall success: several major river systems are presently occupied, so that the species is no longer at risk in France.
• 3 However, beaver recolonization took place in parallel with increasing human impacts on the environment. In addition to natural limiting factors, anthropogenic factors impeded beaver settlement in many areas. Today, beavers often occupy suboptimal habitats and, as a consequence, come into conflict with human activities. Effective solutions for preventing beaver damage include the restoration of riparian habitats to discourage crop damage and the provision of physical barriers to protect crops.
• 4 Beaver populations reintroduced into France all originate from the relict Rhône population. However, in recent years, beavers from populations in neighbouring countries have been expanding into north‐eastern France. Therefore, our review of beaver origin and distribution in these countries may contribute to the development of appropriate national management strategies and towards important decisions, e.g. the decision to try to keep Rhône beavers genetically isolated, or to allow populations to mix.
• 5 The recently discovered presence of North American beavers Castor canadensis in three countries surrounding France has raised an important issue. This species may out‐compete C. fiber in places where the species come into contact. A programme based on field‐trapping sessions and genetic analyses has recently been initiated in some western countries in order to eradicate this non‐native species.

     

Hereditary deafness in Kirov oblast: a genetic epidemiological study

The results of a genetic epidemiological study of hereditary deafness (HD) in ten raions (districts) of Kirov oblast (administrative region), Russia, are presented. A total of 122 075 people have been examined. Segregation analysis of all families with diagnosed HD has demonstrated a good fit to either the autosomal dominant (AD) or autosomal recessive (AR) mode of inheritance. The total prevalence rates of AD and AR HDs, as well as the specific prevalence rates of nonsyndromic and syndromic forms of HD, have been calculated for the population often raions. The HD prevalence rate in Kirov oblast has been found to be 1 : 1043 people (1 : 1453 and 1 : 3699 for the nonsyndromic and syndromic forms, respectively). This value has been found to vary in different raions, which is explained by differences in the genetic subdivision levels of the populations studied; the correlation coefficient between the HD load and random inbreeding (F(ST)) in district populations is r = 0.81 +/- 0.22. The diversity of syndromic hearing disorders is described.     

Variability of allozyme and cpSSR markers in the populations of Siberian spruce

The variability of 21 allozyme and three microsatellite loci of chloroplast DNA (cpDNA) was studied in the populations of Siberian spruce (Picea obovata Ledeb.) from Irkutsk oblast, Magadan oblast, Buryatia, and Mongolia. It was demonstrated that the highest level of genetic diversity among the examined populations at both allozyme and microsatellite loci was observed in the Tulyushka population from Irkutsk oblast. The lowest level of genetic diversity was observed in marginal isolated populations of Bogd Uul and Magadan. In the relict spruce population from Olkhon Island, differing from the other populations in the lowest allelic diversity of both types of markers, no expected decline of expected heterozygosity and haplotype diversity was observed. In this population, the variability parameters mentioned were close to the population mean. The obtained intrapopulation and intraspecific variability parameters of allozyme and microsatellite loci of chloroplast DNA and the data on the population differentiation at these loci indicate that the given markers can be used for the analysis of the population structure of Siberian spruce.     

The gene pool of Belgorod oblast population: study of biochemical gene markers in populations of Ukraine and Belarus and the position of the Belgorod population in the Eastern Slavic gene pool system

The characteristics of the gene pools of indigenous populations of Ukraine and Belarus have been studied using 28 alleles of 10 loci of biochemical gene markers (HP, GC, TF, PI, C'3, ACP1, GLO1, PGM1, ESD, and 6-PGD). The gene pools of the Russian and Ukrainian indigenous populations of Belgorod oblast (Russia) and the indigenous populations of Ukraine and Belarus have been compared. Cluster analysis, multidimensional scaling, and factor analysis of the obtained data have been used to determine the position of the Belgorod population gene pool in the Eastern Slavic gene pool system.