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.     

Genetic differences and similarities in the introduced population of European beaver (Castor fiber L., 1785) in the Kirov and Novosibirsk regions 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.     

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.     

Assessment of genetic management at three specific-pathogen-free rhesus macaque (Macaca mulatta) colonies

Genetic management is required to maintain genetic diversity by minimizing inbreeding and genetic subdivision in colonies of animals bred for biomedical research. Polymorphic short tandem repeat (STR) loci are useful for genetic management because they facilitate parentage assignments, genetic characterization of individuals, and estimates of baseline population genetic parameters. Using highly informative STR loci, we estimated gene diversity and F-statistics to determine the level of genetic heterogeneity and genetic structure of three specific-pathogen-free (SPF) rhesus macaque (Macaca mulatta) colonies. Effective population sizes, variance in male reproductive success, and rate of decrease in genetic variability also were estimated for two of the three colonies. We documented the overall success of genetic management in maintaining genetic diversity in captive colonies. We report that even genetically managed SPF colonies, despite maintaining high and stable levels of gene diversity (over 0.75), are prone to genetic subdivision due to different management strategies, founder effects, genetic isolation, and drift. These processes are accelerated by the high variances in male reproductive success and low adult sex ratios that are typical of captive rhesus macaque breeding groups, both of which reduce the effective population sizes of these groups.     

Assessing genetic variation and population structure of invasive North American beaver (<Emphasis Type="Italic">Castor Canadensis</Emphasis> Kuhl, 1820) in Tierra Del Fuego (Argentina)

The North American beaver (Castor Canadensis) was introduced into Isla Grande de Tierra del Fuego, Argentina in 1946 as a potential source of wild fur. The species showed high growth potential, reaching close to 100,000 individuals from an original founding stock of 25 females and 25 males. Beavers adapted rapidly to their new environment and became invasive, providing an excellent model of successful adaptation of introduced populations to a new habitat. In this study, we used polymorphic mitochondrial (mt) DNA to evaluate genetic variation in the introduced beaver population from Tierra del Fuego. Nucleotide variation in partial sequences of Cytochrome b (500 bp) and 12S rRNA (421 bp) genes and the main non-coding D-loop region (521 bp) were analyzed. Our study allowed to identify 10 different mtDNA lineages in the invasive population, none of them shared among the source populations. The pattern observed is a consequence of cessation of gene flow following expansion of the founding beaver population since the time of its introduction. This approach contributes to the understanding of effects of genetic changes on survival ability and reproductive success of invasive species. It also has important management implications to invasive species.     

Colonization, riparian habitat selection and home range size in a reintroduced population of European beavers in the Loire

1. Colonization in a reintroduced population of European beavers in the Loire Valley was studied between 1974 and 1999. It followed a discontinuous remoteness model and a scattered distribution, beavers occupying only 25% of the river system over the 2800 km explored.
2. After 5 years, the colonization rate reached 104.2% year^–1 of new sites occupied (SD 75% year^–1), before dropping over the next 20 years. Nevertheless, the number of new colonies per km (0.125) remained stable throughout the years.
3.  Populus nigra , Salix alba and Fraxinus angustifolia were the dominant woody species in beaver sites, often associated with some herbaceous species.
4. The length of willow grove dominated by S. alba and P. nigra ( x ) was the best predictor of beaver home range ( y ), fitting the equation y =−0.742 x  + 5.9. Long-term maintenance of the population requires a minimum of 1.79-km of willows per colony.
5. In cutting tree trunks, beavers stimulate shoot development from the remaining stumps. They rejuvenate riparian forests, increase the number of tree stems and help stabilize the banks. Their effect on woody plant morphogenesis may have consequences for the helophyte communities used as food or habitat by other aquatic species.     

Protein variation in wild Atlantic salmon, with particular reference to southern Ireland

The extent of genetic variation in wild Atlantic salmon parr, Sulmo salur L., from river systems in Ireland, Iceland and eastern Canada, was investigated using starch gel electrophoresis. Within Ireland, seven polymorphic enzyme loci ( sAAT-4 *, GPI-1 *, IDDH-1 *, IDDH-2 *, IDHP-3 *, MDH-3 * and mMEP-2 *) were screened in nine different rivers and nine tributaries from the River Blackwater. Significant heterogeneity in gene frequencies occurred between riverine samples and between samples from tributaries of the River Blackwater. Variation between tributaries was as great as between rivers elsewhere in the country. Levels of population differentiation were comparable to those found in other regions throughout the range of the species, and temporal stability in gene frequencies was apparent when the results were compared with previously published data. Screening of riverine samples from Iceland and eastern Canada (Newfoundland and New Brunswick) allowed the Irish results to be considered in a broader context. Irish salmon cluster in the western European group, to which may be added Icelandic populations. Salmon from eastern Canada show a high level of genetic distinctiveness from the European group.     

Population trends and reproductive success of the European shag Phalacrocorax aristotelis on the Iberian Peninsula following the Prestige oil spill

In 2003, immediately following the Prestige oil spill in Galicia, Spain, we studied the population trends and reproductive performance of European shags (Phalacrocorax aristotelis) at oiled and unoiled colonies. This bird is an important member of the nearshore marine community, breeding in the area affected by the Prestige oil spill. The European shag feeds around the breeding colonies throughout the year, making it a useful indicator of environmental change. Before the oil spill, population trends were similar between oiled and unoiled colonies. Nevertheless, colonies located within the path of the oil suffered greater declines (ca. 10%) compared with pre-spill trends and with population trends at unoiled colonies. In 2003, the breeding success was 50% lower in oiled colonies compared with unoiled colonies. The data available from pre-spill years suggest that the annual reproductive success did not differ among colonies before the impact. European shags breeding at colonies affected by oil showed a negative initial impact from the Prestige oil spill. The reduction in reproductive success at oiled colonies may be due to sub-lethal effects of oil exposure or low food availability after the oil spill.     

Blood-protein allele frequencies and phylogenetic relationships in Macaca: A review

Allele-frequency data have been assembled for 35 blood-protein loci in 17 of 19 recognized species of Macaca based on 29 published electrophoretic studies; studies of inbred captive colonies have been excluded. Data for 22 polymorphic loci are tabulated in detail for 43 geographic populations of these species. Calculated F_ST values provide a measure of intergroup genetic differentiation at various hierarchical levels—troop, locality, province, country or island, species, species group; polymorphism indices measure genetic variation. The greatest intraspecific genetic differentiation occurs at the level of island populations within species. The pattern of genetic variation among island populations appears to be relictual, suggesting that the reduced genetic variability of island populations of macaques is a result of postisolation genetic drift rather than founder effect. Interspecific relationships were investigated by means of a jackknifed Fitch-Margoliash algorithm, using Papio as outgroup. Phylogenetic inferences based on morphology and zoogeography. The reduced genetic variability that frequently characterizes insular macaque populations complicates phylogenetic interpretation of blood-protein evidence.     

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.     

Fish in the tributaries of the Novosibirsk reservoir

Data on the species composition of the fish population inhabiting the tributaries of the Novosibirsk reservoir are given in the article. The structure of fish communities and possible environmental factors determining it have been revealed. The relation between the heterogeneity of fish communities and the river depth and length, relief, current velocity, floodplain, and estuarial areas, presence of sand-pebble shallow waters and channel order, has been studied.     

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.     

Contrasting age-specific recruitment and survival at different spatial scales: a case study with the European storm petrel

Evolutionary studies on optimal decisions or conservation guidelines are often derived by generalising patterns from a single population, while inter‐population variability in life‐history traits is seldom considered. We investigated here how survival and recruitment probabilities changed with age at different geographical scales using the encounter histories of 5523 European storm petrels from three Mediterranean colonies, and also how our estimates of these parameters might be expected to affect population growth rates using population matrix models. We recorded similar patterns among colonies, but also important biological differences. Local survival, recruitment and breeding success increased with age at all colonies; the most distant of three colonies (Marettimo Is.) showed the largest differences. Strikingly, differences in recruitment were also found between two adjacent colonies (two caves from Benidorm Is.). Birds marked as adults from Marettimo and Benidorm colonies had a different survival, whereas we found no differences within Benidorm. Differences in survival were no longer apparent between the two islands at the end of the study following a reduction in predation by specialist gulls at Benidorm. Since birds marked as fledglings mostly recruited near the end of the study, their overall survival was high and in turn similar among colonies. Results from our population matrix models suggested that different age‐dependent patterns of demographic parameters can lead to similar population growth rates. Variability appeared to be greater for recruitment and the most sensitive parameter was adult survival. Thus conservation actions targeting this vulnerable species should focus on factors influencing adult survival. Differences in survival and recruitment among colonies could reflect the spatial heterogeneity in mortality due to predation and colony‐specific density dependent processes. Results highlight the importance of taking into account the potential spatio‐temporal heterogeneity among populations in vital rates, even in those traits that life‐history theory considers less important in driving population dynamics.     

Biology of the people of Sikkim, India. 2. Colour blindness, ear lobe attachment, mid-phalangeal hair and behavioural traits

14 population groups of Sikkim (India)--Lepchas (2), Bhutias (2), Sherpas, Tamangs, Gurungs, Mangars, Rais, Limboos/Subbas, Pradhans (Newars), Brahmans, Chhetris, Scheduled Castes--have been studied in regard of the intra- and intergroup variability of colour blindness, ear lobe attachment, mid-phalangeal hair and behavioural traits (tongue folding, hand clapsing, arm folding, leg folding, handedness). Some of these variables show a considerable distribution heterogeneity, which is discussed considering history and marriage patterns of these populations. As most of them are highly endogamous one can assume that this heterogeneity is caused by locally acting factors such as drift and/or founder effects, which could be preserved due to as good as lacking gene flow among the populations under study. Beyond that the Sikkim data are compared briefly with those reported for other Indian and Asiatic populations.     

Individual heterogeneity in reproductive rates and cost of reproduction in a long‐lived vertebrate

Individual variation in reproductive success is a key feature of evolution, but also has important implications for predicting population responses to variable environments. Although such individual variation in reproductive outcomes has been reported in numerous studies, most analyses to date have not considered whether these realized differences were due to latent individual heterogeneity in reproduction or merely random chance causing different outcomes among like individuals. Furthermore, latent heterogeneity in fitness components might be expressed differently in contrasted environmental conditions, an issue that has only rarely been investigated. Here, we assessed (i) the potential existence of latent individual heterogeneity and (ii) the nature of its expression (fixed vs. variable) in a population of female Weddell seals (Leptonychotes weddellii), using a hierarchical modeling approach on a 30‐year mark–recapture data set consisting of 954 individual encounter histories. We found strong support for the existence of latent individual heterogeneity in the population, with “robust” individuals expected to produce twice as many pups as “frail” individuals. Moreover, the expression of individual heterogeneity appeared consistent, with only mild evidence that it might be amplified when environmental conditions are severe. Finally, the explicit modeling of individual heterogeneity allowed us to detect a substantial cost of reproduction that was not evidenced when the heterogeneity was ignored.     

Limited genetic differentiation of Mycetomoellerius mikromelanos in Parque National Soberanía,Panama: Implications for queen dispersal

The coevolutionary relationship between fungus-growing ants (Formicidae: Attini: Attina) and their symbionts has been well studied in the Panamanian rain forests. To further understand the ecological context of these evolutionary relationships, we have examined the population-genetic structure of the fungus-growing ant species Mycetomoellerius mikromelanos Cardenas, Schultz, Adams 2021 in the Panama Canal Zone. We specifically investigated the presence of population structure, the significance of geographic features (i.e., creeks) limiting gene flow, and relatedness between ant colonies. To accomplish this, we genotyped 85 ant colonies from nine creeks across an approximately 30 km transect in Parque National Soberanía, Panama, using double-digest restriction-site-associated DNA sequencing. We did not find distinct population structure using two genetic clustering methods; however, we did detect an effect of isolation by distance. Furthermore, related colonies were frequently detected on the same creek or neighboring creeks, and some at further geographic distances. Collectively, these findings demonstrate that new colonies tend to establish on natal creeks and occasionally on distant creeks following long-distance dispersal events. We discuss how population-genetic patterns reveal the natural history of M. mikromelanos in Parque National Soberanía and how these results fit into the context of fungus-growing ant mutualisms. Abstract in Spanish is available with online material.     

Dispersal and individual quality in a long lived species

The idea of differences in individual quality has been put forward in numerous long-term studies in long-lived species to explain differences in lifetime production among individuals. Despite the important role of individual heterogeneity in vital rates in demography, population dynamics and life history theory, the idea of "individual quality" is elusive. It is sometimes assumed to be a static or dynamic individual characteristic. When considered as a dynamic trait, it is sometimes assumed to vary deterministically or stochastically, or to be confounded with the characteristics of the habitat. We addressed heterogeneity in reproductive performance among individuals established in higher-quality habitat in a long-lived seabird species. We used approaches to statistical inference based on individual random effects permitting quantification of heterogeneity in populations and assessment of individual variation from the population mean. We found evidence of heterogeneity in breeding probability, not success probability. We assessed the influence of dispersal on individual reproductive potential. Dispersal is likely to be destabilizing in species with high site and mate fidelity. We detected heterogeneity after dispersal, not before. Individuals may perform well regardless of quality before destabilization, including those that recruited in higher-quality habitat by chance, but only higher-quality individuals may be able to overcome the consequences of dispersal. Importantly, results differed when accounting for individual heterogeneity (an increase in mean breeding probability when individuals dispersed), or not (a decrease in mean breeding probability). In the latter case, the decrease in mean breeding probability may result from a substantial decrease in breeding probability in a few individuals and a slight increase in others. In other words, the pattern observed at the population mean level may not reflect what happens in the majority of individuals.     

Do introduced North American beavers Castor canadensis engineer differently in southern South America? An overview with implications for restoration

• 1 Twenty‐five pairs of North American beavers Castor canadensis Kuhl were introduced to Tierra del Fuego Island in 1946. The population has expanded across the archipelago, arriving at the Chilean mainland by the mid‐1990s. Densities range principally between 0.5–2.05 colonies/km. They have an impact on between 30–50% of stream length and occupy 2–15% of landscape area with impoundments and meadows. Beaver impacts constitute the largest landscape‐level alteration in subantarctic forests since the last ice age.
• 2 The colonization pattern, colony densities and impacted area indicate that habitat in the austral archipelago is optimal for beaver invasion, due to low predator pressure and suitable food resources. Nothofagus pumilio forests are particularly appropriate habitat, but a more recent invasion is occurring in adjacent steppe ecosystems. Nonetheless, Nothofagus reproductive strategies are not well adapted to sustain high beaver population levels.
• 3 Our assessment shows that at the patch‐scale in stream and riparian ecosystems, the direction and magnitude of exotic beaver impacts are predictable from expectations derived from North American studies, relating ecosystem engineering with underlying ecological mechanisms such as the relationships of habitat heterogeneity and productivity on species richness and ecosystem function.
• 4 Based on data from the species' native and exotic range, our ability to predict the effects of beavers is based on: (i) understanding the ecological relationships of its engineering effects on habitat, trophic dynamics and disturbance regimes, and (ii) having an adequate comprehension of the landscape context and natural history of the ecosystem being engineered.
• 5 We conclude that beaver eradication strategies and subsequent ecosystem restoration efforts, currently being considered in southern Chile and Argentina, should focus on the ecology of native ecosystems rather than the biology of this invasive species per se. Furthermore, given the nature of the subantarctic landscape, streams will probably respond to restoration efforts more quickly than riparian ecosystems.

     

High regional differentiation in a North American crab species throughout its native range and invaded European waters: a phylogeographic analysis

Rhithropanopeus harrisii (Gould 1841) has a native distribution from New Brunswick (Canada) to Veracruz (Mexico) and is considered an invasive species in northwestern North American (Oregon and California), South American (Brazil) and European estuaries and rivers. In Europe, it was observed for the first time in 1874, in The Netherlands. We sequenced and analyzed part of the cytochrome oxidase subunit I gene (mitochondrial DNA) of eight populations, three from the east coast of the United States of America (USA) and five from Europe, in order to assess their genetic diversity and to determine a potential founder population. European populations are characterized by a lower number of haplotypes than the whole native region of the eastern USA, suggesting that genetic bottlenecks occurred during the European colonisation. Along the North American East Coast, there is evidence of clearcut genetic heterogeneity, New Jersey being the most similar population in its genetic structure to the postulated Europe-founding population. Also the different European populations are heterogeneous and there is a tendency of higher genetic diversity in the populations founded earlier. R. harrisii is still in the process of expansion in Europe and may have been introduced once or repeatedly by different invasion mechanisms. The pronounced lack of gene flow among populations is of great ecological significance, since it may facilitate rapid adaptation and specialization to local conditions within single estuarine systems.     

The genetic dynamics of the rapid and recent colonization of Denmark by <Emphasis Type="Italic">Arion lusitanicus</Emphasis> (Mollusca,Pulmonata, Arionidae)

We describe the genetic dynamics of the recent establishment of the ‘Iberian slug’, Arion lusitanicus J. Mabille 1868, in Denmark and compare its population structure to two other members of the ‘large Arion complex’, Arion ater ater, native to Denmark, and Arion ater rufus, introduced into Denmark in the early 1900s. Assaying allozyme polymorphism at seven enzyme loci, we found that: (1) None of the three taxa reproduce primarily by self-fertilization. Differences among loci and colonies in the pattern of deviation from Hardy–Weinberg equilibrium are most consistent with isolate mixing and perhaps with low amounts of selfing. (2) For both A. lusitanicus and A. a. rufus, gene diversity is lower in Danish colonies than in southern German colonies, implying population bottlenecks in the establishment of Danish colonies. (3) Significant linkage disequilibrium values usually involve the same three loci, viz. PGI, MDH-1 and MDH-2, suggesting physical linkage among these loci. (4) For both A. a. rufus and A. lusitanicus, the overall gene frequencies from Denmark and southern Germany are homogeneous, while variation among colonies within these regions ranges from around 15 to 28% for the three taxa. This indicates strong, local population genetic subdivision but with little restriction to gene flow from possible source areas. The heterogeneity in measures of diversity and differentiation indicates that population structure for all three taxa is dominated by ongoing founder effects, local extinction/colonisation dynamics, and genetic drift processes.