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Studies on hybridization have proved critical for understanding key evolutionary processes such as speciation and adaptation. However, from the perspective of conservation, hybridization poses a concern, as it can threaten the integrity and fitness of many wild species, including canids. As a result of habitat fragmentation and extensive hunting pressure, gray wolf (Canis lupus) populations have declined dramatically in Europe and elsewhere during recent centuries. Small and fragmented populations have persisted, but often only in the presence of large numbers of dogs, which increase the potential for hybridization and introgression to deleteriously affect wolf populations. Here, we demonstrate hybridization between wolf and dog populations in Estonia and Latvia, and the role of both genders in the hybridization process, using combined analysis of maternal, paternal and biparental genetic markers. Eight animals exhibiting unusual external characteristics for wolves - six from Estonia and two from Latvia - proved to be wolf-dog hybrids. However, one of the hybridization events was extraordinary. Previous field observations and genetic studies have indicated that mating between wolves and dogs is sexually asymmetrical, occurring predominantly between female wolves and male dogs. While this was also the case among the Estonian hybrids, our data revealed the existence of dog mitochondrial genomes in the Latvian hybrids and, together with Y chromosome and autosomal microsatellite data, thus provided the first evidence from Europe of mating between male wolves and female dogs. We discuss patterns of sexual asymmetry in wolf-dog hybridization.

In the first continent-wide study of the golden jackal (Canis aureus), we characterised its population genetic structure and attempted to identify the origin of European populations. This provided a unique insight into genetic characteristics of a native carnivore population with rapid large-scale expansion. We analysed 15 microsatellite markers and a 406 base-pair fragment of the mitochondrial control region. Bayesian-based and principal components methods were applied to evaluate whether the geographical grouping of samples corresponded with genetic groups. Our analysis revealed low levels of genetic diversity, reflecting the unique history of the golden jackal among Europe's native carnivores. The results suggest ongoing gene flow between south-eastern Europe and the Caucasus, with both contributing to the Baltic population, which appeared only recently. The population from the Peloponnese Peninsula in southern Greece forms a common genetic cluster with samples from south-eastern Europe (ΔK approach in STRUCTURE, Principal Components Analysis [PCA]), although the results based on BAPS and the estimated likelihood in STRUCTURE indicate that Peloponnesian jackals may represent a distinct population. Moreover, analyses of population structure also suggest either genetic distinctiveness of the island population from Samos near the coast of Asia Minor (BAPS, most STRUCTURE, PCA), or possibly its connection with the Caucasus population (one analysis in STRUCTURE). We speculate from our results that ancient Mediterranean jackal populations have persisted to the present day, and have merged with jackals colonising from Asia. These data also suggest that new populations of the golden jackal may be founded by long-distance dispersal, and thus should not be treated as an invasive alien species, i.e. an organism that is \"non-native to an ecosystem, and which may cause economic or environmental harm or adversely affect human health\". These insights into the genetic structure and ancestry of Baltic jackals have important implications for management and conservation of jackals in Europe. The golden jackal is listed as an Annex V species in the EU Habitats Directive and as such, considering also the results presented here, should be legally protected in all EU member states.

Gray wolf Canis lupus predation on domestic dogs Canis familiaris is a considerable wolf–human conflict issue in several regions of Europe and North America but it has not been well documented in the scientific literature. Livestock depredations by wolves may be related to the abundance of wild prey. Regardless of the presumed motivations of wolves for attacking dogs (likely due to interference competition and predation), the abundance of wild prey populations may also influence the risk of wolf attacks on dogs. We examined whether the annual number of fatal attacks by wolves on dogs was related to the abundance of primary prey, including wild boar Sus scrofa and roe deer Capreolus capreolus in Estonia, as well as the abundance of moose Alces alces in Finland. Statistical models resulted in significant negative relationships, thus providing evidence that the risk of attacks in both house yards (Estonia) and hunting situations (Finland) was highest when the density of wild prey was low. Wild ungulates cause damage to agriculture and forestry, but they seem to mitigate conflicts between wolves and humans; therefore, it is necessary to develop a holistic, multispecies management approach in which the importance of wild ungulates for large carnivore conservation is addressed.

ContextAdjustments in habitat use by large carnivores can be a key factor facilitating their coexistence with people in shared landscapes. Landscape composition might be a key factor determining how large carnivores can adapt to occurring alongside humans, yet broad-scale analyses investigating adjustments of habitat use across large gradients of human pressure and landscape composition are lacking.ObjectivesHere, we investigate adjustments in habitat use by Eurasian lynx (Lynx lynx) in response to varying availability of refuge habitats (i.e., forests and rugged terrain) and human landscape modification.MethodsUsing a large tracking dataset including 434 individuals from seven populations, we assess functional responses in lynx habitat use across two spatial scales, testing for variation by sex, daytime, and season.ResultsWe found that lynx use refuge habitats more intensively with increasing landscape modification across spatial scales, selecting forests most strongly in otherwise open landscapes and rugged terrain in mountainous regions. Moreover, higher forest availability enabled lynx to place their home ranges in more human-modified landscapes. Human pressure and refuge habitat availability also shaped temporal patterns of lynx habitat use, with lynx increasing refuge habitat use and reducing their use of human-modified areas during periods of high exposure (daytime) or high vulnerability (postnatal period) to human pressure.ConclusionsOur findings suggest a remarkable adaptive capacity of lynx towards human pressure and underline the importance of refuge habitats across scales for enabling coexistence between large carnivores and people. More broadly, we highlight that the composition of landscapes determines how large carnivores can adapt to human pressure and thus play an important role shaping large carnivore habitat use and distributions.

Predation on livestock is a cause of serious and long-lasting conflict between farmers and wildlife, promoting negative public attitudes and endangering conservation of large carnivores. However, while large carnivores, especially the grey wolf (Canis lupus), are often blamed for killing sheep and other farm animals, free-ranging dogs may also act as predators. To develop appropriate measures for livestock protection, reliable methods for identifying predator species are critical. Identification of predators from visual examination of livestock wounds can be ambiguous and genetic analysis is strongly preferable for accurate species determination. To estimate the proportion of wolves and dogs implicated in sheep predation, we developed a sensitive genetic assay to distinguish between wolves and domestic dogs. A total of 183 predator saliva samples collected from killed sheep in Estonia were analysed. The assay identified the predator species in 143 cases (78%). Sheep were most often killed by wolves (81%); however, predation by dogs was substantial (15%). We compared the molecular results with field observations conducted by local environmental officials and recorded some disagreement, with the latter underestimating the role of dogs. As predator saliva samples collected from prey are often of poor quality, we suggest using mitochondrial DNA as a primary tool to maximise the number of successfully analysed samples. We also suggest adopting forensic DNA analysis more widely in livestock predation assessments as a legislative measure since misidentification that is biased against wolves can be counterproductive for conservation by enhancing conflict with society and leading to increased culling and poaching.

Spatial genetics is a relatively new field in wildlife and conservation biology that is becoming an essential tool for unravelling the complexities of animal population processes, and for designing effective strategies for conservation and management. Conceptual and methodological developments in this field are therefore critical. Here we present two novel methodological approaches that further the analytical possibilities of STRUCTURE and DResD. Using these approaches we analyse structure and migrations in a grey wolf (Canislupus) population in north-eastern Europe. We genotyped 16 microsatellite loci in 166 individuals sampled from the wolf population in Estonia and Latvia that has been under strong and continuous hunting pressure for decades. Our analysis demonstrated that this relatively small wolf population is represented by four genetic groups. We also used a novel methodological approach that uses linear interpolation to statistically test the spatial separation of genetic groups. The new method, which is capable of using program STRUCTURE output, can be applied widely in population genetics to reveal both core areas and areas of low significance for genetic groups. We also used a recently developed spatially explicit individual-based method DResD, and applied it for the first time to microsatellite data, revealing a migration corridor and barriers, and several contact zones.

Aim: Using sequences of complete mitochondrial genomes, our aims were: (1) to investigate the matrilineal phylogeographical structure, migration patterns and lineage coalescence times in a large, continuous population of brown bears (Ursus arctos); and (2) to develop a novel spatial genetic method to identify migration corridors and barriers. Location: North-western Eurasia: from eastern European Russia to the Baltic Sea. Methods: We sequenced the complete mitochondrial genomes of 95 brown bears. The phylogeographical resolution of complete genomes was compared to that derived from subsets of the genome, including the most commonly used shorter sequences. We conducted network and Bayesian phylogeographical analyses and developed a novel, spatially explicit, individual-based approach (called DResD) for identifying migration corridors and barriers. Results: Analysis of mitogenome sequences revealed five haplogroups, specific to particular geographical areas, exhibiting far greater resolving power than shorter sequences. Estimated coalescence times for the haplogroups ranged from 7.7 to 15.2 ka, suggesting that their divergence took place after the last glaciation. We found several migration trends, including a large westward migration from eastern European Russia towards Finland. We also found evidence of a potential barrier and a migration corridor in the south-west of the study area. Main conclusions: The use of complete mitochondrial genomes from a brown bear population in north-western Eurasia allowed us to identify phylogeographical structure, signatures of demographic history and spatial processes that had not previously been detected using shorter sequences. These findings have implications for studies on other species and populations, especially those exhibiting low mtDNA diversity. The relatively recent divergence estimates for haplogroups highlight the significance not only of the last glaciation but also of climatic fluctuations during the post-glacial period for the divergence of mammal populations in Europe. Our spatial genetic method represents a new tool for the analysis of genetic data in a geographical context and is applicable to any data that yield genetic distance matrices, including microsatellites, amplified fragment length polymorphisms (AFLPs) and single-nucleotide polymorphisms (SNPs).

Due to their high mobility, large terrestrial predators are potentially capable of maintaining high connectivity, and therefore low genetic differentiation among populations. However, previous molecular studies have provided contradictory findings in relation to this. To elucidate patterns of genetic structure in large carnivores, we studied the genetic variability of the Eurasian lynx, Lynx lynx throughout north-eastern Europe using microsatellite, mitochondrial DNA control region and Y chromosome-linked markers. Using SAMOVA we found analogous patterns of genetic structure based on both mtDNA and microsatellites, which coincided with a relatively little evidence for male-biased dispersal. No polymorphism for the cytochrome b and ATP6 mtDNA genes and Y chromosome-linked markers were found. Lynx inhabiting a large area encompassing Finland, the Baltic countries and western Russia formed a single genetic unit, while some marginal populations were clearly divergent from others. The existence of a migration corridor was suggested to correspond with distribution of continuous forest cover. The lowest variability (in both markers) was found in lynx from Norway and Białowieża Primeval Forest (BPF), which coincided with a recent demographic bottleneck (Norway) or high habitat fragmentation (BPF). The Carpathian population, being monomorphic for the control region, showed relatively high microsatellite diversity, suggesting the effect of a past bottleneck (e.g. during Last Glacial Maximum) on its present genetic composition. Genetic structuring for the mtDNA control region was best explained by latitude and snow cover depth. Microsatellite structuring correlated with the lynx's main prey, especially the proportion of red deer (Cervus elaphus) in its diet. Eurasian lynx are capable of maintaining panmictic populations across eastern Europe unless they are severely limited by habitat continuity or a reduction in numbers. Different correlations of mtDNA and microsatellite population divergence patterns with climatic and ecological factors may suggest separate selective pressures acting on males and females in this solitary carnivore.

After a long and deliberate persecution, the grey wolf (Canis lupus) is slowly recolonizing its former areas in Europe, and the genetic consequences of this process are of particular interest. Wolves, though present in mainland Estonia for a long time, have only recently started to recolonize the country's two largest islands, Saaremaa and Hiiumaa. The main objective of this study was to analyse wolf population structure and processes in Estonia, with particular attention to the recolonization of islands. Fifteen microsatellite loci were genotyped for 185 individuals across Estonia. As a methodological novelty, all putative wolf-dog hybrids were identified and removed (n = 17) from the dataset beforehand to avoid interference of dog alleles in wolf population analysis. After the preliminary filtering, our final dataset comprised of 168 \"pure\" wolves. We recommend using hybrid-removal step as a standard precautionary procedure not only for wolf population studies, but also for other taxa prone to hybridization. STRUCTURE indicated four genetic groups in Estonia. Spatially explicit DResD analysis identified two areas, one of them on Saaremaa island and the other in southwestern Estonia, where neighbouring individuals were genetically more similar than expected from an isolation-by-distance null model. Three blending areas and two contrasting transition zones were identified in central Estonia, where the sampled individuals exhibited strong local differentiation over relatively short distance. Wolves on the largest Estonian islands are part of human-wildlife conflict due to livestock depredation. Negative public attitude, especially on Saaremaa where sheep herding is widespread, poses a significant threat for island wolves. To maintain the long-term viability of the wolf population on Estonian islands, not only wolf hunting quota should be targeted with extreme care, but effective measures should be applied to avoid inbreeding and minimize conflicts with local communities and stakeholders.

We studied the relationship between the variability and contemporary distribution of pelage phenotypes in one of most widely distributed felid species and an array of environmental and demographic conditions. We collected 672 photographic georeferenced records of the Eurasian lynx throughout Eurasia. We assigned each lynx coat to one of five phenotypes. Then we fitted the coat patterns to different environmental and anthropogenic variables, as well as the effective geographic distances from inferred glacial refugia. A majority of lynx were either of the large spotted (41.5%) or unspotted (uniform, 36.2%) phenotype. The remaining patterns (rosettes, small spots and pseudo-rosettes) were represented in 11.0%, 7.4%, and 3.9% of samples, respectively. Although various environmental variables greatly affected lynx distribution and habitat suitability, it was the effect of least-cost distances from locations of the inferred refugia during the Last Glacial Maximum that explained the distribution of lynx coat patterns the best. Whereas the occurrence of lynx phenotypes with large spots was explained by the proximity to refugia located in the Caucasus/Middle East, the uniform phenotype was associated with refugia in the Far East and Central Asia. Despite the widely accepted hypothesis of adaptive functionality of coat patterns in mammals and exceptionally high phenotypic polymorphism in Eurasian lynx, we did not find well-defined signs of habitat matching in the coat pattern of this species. Instead, we showed how the global patterns of morphological variability in this large mammal and its environmental adaptations may have been shaped by past climatic change.