Genetic identification of immigrants to the Scandinavian wolf population

Continued gene flow is fundamental to the survival of small, isolated populations. However, geography and human intervention can often act contrary to this requirement. The Scandinavian wolf population is threatened with a loss of genetic variation yet limited in the accessibility to new immigrants by the geographical distance of this peninsular population from its nearest neighbouring population and by human reluctance to allow wolves in the northern reindeer-breeding areas. In this study, we describe the identification of immigrants into this population using autosomal microsatellites, and maternally inherited mtDNA. Samples of 14 wolves collected in the “dispersal corridor” in northern Sweden in 2002–2005 were compared with 185 resident Scandinavian wolves and 79 wolves from the neighbouring Finnish population. We identified four immigrant wolves, suggesting some westward migration, although only one of these is likely to still survive. The integration of such immigrants into the breeding population is necessary to assure the long-term survival of this isolated and inbred population and highlights the importance of genetics techniques to the management of threatened populations.     

Genetic nature of eastern wolves: Past, present and future

Eastern North American wolves have long been recognized as morphologically distinct from both coyotes and gray wolves. This has led to questions regarding their origins and taxonomic status. Eastern wolves are mainly viewed as: (1) a smaller subspecies of gray wolf (Canis lupus lycaon), potentially the result of historical hybridization between gray wolves (C. lupus) and red wolves (C. rufus), (2) a hybrid, the result of gray wolf (C. lupus) and coyote (C. latrans) interbreeding, or (3) a distinct species, C. lycaon, closely related to the red wolf (C. rufus). Although debate persists, recent molecular studies suggest that the eastern wolf is not a gray wolf subspecies, nor the result of gray wolf/coyote hybridization. Eastern wolves were more likely a distinct species, C. lycaon, prior to the eastward spread of coyotes in the late 1800s. However, contemporary interbreeding exits between C. lycaon to both C. lupus and C. latrans over much of its present range complicating its present taxonomic characterization. While hybridization may be reducing the taxonomic distinctiveness of C. lycaon, it should not necessarily be viewed as negative influence. Hybridization may be enhancing the adaptive potential of eastern wolves, allowing them to more effectively exploit available resources in rapidly changing environments.     

Northwest passages: conservation genetics of Arctic Island wolves

Wolves in the Canadian Arctic Archipelago face several challenges to persistence: a harsh habitat, an unstable prey base, and potentially significant anthropogenic influences. These external factors, if combined with genetic constraints common to island populations, could be particularly difficult to withstand. To determine the genetic status of Arctic Island wolves, we used 14 microsatellite loci to estimate population variation and the extent of inter-island and island-mainland gene flow. All island populations were significantly less variable than mainland wolves; although inbreeding is currently insignificant, the two least variable populations, Banks and the High Arctic (Ellesmere and Devon Islands), showed genetic signatures of recent population declines. Recovery after a bottleneck appears to result, in large part, via recolonization from other islands. These extinction-recolonization dynamics, and the degree of similarity among island wolves revealed by Bayesian clustering, suggest that Arctic Island wolves function as a metapopulation. Persistence of the metapopulation may be supported by periodic migration from mainland populations, occurring primarily through two corridors: Baffin Island in the Eastern Arctic, and Victoria Island in the Western Arctic. This gene flow could be compromised or eliminated by loss—due to climatic warming or increased human activity—of sea ice in the Northwest Passage. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Encounter frequencies between GPS-collared wolves (Canis lupus) and moose (Alces alces) in a Scandinavian wolf territory

Over 6,000 GPS fixes from two wolves (Canis lupus) and 30,000 GPS fixes from five moose (Alces alces) in a wolf territory in southern Scandinavia were used to assess the static and dynamic interactions between predator and prey individuals. Our results showed that wolves were closer to some of the moose when inside their home ranges than expected if they had moved independently of each other, and we also found a higher number of close encounters (<500 m) than expected. This suggests that the wolves were actively seeking the individual moose within their territory. Furthermore, the wolves showed a preference for moving on gravel forest roads, which may be used as convenient travel routes when patrolling the territory and seeking areas where the moose are. However, due to the particularly large size of the wolf territory combined with relatively high moose densities, the wolves generally spent a very small proportion of their time inside the home range of each individual moose, and the frequency of encounters between the wolves and any particular moose was very low. We suggest that the high moose:wolf ratio in this large Scandinavian wolf territory compared to that typically occurring in North America, results in a relatively low encounter frequency and a low predation risk for individual moose, as the predation pressure is spread over a high number of prey individuals.     

Seasonal fluctuations of the wolf diet in the Hustai National Park (Mongolia)

Seasonal fluctuations in the wolf diet in the Hustai National Park (Mongolia) were determined by analysing wolf scats that were collected from 1994–1997. Except for July, August and September, at least 50% of the frequency of occurrence of prey items in wolf scats consisted of livestock. The frequency of occurrence of wild ungulates ranged from 8–43%, and wild ungulates were most often consumed in July, August and October, but seldom in June. Our study suggested that most seasonal changes in the wolf diet were effected by the life history of red deer. The occurrence of wild ungulates in the wolf diet coincided with the availability and vulnerability of calves. To reduce livestock losses and poaching of wolves we recommend more wolf research, protection and monitoring of both wild prey populations and wolves, and the reintroduction of livestock guarding dogs and year-round guarding of domestic horses.

Zusammenfassung

Saisonale Fluktuationen von Wolfsnahrung im Hustai Nationalpark (Mongolei)Anhand von Kotproben von Wölfen, die von 1994 bis 1997 gesammelt wurden, konnten saisonale Fluktuationen der Wolfsdiät im Nationalpark Hustai (Mongolei) festgestellt werden. Mit Ausnahme der Monate Juli, August und September bestand mindestens 50% der im Wolfskot vorgefundenen Beutereste aus Nutztieren. Wildungulat–traten in 8 bis 43% der Proben auf und wurden am häufigsten in den Monaten Juli, August und Oktober genutzt, jedoch kaum im Juni. Diese Untersuchung deutet darauf hin, dass saisonale Veränderungen der Wolfsdiät wahrscheinlich von der Biologie der Rothirsche beeinflusst werden, da die Abnahme von Wildungulaten in der Wolfnahrumg in den Zeitraum der Kälbergeburten fiel, während die Zunahme von Wildungulaten mit jener Periode koinzidierte, in der sich neu geborene Kälber den Herden anschlossen und die Brunftzeit stattfand. Um sowohl den Verlust von Nutzvieh als auch das Wildern auf Wölfe zu vermindern, empfehlen wir die Wolfsforschung zu verstärken, das Monitoring und den Schutz von Wolfs- und Wildbeutepopulationen zu verbessern, die Wiedereinführung von Hütehunden sowie die ganzjährige Überwachung von Nutzpferden.     

Sex identification of wolf (Canis lupus) using non-invasive samples

We have developed new specific primers for sex determination from forensic samples of wolves (Canis lupus), such as hair, saliva, faecal, tooth and urine samples. In order to improve molecular sexing, we performed a multiplex semi-nested polymerase chain reaction (PCR) and several replicated amplifications per sample to avoid errors in low quantity DNA samples, such as allelic dropout and false alleles. The sex of individuals is automatically determined by capillary electrophoresis with a fluorescently labelled internal sex-specific primer from each pair. Our method yielded sex identification on 100% of invasive samples and 93% of forensic samples, being one of the highest success rates obtained from wild animals.     

Evidence of wolf dispersal in anthropogenic habitats of the Polish Carpathian Mountains

In the course of their maturation, most young wolves leave their natal pack and disperse in search for mating partners, improved food availability and new territories. We investigated whether this dispersal is affected by anthropogenic infrastructure in a 5,000 km2 area of the eastern region of the Polish Carpathian Mountains occupied by wolves. A radio-collared male wolf covered 230 km while dispersing through forested hills and densely populated valleys. To test if such dispersal is common in the population we analysed by microsatellite genotyping 39 samples taken from live-trapped wolves or wolves found dead in the study area. Although the obtained genotypes were assigned to different clusters in Bayesian tests, we could not ascribe this structure to landscape features, but rather to shared ancestry of wolf individuals found in distant locations. Moreover, we could not detect a spatial genetic structure in the wolf population, indicating a random occurrence of genotypes within the study area. Observation of the dispersing wolf and the absence of spatial genetic structure imply that wolves are still able to roam the entire area despite high densities of roads and a dense human population. Thus, we concluded that the existing anthropogenic infrastructure does not restrict wolf dispersal in the area and the studied wolves represent a coherent part of the Polish Carpathian wolf population.     

Genomic sweep and potential genetic rescue during limiting environmental conditions in an isolated wolf population

Genetic rescue, in which the introduction of one or more unrelated individuals into an inbred population results in the reduction of detrimental genetic effects and an increase in one or more vital rates, is a potentially important management tool for mitigating adverse effects of inbreeding. We used molecular techniques to document the consequences of a male wolf (Canis lupus) that immigrated, on its own, across Lake Superior ice to the small, inbred wolf population in Isle Royale National Park. The immigrant's fitness so exceeded that of native wolves that within 2.5 generations, he was related to every individual in the population and his ancestry constituted 56 per cent of the population, resulting in a selective sweep of the total genome. In other words, all the male ancestry (50% of the total ancestry) descended from this immigrant, plus 6 per cent owing to the success of some of his inbred offspring. The immigration event occurred in an environment where space was limiting (i.e. packs occupied all available territories) and during a time when environmental conditions had deteriorated (i.e. wolves' prey declined). These conditions probably explain why the immigration event did not obviously improve the population's demography (e.g. increased population numbers or growth rate). Our results show that the beneficial effects of gene flow may be substantial and quickly manifest, short-lived under some circumstances, and how the demographic benefits of genetic rescue might be masked by environmental conditions.     

Molecular identification of Taenia spp. in wolves (Canis lupus), brown bears (Ursus arctos) and cervids from North Europe and Alaska

Taenia tapeworms of Finnish and Swedish wolves (Canis lupus) and Finnish brown bears (Ursus arctos), and muscle cysticerci of Svalbard reindeer (Rangifer tarandus platyrhynchus), Alaskan Grant's caribou (Rangifer tarandus granti) and Alaskan moose (Alces americanus) were identified on the basis of the nucleotide sequence of a 396 bp region of the mitochondrial cytochrome c oxidase subunit 1 gene. Two species were found from wolves: Taenia hydatigena and Taenia krabbei. The cysticerci of reindeer, caribou and one moose also represented T. krabbei. Most of the cysticercal specimens from Alaskan moose, however, belonged to an unknown T. krabbei-like species, which had been reported previously from Eurasian elks (Alces alces) from Finland. Strobilate stages from two bears belonged to this species as well. The present results suggest that this novel Taenia sp. has a Holarctic distribution and uses Alces spp. as intermediate and ursids as final hosts.     

Detecting rare introgression of domestic dog genes into wild wolf (Canis lupus) populations by Bayesian admixture analyses of microsatellite variation

Hybridization with free-ranging dogs isthought to threat the genetic integrity ofwolves in Europe, although available mtDNA dataevidenced only sporadic cases of crossbreeding.Here we report results of population assignmentand genetic admixture analyses in 107wild-living Italian wolves, 95 dogs including30 different breeds and feral dogs, andcaptive-reared wolves of unknown or hybridorigins, which were genotyped at 18microsatellites. Two Italian wolves showedunusually dark coats (``black wolves'), and oneshowed a spur in both hindlegs (``fifth fingerwolf'), suggesting hybridization. Italianwolves showed significant deficit ofheterozygotes, positive F_IS values anddeviations from Hardy-Weinberg equilibrium.Genetic variability was significantlypartitioned between groups, suggesting thatwolves and dogs represent distinct gene pools.Multivariate ordination of individual genotypesand clustering of inter-individual geneticdistances split wolves and dogs into twodifferent clusters congruent with the priorphenotypic classification, but hybrids andwolves of unknown origin were not identifiedfrom genetic information alone. By contrast, aBayesian admixture analysis assigned all theItalian wolves and dogs to two differentclusters, independent of any prior phenotypicinformation, and simultaneously detected theadmixed gene composition of the hybrids, whichwere assigned to more than one cluster.Captive-reared wolves of unknown origin wereprevalently assigned to the Italian wolfpopulation. Admixture analyses showed that one``black wolf' had mixed ancestry in the dog genepool and could be a hybrid, while the other twowolves with unusual phenotypes were assigned tothe Italian wolf population.     

Origin and status of the Great Lakes wolf

An extensive debate concerning the origin and taxonomic status of wolf-like canids in the North American Great Lakes region and the consequences for conservation politics regarding these enigmatic predators is ongoing. Using maternally, paternally and biparentally inherited molecular markers, we demonstrate that the Great Lakes wolves are a unique population or ecotype of gray wolves. Furthermore, we show that the Great Lakes wolves experienced high degrees of ancient and recent introgression of coyote and western gray wolf mtDNA and Y-chromosome haplotypes, and that the recent demographic bottleneck caused by persecution and habitat depletion in the early 1900s is not reflected in the genetic data.