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.     

Molecules and beyond: assessing the distinctness of the Great Lakes wolf

The dog family, Canidae, is a widely distributed group of species that have evolved and radiated relatively recently into 16 genera and 36 recognized species ( Nowak 1999 ). Specific taxonomic designations for some canid taxa can be unclear due to frequent interspecific hybridization among species in both historical and contemporary times, and our imperfect molecular genetic approaches for determining among a series of hypotheses regarding hybridization and evolution. In this issue of Molecular Ecology , Koblmüller et al . tackle the difficult topic of Great Lakes wolf taxonomy and present data that suggest this taxon is currently genetically distinct despite a long history of human persecution and hybridization with related taxa.     

Does interference competition with wolves limit the distribution and abundance of coyotes?

Interference competition with wolves Canis lupus is hypothesized to limit the distribution and abundance of coyotes Canis latrans, and the extirpation of wolves is often invoked to explain the expansion in coyote range throughout much of North America. We used spatial, seasonal and temporal heterogeneity in wolf distribution and abundance to test the hypothesis that interference competition with wolves limits the distribution and abundance of coyotes. From August 2001 to August 2004, we gathered data on cause-specific mortality and survival rates of coyotes captured at wolf-free and wolf-abundant sites in Grand Teton National Park (GTNP), Wyoming, USA, to determine whether mortality due to wolves is sufficient to reduce coyote densities. We examined whether spatial segregation limits the local distribution of coyotes by evaluating home-range overlap between resident coyotes and wolves, and by contrasting dispersal rates of transient coyotes captured in wolf-free and wolf-abundant areas. Finally, we analysed data on population densities of both species at three study areas across the Greater Yellowstone Ecosystem (GYE) to determine whether an inverse relationship exists between coyote and wolf densities. Although coyotes were the numerically dominant predator, across the GYE, densities varied spatially and temporally in accordance with wolf abundance. Mean coyote densities were 33% lower at wolf-abundant sites in GTNP, and densities declined 39% in Yellowstone National Park following wolf reintroduction. A strong negative relationship between coyote and wolf densities (beta = -3.988, P < 0.005, r(2) = 0.54, n = 16), both within and across study sites, supports the hypothesis that competition with wolves limits coyote populations. Overall mortality of coyotes resulting from wolf predation was low, but wolves were responsible for 56% of transient coyote deaths (n = 5). In addition, dispersal rates of transient coyotes captured at wolf-abundant sites were 117% higher than for transients captured in wolf-free areas. Our results support the hypothesis that coyote abundance is limited by competition with wolves, and suggest that differential effects on survival and dispersal rates of transient coyotes are important mechanisms by which wolves reduce coyote densities.     

Ageing and environmental factors affect PCR success in wolf (Canis lupus) excremental DNA samples

Individual genotypes determined from noninvasive DNA samples (typically extracted from shed hairs or scats) are used to estimate population size in monitoring projects of elusive species. However, polymerase chain reaction (PCR) success rates usually are lower, and genotyping errors higher than in standard population genetic surveys, due to DNA degradation or contamination in aged field samples. In this study, we evaluate the results of common garden experiments showing that DNA degradation is significant in wolf (Canis lupus) scats older than 3 days, and it is enhanced in scats in direct contact with soil. A storage test showed that samples kept frozen in 95% ethanol performed better compared to other methods. However, variance of PCR success among samples was high, independent on sample age or storage condition. The detrimental consequences of DNA degradation can be avoided by collecting scat samples as fresh as possible, and implementing efficient multitube procedures and stringent quality control of the laboratory results. Efficient multitube procedures can produce reliable data, like in this study, which showed that the consensus genotypes obtained from excremental DNA exactly matched distinct reference genotypes obtained from wolf blood samples.     

Characterization of 59 canine single nucleotide polymorphisms in the Italian wolf (Canis lupus) population

We characterized 59 canine single nucleotide polymorphisms (SNPs) in the endangered Italian wolf (Canis lupus) population, which were discovered by resequencing sequence‐tagged‐site (STS) DNA sequences that are known to contain SNPs in domestic dogs. Dog SNPs were usually found also in wolves. Additional SNPs unique in dogs or wolves were discovered, which is important for detecting hybrids between dogs and wolves. We developed new primer sets and analysed 15 SNPs by Pyrosequencing. The characterized SNPs will provide an important addition to the genetic markers that are currently available for studying wild populations of canids.     

Genetic diversity, population structure, effective population size and demographic history of the Finnish wolf population

The Finnish wolf population (Canis lupus) was sampled during three different periods (1996-1998, 1999-2001 and 2002-2004), and 118 individuals were genotyped with 10 microsatellite markers. Large genetic variation was found in the population despite a recent demographic bottleneck. No spatial population subdivision was found even though a significant negative relationship between genetic relatedness and geographic distance suggested isolation by distance. Very few individuals did not belong to the local wolf population as determined by assignment analyses, suggesting a low level of immigration in the population. We used the temporal approach and several statistical methods to estimate the variance effective size of the population. All methods gave similar estimates of effective population size, approximately 40 wolves. These estimates were slightly larger than the estimated census size of breeding individuals. A Bayesian model based on Markov chain Monte Carlo simulations indicated strong evidence for a long-term population decline. These results suggest that the contemporary wolf population size is roughly 8% of its historical size, and that the population decline dates back to late 19th century or early 20th century. Despite an increase of over 50% in the census size of the population during the whole study period, there was only weak evidence that the effective population size during the last period was higher than during the first. This may be caused by increased inbreeding, diminished dispersal within the population, and decreased immigration to the population during the last study period.     

Predicting occurrence of wolf territories in Scandinavia

We used logistic regression to compare a set of habitat features inside known Scandinavian wolf Canis lupus territories with the same habitat features in areas outside known territories, but still close enough to be available for wolf colonization. In addition, we analysed changes in habitat variables over time within wolf territories. Wolf territories had lower densities of roads, built-up areas and open land than areas outside wolf territories, but there was no difference in the density of the wolves' main prey, elk Alces alces . The logistic regression model classified 79% of Scandinavia outside the reindeer husbandry area as suitable wolf habitat, that is with a probability of wolf occurrence >0.5. The proportion of built-up areas within the wolf territory decreased as the 'borders' of the wolf territory changed over time. Our model had a reasonably high predictive power, with correct classification in 90% (18 of 20) of the observed wolf territories in the study area. Polygons, randomly distributed outside the observed wolf territories, were correctly classified as not being occupied by wolves in 85% of the cases (17 of 20). This allows a more effective use of resources to, for example, prevent wolf depredation on livestock and dogs.     

Assessment of anadromous salmon resources in the diet of the Alexander Archipelago wolf using stable isotope analysis

The Alexander Archipelago wolf (Canis lupus ligoni) is unique to southeast Alaska, occurring on islands south of Frederick Sound and along the mainland between Dixon Entrance and Yakutat Bay. Sitka black-tailed deer (Odocoileus hemionus sitkensis) are an important prey species for wolves across the southern part of the region. Spawning salmon (Onchorynchus sp.) are seasonally available but their presence in wolf diets has not previously been quantified. We examined the range of bone collagen δ¹³C and δ¹⁵N values for wolves throughout southeast (n = 163) and interior (n = 50) Alaska and used a dual-isotope mixing model to determine the relative contribution of salmon-derived marine protein in the diet. Southeast Alaska wolves consumed significantly more salmon (mean ± SE: 18.3 ± 1.2%) than did wolves from interior Alaska (9.1 ± 0.6%, P<0.001). Wolves on the southeast Alaska mainland appeared to have higher marine isotopic signatures than island wolves, although this difference was not significant. Variation among individual wolf diets was higher for southeast than for interior Alaska wolves, and variation was highest in coastal mainland wolf diets (P<0.001). Marine resources may augment the diet of southeast Alaska wolves during seasonal or annual fluctuations in the availability of deer, particularly in those areas on the mainland where densities of terrestrial ungulates are relatively low. Received: 1 July 1998 / Accepted: 23 February 1999     

Mitochondrial DNA restriction-fragment-length monomorphism in the Italian wolf (Canis lupus) population

Mitochondrial-DNA (mtDNA) restriction patterns were studied in 22 wolves (Canis lupus) sampled in central-northern Italy. A total of 60 restriction sites were detected, encompassing about 2 % of the mitochondrial genome of canids. All wolves showed the same restriction pattern. Therefore, a single mtDNA haplotype was detected in the Italian wolf population. Historical information on peninsular isolation and demographic decline suggest that low genetically effective population size and random drift may have strongly reduced the mtDNA variability of wolves in Italy over the last 100–200 years. A different mtDNA restriction pattern in feral dogs sampled from a wolf range in central Italy was detected. These findings suggest that the hybridization and introgression of female dog genomes into the Italian wolf population may be rare or absent.     

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.