Population Genomic Analysis of North American Eastern Wolves (Canis lycaon) Supports Their Conservation Priority Status

The threatened eastern wolf is found predominantly in protected areas of central Ontario and has an evolutionary history obscured by interbreeding with coyotes and gray wolves, which challenges its conservation status and subsequent management. Here, we used a population genomics approach to uncover spatial patterns of variation in 281 canids in central Ontario and the Great Lakes region. This represents the first genome-wide single nucleotide polymorphism (SNP) dataset with substantial sample sizes of representative populations. Although they comprise their own genetic cluster, we found evidence of eastern wolf dispersal outside of the boundaries of protected areas, in that the frequency of eastern wolf genetic variation decreases with increasing distance from provincial parks. We detected eastern wolf alleles in admixed coyotes along the northeastern regions of Lake Huron and Lake Ontario. Our analyses confirm the unique genomic composition of eastern wolves, which are mostly restricted to small fragmented patches of protected habitat in central Ontario. We hope this work will encourage an innovative discussion regarding a plan for managed introgression, which could conserve eastern wolf genetic material in any genome regardless of their potential mosaic ancestry composition and the habitats that promote them.

The Anthropocene’s animal? Coywolves as feral cotravelers

This article considers the irreducible indeterminacy of the coywolf and how this shapes human perceptions of the animal, as well as attempts to manage it. The hybridity of the coywolf matters very much to its interactions with humans, as well as the panic that has ensued over its evolutionary success. They are genetic and morphological intermediaries, an admixture of western coyote, eastern wolf, and dog. They hunt in packs like wolves but demonstrate a fearlessness to humans more common of coyotes. They thrive in urban or semiurban environs, moving along our highway, transit, and green space systems in search of food and shelter. I suggest it is the putative ferality of the coywolf—its margin—dwelling between urban and wild, between wolf and coyote—that disrupts our prevailing narratives about how, and on whose terms, animals can occupy the world. But it is also an animal that offers an opening to think about mutual flourishing. I contend this is a fruitful place to start tackling the questions raised by the Anthropocene, and reimagining all creatures as cotravelers.

Observation of an Eastern Wolf (Canis sp. cf. lycaon) Caching Food in a Sphagnum Bog in Algonquin Provincial Park, Ontario

We report summer caching of a partial carcass of a White-tailed Deer (Odocoileus virginianus) fawn by an Eastern Wolf (Canis sp. cf. lycaon) in a Sphagnum bog in Algonquin Provincial Park, Ontario, Canada. The microhabitat conditions in bogs (i.e., low temperature, acidity, and organochemical compounds) likely inhibit food spoilage, making bogs potentially important sites for food caching. Wolves in Algonquin Park experience low summer food availability and high pup mortality from starvation. Caches likely serve as necessary reserve food stores for adults and pups. Recent research has shown that wetland habitats are important den and rendezvous sites for Algonquin Eastern Wolves based on prey availability and, we suggest, perhaps for food storage and accessibility. This caching behaviour was recorded on video. We recommend that future research investigate Eastern Wolf selection of food-caching sites, as a complement to other spatial ecology studies.

Whole-genome sequence analysis shows that two endemic species of North American wolf are admixtures of the coyote and gray wolf

Protection of populations comprising admixed genomes is a challenge under the Endangered Species Act (ESA), which is regarded as the most powerful species protection legislation ever passed in the United States but lacks specific provisions for hybrids. The eastern wolf is a newly recognized wolf-like species that is highly admixed and inhabits the Great Lakes and eastern United States, a region previously thought to be included in the geographic range of only the gray wolf. The U.S. Fish and Wildlife Service has argued that the presence of the eastern wolf, rather than the gray wolf, in this area is grounds for removing ESA protection (delisting) from the gray wolf across its geographic range. In contrast, the red wolf from the southeastern United States was one of the first species protected under the ESA and was protected despite admixture with coyotes. We use whole-genome sequence data to demonstrate a lack of unique ancestry in eastern and red wolves that would not be expected if they represented long divergent North American lineages. These results suggest that arguments for delisting the gray wolf are not valid. Our findings demonstrate how a strict designation of a species under the ESA that does not consider admixture can threaten the protection of endangered entities. We argue for a more balanced approach that focuses on the ecological context of admixture and allows for evolutionary processes to potentially restore historical patterns of genetic variation.

Ancient hybrid origin of the eastern wolf not yet off the table: a comment on Rutledge et al . (2015)

A recent study of North American canids by Rutledge et al. ( Biol. Lett. 11 , 20150303 ( doi:10.1098/rsbl.2015.0303 )) refutes the hypothesized hybrid origin of the eastern wolf (EW) based on genomic evidence against very recent hybridization. However, the analyses do not rule out the possibility of more ancient hybridization. Claims to have resolved the evolutionary origin of the EW are therefore inappropriate. Importantly, though, we plead that uncertainty about the ancient history of the taxon should not affect current conservation policy.

Recent occurrences of wild-origin wolves (Canis spp.) in Canada south of the St. Lawrence River revealed by stable isotope and genetic analysis

A free-ranging canid killed near Caraquet, New Brunswick, Canada, in 2012 exhibited a mitochondrial DNA sequence of Gray Wolf (Canis lupus) origin and a Y-chromosome haplotype of Eastern Wolf (C. lycaon) origin. The animal, which is the first wolf recorded in New Brunswick since 1862, was identified as a Gray–Eastern Wolf hybrid (C. lupus x C. lycaon) based on analysis of its autosomal microsatellite genotype. Stable carbon isotope values (δ13C) suggest that the Caraquet wolf was of wild origin. Likewise, δ13C analysis suggests that a wolf–coyote hybrid killed in Quebec south of the St. Lawrence River in 2002 was also of wild origin. However, δ13C values for a wolf from the same region in 2006 suggest that this animal spent most of its life feeding predominantly on non-wild-source food items. Recent occurrences of wild-origin animals south of the St. Lawrence River demonstrate that wolves are capable of dispersal to formerly occupied areas in southeastern Canada and the United States. However, limited natural dispersal alone will likely not be sufficient to re-establish wolves in northeastern North America.

RAD sequencing and genomic simulations resolve hybrid origins within North American Canis

Top predators are disappearing worldwide, significantly changing ecosystems that depend on top-down regulation. Conflict with humans remains the primary roadblock for large carnivore conservation, but for the eastern wolf ( Canis lycaon ), disagreement over its evolutionary origins presents a significant barrier to conservation in Canada and has impeded protection for grey wolves ( Canis lupus ) in the USA. Here, we use 127 235 single-nucleotide polymorphisms (SNPs) identified from restriction-site associated DNA sequencing (RAD-seq) of wolves and coyotes, in combination with genomic simulations, to test hypotheses of hybrid origins of Canis types in eastern North America. A principal components analysis revealed no evidence to support eastern wolves, or any other Canis type, as the product of grey wolf × western coyote hybridization. In contrast, simulations that included eastern wolves as a distinct taxon clarified the hybrid origins of Great Lakes-boreal wolves and eastern coyotes. Our results support the eastern wolf as a distinct genomic cluster in North America and help resolve hybrid origins of Great Lakes wolves and eastern coyotes. The data provide timely information that will shed new light on the debate over wolf conservation in eastern North America.

Moose (Alces alces) predation by eastern coyotes (Canis latrans) and eastern coyote × eastern wolf (Canis latrans × Canis lycaon) hybrids

It has been widely assumed that coyotes (Canis latrans Say, 1823) are incapable of killing adult moose (Alces alces (L., 1758)) and previous studies of coyote predation support this assumption. However, eastern coyotes and eastern coyote × eastern wolf (Canis lycaon Schreber, 1775) are larger than western coyotes and appear to rely on larger prey in some areas. We used a combination of GPS telemetry, genetic analysis, and field investigation to test the hypothesis that eastern coyotes and coyote × wolf hybrids are capable of preying on adult moose in central Ontario. Our hypothesis was supported, as we documented four definitive cases of eastern coyotes and (or) eastern coyote × eastern wolf hybrids killing moose ≥1.5 years old. Predation by coyotes and coyote × wolf hybrids probably does not represent a threat to moose population viability in central Ontario, but our results suggest that researchers and managers in other areas with declining moose populations that are sympatric with eastern coyotes and (or) coyote × wolf hybrids should consider coyote predation as a potential source of mortality.

Taxonomic Implications of Morphological and Genetic Differences in Northeastern Coyotes (Coywolves) (Canis latrans × C. lycaon), Western Coyotes (C. latrans), and Eastern Wolves (C. lycaon or C. lupus lycaon)

The eastern Coyote or Coywolf (Canis latrans × C. lycaon) inhabiting northeastern North America resulted from hybridization between the expanding population of the western Coyote (Canis latrans) and the remnant population of Eastern Wolf (C. lycaon) and possibly domestic dogs (C. lupus familiaris) in the early 20th century. This study compares the body mass of eastern (i.e., northeastern) Coyotes, western Coyotes, and Eastern Wolves and synthesizes the recent literature to gain better insight into the taxonomic relations and differences of closely-related Canis species. Northeastern Coyotes (males = 16.5 kg; females = 14.7 kg) were statistically (P < 0.0001) intermediate in mass between western Coyotes (males = 12.2 kg; females = 10.7 kg) and Eastern Wolves (males = 28.2 kg, females = 23.7 kg), consistent with their hybrid origin, but were numerically closer to western Coyotes. Large Cohen’s d (3.00–8.56), (0.915–0.929), and Cohen’s f (3.28–3.62) values indicated large effect sizes from the body mass comparisons. Eastern Wolves were 61–71% heavier than the same sex in the northeastern Coyotes, which in turn were ca. 35–37% heavier than the same sex in the western Coyotes. Alternatively, western Coyotes were 73–74% of the size of the same sex in the northeastern Coyotes, which in turn were 59–62% of the size of the same sex in the Eastern Wolves. I also attempted to relate mitochondrial DNA (mtDNA) haplotypes to body mass. Six of 17 (35.3%) adult female northeastern Coyotes captured in Massachusetts weighed ≥18 kg, heavier than any other described Coyote from outside northeastern North America. Mitochondrial DNA haplotypes associated with these heavy female northeastern canids were C9 = 4, C19 = 1, and C48 = 1. Body mass (kg) and mtDNA haplotype data of 53 northeastern Coyotes (males = 28, females = 25) showed no difference between haplotype and body mass for males (P < 0.852) or females (P < 0.128), suggesting that there is not a particular haplotype (e.g., C1) that is associated with the heavier animals. I propose that the most appropriate name for this hybrid animal is Coywolf (Canis latrans × C. lycaon), rather than a type of Coyote. Coywolves are distinct, being larger than any other population of Coyotes but smaller than Eastern Wolves. I propose that the 5 distinct types of Canis be recognized as: western Coyote, Coywolf (northeastern Coyote), Eastern Wolf (including Red Wolf C. rufus), Gray × Eastern Wolf hybrids (‘Great Lakes’ Wolves; C. lupus × C. lycaon or C. lycaon × C. lupus), and Gray Wolf (C. lupus). The implications for wolf recovery in the northeastern United States is discussed.

An Account of the Taxonomy of North American Wolves From Morphological and Genetic Analyses

The available scientific literature was reviewed to assess the taxonomic standing of North American wolves, including subspecies of the gray wolf, Canis lupus. The recent scientific proposal that the eastern wolf, C. l. lycaon, is not a subspecies of gray wolf, but a full species, Canis lycaon, is well-supported by both morphological and genetic data. This species' range extends westward to Minnesota, and it hybridizes with gray wolves where the two species are in contact in eastern Canada and the Upper Peninsula of Michigan, Wisconsin, and Minnesota. Genetic data support a close relationship between eastern wolf and red wolf Canis rufus, but do not support the proposal that they are the same species; it is more likely that they evolved independently from different lineages of a common ancestor with coyotes. The genetic distinctiveness of the Mexican wolf Canis lupus baileyi supports its recognition as a subspecies. The available genetic and morphometric data do not provide clear support for the recognition of the Arctic wolf Canis lupus arctos, but the available genetic data are almost entirely limited to one group of genetic markers (microsatellite DNA) and are not definitive on this question. Recognition of the northern timber wolf Canis lupus occidentalis and the plains wolf Canis lupus nubilus as subspecies is supported by morphological data and extensive studies of microsatellite DNA variation where both subspecies are in contact in Canada. The wolves of coastal areas in southeastern Alaska and British Columbia should be assigned to C. lupus nubilus. There is scientific support for the taxa recognized here, but delineation of exact geographic boundaries presents challenges. Rather than sharp boundaries between taxa, boundaries should generally be thought of as intergrade zones of variable width.