Synthesis Northeastern coyote / coywolf taxonomy and admixture : A meta-analysis

A flurry of recent papers have attempted to taxonomically characterise eastern canids, mainly grey wolves Canis lupus, eastern wolves Canis lycaon or Canis lupus lycaon and northeastern coyotes or coywolves Canis latrans, Canis latrans var. or Canis latrans x C. lycaon, in northeastern North America. In this paper, we performed a meta-analysis on northeastern coyote taxonomy by comparing results across studies to synthesise what is known about genetic admixture and taxonomy of this animal. Hybridisation or cladogamy (the crossing between any given clades) between coyotes, wolves and domestic dogs created the northeastern coyote, but the animal now has little genetic input from its parental species across the majority of its northeastern North American (e.g. the New England states) range except in areas where they overlap, such as southeastern Canada, Ohio and Pennsylvania, and the midAtlantic area. The northeastern coyote has roughly 60% genetic influence from coyote, 30% wolf and 10% domestic dog Canis lupus familiaris or Canis familiaris. There is still disagreement about the amount of eastern wolf versus grey wolf in its genome, and additional SNP genotyping needs to sample known eastern wolves from Algonquin Provincial Park, Ontario to verify this. Given its mixed species origin and morphological and genetic uniqueness, the most appropriate name for this animal is “coywolf”, which accounts for its two main genetic influences (i.e. coyotes and wolves) in portmanteau order; this name still applies even with the relatively small amount of dog introgression in its genome since dogs are essentially domesticated grey wolves and dog DNA is found in many other wild Canis species including grey wolf populations. It is important for managers to acknowledge that this animal was produced through cladogamy events ~100 years ago, but there is now minimal recent admixture throughout most of its northeastern range. The fact that the coywolf is clearly morphologically and genetically different to any other described population of Canis should qualify the animal for species status. We suggest that they be scientifically classified as Canis oriens, meaning “east”, or more specifically “eastern canid”, in Latin. This nomenclature gives them a distinct stand-alone name separating them from their parental Canis species/types and the associated relative amounts of latrans, lycaon, lupus, and domestic dog genes contributing to their hybrid background. Efforts should be made to monitor the southern wave of western coyote expansion converging with coywolves to document any genetic introgression that might take place between these closely related Canis and whether these two canids will remain distinct. Copyright © 2016 by the IUCN/SSC Canid Specialist Group. ISSN 1478-2677 Way and Lynn Northeastern coyote taxonomy Canid Biology & Conservation | http://www.canids. org/cbc/ 2 Introduction Hybridisation is progressively being more recognised in nature, having been discovered in many taxa (Way 2013). Vargas Pêgas (2013) noted that hybridisation may be a phenomenon misconceived and underestimated by many modern evolutionary biologists and may need revisions in order to respect the new perspectives on hybridisation’s role in evolution, including with humans. Vargas Pêgas (2013) suggested the use of the term “cladogamy” to substitute with “hybridisation” to refer to the crossing between any given clades, due to difficulties from scientists and arbitrary means of separating species from lower taxa. Cladogamy is a term that can apply to crossings between two species given any species concept and any two clades (subspecies or varieties) distinguished by taxonomy, ecology, genetics, geography or phylogeny. Stronen and Paquet (2013) believed that certain hybrids, like northeastern coyotes/coywolves (Canis latrans or Canis latrans x C. lycaon; hereafter northeastern coyotes for consistency purposes), resulting from human actions should be preserved if the animal has replaced the ecological role of extirpated or extinct parent taxa. They argue for conservation policies focusing on protecting the ecological role of taxa affected by hybridisation. Numerous recent papers have attempted to taxonomically characterise eastern canids, mainly grey wolves Canis lupus, eastern wolves Canis lycaon or Canis lupus lycaon, and northeastern coyotes in eastern North America (Chambers et al. 2012, Fain et al. 2010, Monzón et al. 2013, Way 2013, Wilson et al. 2009). Until the publications of Kays et al. (2010) and Way et al. (2010), there had been more of an emphasis on wolf than coyote genetics. But since then, there has been a flurry of additional research on the topic culminating with Way (2013), Monzón et al. (2013) and Wheeldon et al. (2013). At the same time genetic research on coyotes south of the Northeast U.S. has discovered that these animals are more coyote-like compared to northeastern coyotes. Coyotes in the mid-Atlantic region have small amounts of wolf and dog introgression (Bozarth et al. 2011), and southeastern coyotes are more typical western coyotes that have little wolf but some domestic dog Canis lupus familiaris or Canis familiaris (see Dinets 2015 for discussion of domestic dogs being a separate species from grey wolves) admixture (Parker 1995, von Holdt et al. 2011). In this paper, we synthesise contemporary morphological and genetic studies on northeastern coyotes to come to a better understanding of which research findings are largely agreed upon and which have differing conclusions or interpretations of the data. We conclude by making recommendations for northeastern coyote taxonomy.