The Wolf Intelligencer


Wolf Evolution and Taxonomy

Climatic comparison of the gray wolf (Canis lupus) subspecies in North America using niche-based distribution models and its implications for conservation programs. González-Bernal A, Rojas-Soto O, Martínez-Meyer E. Journal of Mammalogy. 2022 Jul


The gray wolf, Canis lupus, once inhabited much of the northern hemisphere worldwide; however, persecution drove its populations almost to extinction. In North America, diverse conservation programs have been implemented in the last decades to recover its populations in the wild, many of them guided by the historical distribution of the gray wolf subspecies. Over time, several authors have proposed different subspecies classifications. Nevertheless, most of them are mutually inconsistent regarding the number and distribution of subspecies, creating controversy when implementing conservation programs. This study used niche-based distribution models and cluster analysis to explore the bioclimatic profiles of C. lupus across North America and compare them with different subspecies classifications to identify environmental correlatives that support the proposed designations. Our cluster analysis results indicate that the optimal number of climatic groups was five, designated as Northern, Eastern, Western, Coastal, and Southern groups, with transitional overlap boundaries located at their peripheries, indicating climatic gradients between them and supporting the idea of intergrading zones. The geographic ranges of these groups mismatched to a different extent with all subspecies delimitations. In general, the boundaries of putative subspecies did not match the climatic patterns of North America. Our results may contribute to the recovery programs underway for this carnivore by identifying suitable areas for the release of individuals from specific lineages. New approaches to characterizing the intraspecific variation of the gray wolf should include all evidence available, including genetic, morphological, and ecological information.

Geographic variation in skull morphology of the wolf (Canis lupus) in relation to prey size across North America (Doctoral dissertation, Laurentian University of Sudbury). Dawson Ketchen, J., 2021.

Dietary reconstruction and evidence of prey shifting in Pleistocene and recent gray wolves (Canis lupus) from Yukon Territory. Landry Z, Kim S, Trayler RB, Gilbert M, Zazula G, Southon J, Fraser D. Palaeogeography, Palaeoclimatology, Palaeoecology. 2021 Mar


We investigate if and how diets of gray wolves from the Yukon Territory, Canada, have changed from the Pleistocene (>52.8 ka BP to 26.5 ka BP [±170 y BP]) to the recent Holocene (1960s) using dental microwear analysis of carnassial teeth and stable isotope analyses of carbonates (δ13CCO3 and δ18OCO3) and collagen (δ13Ccol and δ15Ncol) from bone. We find that dental microwear patterns are similar between the Pleistocene and Holocene specimens, indicating that there has been no change in carcass utilization behaviours, where flesh, not bone, is primarily consumed. Based on minimal changes in δ13CCO3 and δ13Ccol values, we find that, over thousands of years, Yukon gray wolves have remained generalist predators feeding upon several large ungulate species. Interestingly, δ15Ncol values suggest that the extinction of megafaunal species at ~11.7 Ka induced a shift from a diet comprised primarily of horse (Equus sp.) to one based on cervids (i.e. moose and caribou). Survival of large-bodied cervids, such as caribou (Rangifer tarandus), was likely key to wolf survival. Although gray wolves survived the end Pleistocene megafauna extinction and demonstrate a degree of ecological flexibility, we suggest that failure to preserve major elements of their current niche (e.G. caribou) may result in continued population declines, especially in the face of increasing anthropogenic influences.

Considering Pleistocene North American wolves and coyotes in the eastern Canis origin story. Wilson P, Rutledge LY.Authorea Preprints. 2020 Sep

Global Phylogeographic and Admixture Patterns in Grey Wolves and Genetic Legacy of An Ancient Siberian Lineage. Pilot M, Moura AE, Okhlopkov IM, Mamaev NV, Alagaili AN, Mohammed OB, Yavruyan EG, Manaseryan NH, Hayrapetyan V, Kopaliani N, Tsingarska E. Scientific reports. 2019 Nov

The Late Pleistocene Canis lupus (Canidae, Mammalia) from Avetrana (Apulia, Italy): reappraisal and new insights on the European glacial wolves; B Mecozzi, SB Lucenti – Italian Journal of Geosciences, 2018

The Wayward Dog: Is the Australian native dog or Dingo a distinct species?; SM JACKSON, CP GROVES, PJS FLEMING… – Zootaxa, 2017

Extinct Beringian wolf morphotype found in the continental US has implications for wolf migration and evolution. Meachen JA, Brannick AL, Fry TJ. Ecology and evolution. 2016 May

Ecological determinants of clinal morphological variation in the cranium of the North American gray wolf. O’Keefe FR, Meachen J, Fet EV, Brannick A. Journal of Mammalogy. 2013 Dec

An account of the taxonomy of North American wolves from morphological and genetic analyses. Chambers SM, Fain SR, Fazio B, Amaral M. North American Fauna. 2012 Oct

Wolves in Trans-Himalayas: 165 years of taxonomic confusion. Shrotriya S, Lyngdoh S, Habib B. Current Science. 2012 Oct

An account of the taxonomy of North American wolves from morphological and genetic analyses. Chambers SM, Fain SR, Fazio B, Amaral M. North American Fauna. 2012 Oct

Morphometric variations of the skull in the Gray Wolf (Canis lupus) in Iran. Khosravi R, Kaboli M, Imani J, Nourani E. Acta Theriologica. 2012 Oct

Modeling effects of environmental change on wolf population dynamics, trait evolution, and life history. Coulson T, MacNulty DR, Stahler DR, Wayne RK, Smith DW. Science. 2011 Dec

Use of cranial characters in taxonomy of the Minnesota wolf (Canis sp.). Mech LD, Nowak RM, Weisberg S. Canadian journal of zoology. 2011 Dec

Modeling Effects of Environmental Change on Wolf Population Dynamics, Trait Evolution, and Life History. Coulson T, MacNulty DR, Stahler DR, vonHoldt B, Wayne RK, Smith DW. Science. 2011

Wolf body mass, skull morphology, and mitochondrial DNA haplotypes in the Riding Mountain National Park region of Manitoba, Canada. Stronen AV, Forbes GJ, Sallows T, Goulet G, Musiani M, Paquet PC. Canadian Journal of Zoology. 2010 May

Skull variation in Dinaric-Balkan and Carpathian gray wolf populations revealed by geometric morphometric approaches. Milenvić M, Šipetić VJ, Blagojević J, Tatović S, Vujošević M. Journal of Mammalogy. 2010 Apr

Taxonomy, morphology, and genetics of wolves in the Great Lakes region. In Recovery of Gray Wolves in the Great Lakes Region of the United States (pp. 233-250). Springer, New York, NY. Nowak, R.M., 2009

Wolf body mass cline across Minnesota related to taxonomy?. Mech LD, Paul WJ. Canadian Journal of Zoology. 2008 Aug

Taxonomic status and conservation strategy of the endangered red wolf: a response to Kyle et al.(2006). Murray DL, Waits LP. Conservation Genetics. 2007 Dec

Wolf body mass cline across Minnesota related to taxonomy?. Mech LD, Paul WJ. Canadian Journal of Zoology. 2008 Aug

Megafaunal extinctions and the disappearance of a specialized wolf ecomorph. Leonard JA, Vilà C, Fox-Dobbs K, Koch PL, Wayne RK, Van Valkenburgh B. Current Biology. 2007 Jul

Ancient origin and evolution of the Indian wolf: evidence from mitochondrial DNA typing of wolves from Trans-Himalayan region and Pennisular India. Aggarwal RK, Ramadevi J, Singh L. Genome Biology. 2003 Jun

Landscape influence on Canis morphological and ecological variation in a Coyote-Wolf C. lupus× latrans hybrid zone, southeastern Ontario. The Canadian Field-Naturalist. 2003

DNA profiles of the eastern Canadian wolf and the red wolf provide evidence for a common evolutionary history independent of the gray wolf. Wilson PJ, Grewal S, Lawford ID, Heal JN, Granacki AG, Pennock D, Theberge JB, Theberge MT, Voigt DR, Waddell W, Chambers RE. Canadian Journal of zoology. 2000 Dec

Changes in the skull morphology of the Arctic wolf, Canis lupus arctos, during the twentieth century. Clutton‐Brock J, Kitchener AC, Lynch JM. Journal of Zoology. 1994 May

Comments on red wolf taxonomy. Phillips MK, Henry VG. Conservation Biology. 1992 Dec

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