Nowak, R.M. and N.E. Federoff. 1996. Systematics of wolves in Eastern North America. Pages 187-203 in Defenders of Wildlife's "Wolves of America" conference proceedings, Albany, New York, November, 14-16, 1996.
Cranial morphology of Recent wolves throughout northern and western North America is remarkably consistent. Statistical analysis indicates the presence of four subspecies of gray wolf (Canis lupus) there, which are always distinguishable from the sympatric coyote (Canis latrans). A fifth gray wolf subspecies, lycaon, occurs in southeastern Canada, and the red wolf (Canis rufus), is found in the southeastern United States. During the early 1900s, the coyote moved east of the prairies and hybridized with the native wolves, thereby creating much confusion. Nonetheless, analysis of every available specimen of wild Canis, dating from before the coyote invasion in the region east of the Mississippi River and south of Wisconsin, Michigan and New York, does indicate the presence of a small wolf, distinct from the coyote and showing the statistical consistency of other wolf populations. That series also has close affinity to specimens of the red wolf collected in Louisiana and Missouri prior to 1925 and to Pleistocene fossils from the east. There was a sharp line of morphological distinction between the wolves of the eastern United States and those of the prairies, but a closer approach by the former to the subspecies lycaon, which in turn intergrades with gray wolf populations in western Ontario and Minnesota. Although gaps in our knowledge remain, a reasonable hypothesis is that the entire forested region from southeastern Canada to the Gulf Coast originally was inhabited by populations of small wolves, with a subspecific or specific line just south of the eastern Great Lakes. There is no evidence that southeastern North America was ever occupied by large gray wolves and coyotes that hybridized to form the red wolf.