The gray wolf is the second most specialised member of the genus Canis, after the Ethiopian wolf, as demonstrated by its morphological adaptations to hunting large prey, its more gregarious nature, and its highly advanced expressive behavior. It is nonetheless closely related enough to smaller Canis species, such as the eastern wolf, coyote, and golden jackal, to produce fertile hybrids. It is the only species of Canis to have a range encompassing both the Old and New Worlds, and originated in Eurasia during the Pleistocene, colonizing North America on at least three separate occasions during the Rancholabrean. It is a social animal, travelling in nuclear families consisting of a mated pair, accompanied by the pair's adult offspring. The gray wolf is typically an apex predator throughout its range, with only humans and tigers posing a serious threat to it. It feeds primarily on large ungulates, though it also eats smaller animals, livestock, carrion, and garbage.