Intercontinental Migration of Large Mammalian Carnivores: Earliest Occurrence of the Old World Beardog \u3ci\u3eAmphicyon\u3c/i\u3e (Carnivora, Amphicyonidae) in North America

North American amphicyonid carnivorans are prominent members of the mid-Cenozoic terrestrial carnivore community during the late Eocene to late Miocene (Duchesnean to Clarendonian). Species range in size from \u3c5 kg to \u3e200 kg. Among the largest amphicyonids are Old and New World species of the genus Amphicyon: A. giganteus in Europe (18–~15? Ma) and Africa, A. ingens in North America (15.9–~14.2 Ma). Amphicyon first appears in the Oligocene of western Europe, surviving there until the late Miocene. Migration to Africa and North America takes place in the early Miocene. The genus occurs in the Arrisdrift fauna (Namibia) of southwest Africa, indicating migration south through the length of the African continent by the mid-Miocene. Its occurrence in Asia is problematical because of the tendency to place any moderately large Asian amphicyonid in the genus, and because of the fragmentary nature of many fossils. Here I report the earliest North American occurrences of Amphicyon (18.8–~17.5 Ma), assigning these individuals to a new and previously undescribed species, Amphicyon galushai, from early Hemingfordian sediments of western Nebraska and north-central Colorado. In the New World, small early Hemingfordian Amphicyon galushai is probably ancestral to larger late Hemingfordian A. frendens, and to the terminal and largest species of the genus, early to mid-Barstovian A. ingens. Diagnostic basicranial and dental traits place these species in the Amphicyonidae, and demonstrate a close relationship of the North American lineage to the type species of the genus, A. major, from Sansan, France. Amphicyon galushai is known from a complete adult skull, a partial juvenile skull, three mandibles, and the isolated teeth and postcranial elements of ~15 individuals, all from the early Miocene Runningwater Formation of western Nebraska. The species also is represented by a crushed rostrum from the Troublesome Formation, north-central Colorado. Basicranial, dental, and postcranial anatomy distinguish A. galushai from its contemporary in the Runningwater Formation, the large digitigrade beardog Daphoenodon. The Runningwater Formation contains the last occurrence of Daphoenodon in North America and the first occurrence of Amphicyon; the overlap in stratigraphic ranges of these two carnivores provides a useful early Miocene biostratigraphic datum. The two amphicyonids occur together in the same quarries, associated with canid, mustelid, and rare procyonid carnivores, which are much smaller animals. The North American species of Amphicyon (A. galushai, A. frendens, A. ingens) most likely adopted ecological roles similar to the large living felids (in particular, the lion Panthera leo). Their robust skeleton with powerful forelimbs, massive clawed feet, heavily muscled jaws with large canines, and a composite crushing/shearing dentition suggest a mobile predator that most likely stalked and ambushed prey from cover, overpowering its victims through sheer size and strength