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Mammalia, Carnivora): Appearance of the Eurasian Beardog Ysengrinia in North America Robert M University of Nebraska - Lincoln DigitalCommons@University of Nebraska - Lincoln Papers in the Earth and Atmospheric Sciences Earth and Atmospheric Sciences, Department of 2002 Intercontinental Migration of Neogene Amphicyonids (Mammalia, Carnivora): Appearance of the Eurasian Beardog Ysengrinia in North America Robert M. Hunt Jr. University of Nebraska-Lincoln, [email protected] Follow this and additional works at: https://digitalcommons.unl.edu/geosciencefacpub Part of the Earth Sciences Commons Hunt, Robert M. Jr., "Intercontinental Migration of Neogene Amphicyonids (Mammalia, Carnivora): Appearance of the Eurasian Beardog Ysengrinia in North America" (2002). Papers in the Earth and Atmospheric Sciences. 547. https://digitalcommons.unl.edu/geosciencefacpub/547 This Article is brought to you for free and open access by the Earth and Atmospheric Sciences, Department of at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Papers in the Earth and Atmospheric Sciences by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 3384, 53 pp., 24 ®gures, 6 tables December 27, 2002 Intercontinental Migration of Neogene Amphicyonids (Mammalia, Carnivora): Appearance of the Eurasian Beardog Ysengrinia in North America ROBERT M. HUNT, JR.1 CONTENTS Abstract ...................................................................... 2 Introduction ................................................................... 2 Abbreviations ............................................................... 4 Early History of Amphicyonidae in North America ................................ 5 Early Neogene Amphicyonines .................................................. 6 History of Discovery of North American Ysengrinia ............................... 7 Geographic and Geologic Distribution of North American Ysengrinia ................ 8 Systematics ................................................................... 11 Ysengrinia Ginsburg, 1965 .................................................... 11 Ysengrinia americana (Wortman, 1901), new combination ...................... 14 Postcranial Osteology .......................................................... 26 Basicranial Anatomy ........................................................... 43 Dentition and Feeding .......................................................... 47 Acknowledgments ............................................................. 49 References .................................................................... 49 1 Research Associate, Division of Paleontology, American Museum of Natural History; Professor, Geological Sci- ences; and Curator, Vertebrate Paleontology, University of Nebraska, Lincoln, NE 68588-0549. Copyright q American Museum of Natural History 2002 ISSN 0003-0082 2 AMERICAN MUSEUM NOVITATES NO. 3384 ABSTRACT At the beginning of the Neogene a remarkable faunal turnover occurred within the North American carnivore community. The dominant larger Oligocene carnivores (creodonts, nim- ravid cats, the amphicyonid Daphoenus) became extinct during the late Oligocene and were replaced in the early Miocene by amphicyonine amphicyonids and hemicyonine ursids that entered North America from Eurasia. During a ®ve million-year interval from ;23 to 18 Ma, large amphicyonines appear in late Arikareean and early Hemingfordian faunas of the North American midcontinent. Although most fossils are from western Nebraska and southeastern Wyoming, occurrences of amphicyonines at several sites in the eastern United States (Dela- ware, Florida) indicate that they rapidly established a broad geographic distribution in North America during the early Miocene. This report describes and summarizes the North American specimens of the rare immigrant amphicyonine Ysengrinia, the ®rst large amphicyonine to enter the New World. Ysengrinia exists in North America from ;23 to 19 Ma, becoming extinct at the end of the late Arika- reean. A single species of Ysengrinia is recognized in North America: Y. americana (Wortman, 1901), comb. nov., restricted to the late Arikareean of western Nebraska and southeastern Wyoming. The North American hypodigm includes the only known complete skull of the genus, associated with a mandible and partial postcranial skeleton. Because most Eurasian occurrences of Ysengrinia are limited to mandibles or isolated teeth of single individuals, intraspeci®c variation in teeth and skeleton in these carnivores has been dif®cult to determine. The more complete North American specimens provide estimates of dental and osteological variation in Ysengrinia, and suggest that the North American species is dimorphic. Skeletal remains of early Miocene New World Ysengrinia are most often found in riparian and water- hole environments. INTRODUCTION prising large amphicyonine beardogs, hemi- Faunal turnover profoundly altered inte- cyonine ursids, and diverse species of the am- grated mammalian communities at various phicyonid Daphoenodon. times during the Neogene in both North The temnocyonine amphicyonids were the America and Eurasia. These turnover events only major group of large carnivorans to produced changes in faunal composition span the Paleogene-Neogene boundary in throughout the Holarctic region that eventu- North America, apparently unaffected by this ally in¯uenced the southern continents, ®rst North American large carnivore turnover Africa in the early Miocene, and later South event (NALCTE). Temnocyonines ®rst ap- America in the Plio-Pleistocene. On each oc- peared in the early Arikareean (;29±30 Ma) casion, extinction of multiple lineages was and diversi®ed into several lineages (Hunt, followed by the appearance of a new faunal 1998b) that attained their maximum size association. Nowhere is this pattern more ev- (;40±80 kg) in the late Arikareean. The sub- ident than in the turnover of large carnivor- family became extinct at the end of the late ans during the Neogene of North America. Arikareean in North America. Large carnivorans .200 kg did not exist in By the mid-Miocene, creodonts were re- the New World at the beginning of the Neo- duced to relict species in Africa, southern gene. The largest carnivores of the North Asia, and Europe (Barry, 1980, 1988; Gins- American late Paleogene were an aggregation burg, 1980; Morales et al., 1998). Nimravids of diverse, mid-sized (,100 kg in most cases) were represented in the mid- to late Miocene nimravid cats, creodonts, and amphicyonids. of Eurasia and North America by a single This carnivoran assemblage became extinct subfamily, the Barbourofelinae (Geraads and during the late Oligocene with the gradual GuÈlecË, 1997), of which only a few species disappearance of Nimravinae, creodonts (Barbourofelis fricki, North America, to ;6 (Hyaenodontidae), and the daphoenine am- Ma; nimravid, Asian Siwaliks, to ;7.4 Ma; phicyonid Daphoenus. In the early Miocene a Barry and Flynn, 1989) persisted to near the new community of large carnivorans appeared end of the epoch. The numerous genera and in North America (®g. 1, NALCTE), com- species of late Eocene-Oligocene Nimravi- 2002 HUNT: NEOGENE AMPHICYONID YSENGRINIA 3 Fig. 1. Temporal ranges of large Oligocene and early Miocene carnivores in North America, dem- onstrating the faunal turnover event at the Paleogene-Neogene boundary (Global Polarity Time Scale from Berggren et al., 1995). Abbreviations: NALCTE, North American large carnivore turnover event; GPE, last appearance in Great Plains; JDE, last appearance in John Day Formation, Paci®c Northwest; Ma, Mega-annum. 4 AMERICAN MUSEUM NOVITATES NO. 3384 nae found in North America and Eurasia tocene ursid-felid-canid association (Hunt and were extinct by ;24±25 Ma, prior to the Tedford, 1993). Miocene. This report is one in a series of publica- By the end of the early Miocene, amphi- tions that explores the early Neogene radia- cyonids and hemicyonines dominated niches tion of the Amphicyonidae in the New for large carnivores on the northern conti- World. Earlier studies reviewed the Oligo- nents (Hunt, 1998a, 1998b; Ginsburg and cene amphicyonids of North America (Hunt, Morales, 1998). During the mid-Miocene, 1996, 1998b, 2001), the evolution of the these were the ®rst Neogene carnivorans to family in North America (Hunt, 1998b), and attain sizes .200 kg: among the largest of the intercontinental migration of the large these arctoids were Amphicyon ingens of amphicyonine Amphicyon to the New World North America, and the European Amphi- in the early Miocene (Hunt, in press). Here cyon giganteus and hemicyonine Dinocyon I brie¯y review the early history of amphi- thenardi. By the mid-Miocene several am- cyonids in North America (the temnocyoni- phicyonids, including a large species of Am- nes and endemic daphoenines); describe the phicyon, had entered Africa and rapidly pen- faunal turnover event (NALCTE, ®g. 1) at etrated to the southern limit of the continent the Paleogene-Neogene boundary when cre- (Arrisdrift fauna, Namibia: Hendey, 1978; odonts, nimravine cats, Daphoenus, and tem- Morales et al., 1998). nocyonines were replaced by immigrant am- Amphicyonids and hemicyonines contin- phicyonines and hemicyonines; and place on ued to dominate the carnivoran fauna of Ho- record the ®rst of the large immigrant am- larctica into the late Miocene. But within the phicyonines to appear in North America, the late Miocene, during the Clarendonian land Eurasian beardog Ysengrinia. mammal
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