Evolution of Paleocene and Eocene Phenacodontidae (Mammalia, Condylarthra)
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PAPERS ON PALEONTOLOGY--RECENT NUMBERS 18. Ostracods of the Middle Devonian Silica Formation (Volumes I and 11) by Robert V. Kesling and Ruth B. Chilman 19. Late Pleistocene Cold-blooded Vertebrate Faunas from the Mid-Continental United States. I. Reptilia: Testudines, Crocodilia. by Robert E. Preston 20. The Maple Block Knoll Reef in the Bush Bay Dolostone (Silurian, Engadine Group), Northern Peninsula of Michigan by Allan M. Johnson, Robert V. Kesling, Richard T. Lilienthal, and Harry 0.Sorensen 21. A Synopsis of Fossil Grasshopper Mice, Genus Onychomys, and their Relationships to Recent Species by Michael D. Carleton and Ralph E. Eshelman 22. Systematics of the Fissiculate Blastoidea by D. Bradford Macurda 23. Epizoans on the Brachiopod Paraspirifer bownockeri (Stewart) from the Middle Devonian of Ohio by Diana K. Sparks, Richard D. Hoare, and Robert V. Kesling 24. Early Cenozoic Paleontology and Stratigraphy of the Bighorn Basin, Wyoming by Philip D. Gingerich (ed.) and others 25. Dimorphic Middle Devonian Paleocopan Ostracoda of the Great Lakes Region by Robert V. Kesling and Ruth B. Chilrnan 26. The Clarkforkian Land-Mammal Age and Mammalian Faunal Composition across the Paleocene-Eocene Boundary by Kenneth D. Rose 27. Evolutionary History of Microsyopoidea (Mamrnalia, ?h-imates) and the Relationship between Plesiadapiformes and Primates by Gregg F. Gunnel1 28. New Earliest Wasatchian Mammalian Fauna from the Eocene of Northwestern Wyoming: Composition and Diversity in a Rarely Sampled High-Floodplain Assemblage by Philip D. Gingerich 29. Evolution of Paleocene and Eocene Phenacodontidae (Marnrnalia, Condylarthra) by J. G. M. Thewissen Museum of Paleontology The University of Michigan Ann Arbor, Michigan 48109 EVOLUTION OF PALEOCENE AND EOCENE PHENACODONTIDAE (MAMMALIA, CONDYLARTHRA) EVOLUTION OF PALEOCENE AND EOCENE PHENACODONTIDAE (MAMMALIA, CONDYLARTHRA) J. G. M. THEWISSEN Museum of Paleontology The University of Michigan Ann Arbor, Michigan 48109 Present address: Department of Biological Anthropology and Anatomy Duke University Medical Center Durham, North Carolina 277 10 UNIVERSITY OF MICHIGAN PAPERS ON PALEONTOLOGY NO. 29 Papers on Paleontology, No. 29 Museum of Paleontology The University of Michigan Ann Arbor, Michigan 48 109- 1079 Philip D. Gingerich, Director Published December 14, 1990 TABLE OF CONTENTS List of Figures .................................. vi Ectocion cedrus .......................29 ') 1 Ectociort mediotuber ................... JI List of Tables .................................. vii Ectocion osbornianus ..................35 Ectocion major .......................41 Abstract ....................................... ix Ectocion parvus .......................42 Ectocion superstes .....................44 I . INTRODUCTION ..........................1 Phenacodus ............................45 Phenacodus matthewi .................. 47 Identification and Measurement.............. 1 Phenacodus bisonensis .................49 Institutional abbreviations .................. 4 Phenacodus grangeri ..................50 Acknowledgements ....................... 4 Phenacodus magnus ...................55 Phenacodus vortmani .................. 55 GEOCHRONOLOGY AND LOCALITIES .......7 Phenacodus intermedius ................59 North American biochronology and localities ...7 ....................... Phenacodus trilobatus ..................63 Alberta Syncline 9 Phenacodus teilhardi ................... 64 Williston Basin 9 ........................ Phenacodus lemoinei ...................65 Powder River Basin ..................... 9 Phenacodus condali ................... 66 Crazy Mountains Basin ................. 10 Bighorn Basin ........................ 10 Copecion .............................. 66 Wind River Basin ..................... 12 Copecion brachypternus ................66 Green River Basin ..................... 13 Copecion davisi .......................69 Fossil Basin .......................... 13 Bison Basin .......................... 13 IV . THE PATTERN OF PHENACODONTID Great Divide. Washakie. and Sand Wash EVOLUTION ............................. 71 Basins ............................14 North American Phenacodontidae......... 71 Laramie and Shirley Basins ............. 14 Torrejonian ........................71 Piceance Basin ........................ 14 Tiffanian ..........................71 Huerfano Basin .......................15 Clarkforkian and earliest Wasatchian ....73 San Juan Basin ....................... 15 Wasatchian and Bridgerian ............74 Big Bend National Park ................ 15 European Phenacodontidae ..............76 Laudate Canyon ....................... 16 Baja California .......................16 V . CRANIAL OSTEOLOGY OF THE European Biochronology and Localities ...... 16 PHENACODONTIDAE .....................77 Paris Basin ........................... 16 Material .............................77 London Basin ........................ 16 Description ..........................77 Tremp Basin .........................16 Isabena Basin ........................ 17 VI . NOTES ON THE POSTCRANIAL OSTEOLOGY OF PHENACODONTIDAE ..................83 III SYSTEMATIC PALEONTOLOGY 19 . ........... Material .............................83 Condylarthra ............................ 19 Description ..........................83 Phenacodontidae ........................19 Tetraclaemdon .........................20 VlI . CONCLUDING REMARKS .................87 Tetracluenodonpuercensis .............. 21 LITERATURE CITED 93 Tetracluenodon septentrionalis ...........23 ........................... Ectocion ...............................25 APPENDIX ...................................99 Ectocion collinus ......................25 LIST OF FIGURES 1. Method of dental measurements ................. 3 32 . P4 size in Phenacodus ........................ 51 2 . North American phenacodontid localities .......... 8 33 . M, size in Phenacodus .......................51 3 . Dentition of Tetraclaenodon puercensis .......... 22 34 . M2 size in Phenacodus .......................52 4 . Molar size in Tetraclaenodon .................. 24 35 . Principal component 1 in Phenacodus ...........52 5 . Dentition of Tetraclaenodon septentrionalis ......25 36 . Dentition of Phenacodus grangeri ..............53 6 . Dentition of Ectocion collinus ................. 26 37 . Dentition of Phenacodus magnus ...............56 7 . Dentition of Ectocion collinus ................. 27 38 . Dentition of Phenacodus vortmani ..............57 8 . Cheek teeth size in Tiffanian Ectocion ........... 28 39 . Dentition of Phenacodus intermedius ............60 9 . Dentition of Ectocion cedrus .................. 30 40 . Dentition of Phenucodus trilobatus .............63 10. Discriminant function analysis of Ectocion .......31 41 . Dentition of Phenucodus teilhardi ..............65 11. Dentition of Ectocion mediotuber ............... 32 42 . Dentition of Phenacodus lemoinei .............. 65 12. P, size in Ectocion .......................... 33 43 . Dentition of Copecion brachypternus ............67 13. P4 size in Ectocion .......................... 33 44 . P4 size in Copecion ..........................68 14. M, size in Ectocion ..........................34 45 . MI size in Copecion .........................68 15. M size in Ectocion .......................... 34 46 . M2 size in Copecion .........................68 16. M'i size in Ectocion ..........................34 47 . Principal component 1 in Copecion .............68 17. M2 size in Ectocion .......................... 34 48 . Principal component 2 in Copecion .............69 18. Dentition of Ectocion osbornianus .............. 36 49 . Dentition of Copecion davisi .................. 70 19. Dentition of Ectocion osbornianus ..............37 50 . Habitat specialization in Tiffanian 20 . Dental variability in Ectocion .................. 38 phenacodontids .............................72 21 . Canine size in Ectocion ....................... 39 5 1. Abundance shifts in Wasatchian phenacodontids ...75 22 . Discriminant function analysis of Ectocion .......39 52 . Faunal analysis of the Wasatchian ..............76 23 . Dentition of Ectocion major ................... 42 53 . Skull of Phenacodus intermedius ...............78 24 . Molar size in Ectocion major .................. 43 54 . Skull of Phenucodus vortmani .................79 25 . Molar size of Ectocion major and E . superstes ....43 55 . Skull of Ectocion osbornianus .................80 26 . Dentition of Ectocion parvus .................. 44 . 27 . Dentition of Ectocion superstes ................ 46 56 Postcranial osteology of Copecion ..............84 28 . Size of Phenacodus primaevus ................. 47 57 . Clavicle of Phenacodus vortmani ...............85 29 . Dentition of Phenacodus matthewi .............. 47 58 . Tibiae of Wasatchian Phenacodus ..............86 30 . Size in Tiffanian Phenacodus ..................48 59 . Stratigraphic ranges of phenacodontid species .....89 3 1. Dentition of Phenacodus bisonensis ............. 50 LIST OF TABLES 1. North American Land-Mammal Ages ............9 7 . Cheek tooth ratios in Phenacodus .............. 65 2 . Cheek tooth ratios in Ectocion and Copecion ......28 8 . Principal component loadings of Copecion .......69 3 . Cheek tooth ratios in Ectocion and Copecion ......37 9 . Principal component loadings of Faunal Analysis . 76 4 . Molar shape in Ectocion and Copecion .......... 45 10 . Valid phenacodontid taxa .....................88 5 . P4 proportions in Phenacodus .................. 54 11. Weight estimates for phenacodontids