University of Alberta Early late Paleocene mammals from the Roche Percée local fauna, southeastern Saskatchewan, Canada by Brian Daniel Rankin A thesis submitted to the Faculty of Graduate Studies and Research in partial fulfillment of the requirements for the degree of Master of Science in Systematics and Evolution Department of Biological Sciences ©Brian Daniel Rankin Fall, 2009 Edmonton, Alberta Permission is hereby granted to the University of Alberta Libraries to reproduce single copies of this thesis and to lend or sell such copies for private, scholarly or scientific research purposes only. Where the thesis is converted to, or otherwise made available in digital form, the University of Alberta will advise potential users of the thesis of these terms. The author reserves all other publication and other rights in association with the copyright in the thesis and, except as herein before provided, neither the thesis nor any substantial portion thereof may be printed or otherwise reproduced in any material form whatsoever without the author's prior written permission. Examining Committee Dr. Richard C. Fox, Biological Sciences, University of Alberta Dr. Mark V. H. Wilson, Biological Sciences, University of Alberta Dr. Alison M. Murray, Biological Sciences, University of Alberta Dr. Pamela R. Willoughby, Anthropology, University of Alberta For all my friends and family — who still think I research dinosaurs. ABSTRACT The occurrence of vertebrate fossils from the Ravenscrag Formation near Roche Percée, southeastern Saskatchewan, Canada, documents the presence of a large and diverse assemblage of early late Paleocene (approx. 58 million years) mammals. Previous studies of the Roche Percée localities have examined only a small portion of the fauna, with the vast majority of taxa remaining undescribed. The current research centers on the identification, description and, where appropriate, evolutionary relationships of these undescribed mammals. Significant discoveries, to date, include two new species of the rare viverravid carnivoran Raphictis, a new species of the phenacodontid condylarth Ectocion, a large collection of a probable new species of the semi-aquatic pantolestid Palaeosinopa, and only the second known occurrence of the predominantly European lipotyphlan Adapisorex in North America. This research provides an improved understanding of mammalian diversity and evolution in the northern part of the Western Interior during this important time interval. ACKNOWLEDGMENTS My most sincere thanks are owed to Dr. Richard C. Fox for his co-supervision during my thesis research and for the opportunity to study the mammalian remains from the Roche Percée local fauna, one of the largest and most diverse assemblages in the University of Alberta Laboratory for Vertebrate Paleontology collections. Because of the influence and guidance of Dr. Fox, I have gained much experience and a broad knowledge of early Tertiary mammals. The co-supervision and mentoring of Dr. Mark V. H. Wilson was also paramount to the success of this study and to my development as an academic, and I thank him for all the unexpected (but much appreciated) support. I additionally thank Dr. Ruth A. Stockey for her suggestion and encouragement to continue with academics after graduating with my Bachelor’s of Science, and the other members of my examining committee, Drs. Pam R. Willoughby and Alison M. Murray, for their advice and comments on my thesis research. Dr. Craig S. Scott is owed an enormous debt of gratitude for his guidance during my undergraduate and graduate studies, and for sharing his thoughts on the evolutionary history of several early Tertiary mammals. For reading an earlier version of this thesis and providing many helpful comments during the course of my studies, I thank Dr. Eric Snively. Thanks are also extended to Dr. Michael G. Newbrey for advice on photographic techniques and Dr. Michael W. Caldwell for use of photographic equipment. For their friendship, support, and insight, I owe many thanks to my peers and colleagues at the University of Alberta Laboratory for Vertebrate Paleontology: Victoria Arbour, Braden Barr, Phil Bell, Stephanie Blais, Lisa Buckley, Michael Burns, Todd Cook, Jessica Hawthorn, Michael James, Dr. Takuya Konishi, Derek Larson, Aaron LeBlanc, Lindsay MacKenzie, Dr. Erin E. Maxwell, Scott Persons, Miriam Reichel, Bradley Scott, Robin Sissons, and Andrew Wendruff. In addition, I am fortunate to have met many other graduate students from the Department of Biological Sciences at the University of Alberta (past and present) who were helpful and perceptive. In particular, I thank Steve Karafit, Linnea Mowat, Dustin Raab, Christine Robichaud, Barry Robinson, Jeremy Wideman and Drs. Stefan A. Little, Randal A. Mindell and Selena Y. Smith. My friends and family (Dad, Mom, Lorelle, Nigel, Moya, Rankin, Donna, and Brad) have been a constant source of support and a relaxing break from academic life. Their continued patience, encouragement, and general interest with regards to my education is no small reason for the completion of this study - I hope that you are all proud. Finally, I thank Tara Jamison. Without her love and support this project would not have come to fruition. Funding for this project was provided to me by Graduate Teaching Assistantships from the Department of Biological Sciences and minor awards granted to me from several faculties at the University of Alberta. TABLE OF CONTENTS INTRODUCTION 1 REGIONAL GEOLOGICAL SETTING AND LOCATION 3 TECHNIQUES OF STUDY 5 SYSTEMATIC PALEONTOLOGY 7 Order Didelphimorphia 7 Peradectes cf. P. protinnominatus 7 Superorder Leptictida 9 Prodiacodon cf. P. concordiarcensis 9 Order Erinaceomorpha 10 Diacocherus cf. D. meizon 11 “Xynolestes denommei” 12 Litocherus lacunatus 14 Litocherus cf. L. zygeus 19 Adapisorex sp. 21 Litolestes cf. L. ignotus 22 Order Soricomorpha 23 Nyctitherium sp. 23 Order Cimolesta 25 Pararyctes pattersoni 26 Paleotomus cf. P. junior 27 cf. Paleotomus sp. 1 28 cf. Paleotomus sp. 2 31 Unuchinia cf. U. dysmathes 31 Labidolemur soricoides 40 Titanoides cf. T. primaevus 41 Bessoecetor pilodontus 43 Bessoecetor krausei n. sp. 49 Palaeosinopa sp. 53 Order Carnivora 59 Didymictis dellensis 59 Didymictis cf. D. dellensis 63 Protictis cf. P. paralus 67 Raphictis “iota” 68 Raphictis machaera n. sp. 70 Raphictis nanoptexis n. sp. 77 Order Procreodi 81 Chriacus cf. C. baldwini 81 Chriacus cf. C. oconostotae 82 Thryptacodon australis 83 Arctocyon mumak 86 Arctocyon corrugatus 93 Order Condylarthra 95 Dorraletes diminutivus 95 Phenacodus magnus 99 Ectocion borealis n. sp. 103 CONCLUSIONS 107 Age of the Roche Percée local fauna 107 Faunal comparisons 109 Implications for further research 111 REFERENCES 112 TABLES 139 FIGURES 172 APPENDICES 212 Appendix 1: Taxonomic listing of mammals from the Roche Percée local fauna 212 LIST OF TABLES Table 1.—Measurements and descriptive statistics of the dentition of Prodiacodon cf. P. concordiarcensis from the late middle Tiffanian (Ti4) Roche Percée local fauna, Ravenscrag Formation, Saskatchewan. 139 Table 2.—Measurements and descriptive statistics of the upper dentition of Diacocherus cf. D. meizon from the late middle Tiffanian (Ti4) Roche Percée local fauna, Ravenscrag Formation, Saskatchewan. 140 Table 3.—Measurements and descriptive statistics of the lower dentition of Diacocherus cf. D. meizon from the late middle Tiffanian (Ti4) Roche Percée local fauna, Ravenscrag Formation, Saskatchewan . 141 Table 4.—Measurements and descriptive statistics of the lower dentition of “Xynolestes denommei” from the late middle Tiffanian (Ti4) Roche Percée local fauna, Ravenscrag Formation, Saskatchewan. 142 Table 5.—Measurements and descriptive statistics of the upper dentition of Litocherus lacunatus from the late middle Tiffanian (Ti4) Roche Percée local fauna, Ravenscrag Formation, Saskatchewan. 143 Table 6.—Measurements and descriptive statistics of the lower dentition of Litocherus lacunatus from the late middle Tiffanian (Ti4) Roche Percée local fauna, Ravenscrag Formation, Saskatchewan. 144 Table 7.—Measurements and descriptive statistics of the lower dentition of Litocherus cf. L. zygeus from the late middle Tiffanian (Ti4) Roche Percée local fauna, Ravenscrag Formation, Saskatchewan. 145 Table 8.—Measurements and descriptive statistics of the dentition of Adapisorex sp. from the late middle Tiffanian (Ti4) Roche Percée local fauna, Ravenscrag Formation, Saskatchewan. 146 Table 9.—Measurements and descriptive statistics of the lower dentition of Pararyctes pattersoni from the late middle Tiffanian (Ti4) Roche Percée local fauna, Ravenscrag Formation, Saskatchewan. 147 Table 10.—Measurements and descriptive statistics of the lower dentition of cf. Paleotomus sp. 1 from the late middle Tiffanian (Ti4) Roche Percée local fauna, Ravenscrag Formation, Saskatchewan. 148 Table 11.—Measurements and descriptive statistics of the upper dentition of Unuchinia cf. U. dysmathes from the late middle Tiffanian (Ti4) Roche Percée local fauna, Ravenscrag Formation, Saskatchewan . 149 Table 12.—Measurements and descriptive statistics of the lower dentition of Unuchinia cf. U. dysmathes from the late middle Tiffanian (Ti4) Roche Percée local fauna, Ravenscrag Formation, Saskatchewan. 150 Table 13.—Measurements and descriptive statistics of the upper and lower dentition of Bessoecetor pilodontus from the late middle Tiffanian (Ti4) Roche Percée local fauna, Ravenscrag Formation, Saskatchewan. 151 Table 14.—Measurements and descriptive statistics of the