LATE EOCENE SEA COWS (MAMMALIA , SIRENIA) FROM WADI AL HITAN IN THE FAYUM BASIN, EGYPT by IYAD SALEH ZALMOUT A dissertation submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy (Geology) in The University of Michigan 2008 Doctoral Committee: Professor Philip D. Gingerich, Chair Professor Daniel C. Fisher Professor Philip D. Myers Associate Research Scientist Gregg F. Gunnell Assistant Research Scientist William J. Sanders IYAD SALEH ZALMOUT © 2008 All Rights Reserved TO MY FAMILY ii ACKNOWLEDGMENTS I am indebted to my committee members: to Philip D. Gingerich for his unwavering support, insight, enthusiasm, and generosity; for allowing me to describe the best sirenian specimens collected from the Tethys Sea; and for his considerable time and attention devoted to my training in the field of vertebrate paleontology. I thank Daniel C. Fisher for long insightful conversations, guidance, and helpful suggestions; Philip Myers for his experience in studying mammalian evolution and taxonomy ; William J. Sanders, whose interests and substantial contribution to this work are much appreciated; and Gregg F. Gunnell for his many helpful comments and for his help with the identification and management of specimens. I am also grateful to Daryl P. Domning of Howard University and the National Museum of Natural History for his guidance, valued discussion, and access to Eocene Sirenia of Jamaica as well as comparative material at the Smithsonian Institution. This work could not been achieved without the Egyptian government collaborations; thanks are due to the director and staff of Egyptian Environment Affairs Agency Dr. Mustafa Fouda, Mohammad S. A. Abdulhamid, Mohammed Talaat El- Hennawy; Mohammad Al Hakeem, Yusri Attia, Majdi Zakaria, Ehab El-Sady, and Medhat Al Said of the Geological Museum of Egypt. I also thank the field crew members who carried out several expeditions to Wadi Al Hitan with Philip D. Gingerich when many fossil sirenians were collected and recorded: B. Holly Smith, William J. Sanders, Ali Barakat, William C. Clyde, Amin Strougo, Alex van Nievelt, M. Hilal, A. A. Abdul Latif, Jeffrey A. Wilson, David J. Ward, Chris King, Shannon Peters, Martin Sander, Carole Gee, T. M. Bown, and Elwyn L. Simons. iii William J. Sanders, Joseph R. Groenke, and John Graf did fabulous work in preparing, molding and casting the sirenian material studied here. I am also deeply thankful to Museum of Paleontology illustrator Bonnie J. Miljour for many of the illustrations included here. Timothy P. Utter and Karl E. Longstreth from the Map Library of the University of Michigan were very helpful and supportive during my research. In addition, I thank Gerald R. Smith, Bruce Wilkinson, Catherine E. Badgley, Jeffrey A. Wilson, Kyger Lohmann, Samuel Mukasa, Abdulmajid Al Nubhani, Daniel j. Miller, Miriam L. Zelditch, John A. Whitlock, Jacques LeBlanc, Aaron A. Wood, Adam Rountrey, Ross Secord, Takehito Ikejiri, Ryan M. Bebej, Kathlyn M. Smith, John I. Bloch, Brian Beatty, and Doug M. Boyer for their help and support during my graduate studies at the University of Michigan. Finally, this research was funded by grants from the U. S. National Science Foundation (EAR 0517773, OISE 0513544), the National Geographic Society (most recently 09-1035), and the University of Michigan. iv TABLE OF CONTENTS DEDICATION .................................................................................................................. ii ACKNOWLEDGMENTS .............................................................................................. iii LIST OF TABLES ........................................................................................................ viii LIST OF FIGURES ....................................................................................................... xii ABSTRACT ................................................................................................................. xviii CHAPTER ONE: INTRODUCTION .............................................................................1 Sirenian Evolution .............................................................................................2 Sirenia of Egypt (Fayum and Cairo) ...............................................................4 New Specimens ..................................................................................................7 Methods ..............................................................................................................8 Objectives ...........................................................................................................9 Tables ................................................................................................................11 Figures ..............................................................................................................14 Literature Cited ...............................................................................................21 CHAPTER TWO: GEOLOGY AND STRATIGRAPHY ..........................................29 Geology and Stratigraphy of the Fayum Basin and Surrounding Areas ...30 Stratigraphic Levels Yielding Sirenia in Wadi Al Hitan .............................39 Figures ..............................................................................................................45 Literature Cited ...............................................................................................58 CHAPTER THREE: REVIEW OF THE CENOZOIC SIRENIA OF AFRICA .....62 Systematic Paleontology .................................................................................63 Order Sirenia Illiger, 1811 ..............................................................................63 Family Protosirenidae Sickenberg, 1934 ..............................................63 Protosiren Abel, 1907 ..................................................................64 v Libysiren (Heal, 1973) .................................................................66 Family Dugongidae Gray, 1821 .............................................................67 Subfamily Halitheriinae (Carus, 1868) .................................................68 Eotheroides Palmer, 1899 ...........................................................68 Eosiren Andrews, 1902 ...............................................................70 Halitherium Kaup, 1838 .............................................................74 Metaxytherium de Christol, 1840 ...............................................75 Subfamily Dugonginae (Gray, 1821) ....................................................77 Rytiodus Lartet, 1866 ..................................................................77 Indeterminate Sirenia of Assorted Ages ........................................................78 Discussion .........................................................................................................81 Summary ..........................................................................................................83 Tables ................................................................................................................85 Figures ..............................................................................................................87 Literature Cited ...............................................................................................91 CHAPTER FOUR: NEW EOCENE SIRENIA FROM THE PRIABONIAN OF WADI AL HITAN .........................................................................................................103 Systematic Paleontology ...............................................................................104 Eotheroides clavigerum sp. nov. ...........................................................104 Eotheroides sandersi sp. nov. ...............................................................133 Comparison and Relationships ....................................................................175 Discussion .......................................................................................................182 Conclusions ....................................................................................................186 Tables ..............................................................................................................187 Figures ............................................................................................................213 Literature Cited .............................................................................................268 CHAPTER FIVE: SEXUAL DIMORPHISM IN EOCENE SIRENIA ..................275 Introduction ...................................................................................................275 Sexual Dimorphism in Eocene Sirenia ........................................................278 Pelvic Morphology of Protosirenidae .................................................280 Pelvic Morphology of Dugongidae ......................................................281 vi Discussion and Conclusion ...........................................................................284 Tables ..............................................................................................................287 Figures ............................................................................................................288 Literature Cited .............................................................................................298 CHAPTER SIX: PALEOENVIRONMENTS AND PALEOECOLOGY ...............303