TAPHONOMIC AND BIOSTRATIGRAPHIC ANALYSIS OF FOSSIL FRESHWATER TURTLES IN THE UPPER CRETACEOUS (CAMPANIAN) KAIPAROWITS FORMATION OF SOUTHERN UTAH, USA by Michael Jo Knell A dissertation submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Earth Sciences MONTANA STATE UNIVERSITY Bozeman, Montana November 2012 ©COPYRIGHT by Michael Jo Knell 2012 All Rights Reserved ii APPROVAL of a dissertation submitted by Michael Jo Knell This dissertation has been read by each member of the dissertation committee and has been found to be satisfactory regarding content, English usage, format, citation, bibliographic style, and consistency and is ready for submission to The Graduate School. David J. Varricchio Approved for the Department of Earth Sciences David Mogk Approved for The Graduate School Dr. Ronald W. Larson iii STATEMENT OF PERMISSION TO USE In presenting this dissertation in partial fulfillment of the requirements for a doctoral degree at Montana State University, I agree that the Library shall make it available to borrowers under rules of the Library. I further agree that copying of this dissertation is allowable only for scholarly purposes, consistent with “fair use” as prescribed in the U.S. Copyright Law. Requests for extensive copying or reproduction of this dissertation should be referred to ProQuest Information and Learning, 300 North Zeeb Road, Ann Arbor, Michigan 48106, to whom I have granted “the exclusive right to reproduce and distribute my dissertation in and from microform along with the non- exclusive right to reproduce and distribute my abstract in any format in whole or in part.” Michael Jo Knell November 2012 iv DEDICATION I dedicate this dissertation to my parents, who have supported me emotionally, and frequently financially, through my undergraduate and graduate education. They both encouraged me to follow my interests in the sciences ever since I became hooked on the subject in grade school. I would like to thank them for constantly pushing me to do my best and for being patient through, what seemed like, endless years of graduate school. Thanks to my Dad for making sure my bills were paid and my car was running smoothly. I also need to acknowledge the extraordinary efforts of my Mom, who always offered helpful advice when I needed it and frequently went above and beyond to ensure that I was healthy and well fed. If she hadn’t taken me on summer trip to Montana to look for dinosaur bones back in 1994, I’m not sure I would have followed the path that I did. Now, nearly 20 years later, I’m continuing to do what I love the most, and it all began with that first taste of paleontology. Thank you, Mom and Dad, for everything! v ACKNOWLEDGEMENTS I would like to first acknowledge my primary advisor, David Varricchio, my dissertation committee, Jack Horner, Frankie Jackson, Jim Schmitt, and my graduate representative Jioanna Carjuzaa for their helpful suggestions, edits, and expertise. I must thank Alan Titus at the Bureau of Land Management, Grand Staircase-Escalante National Monument Field Office, for first introducing me to the fossils of the Kaiparowits Formation and for supporting my project, and also Scott Richardson for assistance with fieldwork and fossil preparation. Thanks to Eric Roberts for greatly assisting with my fieldwork questions, use of published figures, and helpful advice regarding my project. I must also thank his father, Steve Roberts, for allowing me to camp on his property while conducting fieldwork. Thanks to Mike Getty and field crew at the Natural History Museum of Utah for their support during fieldwork and also for assisting with collections visits to the museum. Thanks to Pat Holroyd at the University of California Museum of Paleontology and Rod Sheetz at Brigham Young University for access to their collections. Thanks to Howard Hutchison for sharing his wisdom during multiple conversations regarding the fossil turtles of the Kaiparowits. This project would not have been possible without donations of extant turtle specimens by Bill Belzer, George Phillips, and Bob Thomson. I owe a great amount of gratitude to my roommate, and fellow student, Tom Evans for his valuable advice and assistance with my project and preparation of my dissertation. Finally, I wouldn’t have survived without the support and friendship from my fellow Earth Science graduate students through the years, so I want to share my gratitude and wish you all the best in your future endeavors. Cheers! vi VITA Michael Jo Knell was born in Cedar Rapids, IA on November 14, 1979 to Randolph and Linda Knell. His hometown is Council Bluffs, IA where he graduated from Lewis Central Senior High School in 1998. He attended the University of Nebraska in Lincoln where he graduated with a B.S. in Geology in 2002. He subsequently attended South Dakota School of Mines & Technology where he earned his M.S. in Paleontology in 2004. Michael then attended Montana State University to earn his Ph.D. in Earth Sciences. He currently is teaching as an Assistant Professor in the Department of Earth Sciences at Southern Connecticut State University in New, Haven Connecticut. vii TABLE OF CONTENTS 1. INTRODUCTION ...........................................................................................................1 2. BIOGEOGRAPHIC ANALYSIS OF THE CRETACEOUS FRESHWATER AND TERRESTRIAL TURTLES OF WESTERN NORTH AMERICA ........................................................................................................4 Introduction ......................................................................................................................4 The Turtle.....................................................................................................................5 Freshwater Turtles .......................................................................................................6 Terrestrial Turtles.........................................................................................................7 Cretaceous Freshwater Turtles of Western North America .............................................7 Adocidae ......................................................................................................................8 Baenidae .......................................................................................................................9 Bothremydidae ...........................................................................................................10 Chelydridae ................................................................................................................11 Kinosternidae .............................................................................................................11 Macrobaenidae ...........................................................................................................12 Nanhsiungchelyidae ...................................................................................................12 Pleurosternidae ...........................................................................................................13 Solemydidae ...............................................................................................................13 Trionychidae ..............................................................................................................14 Biogeography of Cretaceous Turtles in Western North America ..................................15 Cretaceous Fossil Turtle Assemblages of Western North America ...............................17 Berriasian-Barremian .................................................................................................18 Aptian-Albian ............................................................................................................18 Albian .........................................................................................................................19 Albian-Cenomanian ...................................................................................................19 Cenomanian ...............................................................................................................21 Turonian-Coniacian ...................................................................................................22 Santonian....................................................................................................................23 Campanian .................................................................................................................24 Campanian-Maastrichtian ..........................................................................................28 Maastrichtian..............................................................................................................29 Discussion ......................................................................................................................33 Changes in Diversity and Faunal Turnover ...............................................................33 Phylogenetics and Diversity ......................................................................................35 Spatial Trends ............................................................................................................36 Literature Cited ..............................................................................................................58 viii TABLE OF CONTENTS ¬ CONTINUED 3. TAPHONOMIC ANALYSIS OF FOSSIL FRESHWATER TURTLES IN THE UPPER CRETACEOUS (CAMPANIAN) KAIPAROWITS FORMATION