UNIVERSITY of CALGARY a Fossilized Turtle Egg Clutch With
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UNIVERSITY OF CALGARY A fossilized turtle egg clutch with embryos from the Upper Cretaceous Oldman Formation, southeastern Alberta: Description, taxonomic identity, and embryonic staging by Amanda Rea McGee A THESIS SUBMITTED TO THE FACULTY OF GRADUATE STUDIES IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE DEPARTMENT OF GEOSCIENCE CALGARY, ALBERTA JANUARY, 2012 © Amanda Rea McGee 2012 Library and Archives Bibliothèque et Canada Archives Canada Published Heritage Direction du Branch Patrimoine de l'édition 395 Wellington Street 395, rue Wellington Ottawa ON K1A 0N4 Ottawa ON K1A 0N4 Canada Canada Your file Votre référence ISBN: 978-0-494-83590-6 Our file Notre référence ISBN: 978-0-494-83590-6 NOTICE: AVIS: The author has granted a non- L'auteur a accordé une licence non exclusive exclusive license allowing Library and permettant à la Bibliothèque et Archives Archives Canada to reproduce, Canada de reproduire, publier, archiver, publish, archive, preserve, conserve, sauvegarder, conserver, transmettre au public communicate to the public by par télécommunication ou par l'Internet, prêter, telecommunication or on the Internet, distribuer et vendre des thèses partout dans le loan, distrbute and sell theses monde, à des fins commerciales ou autres, sur worldwide, for commercial or non- support microforme, papier, électronique et/ou commercial purposes, in microform, autres formats. paper, electronic and/or any other formats. The author retains copyright L'auteur conserve la propriété du droit d'auteur ownership and moral rights in this et des droits moraux qui protege cette thèse. Ni thesis. Neither the thesis nor la thèse ni des extraits substantiels de celle-ci substantial extracts from it may be ne doivent être imprimés ou autrement printed or otherwise reproduced reproduits sans son autorisation. without the author's permission. In compliance with the Canadian Conformément à la loi canadienne sur la Privacy Act some supporting forms protection de la vie privée, quelques may have been removed from this formulaires secondaires ont été enlevés de thesis. cette thèse. While these forms may be included Bien que ces formulaires aient inclus dans in the document page count, their la pagination, il n'y aura aucun contenu removal does not represent any loss manquant. of content from the thesis. UNIVERSITY OF CALGARY FACULTY OF GRADUATE STUDIES The undersigned certify that they have read, and recommend to the Faculty of Graduate Studies for acceptance, a thesis entitled "A fossilized turtle egg clutch with embryos from the Upper Cretaceous Oldman Formation, southeastern Alberta: Description, taxonomic identity, and embryonic staging" submitted by Amanda Rea McGee in partial fulfilment of the requirements of the degree of Master of Science. Supervisor, Darla Zelenitsky, Department of Geoscience François Therrien, Department of Geoscience Jason Anderson, Faculty of Veterinary Medicine External Examiner, Donald Brinkman, Royal Tyrrell Museum External Examiner, Chris Sheil, John Carroll University Date ii Abstract Fossilized turtle clutches containing embryos are heretofore undescribed. Here the description of such a specimen from the Upper Cretaceous of Alberta sheds light on the taxonomic identity and paleobiology of the turtle that laid the clutch. Examination of the clutch reveals 33 rigid-shelled, spherical eggs arranged in a layered cluster. Clutch size and the paleoenvironment of the nest site suggest that the fossil clutch belongs to a large, freshwater turtle. Numerous embryonic bones described from virtual reconstructions of computed tomographic data reveal that the embryonic bones are well-ossified. Comparisons with modern turtles indicate the fossil embryos were at a late stage of development, near hatching, at the time of death. Morphology of the maxillae, dentary, and plastron suggests affinities with Adocidae, an extinct freshwater turtle clade. Additional lines of evidence, including egg and eggshell morphology, the predicted size of the female, and geographic and stratigraphic location, also suggest that the fossil clutch belongs to adocids. iii Acknowledgements I have developed a substantial amount of gratitude to many individuals during my time at the University of Calgary, and would like to thank everyone who has helped me. Firstly, I am grateful to my supervisor, Darla Zelenitsky, for providing me with such a valuable learning experience. Her guidance, knowledgeable comments, and feedback throughout the duration of my project has increased my passion for palaeontology, as well as improved my writing style. I greatly appreciate the time and effort she put into editing and improving each chapter of my thesis. I would also like to thank the members of my defence committee: Christopher A. Sheil, François Therrien, Donald Brinkman, and Jason Anderson for their invaluable advice in their respective areas of expertise, which greatly improved the quality of this thesis. I would like to thank Chris Sheil for providing me the opportunity to visit his lab at John Carroll University, and his continued advice throughout this thesis. Chris is an authority on turtle embryology and his expertise has greatly improved my understanding of the subject. I am very appreciative to Don Brinkman for the valuable discussions we have had, and his knowledgeable advice in regards to fossil turtle taxa from Alberta, as well as access to said specimens. For input into the geological interpretation of the clutch site I thank François Therrien, and for access to unpublished revised radiometric dates I thank Dave Eberth. I am grateful to Walter Joyce for his assistance in helping me navigate through the complicated subject that is turtle phylogeny. I thank Wendy Sloboda for collecting and preparing the fossil clutch that was the focus of this study, and I thank Matthew Colbert from the University of Texas, Austin, for micro-CT scanning the specimen. For their advice and guidance on the software program Amira, I am grateful to Jessica Theodore and Julian Divay. I would iv like to extend my appreciation to the Royal Tyrrell Museum and the Biodiversity Research Institute at the University of Kansas for access to fossil and extant turtle specimens, respectively. Additionally, I would like to thank the Royal Tyrrell Museum for access to their scanning electron microscope. For the preparation of eggshell thin sections, I would like to thank the staff at Calgary Rocks and Material Services Inc. For inspirational discussions, motivation, and unwavering support I would like to thank my parents, Chris and Laura McGee, Kaylee Anderson, Caleb Brown, Ashleigh Cadogan, Shirley Cheng, Greg Dean, Chris Farrow, Dani Fraser, Melissa Freeman, Meg Gavilanez, Kristina Koutras-Virlas, Magdalene Leung, Josh Ludtke, Chris McGarrity, Kelly McGarrity, Heather McGee, Sheila McGee, Katie Quinney, Patty Ralrick, Jessica Theodore, Cam Tsujita, Victoria Walker, and Melissa Zieglar. I would like to especially thank Kate Zubins-Stathopoulos, Annie Quinney, Kohei Tanaka, and Kirstin Brink; you guys rock (pun intended). This research was supported by an Alberta Graduate Student Scholarship, a University of Calgary Thesis Research Grant, the Grant Spratt Graduate Scholarship in Geology, two Queen Elizabeth II Graduate Scholarships, as well as a teaching assistant position. v TABLE OF CONTENTS Approval Page ........................................................................................................................ ii Abstract ................................................................................................................................. iii Acknowledgements ................................................................................................................ iv Table of Contents ................................................................................................................... vi List of Tables ...................................................................................................................... viii List of Figures and Illustrations ............................................................................................. ix List of Plates ........................................................................................................................... x CHAPTER ONE: INTRODUCTION ..................................................................................... 1 CHAPTER TWO: CLUTCH, EGGS, AND EGGSHELLS ................................................... 4 2.1 Introduction ................................................................................................................... 4 2.2 Materials and Methods .................................................................................................. 6 2.2.1 Collection and Preparation of the Clutch ............................................................... 6 2.2.2 Analysis of the Fossil Eggs and Eggshells ............................................................ 6 2.2.3 Comparative Extant Turtle Eggshell ...................................................................... 9 2.2.4 Creation of a Stratigraphic Column ..................................................................... 10 2.3 Geologic Setting and Paleoecology ............................................................................ 11 2.4 Description of Clutch, Eggs, and Eggshell ................................................................. 17 2.4.1 Description of TMP 2008.27.1 ...........................................................................