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Investigating Sexual Dimorphism in Ceratopsid Horncores University of Calgary PRISM: University of Calgary's Digital Repository Graduate Studies The Vault: Electronic Theses and Dissertations 2013-01-25 Investigating Sexual Dimorphism in Ceratopsid Horncores Borkovic, Benjamin Borkovic, B. (2013). Investigating Sexual Dimorphism in Ceratopsid Horncores (Unpublished master's thesis). University of Calgary, Calgary, AB. doi:10.11575/PRISM/26635 http://hdl.handle.net/11023/498 master thesis University of Calgary graduate students retain copyright ownership and moral rights for their thesis. You may use this material in any way that is permitted by the Copyright Act or through licensing that has been assigned to the document. For uses that are not allowable under copyright legislation or licensing, you are required to seek permission. Downloaded from PRISM: https://prism.ucalgary.ca UNIVERSITY OF CALGARY Investigating Sexual Dimorphism in Ceratopsid Horncores by Benjamin Borkovic A THESIS SUBMITTED TO THE FACULTY OF GRADUATE STUDIES IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE DEPARTMENT OF BIOLOGICAL SCIENCES CALGARY, ALBERTA JANUARY, 2013 © Benjamin Borkovic 2013 Abstract Evidence for sexual dimorphism was investigated in the horncores of two ceratopsid dinosaurs, Triceratops and Centrosaurus apertus. A review of studies of sexual dimorphism in the vertebrate fossil record revealed methods that were selected for use in ceratopsids. Mountain goats, bison, and pronghorn were selected as exemplar taxa for a proof of principle study that tested the selected methods, and informed and guided the investigation of sexual dimorphism in dinosaurs. Skulls of these exemplar taxa were measured in museum collections, and methods of analysing morphological variation were tested for their ability to demonstrate sexual dimorphism in their horns and horncores. Skulls and isolated horncores of Triceratops and Centrosaurus were measured in museum collections, and analysed with the methods applied to the exemplar taxa. Considerable morphological variation was found in both species, and was suggestive of dimorphism in Centrosaurus, but was deemed not to be attributable to sexual dimorphism. ii Acknowledgements Since long before I set foot on a university campus, my family has been a steady source of encouragement and enthusiasm for my interest in palaeontology, and I am very grateful for all of their support over the years. My family rocks. Once I became interested in graduate school, it was Dr. Russell’s support and patient guidance that helped build this project from mere ideas. I have learned a great deal from Dr. Russell – especially how to be a better scientist. I would like to thank my supervisory committee of Dr. Theodor and Dr. Brinkman for their creative and critical input, and for their assistance with methods and locating specimens. My labmates have been good friends and able teachers, especially Alex Tinius, Sabrina Hynes, Magdalene Leung, Larry Powell, Hillary Maddin, and Robin Cuthbertson. Jordan Mallon was a mentor for many aspects of this project, and we had many informative discussions while driving to and from museums. Dr. Steeve Côté provided data and information about mountain goats early on in my research. There is a long list of people who provided couches, floors, and beds during my research trips, and I am very grateful to Robert Holmes and Allison Murray, Peter Good and Lisa Hall, Eric Oosenbrug, Auntie Sam, the Maddin family, the Hawkins family, Daniel Field, Brian Rankin, and the Crawford family. Meaghan Crawford was a pillar of encouragement during the writing of this thesis, and Kristina Juul was a great saviour in England. Thank you to all of the museum staff and collections managers who provided access to, and assistance with, their collections: Brandon Strilisky, Darren Tanke, Margaret Currie, Bill Weimann, Kevin Seymour, Susan Woodward, Allison Murray, Eileen Westwig, Susan Hochgraf, Amy Henrici, Dan Brinkman, Carl Mehling, Wes Long, Andrew Locock, and Glen and Ethel. iii Table of Contents Abstract ........................................................................................................................................... ii Acknowledgements ........................................................................................................................ iii Table of Contents ........................................................................................................................... iv List of Tables ................................................................................................................................ vii List of Figures and Illustrations ..................................................................................................... ix List of Symbols, Abbreviations, Nomenclature ............................................................................ xii Chapter 1: Introduction ................................................................................................................... 1 1.1 Preamble ............................................................................................................................... 1 1.2 Ceratopsid Dinosaurs ............................................................................................................ 1 1.2.1 Morphological Variation in Ceratopsids ........................................................................ 2 1.3 Horns ..................................................................................................................................... 4 1.4 Sexual Dimorphism .............................................................................................................. 5 1.5 Prospectus ........................................................................................................................... 10 Chapter 2: Evidence for Sexual Dimorphism as Manifested in the Vertebrate Fossil Record ..... 11 2.1 Introduction ......................................................................................................................... 11 2.2.1 Fishes ........................................................................................................................... 16 2.2.2 Amphibia...................................................................................................................... 21 2.2.3 Testudines .................................................................................................................... 22 2.2.4 Sauropterygia ............................................................................................................... 23 2.2.5 Ichthyosauria ................................................................................................................ 24 2.2.6 Archosauria .................................................................................................................. 24 2.2.7 Mammalia .................................................................................................................... 37 2.3 Discussion ........................................................................................................................... 57 2.3.1 Methods........................................................................................................................ 58 2.4 Conclusions ......................................................................................................................... 60 Chapter 3: Selection of Exemplar Taxa for the Study of Sexual Dimorphism in Ceratopsids ..... 86 3.1 Introduction ......................................................................................................................... 86 3.1.1 The Merit of Models .................................................................................................... 87 3.1.2 The Selection of Bovids and Pronghorn ...................................................................... 87 iv 3.2 Relevant Characteristics of the Exemplar Taxa .................................................................. 89 3.2.1 The Occurrence and Degree of Sexual Dimorphism in Bovids ................................... 89 3.2.2 Exemplar Taxon 1: Oreamnos americanus ................................................................. 92 3.2.3 Exemplar Taxon 2: Bison bison ................................................................................... 95 3.2.3 Exemplar taxon 3: Antilocapra americana .................................................................. 99 3.3 Conclusions ....................................................................................................................... 105 Chapter 4: Sexual Dimorphism in Bovids .................................................................................. 107 4.1 Introduction ....................................................................................................................... 107 4.1.1 Objectives .................................................................................................................. 108 4.2 Materials and Methods ...................................................................................................... 109 4.2.1 Material ...................................................................................................................... 109 4.2.2 Measurements ............................................................................................................ 110 4.2.3 Ageing .......................................................................................................................
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