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Body Size Evolution in Leptomeryx and Rhinocerotinae

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Body Size Evolution in Leptomeryx and Rhinocerotinae BODY SIZE EVOLUTION IN LEPTOMERYX AND RHINOCEROTINAE (SUBHYRACODON AND TRIGONIAS) ACROSS THE EOCENE – OLIGOCENE (CHADRONIAN – ORELLAN) BOUNDARY by WENDY A. SCHULTZ B. S. University of Nebraska – Lincoln, 2006 A thesis submitted to the Faculty of the Graduate School of the University of Colorado in partial fulfillment of the requirement of the degree of Master of Science Museum and Field Studies 2009 UMI Number: 1473682 All rights reserved INFORMATION TO ALL USERS The quality of this reproduction is dependent upon the quality of the copy submitted. In the unlikely event that the author did not send a complete manuscript and there are missing pages, these will be noted. Also, if material had to be removed, a note will indicate the deletion. UMI 1473682 Copyright 2010 by ProQuest LLC. All rights reserved. This edition of the work is protected against unauthorized copying under Title 17, United States Code. ProQuest LLC 789 East Eisenhower Parkway P.O. Box 1346 Ann Arbor, MI 48106-1346 This thesis entitled: Body size Evolution in Leptomeryx and Rhinocerotinae (Subhyracodon and Trigonias) Across the Eocene – Oligocene (Chadronian – Orellan) boundary Written by Wendy A. Schultz has been approved for the Department of Museum and Field Studies _____________________________________ Dr. Jaelyn Eberle, Committee Chair ____________________________________ Dr. Richard Stucky, Committee Member ____________________________________ Dr. Christy McCain, Committee Member Date ______________ The final copy of this thesis has been examined by the signatories, and we find that both the content and the form meet acceptable presentation standards of scholarly work in the above mentioned discipline iii Schultz, Wendy A. (M.S., Museum and Field Studies [Vertebrate Paleontology]) Body size Evolution in Leptomeryx and Rhinocerotinae (Subhyracodon and Trigonias) Across the Eocene – Oligocene (Chadronian – Orellan) boundary Thesis directed by Assistant Professor Jaelyn J. Eberle Research has shown that there was dramatic climate change, specifically a significant drop in temperature, across the Eocene – Oligocene (Chadronian – Orellan) boundary. However, few studies have looked at the effects of this climatic cooling on the terrestrial vertebrate fauna. My study focused on changes in mammalian body size across the Chadronian – Orellan transition, in particular within the small artiodactyl Leptomeryx and the large rhinocerotine perissodactyls, based upon fossils from the White River Group in the Northern Plains (Colorado, Wyoming, Nebraska, and South Dakota). Specifically, I tested whether Bergmann’s Rule (which suggests that in a given taxon, those individuals inhabiting colder climates will be larger-bodied than those living in a warmer climate) occurred with climatic cooling across the Chadronian-Orellan boundary. With regard to the rhinocerotines, I used Trigonias, Subhyracodon, and specimens identified as Rhinocerotinae gen. indet. Rhinocerotineae gen.indet. specimens are missing their anterior dentition (which is what differentiates teeth of Trigonias from Subhyracodon), and consequently were simply identified only as Rhinocerotinae genus indet. Length and width measurements of upper and lower molars were used as a proxy for body size of both Chadronian and Orellan representatives of two lineages of Leptomeryx, (L. speciosus – L. evansi lineage and L. yoderi – L. mammifer – L. exilis lineage). The two lineages of iv Leptomeryx showed a statistically significant decrease in tooth size in the Orellan. The Rhinocerotinae show a significant decrease in tooth size across the Chadronian – Orellan transition. Although other researchers have noted that only one Leptomeryx lineage survived the Eocene-Oligocene transition, this research suggests that both lineages survived this boundary. The decrease in body size with decrease in temperature in Leptomeryx and Rhinocerotinae is opposite of the expected pattern of Bergmann’s Rule. My data suggest that other factors, besides climatic cooling, also need to be considered in mammalian body size evolution across the Eocene-Oligocene boundary. v Acknowledgments I would like to thank my committee members, Richard Stucky and Christy McCain, and my committee chair, Jaelyn Eberle for always being willing to meet with me when I had questions or concerns about localities or my data. Additional thanks to Jaelyn for her guidance and editing through the course of this thesis project. Thank you to the collections managers and curators at the four museums I visited. Bill Simpson allowed me to conduct research while at the Field Museum of Natural History for my internship, and Olivier Rieppel for authorized a loan of Leptomeryx specimens. Sally Shelton gave me access to the collections at South Dakota School of Mines and Technology, and I thank Ed Walsh for his help in the collection and for showing me around town. Logan Ivy at Denver Museum of Nature & Science helped in collections, specifically in tracking down specimens and accessory information, and Richard Stucky granted access to the collections. I thank Toni Culver at the Colorado Museum of Natural History, for not only lending me undergraduates to help find specimens in the collections, but also for answering many questions I had about our collection database. Extra thanks to Toni for being an awesome boss by allowing me a very flexible work schedule so I could work around my thesis. This project could not have been completed without my fiancé, Nathan Beins, who listened to me when I would talk about my research and writing (even if he didn’t understand anything of what I was saying), and put up with all of my complaints during the whole process. Thank you to my friend Nick Walker for discussing statistics and math, and to Hannah Johnson for letting me borrow him. Thank you so much to Hobbes Hayden for helping me put Adobe Photoshop on both of my computers, vi teaching me how to use it, being my tech support when I couldn’t remember what he taught me, and for allowing me to work on my thesis at his place when I needed a change in scenery. A huge thank you to my office mates, Ben Burger, Alex Dutchak, and Laura Wilson for answering my many questions and helping me along this process. Especially a huge thank you to Laura who is the best office mate, co-worker, and good friend a graduate student could ask for. She was always there, even at midnight, to answer my many, many questions, help me with phrasing, and proofread parts of my thesis before I sent them to Jaelyn, even when we were in different states and she was out in the field. She also made sure I still took time to relax and didn’t get too stressed out. Project and research funding came from the University of Colorado Museum of Natural History. Funding for my research trip to the Field Museum in Chicago was provided by the Walker Van Riper fund. Funding for my research trip to South Dakota School of Mines and Technology came from the University of Colorado Museum of Natural History Paleontology Section. vii TABLE OF CONTENTS I. Introduction……………………………………………………………..………….1 1. The EOT Cooling Event………………..………………………………………...5 2. Bergmann’s Rule……………………………………………………………......…7 3. Background to Biostratigraphy………………………………………....…15 4. Sedimentology and Stratigraphy ……………………………………..…..19 a. Nebraska……………………………………………………………………22 b. South Dakota…………………………………………………………...…23 c. Wyoming…………………………………………………………...………24 d. Colorado………………………………………………………...…….……25 5. Mammalian Taxa…………………………………………………..……………25 a. Rhinocerotinae…………………………………………………………...26 i. Subhyracodon………………………………………………….……..….….27 ii. Trigonias………………………………………………………………..……28 b. Leptomeryx……………………………………………………………..….29 II. Materials and Methods……………………………………………………….33 1. Age Categories……………………………………………………………..……..41 2. Methods of Estimating Body Mass…………………………….…………..43 3. Statistical Methods……………………………………………………...………45 III. Results……………………………………………………………………...……….47 1. Leptomeryx……………………………………………………………….….…….47 2. Rhinocerotinae…………………………………………………….……….…….55 IV. Discussion and Conclusions………………………………………….…….64 1. Future Research…………………………………………………………...……..69 V. Bibliography………………………………………………………………………71 VI. Appendix 1: Statistical analysis of dental elements of all adult Leptomeryx used in this study ……………………………………………………………81 VII. Appendix 2………………………………………………………...………………82 1. Table 1: Measurements of Leptomeryx lower dentition……………………..82 2. Table 2: Measurements of Leptomeryx upper dentition…………………….94 VIII. Appendix 3……………………………………………………………………....101 1. Table 1: Measurements of Rhinocerotinae lower dentition……………….101 2. Table 2: Measurements of Rhinocerotinae upper dentition…………...……..107 viii LIST OF TABLES Table 1: Comparison of the right and left sides of the skull and jaws in all Leptomeryx used in this study……………………………………………………………...47 Table 2: Statistical analysis of dental elements of L. speciosus – L. evansi used in this study…………………………………………………………………………..48 Table 3: Statistical analysis of dental elements of L. mammifer – L. exilis lineage used in this study ……………………………………………………………………..49 Table 4: Statistical analysis of dental elements of Colorado Leptomeryx used in this study ………………………………………………………………………………..51 Table 5: Statistical analysis of dental elements of non-Colorado Leptomeryx used in this study ……………………………………………………………………….....52 Table 6: Univariate body mass estimates of Leptomeryx based on m2 length………...…55 Table 7: Statistical analysis of dental elements of Rhinocerotinae used in this study…...57 Table 8: Statistical analysis of dental
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