Stable Isotope and Trace Element Paleoecology of the Rudabánya II Fauna: Paleoenvironmental Implications for the Late Miocene Hominoid, Rudapithecus hungaricus by Laura Campbell Eastham A thesis submitted in conformity with the requirements for the degree of Doctor of Philosophy Graduate Department of Anthropology University of Toronto © Copyright by Laura C. Eastham 2017 Stable Isotope and Trace Element Paleoecology of the Rudabánya II Fauna: Paleoenvironmental Implications for the Late Miocene Hominoid, Rudapithecus hungaricus Laura C. Eastham Doctor of Philosophy Graduate Department of Anthropology University of Toronto 2017 Abstract The Late Miocene extinction of great apes in Europe has generally been regarded as the consequence of environmental changes that occurred in correlation with global Late Miocene cooling. However, given the range of dietary and locomotor adaptations observed among the different hominoid genera it is unlikely that the same environmental factors can account for the decline of the entire group. To better understand the factors that influenced the extinction of European Miocene apes it is necessary to evaluate their paleoecology on a regional scale. This research utilizes stable carbon and oxygen isotope (δ13C and δ18O) and strontium/calcium (Sr/Ca) trace element ratios measured in fossil ungulate tooth enamel to reconstruct the paleoecology and paleoclimate of Rudabánya II (R. II), an early Late Miocene (~10 Ma) hominoid locality in northeastern Hungary. The fossiliferous deposits at R. II preserve abundant samples of the extinct great ape Rudapithecus hungaricus. Primary aims of this research include: 1) evaluating the types of habitats present in terms of forest canopy cover, 2) examining trophic niche dynamics among the diverse ungulate community, and 3) estimating key climatic variables including mean annual temperature (MAT), mean annual precipitation (MAP), and degree of seasonality. Stable isotope and Sr/Ca values indicate the presence of a heterogeneous wetland- ii forest environment, with a gradient of more open to closed canopy habitats. Significant differences in stable isotope and Sr/Ca values were observed among the sampled ungulate fauna, supporting the interpretation of resource specialization and partitioning. A MAP of 1030-1333 mm/yr, and a MAT of 14°C were calculated from the average δ13C and δ18O values of the equid Hippotherium intrans. Intra-tooth δ18O values revealed low amplitudes of variation indicating that seasonal changes in temperature and precipitation were relatively mild. The paleoenvironment that Rudapithecus inhabited was similar to that of contemporaneous hominoids in Western Europe, but strikingly different from that of hominoids in the Eastern Meditterranean. While adaptations to fallback feeding and efficient suspensory arboreality would have allowed Rudapithecus to endure some degree of environmental deterioration, the progressive restriction and fragmentation of humid wetland-forests following the retreat of Lake Pannon would have eventually led to its extinction. iii Acknowledgments I am incredibly grateful to the many people who supported me over the course of my doctoral research. First and foremost, I recognize my supervisor, David Begun, for his tremendous guidance throughout my time at the University of Toronto. I am deeply appreciative of the freedom I was given to develop my thoughts, as well as the continued encouragement and patience while I worked through them. Thank you, David. The DRB abides. I would like to thank Robert Feranec for being a wonderful mentor and friend. You have consistently encouraged me to maintain high expectations of my own abilities. I have learned a great deal through your guidance, expertise, and passion for science. It’s not often that you email someone asking about their paper and they reply offering to teach you all of the geochemical methods required for your doctoral research. I hope to be able to do that for a student in the future. You have shown me the importance of not only being a good scientist, but also a good person. I must also thank my core committee members, Jochen Halfar and Shawn Lehman. Your thoughtful and constructive comments throughout my doctoral program have improved my dissertation each step of the way. Further, your diverse expertise and insights have greatly broadened the scope of my research. I am thankful for the comments and support provided by my internal examiner, Mary Silcox. Your critical appraisal of my work and guidance on future research and career path has been extremely helpful. I also thank my external examiner, Fred Longstaffe, whose exceptionally insightful and encouraging appraisal of my dissertation has reinforced my belief in my own research and enthusiasm for stable isotope paleoecology. I’ll never look at clouds the same way, Fred. I would like to express my sincere gratitude to the entire faculty and staff of the Anthropology Department at U of T. In particular, I would like to acknowledge Bence Viola, Heather Miller, Michael Schillaci, Sherry Fukuzawa, Trevor Orchard, Natalia Krencil, Josie Alaimo, and Sophia Cottrell. I am grateful for my home away from home in the School of the Environment at U of T. My time spent teaching and conducting research at the School has profoundly influenced the way I perceive the human-environment relationship. I thank David Powell, Sarah Finklestein, and Kim Strong for their support. I will never have the words to thank Douglas Macdonald for all he has iv done. Our research has changed the way I look at paleoecology and human evolution. Your genuine concern for the environment and interest in effecting positive change in the world will always inspire me. Thank you so much for your friendship and guidance. I would like to acknowledge the sources of funding for my dissertation: Ontario Graduate Scholarship, General Motors Women in Science and Mathematics Award, Geological Society of America Graduate Research Award, University of Toronto Fellowship, University of Toronto Department of Anthropology Pilot Dissertation Fund, University of Toronto School of Graduate Studies Travel Grant, and University of Toronto Department of Anthropology Doctoral Completion Award. Without this support, I would not have been able to do this research. So many amazing friends have seen me through the ups and downs of graduate school. I would like to thank the following people who have made this journey one to remember: Karyne Nancy Rabey (Mentor), Stephanie Kozakowski (Kozzel), Amber MacKenzie (Unicorn), Peter Bikoulis (Peta Joules), Joel Cahn (HurriCahn), Lelia Watamaniuk, Sarah Ranlett, Anastasia Hervas, Travis Steffens, Keriann McGoogan, Achinie Wijesinghe, Alana Peters, Cooper Campbell, and Nicole Delaney. I would also like to thank past and present members of the Begun Lab for their support and feedback on my work throughout my time at U of T. I am grateful for (Big) Dave Bovee whose humour, geological know-how, and cowboy coffee always made fieldwork highly excellent. I sincerely thank Dave Boutilier for putting up with the Miocene and me for this long. You have helped me far more than you know. I am so thankful that you are in my life. Finally, thank you to my mother, Anne Eastham, whose unwavering support, love, and sense of humour has kept me going throughout the years. You have always been my number one supporter. Without you none of this would have been possible. You’re the best AnneHam! v I dedicate this dissertation to my friend and mentor, Douglas Macdonald, who taught me to look beyond the fossils. vi Table of Contents 1 Context and Objectives……………………………………………………………………1 1.1 Late Miocene Environmental Change and the Extinction of Hominoids in Europe………….………………………………………………………………….1 1.2 Objectives and Organization of this Dissertation…………………………………6 References…………………………………………………………………………………9 2 Stable Isotopes Show Resource Partitioning Among the early Late Miocene Herbivore Community at Rudabánya II: Paleoenvironmental Implications for the Hominoid, Rudapithecus hungaricus…………………………………………………….21 1 Introduction………………………………………………………………………22 2 Background………………………………………………………………………25 2.1 Rudabánya: Geology, Paleontology, and Paleoecology…………………25 2.2 Stable Carbon Isotope Values in Mammalian Enamel…………………..27 2.3 Stable Oxygen Isotope Values in Mammalian Enamel……….…………29 3 Materials and Methods………………………….………………………………..30 4 Results……………………………………………………………………………32 4.1 Stable Carbon Isotope Values……………………………………………32 4.2 Stable Oxygen Isotope Values…………………………………………...32 5 Discussion………………………………………………………………………..33 5.1 Mammalian Paleoecology: Resource Use and Partitioning……………...33 5.1.1 Mammalian Paleoecology in Open Canopy Habitats……………………34 vii 5.1.2 Mammalian Paleoecology in Closed Canopy Habitats…………………..38 5.2 The Paleoenvironment of Rudapithecus hungaricus…………………….39 5.3 Comparative Paleoecology of Late Miocene Hominoids in Western Eurasia……………………………………………………………………42 6 Conclusions………………………………………………………………………44 References……………………………………………………………………..…45 3 Trace Element Analysis Provides Insight into the Diets of early Late Miocene Ungulates from the Rudabánya II Locality (Hungary)…………………………………..68 1 Introduction………………………………………………………………………69 2 Geological Setting………………………………………………………………..71 3 Materials and Methods…………………………………………………………...73 4 Results……………………………………………………………………………76 5 Discussion………………………………………………………………………..77 6 Conclusions………………………………………………………………………81 References…………..……………………………………………………………82 4 Paleoclimate of the early Late Miocene Rudabánya