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A Geometric Morphometric Assessment of the Foot of Oreopithecus

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A Geometric Morphometric Assessment of the Foot of Oreopithecus A geometric morphometric assessment of the foot of Oreopithecus By Kyle Billington A thesis submitted in partial fulfilment of the requirements for the degree of Doctor of Philosophy The University of Sheffield Department of Archaeology July 2016 In loving memory of Dave Lloyd Acknowledgements There are a great number of people I owe a huge amount of gratitude to and unfortunately this list will not be complete. Omission from this list in no way constitutes a lack of gratitude on my part. Thanks are owed, in the first place, to the people who made this work possible: The University of Sheffield and the Department of Archaeology made the project financially viable and provided a place in which to produce the work; Roberto Portela-Miguez of the Natural History Museum, London, for arranging access to the primate collections there and being extremely accommodating to my needs; Loïc Costeur of the Naturhistorisches Museum, Basel, for granting access to the Oreopithecus remains under his care; Marcia Ponce de Léon for arranging permissions for me to access the primate collections and fossil casts at the University of Zürich; Angela Gill for granting access to the primate remains at the Powell-Cotton Museum. I would like to thank Professor Paul Pettitt for convincing me to continue with this work the first time I considered not completing the project. He showed confidence in me which I didn’t have in myself and was a source of great encouragement at a difficult time. Subsequently he offered valuable comments on my work which was outside his area of expertise and beyond his responsibilities. I am also grateful to Doctor Pia Nyström for assuming the role of second supervisor following a change of personnel within the department. A number of students were added to those already under her supervision and she managed to find the time to offer comments and advice when needed despite her incredibly busy schedule. The prompt help and expertise of Paul O’Higgins in making the necessary corrections to the thesis has been invaluable and I am indebted to him for this. Umberto Albarella also provided invaluable comments and Along with Paul made the viva a thoroughly enjoyable experience. My friends and family have been a constant source of encouragement and inspiration to me and it is to them that I offer the most gratitude. Nichola Austen has been incredibly supportive throughout the project and was instrumental in helping me to arrange my trip to v Switzerland to collect data for the project, huge thanks are owed to her and I wish her all the best with her own PhD. Ed has provided a constant supply of laughs even at times when my mood was low. Other friends; James, Rachael, Ben, Mike, Toby, Rich, Mark, Katie, and many more friends whose inclusion is restricted by space, have provided often needed breaks and distractions to help refocus and remind me that there is a life outside of this project. Very special thanks go to J and Isabelle, two incredible friends and colleagues to whom I often turned to vent frustrations, seek advice, and find unswerving support for the duration of my student career. Noémie is deserving of my deepest and sincerest love and thanks. It is thanks to her that I found the confidence and strength to continue on from the second time I seriously considered not completing this project. She has given me the belief that completing this thesis is a worthwhile undertaking regardless of what follows. Because of her I am confident that whatever the outcome I will ultimately be happy and my future is bright. None of this would have been possible without the support of my family. My aunts, uncles and cousins, although we’re together only rarely, never fail to lift my spirits and make me laugh. My parents and brother have constantly believed in me and offered their love and support at every step of the process and throughout my life to this point; I could never do enough to repay their generosity. My grandparents have been particularly helpful throughout my higher education and always advised me to do what makes me happy. It is with their wise words that I turn to the next chapter of my life confident in the knowledge that I couldn’t wish for better friends and family and that whatever the future holds I will always be looked after well. vi Abstract Oreopithecus is an enigmatic primate from the Miocene of Italy. It has been the subject of more than a century of controversial research and debate. Most recently, the claim that Oreopithecus exhibited a substantial degree of bipedal behaviour has permeated the literature. Specifically, the pedal anatomy of Oreopithecus has been suggested to be unique among the hominoids and this has been advanced as evidence for bipedal adaptations in this taxon. The possibility that Oreopithecus was a bipedal ape is examined using geometric morphometric techniques to assess the shape of Oreopithecus pedal remains in comparison to other well-known species, and functional interpretations are drawn from these results. This study has examined the medial column of the pedal skeleton of five extant primate taxa, as well as that of Oreopithecus, Homo habilis, and Nacholapithecus. The possible function of the foot is considered in the context of published information regarding the rest of the postcranial skeleton of Oreopithecus and the known positional behaviour of the extant species used in the study. It is found that Oreopithecus closely resembles the African ape condition in the shape of its pedal skeleton, though there are subtle differences; however, none of the differences found in the pedal skeleton of Oreopithecus offer support to the contention that the foot was especially well-adapted to bipedal behaviour. The morphology of the medial cuneiform suggests that the degree of abduction of the hallux was comparable to that observed in Pan. Similarly, the morphology of the navicular, and the lateral and intermediate cuneiforms, indicate that the orientations of the articulations of the midfoot had a configuration that was more or less the same as that observed for extant African apes, particularly Pan. It is therefore concluded that Oreopithecus was probably not habitually bipedal. But the results presented here do not in and of themselves preclude bipedalism from its locomotory repertoire, as bipedal behaviour is exhibited among the extant apes to which it is similar. However, the finding that the foot of Oreopithecus was significantly smaller than it is in any extant ape casts doubt on the likelihood of any significant level of bipedalism and may indicate that Oreopithecus was adapted to a forelimb dominated locomotory strategy. vii Contents Acknowledgements .................................................................................................................................. v Abstract .................................................................................................................................................. vii List of Figures .......................................................................................................................................... xii List of Tables ........................................................................................................................................... xx 1. Introduction ..................................................................................................................................... 1 1.2. Background to the topic .......................................................................................................... 1 1.3. Structure and aims of the thesis .............................................................................................. 4 1.4. Oreopithecus ............................................................................................................................ 5 1.5. Overview and evolution of Hominoidea .................................................................................. 9 1.5.1. Extant Hominoidea .......................................................................................................... 9 1.5.2. Miocene fossil Hominoidea ........................................................................................... 13 1.5.3. Plio-Pleistocene Hominoidea ......................................................................................... 17 1.6. Functional anatomy of the foot ............................................................................................. 21 1.6.1. Functional anatomy of the human foot ......................................................................... 21 1.6.2. Comparative functional anatomy of the primate foot ................................................... 28 1.7. Positional behaviour of primates in this study ....................................................................... 31 1.8. Specific hypotheses tested in this study ................................................................................ 33 2. Materials and Methods ................................................................................................................. 35 2.1. Materials ..................................................................................................................................... 35 2.2. Methods ................................................................................................................................ 39 2.2.1. Choice of geometric morphometrics ............................................................................. 39 2.2.2. Structured
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