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Copyright by Robert Smith Scott 2004 Copyright by Robert Smith Scott 2004 The Dissertation Committee for Robert Smith Scott certifies that this is the approved version of the following dissertation: The Comparative Paleoecology of Late Miocene Eurasian Hominoids Committee: John Kappelman, Supervisor Claud Bramblett Deborah Overdorff Raymond L. Bernor Liza Shapiro The Comparative Paleoecology of Late Miocene Eurasian Hominoids by Robert Smith Scott, B.A., M.A. Dissertation Presented to the Faculty of the Graduate School of The University of Texas at Austin in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy The University of Texas at Austin August, 2004 Dedication This dissertation is dedicated to my grandmother Alma Dinehart Smith who dedicated her working years to science as a scientific illustrator of considerable skill and who has always been an inspiration to her whole family. Acknowledgements This work was supported by NSF grant BCS-0112659 to J. Kappelman (PI) and R.S. Scott (Co-PI) and a Homer Lindsey Bruce Fellowship from the University of Texas at Austin to R.S. Scott. Thanks to Ray Bernor for the contribution of measurements from his extensive hipparion database funded by NSF grant EAR-0125009. Thanks to Adam Gordon and Dave Raichlen for their many commentaries on and off topic. Additional thanks to my committee John Kappelman, Ray Bernor, Claud Bramblett, Deborah Overdorff, and Liza Shapiro. Much gratitude is due my many hospitable European colleagues who have helped make this work possible. And finally, thanks to my wife Shannon for her patience and support throughout this process. v The Comparative Paleoecology of Late Miocene Eurasian Hominoids Publication No._____________ Robert Smith Scott, Ph.D The University of Texas at Austin, 2004 Supervisor: John Kappelman Abstract: Remains of late Miocene hominoids increasingly indicate both taxonomic and adaptive diversity. Understanding hominoid paleoenvironments may illuminate the selection pressures relevant to this diversity. Previous analyses of bovid ecomorphology have been applied to understanding hominoid environments. In this study, a new analysis of distal and complete bovid metapodials assigned to five habitats was conducted and applied to several fossil sites including Can Llobateres and Y0311 in the Siwaliks. An ecomorphological framework was also developed for hipparion metapodials and applied to many late Miocene fossil sites. The distal and complete discriminant models for bovid metapodials reported here performed between 3.8 and 4.1 times better than chance. Compared to prior discriminant models developed for the bovid femur (Scott et al, 1999; Kappelman, 1988), astragalus (DeGusta and Vrba, 2002) and metapodials (Plummer and Bishop, 1994) all results reported here resulted in more robust values of Press’s Q statistic and better performance when compared to chance. Habitat scores were developed for hipparions based mainly on metapodial length and the medial-lateral dimension of the metapodial diaphysis. These vi scores yielded paleoenvironmental interpretations congruent with those already published for well-sampled sites such as Höwenegg and provide important evidence regarding late Miocene paleoenvironments. In total, the results of this analysis indicate two general patterns relevant to late Miocene hominoids. Sites such as Can Llobateres and Rudabánya are marked by lower hipparion diversity, hipparions adapted to closed habitats, and suspensory, frugivorous hominoids. In contrast, a second pattern is evident for locality 12 of the Sinap Formation and Ravin de la Pluie where a diverse and likely abundant hipparion fauna including open-adapted and closed-adapted forms is associated with pronograde, hard object feeding hominoids. vii Table of Contents List of Tables ....................................................................................................... xvi List of Figures...................................................................................................... xix PART I: INTRODUCTION 1 Chapter 1: Background and Hypotheses..................................................................1 Background.....................................................................................................1 Hypotheses......................................................................................................6 Specific Functional Hypotheses.............................................................7 Paleoenvironmental Hypotheses............................................................7 Hominoids..............................................................................................8 Summary of Goals and General Remarks.......................................................8 PART II: BOVID ECOMORPHOLOGY 14 Chapter 2: Habitat Specific Adaptations of Bovid Metapodials and Late Miocene Hominoid Paleoenvironments.......................................................................14 Introduction...................................................................................................14 Materials and Methods..................................................................................20 Results...........................................................................................................28 Metapodial length ................................................................................28 Metapodial diaphyseal shape ..............................................................29 Distal epiphyses ...................................................................................32 Proximal epiphyses ..............................................................................34 Discriminant function analysis of extant bovids..................................35 Stepwise selection of variables ...................................................35 Complete metatarsal....................................................................36 Complete metacarpal ..................................................................37 Distal metatarsal..........................................................................41 Distal metacarpal ........................................................................43 Analysis of fossil bovids......................................................................45 viii Pikermi........................................................................................45 Ballestar ......................................................................................46 Can Llobateres ............................................................................47 Torrent de Fibulines....................................................................48 Y0496..........................................................................................49 Y0076..........................................................................................49 Y0311..........................................................................................49 Discussion.....................................................................................................51 Reclassification of Extant Bovids........................................................51 Functional Morphology of Extant Bovids ...........................................55 Metapodial-Phalangeal Joint.......................................................55 Metapodial Diaphysis .................................................................57 Metapodial Length......................................................................58 Paleoenvironments...............................................................................58 Pikermi........................................................................................58 Ballestar ......................................................................................61 Can Llobateres ............................................................................61 Torrent de Fibulines....................................................................63 Y0496..........................................................................................63 Y0076..........................................................................................64 Y0311..........................................................................................65 Conclusions...................................................................................................68 PART III: EQUID ECOMORPHOLOGY 129 Chapter 3: Introduction to Equid Ecomorphology...............................................129 Problems and Issues....................................................................................129 Habitat Score...............................................................................................134 Chapter 4: Materials and Methods.......................................................................136 Chapter 5: Extant Equidae ...................................................................................142 Results.........................................................................................................142 Size independent variable computation .............................................142 ix Principal components analysis...........................................................142 Metatarsal III.............................................................................143 Metacarpal III............................................................................143 Habitat score computation .................................................................144
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