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Dental Occlusal Form and Function in Equus Capensis: Evaluating a Controvertible Taxonomic Status

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Dental Occlusal Form and Function in Equus Capensis: Evaluating a Controvertible Taxonomic Status Dental Occlusal Form and Function in Equus capensis: Evaluating a Controvertible Taxonomic Status Megan Malherbe Supervised by Dr. Deano Stynder and Prof. Rebecca Ackermann Town Cape of Dissertation submitted in fulfilment of the requirements for the degree of Master of Philosophy (MPhil) in Archaeology UniversityDepartment of Archaeology University of Cape Town November 2019 The copyright of this thesis vests in the author. No quotation from it or information derived fromTown it is to be published without full acknowledgement of the source. The thesis is to be used for private study or non- commercial research purposes only. Cape Published by the University of Cape Town (UCT) in terms of the non-exclusive licenseof granted to UCT by the author. University Contents Abstract ........................................................................................................................ i Acknowledgements .................................................................................................... iii List of Tables .............................................................................................................. iv List of Figures ............................................................................................................. v List of Appendices ................................................................................................... viii Chapter One: Introduction ....................................................................................... 1 1.1 Uncertainty Around the Taxonomic Status of Equus capensis ............................. 1 1.2 Thesis Outline......................................................................................................... 2 Chapter Two: Background ........................................................................................ 5 2.1 Equid Evolution ...................................................................................................... 5 2.1.1 The Complex 55-Million-Year History .................................................................. 5 2.1.2 Major Trends in Equid Dental Evolution ................................................................ 8 2.1.3 The Origins of the Genus Equus........................................................................... 11 2.1.4 Evaluating Taxonomy in the Fossil Record .......................................................... 12 2.1.5 Issues in Taxonomic Evaluation of Equids ........................................................... 18 2.2 Equus capensis ..................................................................................................... 21 2.2.1 Naming of E. capensis as a New Fossil Equid Species ......................................... 21 2.2.2 Range and Dietary Adaptation ............................................................................. 28 2.2.3 Ancient DNA Studies .......................................................................................... 32 2.3 Global Climate Change Over the Last Five Million Years .................................. 38 2.3.1 Mid-to Late Pleistocene Climatic Fluctuations ..................................................... 38 2.3.2 Effect of the Climatic Shifts on Animal Communities .......................................... 40 2.3.3 Responses of the Equid Family to Pleistocene Climatic and Environmental Shifts..44 2.4 Depositional Contexts of South African Equus Fossils ....................................... 44 2.5 Using Dental Morphology to Investigate the Taxonomic Distinctiveness of E. capensis ...................................................................................................................... 51 2.5.1 Morphometric Approaches for Species Discrimination ....................................... 53 Chapter Three: Materials and Methods ................................................................ 56 3.1 Materials ............................................................................................................... 56 3.2 Methods ................................................................................................................ 58 3.2.1 Linear Measurements ........................................................................................... 61 3.2.2 Geometric Morphometrics – Thin Plate Splines ................................................... 61 3.2.3 Occlusal Enamel Index ........................................................................................ 69 3.2.4 Mesowear ............................................................................................................ 71 Chapter Four: Results.............................................................................................. 78 4.1 Geometric Morphometric Analyses ..................................................................... 78 4.1.1 Relative Warp Analyses ....................................................................................... 78 4.1.2 Centroid Size Analyses ........................................................................................ 85 4.2 Linear Measurements ........................................................................................... 90 4.3 Occlusal Enamel Index ......................................................................................... 92 4.4 Mesowear ............................................................................................................. 94 Chapter Five: Discussion and Conclusions ............................................................ 96 5.1 Complications with the Historical Classification of Equus capensis .................. 98 5.1.1 Lower Dentition ................................................................................................... 99 5.1.2 Upper Dentition ..................................................................................................101 5.2 Equus capensis versus Equus quagga ................................................................ 103 5.2.1 Further Consideration of E. capensis and E. quagga Enamel Patterns .................104 5.2.2 Homogenous Enamel Complexity in E. capensis and E. quagga .........................105 5.2.3 Phenotypic Plasticity in Equus quagga ................................................................106 5.3 Body Size Differences Correlated with Pleistocene Climatic Fluctuations ....... 108 5.3.1 Bergmann’s Rule ................................................................................................109 5.3.2 The Impact of Bergmann’s Rule on Equids .........................................................113 5.3.3 Equus quagga and Bergmann’s Rule in the Pleistocene ......................................115 5.4 Addressing the question of conspecificity ......................................................... 116 5.4.1 Fossil evidence of co-occurrence? .......................................................................117 5.5 Future Research .................................................................................................. 119 5.6 Potential Limitations .......................................................................................... 120 5.6.1 GMM Protocol for Upper Dentition ....................................................................120 5.6.2 No Distinction Between P3 – M2 ........................................................................121 5.6.3 Tooth Size as a Proxy for Body Size ...................................................................121 5.7 Conclusions ........................................................................................................ 122 Reference List ......................................................................................................... 124 Appendices .............................................................................................................. 145 Abstract The Giant Cape zebra Equus capensis is one of the most iconic fossil herbivore species of the South African Quaternary. It is widely believed to have persisted relatively unchanged in its morphology for the duration of its existence, that encompassed the mid-to late Pleistocene. Described as a large-bodied equid, its skeletal remains were originally regarded as distinct and readily distinguishable from those of other fossil equids, especially with respect to their large size. This applied particularly to its cheek teeth, that were described as large with unique occlusal enamel morphology. Palaeontologists have generally accepted the taxonomic status of E. capensis without question. However in reality, its taxonomic status is far from incontrovertible as many of its definitive morphological traits are based on conjecture. Equus capensis was originally described on the basis of a large mandible embedded in limestone that had much of its dental row obscured. The rest of it was reconstructed using unassociated, usually isolated osteological elements from a variety of localities. Perhaps not surprisingly, apart from size, researchers have struggled over the years to distinguish between E. capensis skeletal elements and those of other fossil equids. An ancient DNA study of four E. capensis teeth further compounded the uncertainty around its taxonomic status by exposing a high degree of genetic similarity to E. quagga. The present study represents the first systematic investigation of morphological traits originally used to define E. capensis. Specifically, it considers dental occlusal enamel form and function to i help shed light on the
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