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NATURAL and ENVIRONMENTAL SCIENCE National Oceanography Centre ` UNIVERSITY OF SOUTHAMPTON NATURAL AND ENVIRONMENTAL SCIENCE National Oceanography Centre Diversity and extinction in Aepyornithidae by James Peter Hansford A dissertation submitted to the University of Southampton in accordance with the requirements for award of the degree of Doctor of Philosophy in the Department of Ocean and Earth Science September 2017 ` ` UNIVERISTY OF SOUTHAMPTON ABSTRACT NATURAL AND ENVIRONMENTAL SCIENCE Ocean and Earth Science Thesis for the degree of Doctor of Philosophy DIVERSITY AND EXTINCTION IN AEPYORNITHIDAE by James Peter Hansford The Aepyornithidae are an insular radiation of giant birds from the late Quaternary of Madagascar which have been extinct for c. 1000 years. Complex and conflicting historical taxonomic hypotheses have limited study into these charismatic megafauna and they have been subject to little modern study in comparison to other avian megafauna and the mammalian megafauna of Madagascar. This thesis is the first modern study of Aepyornithidae to quantify the diversity and biogeography of skeletal remains in comparison to putative taxonomic hypotheses. Clarifying the convoluted history of historically proposed taxa underpins a modern framework for studying these enigmatic birds. A novel chronological sequence of high-quality radiocarbon dates provides the most reliable evidence for their species-specific extinction timings. Late survival of these birds contrasts markedly with rapid extinction processes observed in avian megafauna and presents an extended period of co-occurrence with human colonists suggesting complex and poorly understood interactions. Recording and dating evidence of butchery to modern osteological standards has provided unique and extraordinary evidence of human settlement 6000 years earlier than any other evidence suggests, and is the first verifiable information on direct impacts of butchery and hunting of Aepyornithidae. Dietary analysis demonstrates that these morphologically diverse species fulfilled different ecological niches, including extensive grazing behaviour in the central highlands. This promotes the need for new discussions into the pristine landscape of Madagascar, including more expansive investigations into aepyornithid ecosystem functions and the pre-human distribution of grassland savannah versus forested regions. It is hoped that this thesis will lead to novel research into Aepyornithidae and develop understanding of their role in defining the natural state of one of the worlds most threatened ecosystems. ` ` Table of contents Table of contents I List of tables IV List of figures VI List of accompanying materials VIII DECLARATION OF AUTHORSHIP IX Acknowledgements X Institutional abbreviations XII Chapter one: Introduction 1.1 Late Quaternary of Madagascar 3 1.2 Madagascar’s terrestrial avian megafauna: The elephant birds 7 1.2.1 Scientific discovery 7 1.2.2 Taxonomic revisions and confusion 9 1.3 19th and 20th century taxonomy of Aepyornithdae 11 1.4 The biogeography of Aepyornithdae 28 1.5 Co-occurrence and conflict with humans 29 1.5.1 Sinbad’s Rukh: Cryptic evidence for Madagascar’s avian megafauna? 31 1.6 Thesis aims 33 Chapter two: Unexpected diversity within the extinct elephant birds (Aves: Aepyornithidae), and a new identity for the world’s largest bird 2.1 Abstract 37 2.2 Introduction 38 2.2.1 History of research on elephant birds 39 2.2.2 Towards a modern morphometric framework for elephant bird taxonomy 41 2.3 Methods 44 2.3.1 Specimens and measurements 44 2.3.2 Missing data imputation 44 2.3.3 Cluster analysis 45 2.4 Results 48 2.4.1 Morphometric analysis 48 2.4.2 Taxonomy of morphometric clusters 52 2.4.3 Biogeography of Aepyornithidae 55 I ` 2.5 Systematic palaeontology 58 2.6 Discussion 74 Chapter three: Early Holocene human presence in Madagascar evidenced by exploitation of avian megafauna 3.1 Abstract 81 3.2 Introduction 82 3.3 Methods 84 3.4 Results 86 3.5 Discussion 91 Chapter four: Extinction chronology of the elephant birds 4.1 Abstract 95 4.2 Introduction 96 4.3 Methods 102 4.4 Results 104 4.5 Discussion 111 4.5.1 Aepyornis hildebrandti and the central highlands 114 4.5.2 Mullerornis modestus 115 4.5.3 Vorombe titan 115 4.5.4 Thick eggshell 115 4.5.5 Extirpation in the south and south west 116 4.5.6 The “Voroun patra” 117 4.7 Conclusion 118 Chapter five: Reconstructing the dietary ecology of aepyornithids using stable isotope analysis 5.1 Abstract 122 5.2 Introduction 123 5.2.1 Late Quaternary environmental history of Madagascar 123 5.2.2 Stable carbon isotopes and diet 125 5.3 Methods 127 5.4 Results 129 5.6 Discussion 133 II ` Conclusion 140 References 143 Appendix one 158 Appendix two 186 Appendix three 197 Appendix four 203 III ` List of tables Chapter one Table 1: Saint-Hiliarie’s 1851 egg measurements 11 Table 2: Owen’s 1852 measurements of Aepyornis maximus tarsometatarsus vs. other ratites 11 Table 3: Milne-Edwards and Grandidier’s femora measurements for A. medius, A. modestus 12 and A. maximus type specimens Table 4: Burckhardt’s A. hildebrandti hind-limb bone measurements 13 Table 5: Burckhardt’s Aepyornis comparative femora measurements 13 Table 6: Burckhardt’s Aepyornis comparative Tibiotarsus measurements 13 Table 7: Andrews’ Aepyornis comparative Tibiotarsus measurements 14 Table 8: Andrews’ Aepyornis comparative femora measurements 14 Table 9: Milne-Edwards and Grandidier’s A. ingens hind-limb bone measurements 15 Table 10: Milne-Edwards and Grandidier’s A. cursor tarsometatarsus measurements 15 Table 11: Milne-Edwards and Grandidier’s A. lentus tarsometatarsus measurements 16 Table 12: Milne-Edwards and Grandidier’s M. betsilei hind-limb bone measurements 17 Table 13: Milne-Edwards and Grandidier’s M. agilis Tibiotarsus measurements 17 Table 14: Milne-Edwards and Grandidier’s M. rudis Tibiotarsus measurements 18 Table 15: Andrews’ F. rudis Tibiotarsus measurements 18 Table 16: Cranial measurements of Aepyornis species made by Monnier 21 Table 17: Tibiotarsi measurements of Aepyornis species made by Monnier 22 Table 18: Tarsometatarsi measurements of Aepyornis species made by Monnier 22 Table 19: Femora measurements of Aepyornis species made by Monnier 23 Table 20: Lambrecht’s comparative measurements of Aepyornithidae 25 Table 21: Lamberton’s comparative femora measurements of aepyornithids 26 Table 22: Lamberton’s comparative tibiotarsi measurements of aepyornithids 26 Chapter two Table 1: Historically proposed species of elephant birds. Note: M. rudis was subsequently 42 designated as the type species of Flacourtia by Andrews (1895) Table 2: Taxonomic matrix describing morphometric assignment and taxonomic seniority of 56 type material of named elephant bird species to the morphometric clusters identified in this study Table 3: Mass estimates for elephant bird species recognised in this study 70 Table 4: Femoral measurement ranges for elephant bird species recognised in this study 70 IV ` Table 5: Tibiotarsal measurements (in mm) for elephant bird species recognised in this study 71 Table 6: Tarsometatarsal measurement ranges for elephant bird species recognised in this 72 study Chapter three Table 1: Dimensions of tool marks on Aepyornis maximus, USNM A605209 87 Table 2: List of newly recognized elephant bird bones with perimortem anthropogenic 89 modification and AMS radiocarbon dates Chapter four Table 1: Rejected radiocarbon dates 103 Table 2: Aepyornis hildebrandti radiocarbon dates 104 Table 3: Mullerornis modestus radiocarbon dates 105 Table 4: Vorombe titan radiocarbon dates 106 Table 5: Radiocarbon dates from thick eggshell 108 Table 6: GRIWM extinction estimates for elephant birds 110 Chapter five Table 1: Mean isotope values and dietary proportion estimates from ISOERROR 1.04 131 Table 2: Tukey’s pairwise results for 훿13C ‰ in Madagascar’s megaherbivores 131 Table 3: Tukey’s pairwise results: Observed 훿13C isotopes for Mullerornis skeletal elements 132 and eggshells Table 4: Tukey’s pairwise results: Adjusted 훿13C isotopes for Mullerornis skeletal elements and 132 eggshells V ` List of figures Chapter one Figure 1: Ecological zones and topography of Madagascar 3 Figure 2: Aepyornithidae skeletons from the Queens palace, Antananarivo, Madagascar 10 Figure 3: Photographs of Aepyornis crania From Monnier 1913 21 Figure 4: Monnier’s sketches of femoral morphotypes of Aepyornis 23 Figure 5: Lamberton’s Aepyornis hildebrandti skull 24 Figure 6: Photographs of examples from Lamberton’s comparative material 27 Chapter two Figure 1: Diagram of linear measurements taken on aepyornithid leg bones 46 Figure 2: Femora clusters 48 Figure 3: Tibiotarsi clusters 49 Figure 4: Tarsometatarsi clusters 50 Figure 5: Distribution across Madagascar of identified specimens of elephant bird species 55 Figure 6: Newly designated type series of Aepyornis maximus. 60 Figure 7: Vorombe titan, lectotype (femur, NHM A439), Itampulu Vé, Madagascar. 68 Figure 8: Vorombe titan, Tibiotarsus (NHM A437), Itampulu Vé, Madagascar; part of original 69 syntype series Chapter three Figure 1: Sites containing butchered elephant bird bones and AMS radiocarbon dates 85 Figure 2: Aepyornis maximus skeletal reconstruction 87 Figure 3: Tool marks on Aepyornis maximus tarsometatarsus, USNM A605209 89 Figure 4: Tool marks on Aepyornis maximus Tibiotarsus, USNM A605208 90 Chapter four Figure 1: Collection localities across Madagascar of radiocarbon dated specimens in this study 101 Figure 2: Calibrated radiocarbon dates for Aepyornis hildebrandti skeletal remains 105 Figure 3: Calibrated radiocarbon dates for Mullerornis modestus
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