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Systematic and Palaeobiological Implications of Postcranial Morphology in the Diprotodontidae (Marsupialia)

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Systematic and Palaeobiological Implications of Postcranial Morphology in the Diprotodontidae (Marsupialia) Systematic and palaeobiological implications of postcranial morphology in the Diprotodontidae (Marsupialia) Aaron B. Camens School of Earth and Environmental Sciences Discipline of Ecology and Evolutionary Biology The University of Adelaide South Australia A thesis submitted in partial fulfilment of the degree of Doctor of Philosophy at the University of Adelaide February 2010 II Declaration This work contains no material which has been accepted for the award of any other degree or diploma in any university or other tertiary institution to Aaron Camens and, to the best of my knowledge and belief, contains no material previously published or written by another person, except where due reference has been made in the text. I give consent to this copy of my thesis when deposited in the University Library, being made available for loan and photocopying, subject to the provisions of the Copyright Act 1968. The author acknowledges that copyright of published works contained within this thesis (as listed below) resides with the copyright holder(s) of those works. I also give permission for the digital version of my thesis to be made available on the web, via the University’s digital research repository, the Library catalogue, the Australasian Digital Theses Program (ADTP) and also through web search engines, unless permission has been granted by the University to restrict access for a period of time. Publications in this thesis include: Camens, A. B. and Wells, R.T. 2009. Diprotodontid footprints from the Pliocene of Central Australia. Journal of Vertebrate Paleontology 29: 863-869. Copyright held by Taylor and Francis. Camens, A. B. and Wells, R.T. 2010. Palaeobiology of Euowenia grata (Marsupialia: Diprotodontinae) and its presence in Northern South Australia. Journal of Mammalian Evolution 17: 3-19, DOI 10.1007/s10914-009-9121-2. Copyright held by Springer. ………………………… Aaron Camens III Table of Contents CHAPTER 1 Introduction 1 1.1 Introduction to the Diprotodontidae 1 1.2 Systematic relationships in the Diprotodontidae 2 1.2.1 Sub-family incertae sedis 3 1.2.1.1 Nototherium 3 1.2.1.2 Sthenomerus 5 1.2.2 Sub-family Diprotodontinae 5 1.2.2.1 Ngapakaldia 5 1.2.2.2 Pitikantia 6 1.2.2.3 Bematherium 6 1.2.2.4 Pyramios 7 1.2.2.5 Euowenia 7 1.2.2.6 Meniscolophus 8 1.2.2.7 Euryzygoma 8 1.2.2.8 Diarcodon 8 1.2.2.9 Diprotodon 9 1.2.3 Sub-family Zygomaturinae 11 1.2.3.1 Raemeotherium 11 1.2.3.2 Silvabestius 12 1.2.3.3 Alkwertatherium 12 1.2.3.4 Nimbadon 13 1.2.3.5 Neohelos 13 1.2.3.6 Plaisiodon 14 1.2.3.7 Kolopsis 14 1.2.3.8 Kolopsoides 15 1.2.3.9 Maokopia 15 1.2.3.10 Hulitherium 16 1.2.3.11 Zygomaturus 16 1.3 Diprotodontid palaeobiology 18 1.3.1 Sub-family Diprotodontinae 18 1.3.1.1 Ngapakaldia 18 1.3.1.2 Pitikantia 19 1.3.1.3 Pyramios 19 1.3.1.4 Meniscolophus 19 1.3.1.5 Euowenia 19 1.3.1.6 Euryzygoma 19 1.3.1.7 Diprotodon 20 1.3.2 Sub-family Zygomaturinae 23 1.3.2.1 Raemeotherium 23 1.3.2.2 Silvabestius 23 1.3.2.3 Alkwertatherium 23 1.3.2.4 Nimbadon 23 1.3.2.5 Neohelos 23 1.3.2.6 Plaisiodon 24 1.3.2.7 Kolopsis 24 IV 1.3.2.8 Kolopsoides 24 1.3.2.9 Maokopia 24 1.3.2.10 Hulitherium 25 1.3.2.11 Zygomaturus 25 1.4 Thesis aims 27 1.5 References 28 1.6 Appendix 1 35 CHAPTER 2 Camens, A.B., Wells, R.T. (2010). Palaeobiology of Euowenia 37 grata (Marsupialia: Diprotodontinae) and its presence in northern South Australia. Journal of Mammalian Evolution. DOI 10.1007/s10914-009-9121-2. CHAPTER 3 Camens, A.B., Wells, R.T. (2009). Diprotodontid footprints from 55 the Pliocene of Central Australia. Journal of Vertebrate Paleontology 29, 863-869. CHAPTER 4 Camens, A.B., Carey, S.P., Cupper, M.L., Grün, R., Hellstrom, J.C., 63 McKnight, S.W., McLennan, I., Trusler, P. and Aubert, M. (to be submitted) Megamarsupial palaeobiology revealed from fossil trackways of the Victorian Volcanic Plains, Australia. Quaternary Science Reviews. CHAPTER 5 Diprotodontid functional morphology and palaeobiology inferred 116 from postcranial anatomy. 5.1 Introduction 116 5.2 Methods 119 5.2.1 Included taxa 119 5.2.2 Abbreviations 119 5.2.2.1 Institutions 119 5.2.2.2 Hind limb abbreviations 120 5.2.2.3 Fore limb abbreviations 121 5.2.3 Measurements 121 5.3 Results 5.3 Morphological descriptions 122 5.3.1 Subfamily Diprotodontinae 122 5.3.1.1 Ngapakaldia 122 5.3.1.1.1 Morphology of the hindlimb 122 5.3.1.1.2 Morphology of the forelimb 129 5.3.1.2 Euowenia 136 5.3.1.1.2.1 Morphology of the hindlimb 136 5.3.1.1.2.2 Morphology of the forelimb 142 5.3.1.3 Diprotodon 147 5.3.1.3.1 Morphology of the hindlimb 147 5.3.1.3.2 Morphology of the forelimb 155 5.3.2 Subfamily Zygomaturinae 162 5.3.2.1 Nimbadon 162 5.3.2.1.1 Morphology of the hindlimb 162 5.3.2.1.2 Morphology of the forelimb 168 5.3.2.2 Neohelos 176 5.3.2.2.1 Morphology of the hindlimb 176 5.3.2.2.2 Morphology of the forelimb 181 5.3.2.3 Plaisiodon 185 V 5.3.2.3.1 Morphology of the hindlimb 185 5.3.2.3.2 Morphology of the forelimb 191 5.3.2.4 Kolopsis 195 5.3.2.4.1 Morphology of the hindlimb 195 5.3.2.4.2 Morphology of the forelimb 200 5.3.2.5 Zygomaturus 204 5.3.2.5.1 Morphology of the hindlimb 204 5.3.2.5.2 Morphology of the forelimb 212 5.4 Discussion 219 5.4 Functional morphological and palaeobiological interpretations 219 5.4.1 Sub-family Diprotodontinae 221 5.4.1.1 Ngapakaldia 221 5.4.1.1.1 Hindlimb morphological implications 221 5.4.1.1.2 Forelimb morphological implications 223 5.4.1.2 Euowenia 226 5.4.1.2.1 Hindlimb morphological implications 226 5.4.1.2.2 Forelimb morphological implications 227 5.4.1.3 Diprotodon 229 5.4.1.3.1 Hindlimb morphological implications 229 5.4.1.3.2 Forelimb morphological implications 233 5.4.2 Sub-family Zygomaturinae 237 5.4.2.1 Nimbadon 237 5.4.2.1.1 Hindlimb morphological implications 237 5.4.2.1.2 Forelimb morphological implications 239 5.4.2.2 Neohelos 242 5.4.2.2.1 Hindlimb morphological implications 242 5.4.2.2.2 Forelimb morphological implications 244 5.4.2.3 Plaisiodon 246 5.4.2.3.1 Hindlimb morphological implications 246 5.4.2.3.2 Forelimb morphological implications 247 5.4.2.4 Kolopsis 248 5.4.2.4.1 Hindlimb morphological implications 248 5.4.2.4.2 Forelimb morphological implications 250 5.4.2.5 Zygomaturus 252 5.4.2.5.1 Hindlimb morphological implications 252 5.4.2.5.2 Forelimb morphological implications 254 5.5 Conclusion 257 5.6 References 259 5.7 Appendices 269 5.7.1 Appendix 1: List of non-diprotodontid specimens used 269 5.8 Figures 271 CHAPTER 6: A phylogeny of the Diprotodontidae (Marsupialia) based on 295 postcranial morphology. 6.1 Introduction 295 6.1.1 Interfamilial relationships 296 6.1.2 Intrafamilial relationships 299 6.1.3 Postcrania in the Diprotodontidae 302 6.2 Methods 303 VI 6.2.1 Included taxa 303 6.2.2 Institutions 304 6.2.3 Abbreviations 304 6.2.3.1 Hind limb abbreviations 304 6.2.3.2 Fore limb abbreviations 305 6.2.4 Phylogenetic analyses 306 6.2.5 Character description 309 6.2.5.1 Characters relating to the hindlimbs 310 6.2.5.1.1 The Pes 310 6.2.5.1.2 Characters relating to bones of the hind limb 321 6.2.5.2 Forelimb characters used in cladistic analysis 325 6.2.5.2.1 The Manus 325 6.2.5.2.2 Characters relating to the forelimb bones 332 6.3 Results 337 6.3 Phylogenetic analysis 337 6.3.1 Hindlimb character set only 337 6.3.1.1 Intrafamilial analyses 337 6.3.1.2 Interfamilial analyses 343 6.3.1.3 Character states related to functional grouping 354 6.3.2 Forelimb character set only 357 6.3.2.1 Intrafamilial analyses 357 6.3.2.2 Interfamilial analyses 363 6.3.2.3 Character states related to functional grouping 378 6.3.2.4 Character states related to phylogenetic grouping 381 6.3.3 Combined analyses (forelimb and hindlimb) 384 6.3.3.1 Intrafamilial analyses 384 6.3.3.2 Interfamilial analyses 385 6.3.4 Comparison to forelimb-only and hindlimb-only analyses 388 6.4 Discussion 389 6.4 Phylogenetic implications of postcranial anatomy 389 6.4.1 Hindlimb analyses 379 6.4.2 Forelimb analyses 391 6.5 Conclusion 398 6.6 References 399 6.7 Appendices 409 6.7.1 Appendix 1: List of non-diprotodontid specimens used 409 6.7.2 Appendix 2: Character matrix 410 6.7.3 Appendix 3: Supporting characters for various nodes 410 6.8 Figures 421 CHAPTER 7: General Discussion 454 7.1 Thesis summary and aims 454 7.2 Project findings 454 7.2.1 The functional morphology of the hindlimb of E.
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