Taphonomic and Palaeoecological
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TAPHONOMIC AND PALAEOECOLOGICAL INVESTIGATIONS OF RIVERSLEIGH OLIGO-MIOCENE FOSSIL SITES MAMMALIAN PALAEOCOMMUNITIES AND THEIR HABITATS Mina Bassarova Thesis submitted for the degree of Doctor of Philosophy at the University of New South Wales, Australia August, 2005 PLEASE TYPE THE UNIVERSITY OF NEW SOUTH WALES Thesis/Dissertation Sheet Surname or Family name: Bassarova First name: Mina Other name/s: Plamenova Abbreviation for degree as given in the University calendar: PhD School: BEES Faculty: Science Title: Taphonomic and palaeoecological investigations of Riversleigh Oligo-Miocene fossil sites – mammalian palaeocommunities and their habitats Abstract 350 words maximum: (PLEASE TYPE) The palaeoecology of selected fossil sites from the Riversleigh World Heritage Area, northwestern Queensland, Australia, was studied with the aim of describing the palaeohabitats of the sites through the use of characteristics of mammalian community structure. Taphonomic analyses were carried out to determine whether the study sites represent allochthonous or autochthonous assemblages. Subsequently, ecological attributes of the mammalian fossil assemblages were inferred from functional morphology. Trophic and locomotor behaviours were used to describe the adaptive structure of communities and a method was established for inferring the locomotor behaviour of fossil taxa from morphometrics of their calcanea. Such ecological attributes of the assemblages can be used in reconstructing habitats. This is possible because modern mammalian community structure (as represented by ecological diversity/attribute characteristics) has been found to correlate with habitat structure and thus has predictive value, directly applicable in palaeoecology. Modern mammalian faunas from a variety of habitats around the world were used as possible analogues for the Riversleigh fossil faunas. Multivariate statistical techniques were explored for identifying potential similarities between the community structure of the fossil faunas and that of the modern faunas. Annual rainfall was then estimated for the fossil sites through regression analysis allowing climatic inference from the faunal palaeocommunities. On the basis of similarities in community structure, general habitat or vegetation structure was proposed for the fossil assemblages. The results of the analyses undertaken indicate that Riversleigh early-middle Miocene habitats were densely forested. The late Oligocene Quantum Leap Site local fauna and the late Miocene Encore Site local fauna suggest mixed vegetation, or more open environments. The trend of decreasing annual rainfall through the Miocene and the palaeohabitats of the Miocene sites proposed here fit the general pattern of vegetation and climate change during this period for the Australian continent as a whole. Declaration relating to disposition of project thesis/dissertation I hereby grant to the University of New South Wales or its agents the right to archive and to make available my thesis or dissertation in whole or in part in the University libraries in all forms of media, now or here after known, subject to the provisions of the Copyright Act 1968. I retain all property rights, such as patent rights. I also retain the right to use in future works (such as articles or books) all or part of this thesis or dissertation. I also authorise University Microfilms to use the 350 word abstract of my thesis in Dissertation Abstracts International (this is applicable to doctoral theses only). ……………………………………… …………………………………………………… ……….……………………. Signature Witness Date The University recognises that there may be exceptional circumstances requiring restrictions on copying or conditions on use. Requests for restriction for a period of up to 2 years must be made in writing to the Registrar. Requests for a longer period of restriction may be considered in exceptional circumstances if accompanied by a letter of support from the Supervisor or Head of School. Such requests must be submitted with the thesis/dissertation. FOR OFFICE USE ONLY Date of completion of requirements for Award: Registrar and Deputy Principal THIS SHEET IS TO BE GLUED TO THE INSIDE FRONT COVER OF THE THESIS N:\FLORENCE\ABSTRACT Table of contents 1. Introduction 1 1.1. Background 1 1.2. Aims 3 1.3. The sites 4 1.4. Chapter outline 7 1.5. References 10 2. Taphonomy of Oligo-Miocene fossil sites of the Riversleigh World Heritage Area 13 2.1. Introduction 13 2.1.1. Definitions 13 2.1.2. Time-averaging 14 2.1.3. Methods of quantification 17 2.2. Aims 21 2.3. Methods 21 2.4. Results and discussion 30 2.5. Summary 48 2.6. References 50 Appendix A1. Raw data for percentage of various break shapes on limb bones for all size categories 57 Appendix A2. Raw data for skeletal representation for small, medium and large size categories at the fossil sites studied 58 Appendix A3. Specimens used from Camel Sputum Site for age distribution analysis 59 3. The calcaneum – on the heels of marsupial locomotion 61 3.1. Introduction 61 3.1.1. Locomotor behaviour 61 3.1.2. Aims 63 3.1.3. Why calcanea? 63 3.1.4. Ancestry as a confounding factor 65 3.2. Methods 66 3.2.1. Specimens 66 3.2.2. Measurements 66 3.2.3. Locomotor categories 68 3.2.4. Notes on the modern marsupial sample 69 3.2.5. Statistical analysis 70 3.3. Results 71 3.4. Discussion 81 3.5. Conclusions 85 3.6. References 86 Appendix B1. Calcaneum measurements for modern marsupial species represented by more than one specimen 89 Appendix B2. Raw calcaneum measurements for modern marsupial species represented by single specimens 90 ii Appendix B3. Raw calcaneum measurements for modern marsupial species used as ‘unknowns’ in analysis 91 Appendix B4. Raw calcaneum measurements for fossil marsupials 92 Appendix B5. Normal probability plots for all variables 93 4. The diets of Riversleigh fossil mammals 94 4.1. Introduction 94 4.2. Mammalian cranio-dental functional morphology 95 4.3. Trophic categories 99 4.4. References 105 5. Palaeoecology of Riversleigh fossil sites 109 5.1. Introduction 109 5.1.1. Definitions 109 5.1.2. Aims 111 5.1.3. Background 112 5.2. Methods 113 5.2.1. Consideration of potential bias in fossil community representation 113 5.2.2. Modern communities: localities and habitats 118 5.2.3. Ecological diversity analyses 124 5.2.3.1. Variables used 124 5.2.3.2. Mammal taxa 125 5.2.3.3. Statistical analyses – determining habitat types and annual rainfall for fossil communities 129 5.3. Results 132 5.3.1. Consideration of potential bias in fossil community representation 132 5.3.2. Ecological diversity analyses 138 5.3.2.1.Determining broad habitat types from modern communities 138 5.3.2.2.Grouping fossil communities into broad habitat types 142 5.3.2.3.Determining annual rainfall for fossil communities 150 5.4. Discussion 154 5.4.1. Consideration of potential bias in fossil community representation 154 5.4.2. Habitat descriptions based on modern localities 156 5.4.3. Determining habitat types and annual rainfall for fossil communities 165 5.4.4. The influence of abiotic factors on vegetation structure and animal communities 169 5.5. Conclusions 171 5.6. References 173 Appendix C1. Species lists for Australian sites 181 Appendix C2. Species list for New Guinea locality 182 Appendix C3. Species lists for Central American sites 183 Appendix C4. Species lists for South American sites 189 Appendix C5. Species lists for North and Central American sites 191 iii Appendix C6.1. Trophic and locomotor adaptations for modern marsupial and monotreme species 193 Appendix C6.2. Trophic and locomotor adaptations for modern primate and edentate species from Central and South America 196 Appendix C6.3. Trophic and locomotor adaptations for modern species of Carnivora, Perissodactyla, Artiodactyla and Lagomorphia from North, Central and South America 197 Appendix C6.4. Trophic and locomotor adaptations for modern rodent species, listed by area 199 Appendix C7.1. Split-sample cluster analysis for diet categories for modern sites 203 Appendix C7.2. Split-sample cluster analysis for diet and locomotion categories for modern sites excluding Africa 204 Appendix C8.1. Zskewness values for variables from the ‘Diet Only’ data set 205 Appendix C8.2. Zskewness values for variables from the ‘Diet and Locomotion’ data set 205 6. Australia’s changing climate through the Tertiary: review of different lines of evidence 206 6.1. Climate change 206 6.2. Evidence of climate change 207 6.2.1. Moving continents and changing currents 207 6.2.2. Palynology and plant macrofossils 209 6.2.3. Oxygen isotopes 214 6.2.4. Sea creatures 215 6.2.5. Grossplots 216 6.2.6. Terrestrial faunas 217 6.2.7. Other factors relevant to climate and vegetation 219 6.3. Concluding remarks 221 6.4. References 224 7. Conclusions and further work 228 iv List of figures Figure 1.1.1. Map of Australia showing location of Riversleigh 1 Figure 1.3.1. Quantum Leap Site and BitesantennarySite 5 Figure 1.3.2. Camel Sputum and Mike’s Menagerie Sites 6 Figure 1.3.3. Ringtail Site 6 Figure 1.3.4. Encore Site 6 Figure 2.3.1. Relative abundance of taxa at each site 26 Figure 2.3.2. Size variation at Camel Sputum Site 26 Figure 2.4.1. Percentage of various levels of bone weathering 31 Figure 2.4.2. Percentage of various break shapes on limb bones 33 Figure 2.4.3. Fragmentation of particular elements 34 Figure 2.4.4. Extent of fragmentation for all elements and all size categories combined 36 Figure 2.4.5. Nambaroo sp. 3 from Quantum Leap Site 37 Figure 2.4.6. Turtle carapace from Quantum Leap Site 37 Figure 2.4.7. Skeletal representation 39 Figure 2.4.8. Hydraulic transport potential groupings 39 Figure 2.4.9. Mortality profiles for dasyurids and Perameles gunni 43 Figure 2.4.10. Mortality profiles for kangaroos 45 Figure 2.4.11.