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Environmental and Geological Controls on the Diversity and Distribution of the Sauropodomorpha

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Environmental and Geological Controls on the Diversity and Distribution of the Sauropodomorpha 1 ENVIRONMENTAL AND GEOLOGICAL CONTROLS ON THE DIVERSITY AND DISTRIBUTION OF THE SAUROPODOMORPHA PHILIP DAVID MANNION UNIVERSITY COLLEGE LONDON PHD IN PALAEOBIOLOGY 2 I, Philip David Mannion, confirm that the work presented in this thesis is my own. Where information has been derived from other sources, I confirm that this has been indicated in the thesis. 3 ACKNOWLEDGMENTS I would like to express my greatest thanks to my two thesis supervisors, Paul Upchurch (UCL) and Paul M. Barrett (NHM, London). They have helped me throughout my PhD, offering me advice and assistance whenever it was requested and have always made themselves available to me. Their well designed project has also meant that my PhD has run extremely smoothly, with little or no problems during the three years. NERC and UCL are also thanked for risking money on me. Richard J. Butler (Bayerische Staatssammlung für Paläontologie und Geologie) has also helped me enormously over the course of my PhD – patiently explaining to me (several times) how to implement various statistical tests and use GIS, and in general providing extremely useful advice. I would also like to acknowledge the help and advice of (and useful discussions with) Roger B. J. Benson (University of Cambridge), Jon Bielby (Institute of Zoology, London), Chris Carbone (Institute of Zoology, London), Matthew T. Carrano (Smithsonian Institution), Chris McManus (UCL) and Alistair J. McGowan (Museum für Naturkunde), as well as thank my two thesis examiners, Susan E. Evans (UCL) and Oliver W. M. Rauhut (Bayerische Staatssammlung für Paläontologie und Geologie), for their incisive and helpful comments. People in numerous institutions around the world have helped me in my data collection: I would like to express my gratitude to all of these people, with particular thanks to the following people who truly went out of their way to be of assistance: Nils Knötschke (Dinosaurier‐ Freilichtmuseum Münchehagen) and Bernhard Zipfel (Bernard Price Institute), and an especially large thanks to Brent Breithaupt (Bureau of Land Management, Wyoming) and Neffra Matthews (Bureau of Land Management, Denver). My family deserves a large amount of recognition for all their help and support throughout the last 26 years and I doubt that I would be where I am now without them (beyond the fairly significant issue of not having been born in the first place). Several other people have supported 4 me over the years, particularly during some quite difficult times, and so I would like to thank Amy Beddows, Claire Cousins and Barbara Hanson. Lastly, if you felt you deserved thanks but didn’t get it, you obviously didn’t help hard enough… 5 Abstract Sauropodomorph dinosaurs were an important component of Mesozoic terrestrial ecosystems. Their diversity and abundance fluctuated throughout the Mesozoic but whether this reflects genuine biological changes or merely variations in our sampling of the rock record is uncertain. A database of all sauropodomorph individuals (2335) has been compiled, including environmental, geological, taxonomic and taphonomic data. Using a variety of sampling proxies (including a new specimen completeness metric) and a number of analytical techniques (residuals, rarefaction and phylogenetic diversity estimates), this work has demonstrated that sauropodomorph diversity appears to be genuinely high in the Pliensbachian‐Callovian and Kimmeridgian‐Tithonian, while low diversity levels are recorded for the Oxfordian and Berriasian‐Barremian, with the J/K boundary seemingly representing a real diversity crash. Diversity in the remaining Triassic‐Jurassic stages appears to be largely controlled by sampling biases while Late Cretaceous diversity is difficult to elucidate and perhaps remains relatively under‐sampled. Sea level affects diversity and abundance in the Jurassic‐Early Cretaceous, but does not appear to be linked in the Late Cretaceous. Different clades of sauropodomorphs potentially preferred different environments and this may have had an effect on changes in their distribution and diversity. Titanosaurs have been demonstrated to show a preference for inland environments compared to non‐titanosaurs, and it is possible that this led to their success in the Cretaceous when other sauropod clades were in decline. An assessment of the palaeolatitudinal patterns of sauropods and ornithischians reveals a distributional skew in the Late Cretaceous, which may reflect environmental and/or dietary preferences. A study of completeness through historical time contradicts the recent claim that the quality of dinosaurian type material has improved from the 19th century to the present. These studies illustrate that use of a number of techniques is imperative in any attempt to tease apart genuine patterns from the biases of an uneven rock record. 6 THESIS CONTENTS Chapter One – Introduction and Reviews 11 Introduction to the Sauropodomorpha 12 Review of previous environmental associations studies of sauropodomorphs 26 Review of previous dinosaurian distribution, diversity and completeness studies 29 Review of previous taphonomic studies of vertebrates 54 Chapter Two – Materials and Methods 77 Data 78 Geographic Information Systems (GIS) 81 Taxonomic revision 83 Methodological approaches 101 Chapter Three – Environmental Associations 142 Analyses and Results 143 Discussion 165 Chapter Four – Diversity 189 Analyses and Results 190 Discussion 202 Chapter Five – Completeness Metrics 214 The quality of the sauropodomorph fossil record through geological time 215 Analyses and Results 215 Interpretation and Discussion 219 Historical trends in specimen collection and taxonomy 225 Analyses and Results 225 Discussion 232 Completeness metrics in a wider context 239 Chapter Six – Palaeolatitudinal Patterns 243 Analyses and Results 244 Discussion 257 Chapter Seven – Trackway Abundance 260 Analyses and Results 261 Discussion 273 Chapter Eight – Taphonomy 275 Analyses and Results 276 Discussion 284 Chapter Nine – Conclusions and Future Work 286 Conclusions 287 Future Work 295 References 300 Appendix 348 Acknowledgements of Personal Communications 349 7 LIST OF TABLES 1.1. Taphonomic classes of Diictodon (after Smith 1993). 61 1.2. Disarticulation scheme of Canadian dinosaur specimens (after Dodson 1971). 63 1.3. Completeness classes of Plateosaurus specimens (after Sander 1992). 65 1.4. Disarticulation stages of Tendaguru dinosaur skeletons (after Heinrich 1999). 68 1.5. Bone weathering categories of mammal carcasses (after Behrensmeyer 1978). 73 1.6. Bone weathering rates of mammal carcasses (after Behrensmeyer 1978). 74 1.7. Fossil bone weathering categories (after Fiorillo 1988). 74 1.8. Fossil bone abrasion categories (after Fiorillo 1988). 75 2.1. List of museums and repositories visited for study of sauropodomorph material. 79 2.2. List of valid sauropodomorph taxa (and stratigraphic ranges). 83 2.3. Relative abundances of inland and coastal environments. 109 2.4. Completeness percentages attributed to regions of the body for SCM and CCM. 131 2.5. CCM values attributed to regions of the body for each phylogeny. 134 3.1. Environmental analyses of sauropods. 145 3.2. Epoch and period level environmental analyses of sauropods. 148 3.3. Stage level environmental analyses (body fossil individuals). 152 3.4. Stage level environmental analyses (tracksite individuals). 153 3.5. Stage level environmental analyses (body fossil and tracksite individuals). 154 3.6. Stage level environmental analyses (body fossil localities). 155 3.7. Stage level environmental analyses (tracksite localities). 156 3.8. Stage level environmental analyses (tracksite and body fossil localities). 157 3.9. ‘Sensitivity’ environmental analyses. 158 3.10. Environmental analyses of separate sauropod groups 162 4.1. Statistical comparisons of the various diversity curves to one another. 192 8 4.2. Statistical comparisons between diversity and sampling proxies. 195 4.3. Statistical comparisons between ‘corrected’ diversity and sea level. 200 5.1. Statistical comparisons of completeness curves through geological time. 217 5.2. Average completeness percentages for sauropodomorph clades and grades. 223 5.3. Statistical comparisons of completeness curves through historical time. 229 6.1. Statistical comparisons between tracksite abundance and numbers of DBCs. 247 6.2. Statistical comparisons between diversity and numbers of DBCs. 250 7.1. Statistical comparisons between tracksite abundance and diversity. 263 7.2. Statistical comparisons between tracksite abundance and sampling proxies. 267 7.3. Statistical comparisons between European tracksite abundance and rock outcrop 269 8.1. Preserved body regions of sauropodomorph individuals. 277 8.2. Preserved body regions of >=10%/20%complete sauropodomorph individuals. 279 8.3. Preserved body regions of articulated sauropodomorph individuals. 280 8.4. Preserved body regions of sauropodomorph individuals by environment. 281 8.5. Sauropodomorph disarticulation sequence. 282 LIST OF FIGURES 1.1. Sauropodomorph cladogram. 13 1.2. Mesozoic sauropodomorph taxic diversity (after Upchurch and Barrett 2005). 14 1.3. Basal sauropodomorph cladogram of Upchurch et al. (2007a). 16 1.4. Basal sauropodomorph cladogram of Yates (2007). 17 1.5. Late Jurassic distributions of dinosaur and plant taxa (after Rees et al. 2004). 32 1.6. Late Jurassic plant, evaporite, coal and dinosaur localities (after Rees et al. 2004). 33 1.7. Dinosaur taxic diversity by stratigraphic stage (after Taylor 2006). 37 9 1.8. Temporally calibrated phylogeny of the Dinosauria
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