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Underfoot Pressure Equalisation As an Explanation for Extreme Underfoot Pressure Equalisation as an Explanation for Extreme Heteropody ELEANOR CATHERINE STRICKSON A thesis submitted in partial fulfilment of the requirements of Liverpool John Moores University for the degree of Doctor of Philosophy January 2020 Table of Contents Acknowledgements……………………………………………………………………….7 List of Tables and Equations…………………………………………………………….10 List of Figures…………………………………………………………………………….11 Abstract……………………………………………………………………………………14 Declaration………………………………………………………………………………..15 Chapter 1 – Introduction…………………………………………………………………16 1.1 Mass Distribution and Stability of Posture……………………………16 1.2 Studying Locomotion in Extinct Animals…………………………...…22 1.2.1 3D Modelling of Organisms………………………………...….24 1.3 Dinosaur Footprints, Heteropody, and Extreme Heteropody…...…..27 1.3.1 Ichnology and the Study of Footprints……………………...…27 1.3.2 Heteropody and Extreme Heteropody………………………...28 1.4 Quantifying Pressure in Living Animals………………………………..31 1.5 Extreme Heteropody as a Pressure Equalisation Mechanism?.........32 1.6 Thesis Aims……………………………………………………………….34 Chapter 2 – Can Skeletal Surface Area Predict in vivo Foot Surface Area?.............36 2.1 Summary of Chapter……………………………………………………..37 2.2 Introduction………………………………………………………………..38 2.3 Materials & Methods……………………………………………………...41 2.4 Results……………………………………………………………………..47 2.5 Discussion…………………………………………………………………55 2.6 Conclusions……………………………………………………………….62 Chapter 3 – Foot Shape Tests...................................................................................63 2 3.1 Purpose…………….……………………………………………………..63 3.2 Methods…………………………………………………………………..63 3.3 Test Outcomes…………………………………………………………..65 3.3.1 Basic Shapes……………………………………………………65 3.3.2 Animal Foot Outlines……………………………………………66 3.3.3 Dinosauria Predictions………………………………………….68 3.4 Conclusions………………………………………………………………69 Chapter 4 – Are Heteropody and Centre of Mass Position Linked in Skeletons of Extant Tetrapods?................................................................................................................70 4.1 Summary of Chapter……………………………………………………..70 4.2 Introduction………………………………………………………………..71 4.2.1 Mechanical Roles of the Forelimb and Hindlimb……………..72 4.3 Materials & Methods……………………………………………………..74 4.3.1 3D Models and Hulls…………………………………………….74 4.3.2 Anatomical Measurements…………………………………...…77 4.3.3 Analysis……………………………………………………………79 4.4 Results……………………………………………………………………..80 4.5 Discussion………………………………………………………………....87 Chapter 5 – Calibration Tests for Footscan Pressure Mat……………………………...92 5.1 Purpose……………………………………………………………………..92 5.2 Methods…………………………………………………………………….92 5.3 Test Outcomes…………………………………………………………….94 5.3.1 Human Calibration………………………………………………94 3 5.3.2 Weight and Shoe Size Calibrations……………………………96 5.3.3 T-tests……………………………………………………………..96 5.4 Conclusions………………………………………………………………..97 Chapter 6 – Are Underfoot Pressures Reflective of Heteropody and Centre of Mass Position?....................................................................................................................98 6.1 Summary of Chapter……………………………………………………..98 6.2 Introduction………………………………………………………………..99 6.3 Materials & Methods……………………………………………………..101 6.3.1 Pressure Mat Use………………………………………………..104 6.4 Results……………………………………………………………………..105 6.5 Discussion…………………………………………………………………112 Chapter 7 – Does the Fossil Record of Dinosauria Suggest the Use of Extreme Heteropody as an Underfoot Pressure Equalisation Mechanism?...........................117 7.1 Summary of Chapter…………………………………………………....117 7.2 Background for Heteropody and Pressure Equalisation in Palaeobiology…………………..………………………………………………...117 7.2.1 Factors in Footprint Formation………………………………...119 7.2.2 Extreme Heteropody and Manus-Only Trackways as Phenomena………………………………………………………………121 7.2.3 Centre of Mass and Convex Hulls…………………………….121 7.2.4 Aims………………………………………………………………122 7.3 Methods for Examining Heteropody and Its Implications in the Fossil Record……………………………………………………………………………..122 4 7.3.1 Methods Involving Body Fossils……………………………….123 7.3.2 Methods Involving Trace Fossils………………………………125 7.4 Results…………………………………………………………………….129 7.4.1 Trace Fossil Analysis………………………………………...…129 7.4.2 Body Fossil Analysis…………………………………………….135 7.5 Discussion, Implications and Explanations of Dinosaur Analysis...…138 7.5.1 Potential Trackway Issues………………………………………139 7.5.2 Sauropod Foot Posture………………………………………….140 7.5.3 Cursoriality…………………………………………………...…...140 7.5.4 What Can Be Gleaned from Osteological Analysis………..…141 7.5.5 Soft Tissue Implications………………………………………….142 7.5.6 Centre of Mass Diversity, and Explanations for Extreme Heteropody………………………………………………………………....144 Chapter 8 – Discussion…………………………………………………………………….145 8.1 Summary of Preceding Chapters and Wider Implications…………….145 8.1.1 Predictability of in vivo from Skeletal Foot Contact Area……..146 8.1.2 Pressure Measurements and their Utility………………………..146 8.1.3 Linking Centre of Mass Position and Heteropody in Extant Tetrapod Skeletons……………………………………………………………………148 8.1.4 Manus-Pes Disparity During Locomotion in Pressure, Area, and Vertical Force……………………………………………………………….148 8.1.5 Heteropody and Centre of Mass Trends in Quadrupedal Dinosaurs……………………………………………………………………149 5 8.2 Wider Implications of This Work……………………………………….….150 8.2.1 Heteropody as a Potential Equalisation Mechanism……..……150 8.2.2 Implications of Soft Tissue Disparity and Heteropody………….151 8.2.3 Body Plans of Taxonomic Groups and Heteropody…………….152 8.2.4 Why Have Extreme Heteropody?...............................................154 8.3 Potential Future Studies…………………………………………………….155 8.3.1 Larger/More Focused Data Sets…………………………………..155 8.3.2 Foot Posture………………………………………………………….156 8.3.3 Manus Versus Pes Underfoot Soft Tissue Distribution………….156 8.4 Conclusions……………………………………………………………………157 References………………………………………………………………………………………158 Supplementary Material……………………………………………………………………......192 6 Acknowledgements First and foremost, I would like to thank Liverpool John Moores University for awarding me a Faculty of Science PhD Scholarship, without which I would not have been able to move to Liverpool and start this PhD, never mind doing it for three years while keeping myself fed and clothed. Thanks also for awarding me with the oral presentation prize for the school of Natural Sciences and Psychology. It really helped boost my confidence at a pivotal point in my life. I would also like to thank LJMU for allowing me to go to conferences everywhere from Manchester to Australia, where I met and discussed science with many amazing people. Thanks also to the Carl Gans Collections and Charitable Fund, for awarding me a travel grant to go and present my research at ICVM 2019 in Prague, and saving me from having to stay in a 20 person room in an all-girls party hostel for a multi-day conference. Thank you to my co-supervisors, David Wilkinson at Lincoln University, at John Hutchinson at the Royal Veterinary College, Hatfield, for their help and advice, and especially to John for providing the bulk of the CT scan data used for chapter 2 and its corresponding Journal of Anatomy paper. I had a blast working with it. Thanks, on the topic of chapter 2, to David Blackburn at the Florida Museum of Natural History for providing us with frog CT scans to use, and to Diego Sustiata and two anonymous reviewers for reviewing the paper for submission, and ultimately helping us get it published. A special thank you should go out to the NHS, without which I would not be here finishing this PhD. I went in for multiple hospital visits, for surgeries, surgical revisions, road traffic accidents, the works. Thank God for the NHS. Thanks especially to Dr. Rashid and Dr. Morley at Charing Cross Hospital for literally cutting me up and stitching me back together multiple times this year, and to Susan Wilder for giving me CBT at the darkest point in my life since my teenage years. I honestly would not be alive, never mind finishing this PhD, if 7 it wasn’t for your intervention. You guys are literal lifesavers. Thank you too to Jeremy Corbyn for trying to save it as an institution. You tried, bud. On a lighter note, thank you to my friends for helping me through this and putting up with long periods of absence followed by being back in your life sporadically, and my parents, for supporting me and lifting me up when I needed it. Thank you especially to Karin Moejes and Ryan Marek for having my back and being there for me when I needed it most. I love you guys. Honourable mentions go to Alice Manterfield, Naomi Wain, Jack Warburton, and Heather Burton. Thanks also to Marta Zaher, for giving me back my confidence as a scientist when it was completely lost, for giving me somewhere to write up, and for all that other stuff that’s best left out of an acknowledgements section. I love you. Lastly, thanks to the co-operative for spotting me to star in an ad campaign. Without that opportunity I wouldn’t have been able to afford rent, or to finish writing up, never mind getting onto Nottingham’s 30 under 30 list. Thank you for standing up for trans rights, and for the rights of all oppressed peoples, and thank you to everyone else who does too. Solidarity. Together, we shall overcome. Until then, the fight continues. 8 “The philosophers have only interpreted the world, in various ways; the point is to change it.” – Karl Marx “Everyone thinks that courage is about facing death without flinching. But almost anyone can do that. Almost anyone can hold their breath and not scream for as long
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