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The Jurassic/Cretaceous Boundary: a Hidden Mass Extinction in Tetrapods?

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The Jurassic/Cretaceous Boundary: a Hidden Mass Extinction in Tetrapods? The Jurassic/Cretaceous boundary: a hidden mass extinction in tetrapods? Jonathan P. Tennant CID: 00661116 Imperial College London Department of Earth Science and Engineering Thesis submitted to fulfil the requirements for the degree of Doctor of Philosophy and Diploma of Imperial College Image credit: Robert Nicholls (CC BY 4.0). Depicts Sarcosuchus imperator, a giant predatory crocodyliform from the Cretaceous of North Africa. 1 Declaration of originality I declare that the works presented within this thesis are my own, and that all other work is appropriately acknowledged and referenced within. Copyright declaration The copyright of this thesis rests with the author, and it is made available under a Creative Commons Attribution (CC BY 4.0) license. Researchers are free to copy, distribute and transmit this thesis on the condition that it is appropriately attributed. Jonathan Peter Tennant Supervisors: Dr. Philip Mannion (Imperial College London); Prof. Paul Upchurch (University College London); Dr. Mark Sutton (Imperial College London). 2 Acknowledgements First and definitely foremost, I want to extend my greatest thanks to Phil Mannion. As his first PhD student, I am sure he regretted his decision after day one, but stuck with it until the end, and has been a stoic mentor throughout. This project would have been a shadow of what it came to be without his guidance. I am still yet to get him on Twitter though. I am also hugely grateful to Paul Upchurch and Mark Sutton for their input and experience throughout this project. I also could not have completed this project without the encouragement and support from my girlfriend, friends, and family, and am hugely grateful to them. Each of you are amazing, and I love you all in your own special way. I would like to extend my gratitude to NERC for funding this PhD project. Other institutes have provided various degrees of financial support throughout this research, and for that I would like to thank the Palaeontological Association, the Public Library of Science, and the Society of Vertebrate Paleontology. The whole Palaeontology community deserves collective thanks, and I would especially like to acknowledge the London vertebrate palaeontology community for their continuing support and advice throughout this PhD. I’d also like to extend my thanks to Ross Mounce, Mike Taylor and the global ‘open science’ community for their encouragement in encouraging me to ‘open up’ this research project, and for providing advice on the various aspects of publishing and practising open scholarship. Graham Steel has also indirectly influenced the mind-set with which I approached this PhD. He once said at a conference, “Getting younger people to be more open about their science: that's the revolution.” This inspired me to be completely open about all aspects of this PhD, from the research processes through to the outputs, including this thesis. As well as these, I feel that communities such as Wikipaleo and Twitter should be thanked for providing access to hundreds of research papers that I otherwise would not have had access to and without which I would not have been able to complete my research. 3 I am appreciative of anyone else who has ever offered encouragement directly or through social media. A special thanks to Ed Yong and Brian Switek as, although they don't know it, they encouraged me to start writing about science in the first place, which began my development along the path of science communication. I’d also like to extend my thanks to Barbara Ferreira of the European Geosciences Union for hosting my blog on the EGU network, which undoubtedly directly contributed to my development as a science writer, and has been an outlet for me to collect my thoughts. Nic Bilham of the Geological Society, London, has been a strong mentor before and during this PhD, and always an incredible educator on the more social and policy-oriented aspects of scientific research. Without his guidance, I would never have really considered the broader impacts of my research and skills development as a junior researcher, so am indebted to him for this horizon broadening, and the initial risk he took in employing me. He also offered me one of the best [and indirectly targeted] pieces of advice I've ever had in my pursuit for knowledge: “If you don't know what you're talking about, keep your mouth shut.” Needs no further explanation. Thanks to [Dr!] James Lewis as, in spite of being a planetary geochemist, he has always had useful comments to make on various aspects of this project. Also, life at Imperial College would have been a lot duller without the presence of other characters in the department, so thanks to everyone for all the hallway chats, shenanigans, and various offerings of cake, coffee, and cookies. For access to specimens, there are numerous people to thank for accommodating me and providing conversations about my research throughout: John Hutchinson for access to crocodile specimens at the RVC (London); Lorna Steel for access to specimens at the NHM (London), and useful discussion on crocodile anatomy; Didier Berthet at the MNHL (Centre de Conservation, Lyon); Zoltán Csiki-Sava at the FGGUB (Bucharest); Géraldine Garcia at UP (Poitieres); Amy Henrici at the CM (Pittsburgh); Carl Mehling of the AMNH (New York); Oliver Rauhut at the BSPG (Munich); Chris Sagebiel at UT (Austin); Daniela Schwarz at the MfN (Berlin); Herman Voogd from the TM (Haarlem); and Dale Winkler of the 4 SMU (Dallas). For those from non-English speaking countries, thanks for not making me feel any more stupid for not knowing your language. Without specimen access, much of this thesis would not have been possible, and I would like to thank each of you for your time and patience in allowing me to visit your collections. I would also like to thank Greg Price for useful discussion on the environmental changes occurring across the Jurassic/Cretaceous transition. Finally, I would like to offer my thanks to Robert Berner for freely providing his GEOCARBSULF data on request. Finally, I am grateful for the efforts of all those who have collected Jurassic–Cretaceous tetrapod data, and to those who have entered this data into the Paleobiology Database, especially John Alroy, Matt Carrano, Roger Benson, and Richard Butler. I would also like to thank John Alroy and Graeme Lloyd for assistance with R during the Fossilworks Analytical Course in the summer of 2014. J. Alroy also provided the scripts for implementing SQS in R (bio.mq.edu.au/~jalroy/SQS.html) and in Perl. Silhouettes in figures were created by Gareth Monger, Scott Hartman, Michael Keesey, and Nobu Tamura, and are available from phylopic.org/. If I have missed anyone out, it’s either because I didn’t want to thank you, or because writing an exhaustive list of everyone who has contributed in some way to my development as a scientist over the last few years would be a separate thesis in itself. 5 Published works by the author contributing to this thesis Tennant, J. P., Mannion, P. D., and Upchurch, P. U. (in press) Sea level regulated tetrapod diversity through the Jurassic/Cretaceous transition, Nature Communications. Tennant, J. P., Mannion, P. D., and Upchurch, P. (2016) Evolutionary relationships and systematics of Atoposauridae (Crocodylomorpha: Neosuchia): implications for the rise of Eusuchia, Zoological Journal of the Linnean Society, 177, 854-936. Tennant, J. P., Mannion, P. D., and Upchurch, P. (2016) Environmental drivers of crocodyliform diversity across the Jurassic/Cretaceous transition, Proceedings of the Royal Society Series B: Biological Sciences, 283, 20152840. (link) Tennant, J. P. Mannion, P. D., Upchurch, P., Sutton, M. and Price, G. (2016) Biotic and environmental dynamics across the Jurassic/Cretaceous boundary: evidence for a protracted period of faunal and ecological turnover, Biological Reviews (advanced online publication). (link) Young, M. T., Tennant, J. P., Brusatte, S. (2016) The first definitive Middle Jurassic atoposaurid (Crocodylomorpha, Neosuchia), and a discussion on the genus Theriosuchus, Zoological Journal of the Linnean Society, 176(2), 443-462. (link) Mannion, P. D., Benson, R. B. J., Carrano, M. T., Tennant, J. P., Judd, J. and Butler, R. J. (2015) Climate constrains the evolutionary history and biodiversity of crocodylians, Nature Communications, 6, 8438. (link) Tennant, J. P. and Mannion, P. D. (2014) Revision of the Late Jurassic crocodyliform Alligatorellus, and evidence for allopatric speciation driving high diversity in western European atoposaurids, PeerJ, 2, e599. (link) 6 Young, M. T., Steel, L., Foffa, D., Price, T., Naish, D., and Tennant, J. P. (2014) Marine tethysuchian crocodyliform from the ?Aptian-Albian (Early Cretaceous) of the Isle of Wight, England, Biological Journal of the Linnean Society, 113(3), 854-871 (link). Yi, H., Tennant, J. P., Young, M. T., Foffa, D., Hudson, J. D., Challands, T. J,. Ross, D. A. and Brusatte, S. L. (in review) An unusual small-bodied crocodyliform from the Middle Jurassic of Scotland, UK, and potential evidence for an early diversification of ‘advanced’ neosuchians. Transactions of the Geological Society of Edinburgh. Tennant, J. P. and Evers, S. (in prep.) How does diversity through time change through time? 7 “An approximate answer to the right question is worth a great deal more than a precise answer to the wrong question.” – John Tukey. “You can come up with statistics to prove anything, Kent.” – Homer Simpson. 8 Table of Contents Declaration of originality .......................................................................................................................
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