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This Electronic Thesis Or Dissertation Has Been Downloaded from Explore Bristol Research This electronic thesis or dissertation has been downloaded from Explore Bristol Research, http://research-information.bristol.ac.uk Author: Apostolaki, Naomi Title: The evolution of pneumaticity in Sauropodomorpha and its correlation to body size General rights Access to the thesis is subject to the Creative Commons Attribution - NonCommercial-No Derivatives 4.0 International Public License. A copy of this may be found at https://creativecommons.org/licenses/by-nc-nd/4.0/legalcode This license sets out your rights and the restrictions that apply to your access to the thesis so it is important you read this before proceeding. Take down policy Some pages of this thesis may have been removed for copyright restrictions prior to having it been deposited in Explore Bristol Research. However, if you have discovered material within the thesis that you consider to be unlawful e.g. breaches of copyright (either yours or that of a third party) or any other law, including but not limited to those relating to patent, trademark, confidentiality, data protection, obscenity, defamation, libel, then please contact [email protected] and include the following information in your message: •Your contact details •Bibliographic details for the item, including a URL •An outline nature of the complaint Your claim will be investigated and, where appropriate, the item in question will be removed from public view as soon as possible. This electronic thesis or dissertation has been downloaded from Explore Bristol Research, http://research-information.bristol.ac.uk Author: Apostolaki, Naomi Title: The evolution of pneumaticity in Sauropodomorpha and its correlation to body size General rights Access to the thesis is subject to the Creative Commons Attribution - NonCommercial-No Derivatives 4.0 International Public License. A copy of this may be found at https://creativecommons.org/licenses/by-nc-nd/4.0/legalcode This license sets out your rights and the restrictions that apply to your access to the thesis so it is important you read this before proceeding. Take down policy Some pages of this thesis may have been removed for copyright restrictions prior to having it been deposited in Explore Bristol Research. However, if you have discovered material within the thesis that you consider to be unlawful e.g. breaches of copyright (either yours or that of a third party) or any other law, including but not limited to those relating to patent, trademark, confidentiality, data protection, obscenity, defamation, libel, then please contact [email protected] and include the following information in your message: •Your contact details •Bibliographic details for the item, including a URL •An outline nature of the complaint Your claim will be investigated and, where appropriate, the item in question will be removed from public view as soon as possible. The evolution of pneumaticity in Sauropodomorpha and its correlation to body size Naomi-Eva Apostolaki A dissertation submitted to the University of Bristol in accordance with the requirements of the degree of Doctor of Philosophy in the Faculty of Science. Department of Earth Sciences University of Bristol October 2018 Supervisor Prof. Michael J. Benton Word count: 105,881 Abstract The superfamily Sauropodomorpha comprised some of the largest terrestrial herbivorous vertebrates that have ever walked on this planet. The study of these fascinating beings has been at the forefront of palaeontological research throughout the years. Especially in the last 20 years, due to the advent of modern techniques and technologies, significant steps have been made towards our understanding of their physiology. One of the sauropodomorphs’ central morphological characteristics is the expression of postcranial skeletal pneumaticity (PSP), a condition that is also expressed in the other superfamily of the order Saurischia, the Theropoda, as well as in their extant relatives, the Avians. This condition remodels the vertebrae and, occasionally the girdles and appendicular elements, producing perforations, depressions, excavations, cavities and internal chambers. This is a result of bone invasion and resorption from the development of the lung-air sac diverticula of the respiratory system. The various forms of this expression have been addressed across all subfamilies of sauropodomorphs mostly from an evolutionary aspect and, recently, from a developmental scope too. The resulting hypotheses state that PSP may have acted as a mechanism for weight reduction, allowing for sauropods to attain large sizes without having to suffer analogous gravitational constraints from equally attained masses. Any possible associations, though, between the expression of pneumaticity and body size have not been put under test. Through this study, a method of quantification and categorisation of PSP, and therefore, classification of the sauropodomorphs which express it, is created from the data retrieved from 61 taxa across all subfamilies, permitting us not only to trace any correlation between PSP and metric size data (body mass and body length) but also to visualise the evolution of PSP throughout Sauropodomorpha. This classification scheme from highest to lowest expression of PSP, ‘Alpha’, ‘Beta’, ‘Gamma’, ‘Delta’, and ‘Epsilon’ stems from the numerical estimate of pneumaticity in terms of a percentage, called the Pneumaticity Degree Index (PDI%). The revised scheme, Pneumaticity Degree Index (PDI%), takes account of the number of vertebral elements that are pneumatized in a single vertebra, the nature of pneumaticity traits, as well as the intensity of pneumatization in different body regions of interest (e.g. vertebral column), resulting in an integral and comprehensive measure of PSP. The proposed method ranks each pneumaticity trait with a value from 1 to 5 with 1 (100%) representing the most invasive unambiguous trait (e.g. camellae) and 5 (~ 10%) the least invasive and most ambiguous trait (i.e. fossa). By adding and dividing by their number all of the observed traits of every available vertebra of a region we retrieve a decimal numeric outcome and this outcome is translated to a percentage. The total average pneumatization of any taxon is calculated by retrieving the total average of the pneumaticity from all available body regions. In this study, only vertebral and pelvic elements were used for the retrieval of ii pneumaticity data since they are the most frequently exhibiting pneumatic traits in comparison to pectoral and appendicular elements. Results show that the total average PDI% range of basal Sauropodomorpha is 0%-59%, of the non-neosauropod Eusauropoda is 23%-90%, of Diplodocoidea is 45%-73%, of Macronaria is 45%-92% and of Somphospondyli is 46%-94%. The most pneumatized vertebral landmarks are the centrum, neural arch and neural spine. The vertebral regions most commonly pneumatized are the cervicals and dorsals. No genus or subfamily of sauropodomorphs ever reaches 100% pneumatization in all vertebral regions. Furthermore, PSP is not always positively correlated with mass, rather its expression is mostly correlated with length. Taxa with low or high masses may exhibit either low or high PDI’s. In addition, increasing progression of the extent and expression of PSP occurs mostly on a subfamilial level and less throughout the entire lineage. Finally, modelling calculations result in an Ornstein-Ulenbeck with an early burst progress of the expression of pneumatisation in sauropodomorphs. After that, the expression proceeds relatively steadily throughout the entire superfamily. The biotic causes of PSP are still unclear, as it could be an artefact of inheritance and genetic drift throughout speciation events. The pneumatization degree index (PDI%) is a means of quantifying and categorising pneumatization in any archosaurian taxon that is faster and cheaper, though less accurate, than CT scanning. It is more precise than the Pneumaticity Index since it integrates a wider selection of the qualities and osteological characteristics of pneumaticity we want to measure, provided that the bone under study is at least 50% free of matrix. iii Dedicatory I dedicate this PhD thesis and its journey to my beloved parents, Kalliopi Voikou and Theocharis Mike Apostolakis, to whom I am immensely grateful for their enormous dedication and precious support towards the fulfilment of my childhood dream to become a dinosaur palaeontologist. Acknowledgements First and foremost, I want to wholeheartedly thank my dear supervisor and mentor, Mike Benton, for his untiring patience in providing me with advice, guidance, aid, feedback and support whenever I needed it most. His exemplary presence was inspirational to my academic ‘upbringing’ throughout my years of studying in the University of Bristol. I am grateful to Mary Benton, Ember Kelly and Guy Molloy for their aid and support, especially in giving me the opportunity to work in a rewarding part-time job as a Disabilities Support Worker which enabled me to cover my expenses during the course of my studies in Bristol. I also want to thank the people in the Exams Office for providing me with additional work opportunities as an examinations invigilator. I am also grateful to Emily Rayfield, Manabu Sakamoto and Gavin Thomas as well as to my progress reviewers Phil Donoghue, Phil Anderson and Davide Pisani for their helpful input as well as for driving me to develop confidence and skills as a ‘growing’ academic. Some important parts of this thesis would not have been completed without
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