UNIVERSIDADE DE SÃO PAULO FFCLRP - DEPARTAMENTO DE BIOLOGIA PROGRAMA DE PÓS-GRADUAÇÃO EM BIOLOGIA COMPARADA Patterns of morphological evolution in the skull of turtles: contributions from digital paleontology, neuroanatomy and biomechanics Padrões de evolução morfológica no crânio das tartarugas: contribuições da paleontologia digital, neuroanatomia e biomecânica Gabriel de Souza Ferreira RIBEIRÃO PRETO - SP 2019 UNIVERSIDADE DE SÃO PAULO FFCLRP - DEPARTAMENTO DE BIOLOGIA PROGRAMA DE PÓS-GRADUAÇÃO EM BIOLOGIA COMPARADA Patterns of morphological evolution in the skull of turtles: contributions from digital paleontology, neuroanatomy and biomechanics Padrões de evolução morfológica no crânio das tartarugas: contribuições da paleontologia digital, neuroanatomia e biomecânica Gabriel de Souza Ferreira Supervisor: Prof. Dr. Max Cardoso Langer Co-supervisor: Profa. Dra. Madelaine Böhme Tese apresentada à Faculdade de Filosofia, Ciências e Letras de Ribeirão Preto da USP, como parte das exigências para a obtenção do título de Doutor em Ciências, Área: BIOLOGIA COMPARADA. RIBEIRÃO PRETO - SP 2019 Autorizo a reprodução e divulgação total ou parcial deste trabalho, por qualquer meio convencional ou eletrônico, para fins de estudo e pesquisa, desde que citada a fonte FICHA CATALOGRÁFICA Ferreira, Gabriel de Souza Patterns of morphological evolution in the skull of turtles: contributions from digital paleontology, neuroanatomy and biomechanics. 190 p. : il. ; 30cm Tese de doutorado, apresentada ao Departamento de Biologia da Faculdade de Filosofia, Ciências e Letras de Ribeirão Preto/USP – Área de concentração: Biologia Comparada. Orientador: Langer, Max Cardoso. Co-orientadora: Böhme, Madelaine 1. Computed tomography. 2. Digital endocast. 3. Finite-Element Analysis. 4. Testudinata. 5. Skull. Name: Ferreira, Gabriel de Souza Title: Patterns of morphological evolution in the skull of turtles: contributions from digital paleontology, neuroanatomy and biomechanics. Thesis presented to the Faculdade de Filosofia, Ciências e Letras de Ribeirão Preto da Universidade de São Paulo to obtain a doctorate in Sciences, Comparative Biology area, and to the Mathematisch- Naturwissenschaftlichen Fakultät der Eberhard Karls Universität Tübingen to obtain a doctorate in Natural Sciences (Dr. rer. Nat.) Approved in: Examination board Prof. Dr. ________________________ Institution: _________________________ Verdict: _____________________ Prof. Dr. ________________________ Institution: _________________________ Verdict: _____________________ Prof. Dr. ________________________ Institution: _________________________ Verdict: _____________________ Prof. Dr. ________________________ Institution: _________________________ Verdict: _____________________ Prof. Dr. ________________________ Institution: _________________________ Verdict: _____________________ In nature we never see anything isolated, but everything in connection with something else which is before it, beside it, under it and over it Johann Wolfgang von Goethe Doubt is not a pleasant condition, but certainty is absurd François Voltaire i Ferreira – Patterns of morphological evolution in the skull of turtles Acknowledgements I am very grateful to my supervisor Max Langer, who offered me a space in his lab for the past ten years and immensily contributed to shape my career path until now. Max not only helped me think through paleo-problems, but also about career options and personal matters, always being present and giving support when I needed. I also thank my PhD co- supervisor in Tübingen, Prof. Dr. Madelaine Böhme, who accepted and welcomed me at the Senckenberg Institute and Universität Tübingen for a whole year, offering me not only a space to work, but also interesting discussions on various subjects. I am very grateful to my “unofficial” co-supervisor, Dr. Ingmar Werneburg, who has supported me from the beginning of my PhD, helping already when I was writing my doctoral research project and now, during this agitated last year. He granted me an invaluable amount of time during my stay in Tübingen, and also after I returned to Brazil, through several Skype calls and email exchange. All the conversations and discussions with them were very productive and insightful and, no doubt, they shaped not only the way I think about paleontology, evolutionary biology and science, but also about my life. I would also like to thank Juliana Sterli, who was my co-supervisor during my Master dissertation and with whom I spent six months working in Trelew. I learned a lot with you and you also had an importat role in defining my career. To you, my sincere thanks! I gratefully acknowledge the agreement FAPESP (São Paulo Research Foundation) and CAPES (Coordenação de Aperfeiçoamento de Pessoal de Nível Superior) for granting me the funding for my PhD project and my stay at the Universität Tübingen (grant numbers 2014/2539-5 and 2016/03934-2) and support by the Center for Biodiversity Documentation (Centro para Documentação da Biodiversidade), Department of Biology, FFCLRP, University of São Paulo, Brazil. This thesis constitutes part of the project 'Core-facility for the conservation of scientific documentation: biological collections and high technology research in comparative morphology' (CT-INFRA 01/2013), financed by the Funding Authority for Studies and Projects (FINEP), Ministry of Science, Technology, Innovations and Communication, Brazilian Federal Government. ii Ferreira – Patterns of morphological evolution in the skull of turtles I acknowledge the contribution of the following colleagues for discussion, collaboration, reviewing, collection or facility access, etc.: A. C. Arruda-Campos (Monte Alto, Brazil), A. Tröscher (Tübingen, Germany), A. Kupfer (Stuttgart, Germany), A. Rincón (Caracas, Venezuela), A. Solórzano (Caracas, Venezuela), B. Simpson (Chicago, USA), C. A. Francisco (Monte Alto, Brazil), C. Mehling (New York, USA), C. Pfaff (Viena, Austria), D. J. Button (London, UK), D. Silva, (Monte Alto, Brazil), D. Brinkman (New Haven, USA), D. Schwarz (Berlin, Germany), D. Kizirian (New York, USA), E. Cadena (Bogotá, Colombia), E. Almeida (Ribeirão Preto, Brazil), F. A. Carbonaro (Bauru, Brazil), F. V. Iori (Monte Alto, Brazil), G. S. Bever (New York, USA), I. G. Danilov (St. Petersburg, Russia), I. Ruf (Frankfurt, Germany), J. Prochel (Tübingen, Germany), J. Ziermann (Washington DC, USA), J. Cundiff (Boston, USA), J. Carrillo (Zürich, Switzerland), J. Hinz (Tübingen, Germany), J. Kriwet (Viena, Austria), K. Tighe (Washington DC, USA), K. Mahlow (Berlin, Germany), L. E. Sabino (Araraquara, Brazil), L. Vonnahme (New York, USA), M. S. Magalhães (Manaus, Brazil), M. de la Fuente (San Rafael, Argentina), M. Sánchez-Villagra (Zürich, Switzerland), P. S. R. Romano (Viçosa, Brazil), R. Garbin (Fribourg, Switzerland), R. P. Ghilardi (Bauru, Brazil), R. Schoch (Stuttgart, Germany), R. Diaz Jr. (Riverside, USA), R. Hirayama (Tokyo, Japan), R. Butler (Birmingham, UK), R. Benson (Oxford, UK), R. Diogo (Washington DC, USA), S. Bandyopadhyay (Kolkata, India), S. Evers (Fribourg, Switzerland), S. Chapman (London, UK), S. Lautenschlager (Birmingham, UK), S. S. Nihei (São Paulo, Brazil), S. Walsh (Edinburgh, UK), T. Kohlsdorf (Ribeirão Preto, Brazil), T. Scheyer (Zürich, Switzerland), V. Volpato (Frankfurt, Germany), W. G. Joyce (Fribourg, Switzerland), W. Maier (Tübingen, Germany). To all the PaleoLab past and present members for the uncountable hours of coffee breaks, Cantina do Seu Zé, bandejões, PaleoNatais and the endless and coolest (and sometimes also useless) discussions on an increadible diversity of topics. In particular, to my friends Tomate, Wafa, Julio, Simone, Mariela, Squirtle, Bruninho, Schumy, Fezão, Marquinho, Roque, Annie, who were part of my life during (very) different stages, and to the more recent coming, Bruna, Gustavo, Fellipe, Silvio, João, Chico, Gabriel, you became very rapidly an important part of my life. To all my Tübingen colleagues and friends, in particular Márton, Eme, Alex, Antonio, Anna, Priscila, Zeina, Melina, you helped me to have a great time during my German year. I am very thankful to all of you! iii Ferreira – Patterns of morphological evolution in the skull of turtles To all my friends in Americana, Nacim, Diógenes, Giu, Danilo, Gui, and in Ribeirão Preto, from Tiana’s lab, in particular Melissa, Gäelle, Nathalia, Aline, Gaga. To Gabriela, for your partnership and love during all these years, you have a special place in my heart, no matter what. To all 45ers, in particular Frango, Mestre, Ursa, Ket, Luigi, Zuado, Canjica, Garfo, MX, Paulinha, all of you. To Letícia, who notwithstanding I’ve found (or you found me?) in this troubled period, readly offered me happiness, companionship and love, pure love. Finally, to all my family, in particular my parents Ana and Gil, my grandparents, Vó Cida, Vó Peê and Vô Nésio, and my tia Cristina, who always supported and believe in me, since I was a little kid, with a lot of hair on my head. Thank you all, very much, for your companionship, friendship, and love, lots of love, through all those years! O amor da gente é como um grão, morre e nasce trigo, vive e morre pão iv Ferreira – Patterns of morphological evolution in the skull of turtles English Abstract In the current framework of Evolutionary Biology, Paleontology has an important role to play. The fossil record represents a fundamental aspect in studies on the evolution of morphology, since from its study it is possible to retrieve reliable data on many pertinent aspects, e.g.,