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Universidade De São Paulo Faculdade De Filosofia UNIVERSIDADE DE SÃO PAULO FACULDADE DE FILOSOFIA, CIÊNCIAS E LETRAS DE RIBEIRÃO PRETO PROGRAMA DE PÓS-GRADUAÇÃO EM BIOLOGIA COMPARADA Characterization of macroevolutionary patterns of body size in Testudinata Caracterização dos padrões macroevolutivos do tamanho corporal em Testudinata Bruna de Mattos Farina Dissertação apresentada à Faculdade de Filosofia, Ciências e Letras de Ribeirão Preto da Universidade de São Paulo, como parte das exigências para obtenção do título de Mestre em Ciências, obtido no Programa de Pós-Graduação em Biologia Comparada Ribeirão Preto - SP 2021 UNIVERSIDADE DE SÃO PAULO FACULDADE DE FILOSOFIA, CIÊNCIAS E LETRAS DE RIBEIRÃO PRETO PROGRAMA DE PÓS-GRADUAÇÃO EM BIOLOGIA COMPARADA Characterization of macroevolutionary patterns of body size in Testudinata Caracterização dos padrões macroevolutivos do tamanho corporal em Testudinata Bruna de Mattos Farina Dissertação apresentada à Faculdade de Filosofia, Ciências e Letras de Ribeirão Preto da Universidade de São Paulo, como parte das exigências para obtenção do título de Mestre em Ciências, obtido no Programa de Pós-Graduação em Biologia Comparada Orientador: Prof. Dr. Max Cardoso Langer Ribeirão Preto - SP 2021 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. I authorize the reproduction and total or partial disclosure of this work, via any conventional or electronic medium, for aims of study and research, with the condition that the source is cited. FICHA CATALOGRÁFICA Farina, Bruna de Mattos Characterization of macroevolutionary patterns of body size in Testudinata, 2021. 65 páginas. Dissertação de Mestrado, apresentada à Faculdade de Filosofia, Ciências e Letras de Ribeirão Preto/USP – Área de concentração: Biologia Comparada. Orientador: Langer, Max Cardoso 1. Turtles. 2. Body size. 3. Macroevolution. 4. Phylogenetic comparative methods. 5. Cope’s rule. Acknowledgements During my trajectory so far, I had the chance to meet different people, which I am extremely grateful. Therefore, even though it is difficult to put into words, I cannot help but thank to: Prof. Max Langer, my supervisor, who gave the chance to be a member of the best lab I have ever been to. I came to the Laboratório de Paleontologia because I really admire you as a researcher and I could not be more grateful about the chance you gave me and the time I spend there. Dr. Gabriel Ferreira, my almost/non-official supervisor and also a great friend. You taught so much, since I first stepped into the lab. There were countless times that you helped and supported me during these almost three years. This project definitively would not have been possible without you. But more than a mentor and an excellent researcher, you are a great friend, and I was lucky enough to share so much time with you. Dr. Pedro Godoy, also a non-official supervisor, who, ironically, I never had the chance to meet personally, but helped me tireless times, always extremely kind. Again, this project would not be the same without your support. Prof. Roger Benson, my supervisor during MSc internship at the Earth Sciences Department, who is probably the most patient person in this world for explaining things to me so many times and also encouraging me to learn coding. All members of the Laboratório de Paleontologia. Each one of you has a part in my professional growth and I will always be grateful for that. I must especially thank Gustavo Darlim, Silvio Onary, Fellipe Muniz, and Guilherme Hermanson, for being by my side all this time, and sharing so many unforgettable moments with me. Programa Pós-Graduação em Biologia Comparada, for the opportunity to do my master’s degree here, and for being so competent and attentive with the students. FAPESP, for awarding my master’s scholarship and also my scholarship abroad. Earth Sciences Department at the University of Oxford, for welcoming me so well. The friends I made during my time in Oxford, especially Victor, Felipe, Svenja, Baran, Davide, Lena, and Jurgen. You made my time there really special, and I will never forget those game nights, our dinners togethers, and the conversations in the kitchen. The North Oxford Overseas Centre, which welcomed me very well and made my stay in Oxford possible. João Pedro Drudi, my boyfriend, who always support my choices and for being so patient all the time (especially living together during this never-ending quarantine). You already know how much you inspire me. And your participation in my life, be it professional or personal, has always been essential. I also thank the amazing moments we had the chance to share during these 6 years. Last, but not least, my parents, Alessandra Farina and Luiz Cesar Farina, and my sister, Beatriz Farina, for their unconditional love and support. Thank you for taking care of me so well, even now. I also need to apologize for not being able to be present in so many special moments, but I am always thinking about you. [...] A gente vive repetido, o repetido, e, escorregável, num mim minuto, já está empurrado noutro galho. Acertasse eu com o que depois sabendo fiquei, para de lá de tantos assombros…um está sempre no escuro, só no último derradeiro é que clareiam a sala. Digo: o real não está na saída nem na chagada: ele se dispões para a gente é no meio da travessia. [...] – João Guimarães Rosa, em “Grande Sertão: Veredas” Resumo Organismos, sejam eles vertebrados ou invertebrados, apresentam uma grande variedade de tamanhos e formas. A explicação para isso, entretanto, sempre foi conturbada, e diversas hipóteses, incluindo tanto tendências intrínsecas, quanto extrínsecas (e.g., termorregulação), foram propostas para explicar tais variações. Nesse contexto, os Testudinata apresentam grande disparidade de tamanhos corpóreos, especialmente se levado em conta o seu rico registro fóssil. No presente trabalho, foi explorada a evolução do tamanho corpóreo no grupo, analisando fatores relacionados aos padrões encontrados e a possível presença de tendências evolutivas (e.g., Lei de Cope). Para tal, duas superárvores foram construídas e calibradas no tempo geológico, baseadas em duas hipóteses filogenéticas independentes, e foram coletados dados de tamanho corpóreo, bem como outras informações como habitat e idade de cada táxon. Para a análise dos dados, foram realizadas regressão linear entre paleotemperatura e tamanho corpóreo, reconstrução de estados ancestrais, ajuste a diferentes modelos evolutivos, além da análise dos dados de tamanho ao longo do tempo. Os resultados obtidos indicam que não há um padrão geral que explique o tamanho das tartarugas ao longo do tempo; pelo contrário, tartarugas de diferentes hábitos ecológicos e diferentes linhagens apresentam padrões diferentes de variação de tamanho. Enquanto tartarugas de água doce apresentam pouca variação de disparidade ao longo tempo, possivelmente relacionada com os diferentes fatores ambientais característicos dos mais variados ambientes dulcícolas, tartarugas terrestres e marinhas apresentam grande variação, sendo o tamanho das terrestres possivelmente relacionado com certas condições ambientais e colonização de ilhas, e o das marinhas com adaptações para o ambiente pelágico, o que possibilitou maior dispersão do grupo. Mesmo tendo sido observada uma possível tendência de aumento ou diminuição de tamanho em certas linhagens, isso não foi confirmado pelos modelos evolutivos ajustados. Abstract Organisms, vertebrates or invertebrates, display an amazing variation of body size and shape. The explanation for that, however, has always been debated, and different hypotheses, including trends of body size and thermoregulation, were proposed to explain this variation. In this sense, Testudinata shows great disparity of body size, especially if its rich fossil record is accounted. In this study, I explore the body size evolution in turtles, analyzing which factors could influence the observed patterns, and the presence of long- term evolutionary trends (e.g., Cope’s Rule). For this, I constructed and time-calibrated two supertrees, based on two independent phylogenetic hypotheses, and collected body size data, as well as habitat information and age for each sampled taxon. To analyze body size data through time, I performed linear regression between paleotemperature and body size, ancestral state reconstructions, and macroevolutionary models fitting. The results suggest there is no general pattern that explains body size evolution in the group; instead, different habitats and lineages show different patterns of body size variation. Whereas freshwater turtles display a homogenous body size variation through time, possibly related to the uneven environmental factors of these habitats, terrestrial and marine turtles show more pronounced variation. Terrestrial turtles body size is likely related to environmental conditions and island colonization, and that of sea turtles to their pelagic lifestyle, which allowed great dispersal. Even though a few groups seemed to show some trends of body size, the evolutionary models indicate the opposite. Summary Introduction .................................................................................................................... 1 Body size, evolutionary trends, and thermoregulation hypotheses ...................... 1 Turtles and their morphological disparity ............................................................. 2 Aims ................................................................................................................................
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