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João Fabrício Mota Rodrigues ECOLOGIA GEOGRÁFICA E UNIVERSIDADE FEDERAL DE GOIÁS INSTITUTO DE CIÊNCIAS BIOLÓGICAS PROGRAMA DE PÓS-GRADUAÇÃO EM ECOLOGIA E EVOLUÇÃO João Fabrício Mota Rodrigues ECOLOGIA GEOGRÁFICA E EVOLUÇÃO DE QUELÔNIOS CONTINENTAIS Orientador: José Alexandre Felizola Diniz Filho Goiânia Maio 2017 TERMO DE CIÊNCIA E DE AUTORIZAÇÃO PARA DISPONIBILIZAR VERSÕES ELETRÔNICAS DE TESES E DISSERTAÇÕES NA BIBLIOTECA DIGITAL DA UFG Na qualidade de titular dos direitos de autor, autorizo a Universidade Federal de Goiás (UFG) a disponibilizar, gratuitamente, por meio da Biblioteca Digital de Te- ses e Dissertações (BDTD/UFG), regulamentada pela Resolução CEPEC nº 832/2007, sem ressarcimento dos direitos autorais, de acordo com a Lei nº 9610/98, o documento conforme permissões assinaladas abaixo, para fins de leitura, impres- são e/ou download, a título de divulgação da produção científica brasileira, a partir desta data. 1. Identificação do material bibliográfico: [ ] Dissertação [x] Tese 2. Identificação da Tese ou Dissertação: Nome completo do autor: João Fabrício Mota Rodrigues Título do trabalho: Ecologia geográfica e evolução de quelônios continentais 3. Informações de acesso ao documento: Concorda com a liberação total do documento [x] SIM [ ] NÃO1 Havendo concordância com a disponibilização eletrônica, torna-se imprescin- dível o envio do(s) arquivo(s) em formato digital PDF da tese ou dissertação. Assinatura do(a) autor(a)2 Ciente e de acordo: 1 Neste caso o documento será embargado por até um ano a partir da data de defesa. A extensão deste prazo suscita justificativa junto à coordenação do curso. Os dados do documento não serão disponibilizados durante o período de embargo. Casos de embargo: - Solicitação de registro de patente; - Submissão de artigo em revista científica; - Publicação como capítulo de livro; - Publicação da dissertação/tese em livro. 2 A assinatura deve ser escaneada. Assinatura do(a) orientador(a)² Data: 11 / 07 / 2018 Versão atualizada em setembro de 2017. UNIVERSIDADE FEDERAL DE GOIÁS INSTITUTO DE CIÊNCIAS BIOLÓGICAS PROGRAMA DE PÓS-GRADUAÇÃO EM ECOLOGIA E EVOLUÇÃO João Fabrício Mota Rodrigues ECOLOGIA GEOGRÁFICA E EVOLUÇÃO DE QUELÔNIOS CONTINENTAIS Orientador: José Alexandre Felizola Diniz Filho Tese apresentada à Universidade Federal de Goiás como parte das exigências do Programa de Pós-Graduação em Ecologia e Evolução para obtenção do título de Doutor em Ecologia e Evolução Goiânia Maio 2017 João Fabrício Mota Rodrigues ECOLOGIA GEOGRÁFICA E EVOLUÇÃO DE QUELÔNIOS CONTINENTAIS Tese apresentada à Universidade Federal de Goiás como parte das exigências do Programa de Pós-Graduação em Ecologia e Evolução para obtenção do título de Doutor em Ecologia e Evolução Banca avaliadora: Membros titulares: Dr. Franco Leandro de Souza (UFMS) Dr. Nelson Jorge da Silva Jr. (PUG-GO) Dr. Leo Caetano Fernandes da Silva (IBAMA-GO) Dra. Levi Carina Terribile (UFG) Dr. José Alexandre Felizola Diniz-Filho (UFG – Orientador) Membros suplentes Dr. Natan Medeiros Maciel (UFG) Dr. Luis Maurício Bini (UFG) Goiânia Maio 2017 Agradecimentos Ao professor e orientador José Alexandre Felizola Diniz Filho por ter aceitado me orientar durante a divertida e produtiva jornada do doutorado, assim como por todos os ensinamentos, paciência, confiança e suporte oferecidos. Foi um grande prazer contar com a sua orientação nessa jornada. Aos meus pais (José e Irene) por todo o carinho, apoio, amor, atenção e preocupação mesmo nos períodos em que estivemos distantes geograficamente. Às minhas irmãs (Lívia e Débora) por todo o carinho, conversas, brincadeiras e descontrações que tornavam os momentos de descansos mais reparadores. À Leticia por todo o apoio, amor, confiança e companheirismo, ajudando a deixar divertido e leve mesmo os períodos mais corridos. Aos amigos do Laboratório de Ecologia Teórica e Síntese (LETS) por toda a descontração, almoços no RU e discussões produtivas acerca de temas diversos de ecologia e evolução. Aos professores do Programa de Pós-Graduação em Ecologia e Evolução (especialmente, Thiago Rangel, Luis Maurício Bini, Adriano Melo e Matheus Ribeiro) pelas disciplinas cursadas e principalmente pela atenção e disponibilidade em ajudar quando perturbados nos corredores da UFG ou mesmo por email. Aos professores Thiago Rangel e Natan Maciel por terem aceitado compor minha banca de qualificação e por terem me incentivado a olhar com mais atenção as importantes informações que os fósseis podem nos trazer. Aos membros da Banca de Defesa da Tese (Franco Leandro, Nelson Jorge, Levi Carina e Leo Caetano) por terem aceitado investir um pouco do seu precioso tempo em minha tese e na minha formação. III Ao Marco Túlio pela amizade, companheirismo, hospitalidade e, claro, pelas inúmeras conversas científicas e profissionais que tivemos ao longo dessa jornada. A todas as pessoas com quem morei/convivi mais rotineiramente durante minha estadia em Goiânia (Dona Orozina, Victor, Jean, Marco, Bruno, Seu João, Geizi, Leila, Marga e Herlander) por terem dado seu melhor para tornar minha estadia em Goiânia algo cada vez melhor. Ao Luciano e ao Jesus por sempre me receberem tão bem em sua casa e por terem me proporcionado muitas risadas e felicidades nessas visitas. Aos amigos de Fortaleza pelas brincadeiras e descontrações durante os momentos de lazer em Fortaleza. Ao Programa de Pós-Graduação em Ecologia e Evolução e à Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) por terem me disponibilizado uma bolsa de doutorado a qual me sustentou financeiramente durante as pesquisas do doutorado. A Deus por ter me dado saúde e força de vontade para encarar os desafios que apareceram ao longo dessa jornada da melhor forma possível. IV RESUMO Compreender os processos responsáveis pelos padrões de distribuição atuais das espécies é um dos principais objetivos da Ecologia. Nesta tese, visamos entender quais fatores estão relacionados à distribuição da diversidade de quelônios, grupo de animais ectotérmicos cujos padrões de diversidade ainda são pouco conhecidos, ao longo do tempo e do espaço. Para esse fim, usamos dados de mapas de distribuição das espécies de quelônios continentais, além de informações de história de vida (tipo de hábitat e tamanho corporal), e reconstruímos uma hipótese filogenética para o grupo usando dados moleculares. O grupo dos quelônios apresentou uma explosão de diversificação durante a irradiação da família Emydidae, o que está provavelmente ligado a um evento de oportunidade ecológica. Além disso, animais aquáticos apresentaram taxas de diversificação mais elevadas que os animais terrestres, o que ajuda a explicar a maior diversidade de animais aquáticos atuais. A distribuição da riqueza de quelônios ao longo dos continentes é influenciada principalmente por variáveis climáticas tais como temperatura e precipitação, porém o intervalo de tempo desde o qual as áreas foram colonizadas também influencia nesse padrão. A diversidade beta entre as comunidades de quelônios é influenciada principalmente pela distância geográfica entre as comunidades, e comunidades de diferentes de domínios biogeográficos são estruturadas de modo diferenciado. Finalmente, a diversidade de tamanhos corporais dos quelônios também é influenciada pela temperatura, de modo que animais menores são mais comuns em áreas mais frias. Palavras-chave: Diversidade Beta; Diversificação; Métodos Filogenéticos Comparativos; Oportunidade Ecológica; Quelônios; Riqueza de Espécies; Tamanho Corporal. V ABSTRACT Understanding the processes that shape the current distribution patterns of species is one of the main goals of Ecology. In this thesis, we aimed to understand which factors are related to the distribution of the turtle diversity, a group of ectothermic animals whose diversity patterns are still little known, over time and space. To that end, we used distribution data from continental chelonian species, as well as life history information (habitat type and body size), and reconstructed a phylogenetic hypothesis for the group using molecular data. Turtles had a burst of lineage diversification during the irradiation of the Emydidae family, which is probably linked to an event of ecological opportunity. In addition, aquatic animals had higher rates of diversification than terrestrial animals, which helps to explain the current greater diversity of aquatic animals. Turtle richness distribution along the continents is mainly influenced by climatic variables such as temperature and precipitation, but the time when lineages first colonized the continental regions also influences this pattern. Beta diversity among chelonian communities is mainly influenced by the geographical distance between communities, and communities from different biogeographic realms are structured by different drivers. Finally, body size diversity of turtles is also influenced by temperature, and small animals are more common in cold areas. Keywords: Beta Diversity; Body Size; Diversification; Ecological Opportunity; Phylogenetic Comparative Methods; Species Richness; Turtles. VI SUMÁRIO INTRODUÇÃO GERAL___________________________________________ 1 CAPÍTULO 1____________________________________________________ 12 Rodrigues, J.F.M., Diniz-Filho, J.A.F. (2016): Ecological opportunities, habitat, and past climatic fluctuations influenced the diversification of modern turtles. Molecular Phylogenetics and Evolution 101: 352–358. CAPÍTULO 2____________________________________________________ 65 Rodrigues, J.F.M., Olalla-Tárraga, M.Á., Iverson, J.B., Akre, T.S.B., Diniz-Filho, J.A.F. (2017): Time and environment explain
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