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FELIPE BANDONI DE OLIVEIRA Evolução Do Crânio Dos Macacos

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FELIPE BANDONI DE OLIVEIRA Evolução Do Crânio Dos Macacos FELIPE BANDONI DE OLIVEIRA Evolução do crânio dos macacos do Velho Mundo: uma abordagem de genética quantitativa SÃO PAULO 2009 FELIPE BANDONI DE OLIVEIRA Evolução do crânio dos macacos do Velho Mundo: uma abordagem de genética quantitativa Tese apresentada ao Instituto de Biociências da Universidade de São Paulo para a obtenção do Título de Doutor em Ciências, na Área de Genética e Biologia Evolutiva. Orientador: Dr. Gabriel Marroig SÃO PAULO 2009 Oliveira, Felipe Bandoni de. Evolução do crânio dos macacos do Velho Mundo: uma abordagem de genética quantitativa. 225 páginas. Tese (Doutorado) - Instituto de Biociências da Universidade de São Paulo. Departamento de Genética e Biologia Evolutiva. Palavras-chave: 1. Evolução morfológica. 2. Integração morfológica. 3. Seleção natural. 4. Deriva genética. I. Universidade de São Paulo. Instituto de Biociências. Departamento de Genética e Biologia Evolutiva. Comissão Julgadora : Prof(a). Dr(a). Prof(a). Dr(a). Prof(a). Dr(a). Prof(a). Dr(a). Prof. Dr. Gabriel Marroig Orientador Para meus pais, raiz, abrigo e espelho. Para Ana Elisa, com amor. Tinha eu que ser doutor... Paulinho da Viola Agradecimentos “Nada pode ser mais proveitoso para um jovem naturalista que uma viagem a um país distante.” Charles Darwin, A viagem do Beagle Essa é a lista dos gigantes, donos dos ombros em que me apoiei durante esses quatro anos de trabalho. Agradeço ao Dr. Gabriel Marroig pela orientação, pelo rigor nas críticas, pela ajuda decisiva nas análises e, acima de tudo, pela liberdade que me deu de conduzir o trabalho. Agradeço imensamente as várias oportunidades que vieram junto com esse projeto; aprendi muitas coisas além da Biologia. Montamos uma parceria extremamente produtiva, e levo comigo a certeza de um trabalho bem feito. À FAPESP e à CAPES, pelas bolsas concedidas. À Dra. Célia Koiffmann, que me auxiliou a pleitear a bolsa da CAPES. Ao Fernando Gomes, pelo empurrão inicial que faltava para eu embarcar neste projeto. Aos curadores e responsáveis por coleções zoológicas que garantiram meu acesso e o bom andamento dos trabalhos nos museus: Mário de Vivo, Juliana Gualda, Rogério Rossi e Carol Ayres (MZUSP); R. Voss e R. MacPhee (AMNH); L. Tomsett, P. Jenkins e D. Hills (BMNH); B. Paterson e B. Stanley (FMNH); J. Chupasko (MCZ); M. Godinot, C. Lefrève e J. Cuisin (MNHN); H. van Grouw (Naturalis); R. Thorington e L. Gordon (NMNH); M. Harman (Powell-Cotton Museum); G. Lenglet (RBINS); E. Gilissen e W. Wendelen (RMCA); I. Thomas e D. Willborn (ZMB); T. Jashashvili (Zürich Universität). Agradecimentos especiais a Eileen Westwig, Mark Omura, François Renoult, Emmanuel Gillisen, Wim Wendelen, Rob Asher, Hein van Grouw, Behnaz Bekkum-Ansari, Cristoph Zollikoffer, Márcia Ponce de León e Tea Jashashvili, por terem feito muito mais que o seu trabalho, ajudando nas horas em que ser estrangeiro é difícil. A todas as pessoas que me acolheram durante as visitas a museus: a turma de Kensal Rise, Tania e a família Sanchez, Benny, Marcia, Stephan e Thomas, Carla Meertens, Alejandra, James, Gabriel Perez e Cathi. Ao Fernando Santomauro, pelo “timing” perfeito em alugar apartamentos. Agradecimentos redobrados à querida Maria Tereza, pelo primeiro café da manhã em Londres, e ao Oscar, pela conta de gás mais valiosa da minha vida. Vocês são os responsáveis pela experiência mais interessante de todo este doutorado. Ao Dr. Marc Godinot, pela prontidão com que assumiu todas as responsabilidades e burocracias de orientador estrangeiro. Obrigado por me ajudar a trabalhar no exterior e pelas dicas para conseguir viver em uma das cidades mais caras do mundo. À Dra. Marta Lahr, pela confiança no meu trabalho, mesmo sem conhecê-lo. Ao Dr. Arne Mooers e Dr. Rutger Vos, que me enviaram a sua proposta de árvore filogenética de Catarrhini antes que fosse publicada. Aos funcionários e amigos do Departamento de Genética e Biologia Evolutiva, em especial à Elzi, Érika, Suzi, Maria, Helenice e Deisy por facilitarem imensamente o trabalho. Aos amigos da Seção de Pós-graduação, Vera, Helder e Erika, que sempre solucionaram minhas dúvidas. Aos grandes amigos da Biologia, que nesses anos descobri serem ainda mais especiais do que eu pensava. Um agradecimento especial ao Gustavinho, que me apresentou ao Matlab. Obrigado pelo impulso inicial e pelas várias dicas; devo a você a precisão e a agilidade das análises deste trabalho. A todos os participantes do Grupo de Discussão de Biogeografia, por discussões estimulantes e por manter aceso o verdadeiro espírito acadêmico, o de falar sobre coisas fundamentais. Aos amigos do choro, pelas comunhões musicais das noites de terça, que “flertam com o sublime” e “enlevam o espírito”. Espero que cheguemos à música dez mil em breve. Agradeço especialmente aos amigos que fiz no laboratório: Arthur, Leila, Tania, Val, Harley, Karina Tatit, Roberta, Ana Paula, Sebastien, Diogo, Guilherme e Hana, por dividirem comigo a dor e a delícia da pós-graduação. Mais recentemente, à Dani, Bárbara e Karina Bornia, a quem desejo um bom trabalho. Agradecimentos especiais a Arthur e Leila, com quem compartilhei o laboratório durante todo o doutorado, e que foram grandes parceiros. Aprendi muito com vocês e espero que tenhamos a chance de trabalhar juntos no futuro. Aos grandes amigos que me acompanham desde antes da Biologia entrar em minha vida, principalmente Fabricio, Lira, Marcão, Felipão e Bia, por manterem meus pés no chão e sempre me perguntarem, afinal, qual é o meu trabalho. As perguntas de vocês me ajudam a entender melhor o que eu faço e porque estou fazendo (embora eu ainda não tenha conseguido explicar). Aos pequenos Gustavo e Flávia, que fazem qualquer tese parecer pouco importante. A minha avó, Cida, que me influencia mais do que ela sabe. A minha irmã, Andrea, a quem admiro cada dia mais. A meus pais, Pedro e Laura, que lançaram a semente deste trabalho há muito tempo, quando “tinha eu quatorze anos de idade”, e cujo papel em tudo isso é maior do que consigo avaliar. Obrigado por estarem sempre aí, me apoiando nas horas em que o mundo inteiro joga contra. Obrigado, muito obrigado. À Ana Elisa, companheira demais, que me ajudou em todos os instantes do trabalho, bancou as horas de distância, me acalmou nos momentos de desespero, e com quem espero dividir as coisas boas da vida. Índice Resumo...................................................................................................................................................................13 Abstract..................................................................................................................................................................14 Introdução Geral: genética quantitativa e integração morfológica .............................................15 Introdução Geral ..................................................................................................................................................17 Integração morfológica: uma constatação empírica.................................................................................................................... 18 Genética Quantitativa: estimando o caminho da evolução........................................................................................................ 19 A equação de Lande.......................................................................................................................................................................... 22 Juntando as peças.............................................................................................................................................................................. 23 Modelo de estudo: o crânio dos macacos do Velho Mundo..................................................................................................... 24 Objetivos................................................................................................................................................................30 Referências.............................................................................................................................................................31 Capítulo 1: Estrutura decovariação nocrânio dos macacos do Velho Mundo: estase do padrão e evolução da magnitude .............................................................. 35 Introdução .............................................................................................................................................................37 Métodos .................................................................................................................................................................39 Amostra............................................................................................................................................................................................... 39 Taxonomia.......................................................................................................................................................................................... 40 Estimativa das matrizes de correlação e de variância/covariância ........................................................................................... 42 Comparação das matrizes de correlação e de variância/covariância........................................................................................ 43 Repetibilidade das matrizes e ajuste das comparações ............................................................................................................... 45 Magnitude geral das correlações
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