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Filogenia Dos Principais Grupos De Chondrichthyes Baseada Na Anatomia Comparada Do Esqueleto Das Nadadeiras Pares E Suas Cinturas

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Filogenia Dos Principais Grupos De Chondrichthyes Baseada Na Anatomia Comparada Do Esqueleto Das Nadadeiras Pares E Suas Cinturas João Paulo Capretz Batista da Silva Filogenia dos principais grupos de Chondrichthyes baseada na anatomia comparada do esqueleto das nadadeiras pares e suas cinturas Phylogeny of the major groups of Chondrichthyes based on the comparative anatomy of the skeleton of the paired fins and girdles (Volume 1) São Paulo 2014 João Paulo Capretz Batista da Silva Filogenia dos principais grupos de Chondrichthyes baseada na anatomia comparada do esqueleto das nadadeiras pares e suas cinturas Phylogeny of the major groups of Chondrichthyes based on the comparative anatomy of the skeleton of the paired fins and girdles Tese apresentada ao Instituto de Biociências da Universidade de São Paulo, como requisito para a obtenção do Título de Doutor em Ciências Biológicas, na Área de Zoologia. Orientador(a): Prof. Dr. Marcelo R. de Carvalho São Paulo 2014 ii Ficha Catalográfica Silva, João Paulo Capretz Batista Filogenia dos principais grupos de Chondrichthyes baseada na anatomia comparada do esqueleto das nadadeiras pares e suas cinturas xxv + 703 páginas. Tese (Doutorado) - Instituto de Biociências da Universidade de São Paulo. Departamento de Zoologia. 1. Chondrichthyes 2. Morfologia 3. Esqueleto apendicular Universidade de São Paulo. Instituto de Biociências. Departamento de Zoologia. Comissão Julgadora: ________________________ _____ _______________________ Prof(a). Dr(a). Prof(a). Dr(a). _________________________ ____________________________ Prof(a). Dr(a). Prof(a). Dr(a). Prof. Dr. Marcelo R. de Carvalho Orientador iii AGRADECIMENTOS Agradeço primeiramente aos meus pais, Cacilda Capretz e João Paulo Batista da Silva por seu amor, companheirismo, amizade e todo suporte necessário, imprescindíveis para realização deste trabalho. Agradeço também as minhas irmãs Nancy Capretz Batista da Silva e Juliana Capretz Batista da Silva por seu afeto e apoio em todas as etapas do meu trabalho. Aos “meninos” Conan e Pinguinho por dez anos de amizade e pela receptividade em todas as minhas voltas pra casa. Vocês estão sempre comigo! Agradeço a Bruna, meu amor, minha companheira, por todo seu amor, carinho, dedicação, atenção, compreensão pela minha ausência em momentos importantes e por ter sido minha consciência em períodos extremamente difíceis. Sua ajuda na organização do segundo volume da tese e na revisão do texto foram extremamente importantes. Agradeço também sua família, Sônia, Márcio, Manoel, Vitor, César e Mônica por me receberem e me acolherem tão bem, além de me ajudarem em vários aspectos na minha vida paulistana. Ao meu orientador Prof. Dr. Marcelo R. de Carvalho pela confiança depositada em mim durante o doutorado, por todo tempo de convivência desde minha iniciação científica e por me ceder a oportunidade de trabalhar com um grupo tão intrigante de peixes. Aos amigos de Laboratório, Thiago Silva Loboda, João Pedro F. A. F. Silva, Diego Francisco Biston Vaz, André Luis da Silva Casas, Flavia Petean, Renan Andrade Moreira, Karla Soares, Andrea Paixão, Leandro Yokota, Victor Giovannetti, Caio Gomes, Akemi Shibuya, Carolina Laurini, Maíra Portella, Mateus Soares, Murilo Carvalho e Sarah Viana, pela sua amizade, sugestões e ajuda no meu trabalho. Thiago, Diego, Joao Pedro e André ainda são especialmente agradecidos pela ajuda em diversos aspectos importantes durante o meu doutorado. Wilson Soares é agradecido pelas ilustrações. iv Diego Vaz, Carolina Laurini, Mariane Targino Rocha e o Prof. Dr. Hussam Zaher ainda são agradecidos pela ajuda com a utilização do TNT. Ao meu amigo Eduardo Pinto (Cabra), por toda sua ajuda, companheirismo, pelas ótimas risadas e ótima convivência nesses quatro anos dividindo apartamento em São Paulo. Ao docente do Departamento de Zoologia do Instituto de Biociências, Prof. Dr. Fernando Portella de Luna Marques pelo material de coleta importante para o presente projeto. À Prof. Dra. Mônica de Toledo Piza Ragazzo do Departamento de Zoologia do Instituto de Biociências da USP por toda a ajuda, sugestões importantes envolvendo meu projeto e com literatura pertinente ao presente estudo. Ao Prof. Dr. George Mattox, do Departamento de Biologia da Universidade Federal de São Carlos (UFSCar, Sorocaba), por sua ajuda e sugestões valiosas durante meu doutorado. Aos pesquisadores Profs. Drs. José Lima de Figueiredo e Aléssio Datovo do Museu de Zoologia da Universidade de São Paulo (MZUSP, São Paulo). O Prof. Dr. José Lima de Figueiredo é especialmente agradecido por todo seu esforço e suporte durante meu período de pós-graduação na USP em São Paulo. Aos amigos do Museu de Zoologia da USP, Túlio Franco Teixeira, Fernando Dagosta, Marina Loeb, Henrique Rosa Varella e Priscila Camelier por toda a ajuda e pelas boas risadas. Ao técnico do Museu de Zoologia da USP, Michel Gianeti pela ajuda no tombamento de material ictiológico vinculado ao presente estudo e a outros projetos. v Aos pesquisadores e amigos do Instituto de Biociências da USP, Marco Sena, Orlemir Carrerette, Karla Paresque, Marcelo Fukuda, Julia Beneti, Renato Nagata, Nilvea Ramalho, Cristiano Costa, Thais Miranda, Pedro Dias, Mariane Targino Rocha e Adriana Jeckel por toda sua ajuda durante o desenvolvimento do meu trabalho. À coordenação do Programa de Pós-Graduação em Zoologia do Instituto de Biociências da USP por financiar viagem para visita a coleção da UERJ e MNRJ no Rio de Janeiro. Ao Prof. Dr. Marcelo Britto do Museu Nacional do Rio de Janeiro (MNRJ), por sua receptividade e suporte durante minha visita a coleção de peixes. Aos membros do Laboratório de Taxonomia de Elasmobrânquios da Universidade do Estado do Rio de Janeiro (UERJ), Prof. Dr. Ulisses leite Gomes, Dr. Hugo Ricardo Secioso Santos e Rafael Fernandes Lopes por toda a receptividade e ajuda durante minha visita. Aos membros da Academy of Natural Sciences, Philadelphia (ANSP), Mark Henry Sabaj Pérez, John Lundberg, Kile Luckenbill e Mariangeles Arce por toda a ajuda e receptividade durante minha visita à coleção de peixes. Aos membros do National Museum of Natural History (USNM), Washington, DC, Jeff Williams, Richard Vari, Jeff Clayton, Sandra Raredon, Kris Murphy e Diane Pitassy por toda a ajuda e receptividade durante visita a coleção de peixes. Os amigos e pesquisadores brasileiros Prof. Dr. Fernando Jerep, Guilherme Dutra e Carlos David de Santana são agradecidos pela amizade e ótimo período de convivência durante minha visita à Washington. Aos membros do California Academy of Sciences (CAS), San Francisco, Dave Catania, Luiz Rocha, Jon Fong, Mysi Hoang, Benito Santos e Moises Bernal por toda o vi suporte e receptividade durante minha visita à coleção de peixes. Agradeço também a amiga Sasha Li por toda a ajuda durante minha visita a San Francisco. Aos membros do American Museum of Natural History (AMNH), Nova Iorque, John Sparks, Barbara Brown, Radford Arrindell e Robert Schelly por todo o suporte durante minha visita à coleção de peixes. Aos técnicos do setor de diagnóstico por imagem Reginaldo Barboza Silva e Hugo Idalgo por toda sua ajuda durante os eventos de radiografia de espécimes de Chondrichthyes. Às secretárias do Departamento de Zoologia do Instituto de Biociências da USP, Marly Salviano de Almeida, Maria Lúcia Vieira e Luzineia Ongaro Juquer. Aos funcionários do Departamento de Zoologia do Instituto de Biociências da USP, Ênio Mattos, Phillip Lenktaitis e Claudemir A. Lopes. Aos membros da Secretaria de Pós-Graduação do Instituto de Biociências da USP, Lilian Parpinelli, Erika Harumi Takamoto de Camargo, Helder Rossi Santos Souza e Vera Lúcia Barboza Lima. A realização deste projeto foi possível graças ao financiamento direto da Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP) na forma de bolsa de doutorado (FAPESP 2010/52677-6) e BEPE (FAPESP 2012/22692-9). vii ÍNDICE AGRADECIMENTOS.................................................................................................... iv ÍNDICE............................................................................................................................ viii 1. INTRODUÇÃO........................................................................................................... 01 1.1. Padrões esqueléticos das nadadeiras pares nos Gnatostomados............................ 01 1.2. Esqueleto das nadadeiras pares e cinturas em Chondrichthyes.............................. 06 1.3. Questões filogenéticas em Chondrichthyes........................................................... 11 2. OBJETIVOS................................................................................................................ 14 3. MATERIAIS E MÉTODOS........................................................................................ 15 3.1. Material................................................................................................................. 15 3.2. Análise anatômica................................................................................................. 27 3.3. Analise filogenética .............................................................................................. 28 4. RESULTADOS........................................................................................................... 30 4.1. Subclasse Elasmobranchii....................................................................................... 30 4.1.1.Ordem Hexanchiformes............................................................................. 30 4.1.1.1. Família Chlamydoselachidae................................................................. 30 4.1.1.2. Família Hexanchidae.............................................................................
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