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Asteroidea:Echinodermata) Filogenia molecular, delimitação de espécies, desenvolvimento inicial e história evolutiva do gênero Echinaster (Asteroidea:Echinodermata) Elinia Medeiros Lopes Rio de Janeiro 2016 i Filogenia molecular, delimitação de espécies, desenvolvimento inicial e história evolutiva do gênero Echinaster (Asteroidea:Echinodermata) Elinia Medeiros Lopes Tese apresentada ao Instituto de Biologia da Universidade Federal do Rio de Janeiro, como parte dos requisitos necessários à obtenção do título de Doutor em Ciências Biológicas na área de Biodiversidade e Biologia Evolutiva. Orientador: Dr. Carlos Renato Rezende Ventura Co-orientador:Dr. Paulo Cesar de Paiva Rio de Janeiro 2016 ii Banca examinadora: ____________________________________________________ Dra. Michelle Regina Lemos Klautau (Presidente) Departamento de Zoologia/Universidade Federal do Rio de Janeiro _____________________________________________________ Dra. Haydée Andrade Cunha Departamento de Oceanografia biológica/Universidade do Estado do Rio de Janeiro _____________________________________________________ Dr. Anderson Vilasboa de Vasconcellos Departamento de Genética//Universidade do Estado do Rio de Janeiro _____________________________________________________ Dr. Marcelo Weksler Fundação Oswaldo Cruz, Instituto Oswaldo Cruz _____________________________________________________ Dr. Fábio Bettini Pitombo Departamento de Biologia Marinha/Universidade Federal Fluminense _____________________________________________________ Dra. Leila Maria Pessôa (Suplente) Departamento de Zoologia/Universidade Federal do Rio de Janeiro _____________________________________________________ Dr. Cristiano Valentim da Silva Lazoski (Suplente) Departamento de Genética//Universidade Federal do Rio de Janeiro Rio de Janeiro 2016 iii Ficha Catalográfica Lopes, Elinia Medeiros Filogenia molecular, delimitação de espécies, desenvolvimento inicial e história evolutiva do gênero Echinaster (Asteroidea:Echinodermata). 111p. Tese de doutorado em Biodiversidade e Biologia Evolutiva, 2016. 1. Echinodermata. 2. Asteroidea. 3. Variação morfológica. 4. Filogenia. 5. Filogeografia 6. Genética populacional 7. Desenvolvimento inicial. I – Ventura, Carlos Renato Rezende. II. Universidade Federal do Rio de Janeiro III – Doutorado Rio de Janeiro 2016 iv Agradecimentos Aos meus pais e irmãos por todo amor, apoio e incentivo. Ao Bruno, pelo companheirismo e apoio em todos os momentos. Obrigada por estarem ao meu lado sempre. Aos orientadores Renato Ventura e Paulo Paiva pela orientação, dedicação e confiança no meu trabalho. Por toda amizade e carinho durante esses anos que estamos trabalhando juntos e também pelo estímulo e colaboração ao meu enriquecimento profissional e pessoal. A Rocío Pérez-Portela, orientadora durante o doutorado sanduiche na Espanha, pela contribuição ao trabalho e todo aprendizado que tive durante o período de estágio. Agradeço ao CEAB (centro de estudos avançados de Blanes-Espanha), em especial Xavier Turon, pelo acolhimento e todo suporte dado durante o estágio sanduiche. Agradeço todos que me contribuíram com o envio e coleta de amostras para o desenvolvimento deste trabalho: Dr. Gustav Paulay (Florida Museum of Natural History/USA), Dr. Timothy O'Hara (Museum Victoria/Austrália), Dr. Bernard Picton (National Museums of Northern Ireland/Irlanda), Dr. Francisco A Solis Marin e Mrs. Carolina Martín (Instituto de Ciencias del Mar y Limnologia/México), equipe do projeto Coral Vivo pelas amostras de Arraial'dAjuda, Nataly Slivak pelas amostras de Florianópolis e Anne Gondim pela ajuda na coleta em João Pessoa. A toda equipe do laboratório de Echinodermata e Polychaeta, por toda ajuda na elaboração da tese e pelo ótimo relacionamento que mantemos no laboratório e que torna o trabalho muito agradável. Um agradecimento especial a Mariana, Bárbara, Alanna, Marcela, Fernanda, Monique, Carlos, Victor e Gustavo, companheiros de trabalho e grandes amigos que conquistei e que certamente estarão para sempre no meu coração. Obrigada pela ajuda e apoio durante o doutorado e pelas conversas diárias, gargalhadas, fofocas, desabafos emocionais, cervejas no china e mangue. Com vocês o trabalho se torna muito mais prazeroso. A todos outros os amigos que me acompanharam nesse período e que me deram força, nos momentos tristes e felizes. Obrigada a todos! Ao programa de Pós Graduação em Biodiversidade e Biologia Evolutiva pelo suporte ao longo do doutorado, disciplinas e pelas sugestões e correções durante os seminários de avaliação interno e exame de qualificação. Em especial a Dr. Cristiano Lazoski, Dr Joana Zanol, Dr Cláudia AM Russo, Dr Daniela M Takiya, Dr Antônio M Solé- Cava pelas sugestões no artigo referente ao capítulo 1 desta tese. Ao CAPES pela bolsa de doutorado e ao projeto CAPES-DGU pela bolsa de doutorado Sanduiche. v Índice Resumo ----------------------------------------------------------------------------------------------- 1 Introdução Geral ----------------------------------------------------------------------------------- 2 Referências ----------------------------------------------------------------------------------------- 6 Capítulo 1 - The molecular phylogeny of the sea star Echinaster (Asteroidea: 7 Echinasteridae) provides insights for genus taxonomy ------------------------------------- Abstract --------------------------------------------------------------------------------------------- 9 Introduction ---------------------------------------------------------------------------------------- 9 Materials and Methods ---------------------------------------------------------------------------- 11 Sampling------------------------------------------------------------------------------------------ 11 DNA extraction, amplification and alignment ----------------------------------------------- 12 Phylogenetic analysis --------------------------------------------------------------------------- 13 Results ---------------------------------------------------------------------------------------------- 15 Sequences ----------------------------------------------------------------------------------------- 15 Phylogenetic analysis --------------------------------------------------------------------------- 15 Discussion ------------------------------------------------------------------------------------------ 18 Acknowledgments --------------------------------------------------------------------------------- 21 References ------------------------------------------------------------------------------------------ 22 Capítulo 2 - Incipient speciation revealed by morphological, morphometric and genetic approaches: a study case in sea stars Echinaster (Othilia) along the Brazilian coast -------------------------------------------------------------------------------------- 49 Abstract --------------------------------------------------------------------------------------------- 51 Introduction ---------------------------------------------------------------------------------------- 51 Material and Methods ----------------------------------------------------------------------------- 54 Samples-------------------------------------------------------------------------------------------- 54 Geometric morphometrics --------------------------------------------------------------------- 55 Morphological character analysis ------------------------------------------------------------ 55 DNA extraction, polymerase chain reaction (PCR), sequencing and alignment ------- 56 Phylogenetic analysis and genetic distance ------------------------------------------------- 56 Phylogeographic analysis ---------------------------------------------------------------------- 58 Population and demographic analysis ------------------------------------------------------- 59 Results ----------------------------------------------------------------------------------------------- 59 Morphometric geometrics ---------------------------------------------------------------------- 59 Morphological analysis ------------------------------------------------------------------------ 60 Phylogenetic analysis --------------------------------------------------------------------------- 62 Phylogeographic analysis ---------------------------------------------------------------------- 62 Population and demographic analysis -------------------------------------------------------- 63 Discussion ------------------------------------------------------------------------------------------ 65 Conclusion ------------------------------------------------------------------------------------------ 68 Acknowledgments --------------------------------------------------------------------------------- 69 References ------------------------------------------------------------------------------------------ 69 Capítulo 3 - Development of the sea star Echinaster (Othilia) brasiliensis (Asteroidea:Echinasteridae): a comparative analysis making inference about the 91 evolution of developmental modes and skeletal plates in Asteroidea--------------------- vi Abstract --------------------------------------------------------------------------------------------- 92 Introduction ---------------------------------------------------------------------------------------- 93 Material and Methods ----------------------------------------------------------------------------- 95 Results ----------------------------------------------------------------------------------------------- 95 Discussion ------------------------------------------------------------------------------------------ 97 Acknowledgments
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