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Salema, Sarpa Salpa (Linnaeus 1758): Stock Structure in the Eastern Atlantic and Biological Characterization Off the Portuguese Coast

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Salema, Sarpa Salpa (Linnaeus 1758): Stock Structure in the Eastern Atlantic and Biological Characterization Off the Portuguese Coast UNIVERSIDADE DE LISBOA FACULDADE DE CIÊNCIAS Salema, Sarpa salpa (Linnaeus 1758): stock structure in the eastern Atlantic and biological characterization off the Portuguese coast Doutoramento em Biologia Especialidade em Biologia Marinha e Aquacultura Rafaela Barros Paiva Tese orientada por: Prof. Doutor Leonel Serrano Gordo e pela Prof.ª Doutora Maria José Costa Documento especialmente elaborado para a obtenção do grau de doutor 2015 UNIVERSIDADE DE LISBOA FACULDADE DE CIÊNCIAS Salema, Sarpa salpa (Linnaeus 1758): stock structure in the eastern Atlantic and biological characterization off the Portuguese coast Doutoramento em Biologia Especialidade em Biologia Marinha e Aquacultura Rafaela Barros Paiva Tese orientada por: Prof. Doutor Leonel Serrano Gordo e pela Prof.ª Doutora Maria José Costa Júri: Presidente: ● Henrique Manuel Roque Nogueira Cabral Vogais: ● Nuno Eduardo Malheiro Magalhães Esteves Formigo ● Pedro Miguel Raposo de Almeida ● Jorge Manuel Santos Gonçalves ● Henrique Manuel Roque Nogueira Cabral ● Leonel Paulo Sul de Serrano Gordo Documento especialmente elaborado para a obtenção do grau de doutor Instituição Financiadora: Fundação para a Ciência e a Tecnologia (bolsa de doutoramento com a referência SFRH/BD/80268/2011) 2015 Doctoral dissertation in Biology (scientific area of Marine Biology and Aquaculture) presented to the University of Lisbon Dissertação apresentada à Universidade de Lisboa para obtenção do grau de Doutor em Biologia (especialidade Biologia Marinha e Aquacultura) Rafaela Carla Viseu Barros Paiva 2015 Page | i Page | ii TABLE OF CONTENTS Table of contents Agradecimentos (PT) ..................................................................................................... 1 Resumo e palavras-chave ............................................................................................. 5 Abstract and keywords ................................................................................................. 7 Resumo alargado .......................................................................................................... 9 List of papers ............................................................................................................... 15 CHAPTER 1................................................................................................................... 19 General introduction ................................................................................................... 21 Objectives and thesis structure .................................................................................. 27 PART I - Stock structure .............................................................................................. 31 CHAPTER 2 - Morphometric analysis .......................................................................... 33 Body geometric morphometric and otolith shape analysis: Complementary tools for stock discrimination of Sarpa salpa in the Northeast Atlantic Ocean ........................ 35 CHAPTER 3 - Genetic analysis ..................................................................................... 57 Post glacial expansion and current reduced gene flow in salema population Sarpa salpa (Linnaeus 1758) (family Sparidae) ..................................................................... 59 PART II - Biology .......................................................................................................... 91 CHAPTER 4 - Age, growth and reproduction .............................................................. 93 Page | iii TABLE OF CONTENTS Age, growth and reproduction of the protandrous hermaphrodite fish, Sarpa salpa, from the Portuguese continental coast ...................................................................... 95 CHAPTER 5 - Cystic structures ................................................................................... 125 Cystic structures in fish ovaries: more common than we think - the case study of Sarpa salpa (Sparidae) .............................................................................................. 127 CHAPTER 6................................................................................................................. 135 General discussion .................................................................................................... 137 Conclusions ............................................................................................................... 146 CHAPTER 7................................................................................................................. 149 References ................................................................................................................ 151 CHAPTER 8................................................................................................................. 169 Supplementary Material ........................................................................................... 171 Page | iv Page | v Page | vi AGRADECIMENTOS (PT) Agradecimentos (PT) Nestes quatro anos que passaram tive a oportunidade de conhecer e de conviver com diversas pessoas. Tudo o que aprendi e realizei corresponde ao reflexo de toda a ajuda, apoio e partilha de experiências que recebi da vossa parte, muito obrigada! Gostaria de agradecer especialmente: - Ao Professor Doutor Leonel Serrano Gordo, MUITO OBRIGADA por TODO o apoio, interesse, disponibilidade, paciência, por partilhar comigo a Sua experiência e conhecimentos e sobretudo por me deixar aprender Consigo; - À Professora Doutora Maria José Costa, MUITO OBRIGADA por ser minha orientadora e por TODO o apoio disponibilizado; - Ao pessoal do Laboratório de Biologia Pesqueira, nomeadamente à Ana Neves, Vera Sequeira, Ana Rita Vieira e Catarina Vendrell obrigada por TUDO durante as diversas fases do trabalho e sobretudo por aturarem o “aroma” das salemas… vocês são pessoas incríveis! Cada uma de vocês teve a capacidade de partilhar comigo conhecimentos e experiências distintas, independentemente do âmbito profissional ou pessoal; - À Ana Sofia Rodrigues, Carla Ribeiro da Silva e ao Francisco Pina-Martins pela preciosa ajuda nas diversas tarefas incluídas na análise genética; - Ao Professor Doutor Octávio Paulo, que mesmo sendo uma pessoa extremamente ocupada conseguiu sempre encaixar tempo para mim no seu horário; Page | 1 AGRADECIMENTOS (PT) - A todas as pessoas que me ajudaram a obter as amostras: Domingo Lago, Dimitrios Moutopoulos, João Barreiros, Jorge Gonçalves, Karim Erzini, Luciana Sola, Mafalda Freitas, Sanja Matid-Skoko, Sérgio Barbosa e Tibério Simões. Muito obrigada também ao Senhor de etnia cigana que me ajudou a “passar o tempo” enquanto eu esperava que as minhas amostras vindas da Madeira estivessem prontas para levantar na área de cargas do aeroporto de Lisboa…estávamos ambos à espera que a mercadoria estivesse disponível; - A todos os pescadores que me permitiram obter amostras, em especial aos Mestres Zé Neves, Zé Oliveira e Jacinto pois durante o período de amostragem, rara foi a semana em que não falámos ao telefone; - A todos os amigos que tiveram um papel essencial nas gargalhadas…em todos os almoços, lanches e jantares que organizámos; - Um obrigada especial à Maria e à Filipa pela companhia na hora do almoço e do café; - À família, por TUDO E MAIS ALGUMA COISA e também por estarem sempre dispostos a provar salema confecionada nas mais diversas formas; - Ao Tiago, por estar presente na MINHA VIDA….e ao Baltazar, que embora tenha vindo alterar completamente as nossas rotinas é o melhor cão que podíamos ter! Page | 2 AGRADECIMENTOS (PT) A autora desta tese foi financiada com uma bolsa de doutoramento da Fundação para a Ciência e a Tecnologia (Referência SFRH/BD/80268/2011). The author of this thesis was funded by Fundação para a Ciência e a Tecnologia with a PhD grant (Reference SFRH/BD/80268/2011). Page | 3 Page | 4 RESUMO E PALAVRAS-CHAVE (PT) Resumo e palavras-chave Resumo: Esta tese contempla o primeiro estudo multidisciplinar de salema, Sarpa salpa, no sul do Nordeste Atlântico porque analisa a estrutura do seu manancial e carateriza alguns dos parâmetros biológicos da espécie. Para analisar a estrutura do manancial, estudaram-se indivíduos de três regiões de Portugal continental e do Arquipélago da Madeira, complementados com informações de indivíduos do Mar Mediterrâneo para a análise genética, enquanto a análise biológica da espécie foi baseada em exemplares capturados na região centro do País. Considerando a análise da estrutura do manancial, técnicas morfo-geométricas aplicadas à forma do corpo e a análise elíptica de Fourier aplicada à forma do otólito indicaram alguma sobreposição dos indivíduos de Portugal continental, especialmente entre o centro e o sul, e uma forte separação dos indivíduos do Arquipélago da Madeira. Ambas as técnicas indicaram que devem existir pelo menos dois mananciais, um na Madeira e outro em Portugal continental. No entanto, os resultados não foram claros quanto à existência de mais um manancial no norte do país deixando em aberto a questão se realmente existe um manancial no norte e outro que contempla o centro e sul de Portugal. A existência de mananciais, colocada em evidência pelas análises morfológicas, indica que os indivíduos foram sujeitos à acção de divergência genética ou à adaptação em locais com condições ambientais particulares, sendo este último o mais plausível no presente caso. Em relação à genética, a análise dos dados mitocondriais e nucleares suportam as seguintes hipóteses: a) não há evidência de estruturação
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