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Universidade Estadual De Campinas Instituto De Biologia 1 UNIVERSIDADE ESTADUAL DE CAMPINAS INSTITUTO DE BIOLOGIA SUELLEN APARECIDA ZATTI SISTEMÁTICA E DISTRIBUIÇÃO DE MIXOSPORÍDEOS (CNIDARIA: MYXOZOA) PARASITOS DE TUCUNARÉ (Cichla spp.) E DOURADA (Brachyplatystoma rousseauxii) NA BACIA AMAZÔNICA SYSTEMATIC AND DISTRIBUTION OF MYXOSPOREANS (CNIDARIA: MYXOZOA) PARASITES OF TUCUNARÉ (Cichla spp.) AND DOURADA (Brachyplatystoma rousseauxii) IN THE AMAZON BASIN CAMPINAS 2017 2 SUELLEN APARECIDA ZATTI SISTEMÁTICA E DISTRIBUIÇÃO DE MIXOSPORÍDEOS (CNIDARIA: MYXOZOA) PARASITOS DE TUCUNARÉ (Cichla spp.) E DOURADA (Brachyplatystoma rousseauxii) NA BACIA AMAZÔNICA SYSTEMATIC AND DISTRIBUTION OF MYXOSPOREANS (CNIDARIA: MYXOZOA) PARASITES OF TUCUNARÉ (Cichla spp.) AND DOURADA (Brachyplatystoma rousseauxii) IN THE AMAZON BASIN Tese apresentada ao Instituto de Biologia da Universidade Estadual de Campinas como parte dos requisitos exigidos para a obtenção do título de Doutora em Biologia Animal, na área de Biodiversidade Animal. Thesis presented to the Institute of Biology of the University of Campinas in partial fulfillment of the requirements for the degree of Doctor in Animal Biology, in the area of Animal Biodiversity. ESTE ARQUIVO DIGITAL CORRESPONDE À VERSÃO FINAL DA TESE DE DOUTORADO DEFENDIDA PELA ALUNA SUELLEN APARECIDA ZATTI E ORIENTADA PELO PROF. DR. EDSON APARECIDO ADRIANO. Orientador: Prof. Dr. Edson Aparecido Adriano CAMPINAS 2017 3 4 Campinas, 06 de outubro de 2017 COMISSÃO EXAMINADORA Prof. Dr. Edson Aparecido Adriano (Presidente) Prof. Dr. Eduardo Makoto Onaka Profa. Dra. Márcia Mayumi Ishhikawa Prof. Dr. Anderson Luis Alves Prof. Dr. Adriano Cappellazzo Coelho Os membros da Comissão Examinadora acima assinaram a Ata de Defesa, que se encontra no processo de vida acadêmica do aluno. 5 AGRADECIMENTOS À Deus por me conceder o dom da vida, por colocar sonhos em meu coração e me dar força para conquistá-los. À minha família pelo amor e suporte. Especialmente à minha mãe Nice, que não me deixou desistir do doutorado ainda no começo frente as dificuldades de financiamento da pesquisa e bolsa. À minha avó Vita, que partiu neste meu último ano de doutorado, e que tanto cuidou de mim. À CAPES (Coordenação de Aperfeiçomento de Pessoal de Nível Superior), que concedeu a bolsa de Doutorado (2014/2017) e a bolsa de Doutoramento Sanduíche no Exterior (PDSE), que permitiu a condução de parte desta pesquisa na Oregon State University-OSU, Corvallis, OR, EUA (2015/2016) (BEX: 6729/2015-00). Ao Prof. Dr. Edson Adriano, pela orientação e suporte desde o Mestrado, pela oportunidade de estudar este grupo de organismos tão particular que me fez crescer imensamente como pesquisadora e por todas as oportunidades que me ofereceu, talvez a maior delas, a de trabalhar com organismos da Bacia Amazônica. Ao Prof. Dr. Antônio Augusto Mendes Maia (USP-Pirassununga), por todos esses anos de co-orientação, suporte e amizade. Obrigada por tudo, pelas longas conversas, pelas coletas, pelas várias ligações feitas para mim quando eu estava estagiando no exterior ou em Minas Gerais. Mais que um professor, ganhei um grande amigo, um exemplo que quero seguir! Aos professores, funcionários e alunos do Departamento de Biologia Animal da Unicamp. Em especial à Profa. Dra. Silmara Alegrette, pelo suporte em todas as fases desse doutorado, desde as mais difíceis. À Profa. Dra. Maura Franco pela confiança em abrir as portas do seu laboratório para que eu pudesse realizar minhas análises. À Profa. Dra. Marlene, umas das pessoas mais queridas que já encontrei neste meio acadêmico. Sempre disposta a me ajudar, me dar bons conselhos e a me fornecer todo o chocolate necessário nos dias que as “PCRs” insistiam em dar errado. Ao Prof. Dr. Danilo Miguel, pela amizade, torcida e pela confiança. Me senti muito honrada em ser sua aluna de estágio de docência, minha primeira experiência nesta área. Quero ser um dia uma professora como você. 6 Toda a minha gratidão aos professores, alunos e funcionários com os quais eu convivi e trabalhei no Department of Microbiology da Oregon State University, Oregon, Estados Unidos. Hoje posso me considerar uma pessoa privilegiada por ter realizado esse sonho que tinha desde que comecei minha carreira acadêmica. Foi difícil, era uma época de grande crise financeira no país, mas eu nunca desisti. À Profa. Dra. Jerri Bartholomew, umas das pesquisadoras mais importantes na minha área, por ter aberto as portas do seu laboratório para as minhas pesquisas, por todo carinho, incentivo, motivação e conhecimento e ao seu marido Dr. John pelo carinho e hospitalidade. Ao Dr. Stephen Atkinson pela orientação durante e após minha passagem pela OSU, por ter aceitado o desafio de me ajudar com minhas análises moleculares e filogenéticas, pelas longas discussões e comentários acerca da minha pesquisa, artigos e tese, pela enorme generosidade. Ao Prof. Dr. Maurício Laterça que tive o grande prazer de conhecer durante minha estadia na OSU e compartilhar o mesmo office, expectativas, alegrias e algumas “PBRs”. Obrigada por tudo, nunca esquecerei seus conselhos. À Dr. Sascha Hallett, Dr. Rich Holt, Dr. Julie Alexander, Ruth Milston-Clements, Sarah Vojnovich, Gema Alama Bermejo e Claire Katherine Howell. Thank you, guys, for sharing all yours experience with me, to be always so kind and helpful in all my moment in Corvallis and at OSU. It was one of the most memorable moments of my life and you have become everything easier! Hope to see you soon again! Aos amigos "brazilians" e “curva de rio” que conheci em Corvallis. Quando viajei não tinha expectativa de encontrar brasileiros pela cidade, mas Deus me presenteou com uma enorme família. Com vocês tudo era mais fácil, mesmo passar o Natal tão longe de nossas famílias. Muito obrigada, não poderia ter encontrado pessoas melhores! Leila, Déborah, Celinho, Lina, Bruno, Aninha e as gêmeas, André, Clarisse, Fernanda, Francine, Luciana, Mauro, Silvania, Gustavo, Rodrigo, Taynara, Carla, Dani, Rodrigo, Aninha Barros e Pedro. À Juliana Jordão, amiga desde a época da graduação. Obrigada Ju, pelo companheirismo, amizade vencida pela distância, pelo incentivo, por aquele abraço no aeroporto que nunca vou esquecer. Você reflete o verdadeiro significado da palavra amizade! 7 Uma pesquisa que envolve um extenso trabalho de campo, não poderia deixar de agradecer a todos que colaboraram com esse processo. Ao Dr. Lincon Corrêa, pela ajuda nas coletas e a todos os pescadores e moradores da região Amazônica que de alguma forma colaboraram para o desenvolvimento deste trabalho. Muito obrigada pela ajuda, recepção e por generosamente compartilharem tamanho conhecimento. Aos meus amigos pesquisadores da USP de Pirassununga, Dra. Márcia, Tiago, Juliana, Kássia, Gabriel e Júlio e à Dra. Isabel Müller, pela amizade e conselhos. Aos amigos que estão sempre a nos apoiar e levantar. Sabemos que as dificuldades neste nosso meio são enormes, as alegrias são muitas, mas muitas são as preocupações, dificuldades e frustrações. À turma de graduação da Universidade Federal de Alfenas, Bio 2006 barra 1 e às mestras da Unifesp, Camila Passos, Camila Uzam, Carina Uliam, Thábata, Tatiana e Valéria. À Vanessa Perdigão e Taís Bonatti, mais que amigas, vocês foram meu suporte dentro e fora da Unicamp. Muito obrigada pela torcida e carinho em cada momento. Sem o apoio e suporte diários de vocês eu não teria conseguido. Vocês são a melhor parte de Campinas. Ao César, pela ajuda com os softwares, amizade e conselhos, Gabriela, Júlia Molina, Diego e demais alunos do Departamento de Parasitologia, pela grande amizade compartilhada neste tempo de convívio. À minha amiga e roommate Juliana Lorensi (Ju), pela amizade sincera, pelo suporte e torcida em todos os momentos bons e difíceis (que foram muitos). Obrigada demais, você é parte de tudo isso! A todos que direta ou indiretamente contribuíram para que esse sonho se tornasse realidade. O grande aprendizado aqui, certamente não está nos resultados e nem nos artigos publicados, mas na persistência, resiliência e acreditar que vai dar tudo certo! “In the middle of difficulty lies opportunity”. Muito obrigada! 8 RESUMO Myxozoa são cnidários caracterizados por possuírem esporos multicelulares. Até o presente momento, mais de 2.400 espécies foram descritas parasitando principalmente peixes e invertebrados aquáticos. Esse grupo de endoparasitos vem atraindo atenção de pesquisadores não somente pelas patologias e danos causados aos peixes de ambiente natural e de cultivo, mas também pela complexidade do seu ciclo biológico, simplificação morfológica e sua inigmática evolução a partir dos cnidários de vida livre. Com o recente avanço das técnicas de biologia molecular no estudo de taxonomia e sistemática destes organismos, divergências taxonômicas surgiram em relação às metodologias morfológicas tradicionais. Além disso, as análises filogenéticas com base em dados moleculares se mostram incongruentes com as classificações atuais, o que têm levado a importantes revisões taxonômicas do grupo. A Bacia Amazônica é a região com maior diversidade de peixes de água doce do mundo, porém os estudos de taxonomia de mixosporídeos na região são escassos e a maioria das pesquisas até aqui realizadas foram baseadas apenas em dados morfológicos. Dos peixes que compõem a ictiofauna da Bacia Amazônica, merecem destaque pela importância ecológica e econômica os cicliformes do gênero Cichla (tucunarés) e o siluriforme Brachyplatystoma rousseauxii (dourada). Estes são peixes que possuem em comum a ampla distribuição geográfica ao longo de toda a bacia e elevado valor comercial. Diante deste
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