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Universidade Do Estado Do Rio De Janeiro Centro De Tecnologia E Ciências Faculdade De Oceanografia Universidade do Estado do Rio de Janeiro Centro de Tecnologia e Ciências Faculdade de Oceanografia Humberto Freitas de Medeiros Fortunato A diversidade da Ordem Suberitida (Filo Porifera) sob a luz dos caracteres morfológicos, assinaturas metabólicas e sistemática molecular Rio de Janeiro 2019 Humberto Freitas de Medeiros Fortunato A diversidade da Ordem Suberitida (Filo Porifera) sob a luz dos caracteres morfológicos, assinaturas metabólicas e sistemática molecular Tese apresentada, como requisito parcial para obtenção do título de Doutor, ao Programa de Pós- Graduação em Oceanografia, da Universidade do Estado do Rio de Janeiro. Área de concentração: Caracterização, Diagnóstico e Evolução de Ambientes Marinhos Orientadora: Prof.a Dra. Gisele Lôbo Hajdu Coorientador: Prof. Dr. Thierry Pérez Rio de Janeiro 2019 CATALOGAÇÃO NA FONTE UERJ / REDE SIRIUS / BIBLIOTECA CTC/C F745d Fortunato, Humberto Freitas de Medeiros. A diversidade da Ordem Suberitida (Filo Porifera) sob a luz dos caracteres morfológicos, assinaturas metabólicas e sistemática molecular / Humberto Freitas de Medeiros Fortunato. – 2019. 266 f. : il. Orientador: Gisele Lôbo Hajdu. Coorientador: Thierry Pérez. Tese (Doutorado) - Universidade do Estado do Rio de Janeiro, Faculdade de Oceanografia. 1. Esponjas – Filogenia – Teses. 2. Biologia marinha – Teses. 3. Metabolômica – Teses. 4. Oceanografia – Taxonomia integrativa – Teses. 5. Atlântico, Oceano – Descoberta e explorações – Teses. I. Hajdu, Gisele Lôbo. II. Pérez, Thierry III. Universidade do Estado do Rio de Janeiro. Faculdade de Oceanografia. IV. Título. CDU 593.4 Bibliotecária responsável: Fernanda Lobo / CRB-7: 5265 Autorizo, apenas para fins acadêmicos e científicos, a reprodução total ou parcial desta tese, desde que citada a fonte. ______________________________________ ______________________ Assinatura Data Humberto Freitas de Medeiros Fortunato A diversidade da Ordem Suberitida (Filo Porifera) sob a luz dos caracteres morfológicos, assinaturas metabólicas e sistemática molecular Tese apresentada, como requisito parcial para obtenção do título de Doutor, ao Programa de Pós- Graduação em Oceanografia, da Universidade do Estado do Rio de Janeiro. Área de concentração: Caracterização, Diagnóstico e Evolução de Ambientes Marinhos Aprovada em 8 de Agosto de 2019. Orientadora: Prof.ª Dra. Gisele Lôbo Hajdu Instituto de Biologia Roberto Alcantara Gomes – UERJ Coorientador: Prof. Dr. Thierry Pérez Station Marine d’Endoume, Aix-Marseille Université - AMU Banca Examinadora: _____________________________________________________ Profª. Dra. Helena Passeri Lavrado Universidade Federal do Rio de Janeiro _____________________________________________________ Profª. Dra. Nicole Boury-Esnault Aix-Marseille Université _____________________________________________________ Profª. Dra. Valéria Laneuville Teixeira Universidade Federal do Estado do Rio de Janeiro _____________________________________________________ Prof. Dr. Cristiano Valentim da Silva Lazoski Universidade Federal do Rio de Janeiro _____________________________________________________ Prof. Dr. Luís Felipe Skinner Faculdade de Formação de Professores – UERJ Rio de Janeiro 2019 DEDICATÓRIA Dedico este trabalho às mulheres da minha família. Onde quer que estejam, olhem por mim. Vocês são exemplo de amor e dignidade: Vó Aglayr, mãe Maritza, irmãs Kharine, Vitória e Tamyris (in memorian). AGRADECIMENTOS Agradeço aos meus orientadores Gisele e Thierry por terem se disponibilizado a embarcar nessa aventura ao meu lado. Gi, obrigado pela confiança, suporte intelectual, financeiro e amizade extrema. Thierry, merci d'avoir essayé de faire de Marseille ma maison. Merci pour les enseignements scientifiques, les menus varies et délicieux, le football et l'amitié. Ao Thiago, um verdadeiro companheiro de trabalho, um gênio escondido sob sua timidez, um orientador sem igual! Sem você esse trabalho não teria sido possível. Obrigado pelas dicas, pelos puxões de orelha, pelos cafés e almoços. Obrigado por acreditar em mim e nesse projeto. Au Stéphane, pour votre soutien dans Le développement de tous lês travaux de métabolomique. Merci pour votre patience, pour Le temps investi en moi et pour être um compagnon quotidian dans um pays lointain. À CAPES por ter concedido a minha bolsa de doutorado, a bolsa PDSE para a realização do doutorado sanduíche e por investir na ciência do país. Ao professor Dr. Eduardo Hajdu por ter me incluído em diversas expedições no litoral nordestino, permitindo assim abranger e aumentar o número de espécimes coletados. Sem contar a oportunidade de admirar lugares paradisíacos tupiniquins. Au professeur Jean Vacelet, Pope Sponge. Merci de m'avoir accueilli à Marseille, de votre attention lorsque vous m'avez appris à préparer des échantillons pour le microscope électronique, dês explications sur la biologie dês éponges, dês histories merveilleuses et pour votre grand copain du football. A professora Nicole Boury-Esnault pelo carinho, ensinamentos e disponibilidade em atravessar o oceano para participar da minha banca de tese, atendendo ao meu pedido. Au professeur Alexander Ereskovski pour m'aider à visualiser lês larves et lês embryons présents dans certains échantillons et de m'avoir brièvement parlé de La reproduction dês éponges. Às(aos) professora(e)s Helena Lavrado, Suzi Ribeiro, Valéria Laneuville Teixeira, Anderson Villas-Bôas e Luís Felipe Skinner pelas correções, direcionamento e incentivo no trabalho, durante os seminários internos. Ao secretário do PPG-Ocn, André Felippe por toda presteza e camaradagem durante esses quatro anos de labuta. Obrigado por esclarecer todas as dúvidas e lembrar a todos os discentes dos nossos deveres nessa caminhada. Você é 10! Aos doutores Eduardo Hajdu, Eduardo Esteves, Ulisses Pinheiro, Thierry Pérez, Joana Sandes e a mestre Diana Ugalde por terem concedido diversos espécimes comparativos, tanto das Américas quanto da França. Ao Laboratório Central (LC) da Universidade Federal de Pernambuco (UFPE), pela de prestação serviço de caráter multiusuário e interdisciplinar, no âmbito da Plataforma Tecnológica de Genômica e Expressão Gênica, o qual permitiu o sequenciamento das amostras de DNA. Ao Laboratório de Microscopia Eletrônica de Varredura do Instituto de Química da UERJ pela disponibilidade nos serviços de microscopia de varredura eletrônica. Ao Laboratório de Contracorrente do Instituto de Pesquisas de Produtos Naturais da UFRJ por disponibilizar o liofilizador para secar as amostras para a realização dos estudos de metabolômica. Levy, obrigado por fazer o meio de campo e pela camaradagem. Ao técnico Eduardo do Núcleo de Biotecnologia Vegetal da UERJ por ajudar na liofilização das amostras e pelos ensinamentos sobre as metodologias de preservação de amostras para trabalho de metabolômica. Aos membros do Projeto Grael, Thiago Marques e Milton Araújo, que receberam a proposta de desenvolver uma atividade extensionista com tanto carinho. Yoyo, obrigado por dividir comigo essa ideia! Aos amigos cientistas que ajudaram nos campos e no laboratório. Obrigado por fazer destes quatro anos de ralação, um tempo de divertimento, reflexão e muito aprendizado sobre esponjas e sobre amizade. Obrigado, Anaíra Lage, Ester Gomes, Marcinha Motta, Camille Leal, Luiz Duarte, Cristiana Castello-Branco, Joana Sandes, Dora Leite, Sula Salani, André Bispo, Júlio Fernandez, Merry Grenier, César Ruiz, Renan Fructuoso, Felipe Sciamarella, Val Aguiar, Maurício Fernandes, Levy Domingos. Aos amigos da vida (de dentro e de fora da Universidade) que me mantém firme e forte. Aqueles que fazem minha vida ganhar sentido, em tempos nefastos de sucateamento da educação brasileira. Por mais distante que estivéssemos meu coração e pensamento estão sempre com vocês. Sou grato pela amizade e cumplicidade de todos. Juliana Magalhães, Amanda Guilherme, Larissa Pires, Yollanda Vançato, Fernanda França. Aos amigos da minha vida toda Pablo, Sina, Fê, Muralha, Gustavo, Júnior, obrigado pelos finais de semana descompromissados e leves. Nat, Liv e Rafa, o que a universidade nos deu, nada pode tirar. Amo todos vocês incondicionalmente! A galera do Tigres FC por me acolher e proporcionar momentos únicos de alegria e solidariedade, sobretudo nos finais de semana. À minha família, que sempre acreditou, torceu e apoiou minhas vontades. À minha mãe pela força, independente do momento, à minha vó pelo espírito de liberdade e conhecimento, à minha irmã pelo carinho. Amo vocês eternamente. À Breylla, minha parceira de todo dia. Quem tem me aturado nos momentos mais tensos e proveitosos da vida, durante esses quatro anos. A pessoa que sempre me salva nas questões referentes à diagramação, formatação e produção de ilustrações, principalmente mapas. Obrigado por me escolher e me querer todo dia. Te amo! O sonho da criação e a criação do sonho: a arte da ciência no tempo do impossível. Jurandir, Wanderlei, Sereno e Enilson G.R.E.S. Unidos da Tijuca, 2004 (Samba Enredo) Beaucoup de gens attaquent la mer, je lui fais l’amour. Jacques-Yves Cousteau While I pass with my spongiological work the columns of Hercules, I am facing a task which, toits full extent, can only be performed with the strength of a hero. O. E. Schmidt RESUMO FORTUNATO, Humberto Freitas de Medeiros. A diversidade da Ordem Suberitida (Filo Porifera) sob a luz dos caracteres morfológicos, assinaturas metabólicas e sistemática molecular. 2019. 266 f. Tese (Doutorado em Oceanografia) – Faculdade de Oceanografia, Universidade do Estado do Rio de Janeiro, Rio de Janeiro, 2019. As esponjas representam
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