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Neuroanatomia E Filogenia Da Família Cetopsidae (Osteichthyes, Ostariophysi, Siluriformes) Com Análises Simultâneas De Dados Morfológicos E Moleculares

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Neuroanatomia E Filogenia Da Família Cetopsidae (Osteichthyes, Ostariophysi, Siluriformes) Com Análises Simultâneas De Dados Morfológicos E Moleculares MUSEU DE ZOOLOGIA DA UNIVERSIDADE DE SÃO PAULO VITOR PIMENTA ABRAHÃO NEUROANATOMIA E FILOGENIA DA FAMÍLIA CETOPSIDAE (OSTEICHTHYES, OSTARIOPHYSI, SILURIFORMES) COM ANÁLISES SIMULTÂNEAS DE DADOS MORFOLÓGICOS E MOLECULARES SÃO PAULO 2018 VITOR PIMENTA ABRAHÃO NEUROANATOMIA E FILOGENIA DA FAMÍLIA CETOPSIDAE (OSTEICHTHYES, OSTARIOPHYSI, SILURIFORMES) COM ANÁLISES SIMULTÂNEAS DE DADOS MORFOLÓGICOS E MOLECULARES TESE DE DOUTORADO MUSEU DE ZOOLOGIA DA UNIVERSIDADE DE SÃO PAULO TESE DE DOUTORADO APRESENTADA COMO UM DOS REQUISITOS PARA A OBTENÇÃO DO TÍTULO DE DOUTOR PELO MUSEU DE ZOOLOGIA DA UNIVERSIDADE DE SÃO PAULO. ÁREA DE CONCENTRAÇÃO: ZOOLOGIA ORIENTADOR: MÁRIO CÉSAR CARDOSO DE PINNA SÃO PAULO 2018 ii AUTORIZO A REPRODUÇÃO E DIVULGAÇÃO TOTAL OU PARCIAL DESTE TRABALHO, POR QUALQUER MEIO CONVENCIONAL OU ELETRÔNICO, PARA FINS DE ESTUDO E PESQUISA, DESDE QUE CITADA A FONTE. I AUTHORIZE THE REPRODUCTION AND DISSEMINATION OF THIS WORK IN PART OR ENTIRELY BY ANY MEANS ELETRONIC OR CONVENTIONAL, FOR STUDY AND RESEARCH, PROVIDE THE SOURCE IS CITED. ii Abrahão, Vitor Pimenta Neuroanatonia e filogenia da família Cetopsidae (Ostariophysis, Siluriformes) com análises simultâneas de dados morfológicos. / Vitor Pimenta Abrahão ; orientador Mário César Cardoso de Pinna. São Paulo, 2018. 367 p. Tese de Doutorado – Programa de Pós-Graduação em Sistemática, Taxonomia e Biodiversidade, Museu de Zoologia, Universidade de São Paulo, 2018. Versão original 1. Neuroanatomia – Cetopsidae 2. Cetopsidae - Filogenia. I. Pinna, Mário César Cardoso de, orient. II. Título. CDU 597.55 CDU 597.551.4 BANCA EXAMINADORA Prof. Dr.__________________________ Instituição:____________________________ Julgamento: _______________________ Assinatura: ___________________________ Prof. Dr.__________________________ Instituição:____________________________ Julgamento: _______________________ Assinatura: ___________________________ Prof. Dr.__________________________ Instituição:____________________________ Julgamento: _______________________ Assinatura: ___________________________ Prof. Dr.__________________________ Instituição:____________________________ Julgamento: _______________________ Assinatura: ___________________________ Prof. Dr.__________________________ Instituição:____________________________ Julgamento: _______________________ Assinatura: ___________________________ ii Aos meus pais Gilmar e Mônica, minha irmã Camila e minha esposa Natália ii AGRADECIMENTOS Este projeto foi financiado diretamente através de Bolsas de Doutorado concedidas pela Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES; vigência 01/05/2014 a 01/10/2016), pela Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP 2014/11397-1; vigência 01/11/2016 a 02/08/2018), e também pela Bolsa Estágio de Pesquisa no Exterior (BEPE-FAPESP 2017/17957-7; vigência 01/01/2018 a 31/03/2018). A bolsa de doutorado foi vinculada ao projeto de auxílio “Integrando dados morfológicos e moleculares na sistemática de três subgrupos de Siluriformes: subfamília Copionodontinae (Trichomycteridae), gênero Pimelodella (Heptapteridae) e família Cetopsidae” (FAPESP 2015/26804-4; vigência 01/08/2016 a 31/07/2018), sob coordenação de Mário C. C. de Pinna, que forneceu grande parte da infraestrutura utilizada na condução do presente estudo. A falsa impressão de que uma tese é construída somente pelo esforço próprio é superada nos primeiros obstáculos encontrados desse novo e desafiador universo de possibilidades. Esse processo rapidamente passa a ser compartilhado por um número cada vez maior de pessoas. E esses agradecimentos não poderiam ser dedicados se não à elas que se fizeram presentes e fundamentais durante toda a construção desse manuscrito. Eu agradeço imensamente e sinceramente ao meu orientador Mário C. C. de Pinna. Não agradeço somente pela sua orientação, mas também pela sua amizade, apoio, liberdade e confiança em mim depositada durante todos esses anos de excelente convívio. Mário é um brilhante professor e um grande amigo. Agradeço a Mark H. Sabaj por ter me recebido na The Academy of Natural Sciences of Drexel University (ANSP), e ter me dado todo o suporte para a condução das minhas pesquisas. Além disso, também agradeço sinceramente a Maria Angeles A. Hernández por ter me recebido tão gentilmente também na mesma instituição e por todo o apoio possível e imaginável para o desenvolvimento de minhas pesquisas. A generosidade de alguns pesquisadores também foi fundamental durante todo esse processo. Agradeço a Aléssio Datovo, Heraldo Britski, Mônica Toledo-Piza, Naércio Menezes e Oscar Shibatta pelas inúmeras discussões sobre anatomia, ictiologia, filogenia, além de incontáveis outros temas. ii Além disso, não poderia deixar de agradecer a todo o apoio e ajuda dos curadores, pesquisadores e técnicos de outras instituições que foram fundamentais para aquisição de todo o material examinado no presente estudo: Melanie Stiassny, Barbara Brown e Radford Arrindel do American Museum of Natural History, New York (AMNH), Mark Sabaj e Maria Angeles Arce da The Academy of Natural Sciences of Drexel University, Philadelphia (ANSP), Jonathan Armbruster e David Werneck do Auburn University Natural History Museum, Auburn (AUM), Dave Catania da California Academy of Sciences, San Francisco (CAS), Kevin Swagel do Field Museum of Natural History, Chicago (FMNH), Carlos DoNascimiento Instituto de Investigación de Recursos Biológicos Alexander van Humboldt, Villa de Leyva (IAvH), Ivan Mojica Instituto de Ciencias Naturales, Museo de Historia Natural, Universidad Nacional de Colombia, Bogotá (ICNMHN), Christopher Taylor do Illinois Natural History Survey, Champaign (INHS), Lúcia Py-Daniel do Instituto de Pesquisas da Amazônia, Manaus (INPA), Christine Thacker do Los Angeles County Musuem of Natural History, Los Angeles (LACM), Cláudio Oliveira do Laboratório de Biologia e genética de Peixes, Universidade Estadual Paulista “Júlio de Mesquita Filho”, UNESP-Botucatu, Botucatu, (LBP), Museo de Ciencias Naturales de UNELLEZ, Guanare (MCNG), Roberto Reis do Museu de Ciências e Tecnologia da PUCRS, Porto Alegre (MCP), Karsten Hartel do Museum of Comparative Zoology, Harvard University, Cambridge (MCZ), Ramiro Barriga do Museo de La Escuela Politecnica Nacional, Quito (MEPN), Marcelo Britto e Cristiano Moreira do Museu Nacional, Universidade Federal do Rio de Janeiro, Rio de Janeiro (MNRJ), Wolmar Wosiack do Museu Paraense Emílio Goeldi, Belém (MPEG), Saúl Prada e Javier Maldonado-Ocampo do Museo Javeriano de Historia Natural “Lorenzo Uribe, S.J”, Pontifica Universidad Javeriana, Bogotá (MPUJ), Hernán Ortega do Museo de Historia Natural de la Universidad Nacional Mayor de San Marcos, Lima (MUSN), Michel Gianetti e Osvaldo Oyakawa do Museu de Zoologia da Universidade de São Paulo, São Paulo (MZUSP), Oscar Shibatta, José Birindelli e Fernando Jerep do Museu de Zoologia da Universidade Estadual de Londrina, Londrina (MZUEL), Sven Kullander do Swedish Museum of Natural History, Stockholm (NRM), Carla Pavanelli e Andrea Bialetzki do Núcleo de Pesquisas em Limnologia, Ictiologia e Aquicultura, Universidade Estadual de Maringá, Maringá (NUPELIA), Hernán Lopez-Fernandez e Erling Holm do Royal Ontario Museum, Toronto (ROM), Luz Jimenez da Colección de Ictiologia del Instituto de Biologia da Universidad de Antioquia, Medellín (UDEA:CIUA), Larry Page iii do University of Florida, Florida State Museum, Gainesville (UF), Douglas Nelson University of Michigan, Museum of Zoology, Ann Arbor (UMMZ), Carolina Doria da Universidade Federal de Rondônia, Porto Velho (UNIR), e Richard Vari† e Lynne Parenti do National Museum of Natural History, Smithsonian Institution, Washington (USNM). Também agradeço sinceramente aos técnicos do Museu de Zoologia, Universidade de São Paulo, Jaqueline Battilana do laboratório de biologia molecular, Alberto Carvalho do laboratório de Micro Tomografia, e Dione Seripieri e todo a equipe da biblioteca do MZUSP pela enorme ajuda e colaboração intensa durante o andamento do presente manuscrito. Agradeço imensamente meus companheiros de laboratório Fábio Pupo, Fernando D’agosta, Gustavo Ballen, Henrique Varella, Igor Mourão, João Genova, Luiz Peixoto, Manoela Marinho, Marina Loeb, Murilo Pastana, Paulo Presti, Priscila Camelier, Thiago Loboda, Túlio Teixeira, Verônica Slobodian, Vinícius Espíndola, Vinícius Carvalho e William Ohara pelas inúmeras discussões e auxílios, além de proporcionarem os momentos mais especiais durante todo meu doutoramento. Todas essas pessoas foram importantes de alguma forma para a construção dessa tese, com discussões gerais sobre ciência, discussões sobre assuntos específicos de anatomia, filogenia, taxonomia entre tantas outras coisas, e também discussões sobre tantos outros assuntos da vida. Também agradeço pessoas de outros laboratórios e outras instituições que se fizeram muito importantes durante todo esse tempo, como Natália Friol, Rafael Penha, Luz Ochoa, Marina Mendonça, Gabriel Silva, Guilherme Dutra, David Santana e Guilherme Silva. Agradeço novamente e demasiadamente a Gustavo Ballen e Luiz Peixoto por toda ajuda com as análises filogenéticas, a Fábio Pupo pelas inúmeras discussões sobre sistema nervoso e a Natália Friol e Vinícius Espíndola pela leitura, correção e discussão sobre essa tese. Por fim, quero expressar minha gratidão para pessoas que não são necessariamente ligadas a academia, mas que contribuíram muito durante meu doutoramento. Quero agradecer toda a minha família, mas em especial algumas pessoas. Meus tios Milton e Neide Mauerwerk que me abrigaram
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