DINÂMICA EVOLUTIVA DO Dnar EM CROMOSSOMOS DE PEIXES DA FAMÍLIA ELEOTRIDAE E REVISÃO CITOGENÉTICA DA ORDEM GOBIIFORMES(OSTEICHTHYES, TELEOSTEI)

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DINÂMICA EVOLUTIVA DO Dnar EM CROMOSSOMOS DE PEIXES DA FAMÍLIA ELEOTRIDAE E REVISÃO CITOGENÉTICA DA ORDEM GOBIIFORMES(OSTEICHTHYES, TELEOSTEI) DINÂMICA EVOLUTIVA DO DNAr EM CROMOSSOMOS DE PEIXES DA FAMÍLIA ELEOTRIDAE E REVISÃO CITOGENÉTICA DA ORDEM GOBIIFORMES(OSTEICHTHYES, TELEOSTEI) SIMIÃO ALEFE SOARES DA SILVA ________________________________________________ Dissertação de Mestrado Natal/RN, Fevereiro de 2019 UNIVERSIDADE FEDERAL DO RIO GRANDE DO NORTE CENTRO DE BIOCIÊNCIAS PROGRAMA DE PÓS-GRADUAÇÃO EM SISTEMÁTICA E EVOLUÇÃO DINÂMICA EVOLUTIVA DO DNAr EM CROMOSSOMOS DE PEIXES DA FAMÍLIA ELEOTRIDAE E REVISÃO CITOGENÉTICA DA ORDEM GOBIIFORMES (OSTEICHTHYES, TELEOSTEI) Simião Alefe Soares da Silva Dissertação apresentada ao Programa de Pós-Graduação em Sistemática e Evolução da Universidade Federal do Rio Grande do Norte, como parte dos requisitos para obtenção do título de Mestre em Sistemática e Evolução. Orientador: Dr. Wagner Franco Molina Co-Orientador: Dr. Paulo Augusto de Lima Filho Natal/RN 2019 Universidade Federal do Rio Grande do Norte - UFRN Sistema de Bibliotecas - SISBI Catalogação de Publicação na Fonte. UFRN - Biblioteca Central Zila Mamede Silva, Simião Alefe Soares da. Dinâmica evolutiva do DNAr em cromossomos de peixes da família Eleotridae e revisão citogenética da ordem gobiiformes (Osteichthyes, Teleostei) / Simião Alefe Soares da Silva. - 2019. 69 f.: il. Dissertação (mestrado) - Universidade Federal do Rio Grande do Norte, Centro de Biociências, Programa de Pós-Graduação em 1. Evolução cromossômica - Dissertação. 2. Diversificação cariotípica - Dissertação. 3. Rearranjos cromossômicos - Dissertação. 4. DNAr - Dissertação. 5. Microssatélites - Dissertação. I. Lima Filho, Paulo Augusto de. II. Molina, Wagner Franco. III. Título. RN/UF/BCZM CDU 575:597.2/.5 SIMIÃO ALEFE SOARES DA SILVA DINÂMICA EVOLUTIVA DO DNAr EM CROMOSSOMOS DE PEIXES DA FAMÍLIA ELEOTRIDAE E REVISÃO CITOGENÉTICA DA ORDEM GOBIIFORMES (OSTEICHTHYES, TELEOSTEI) Dissertação apresentada ao Programa de Pós-Graduação em Sistemática e Evolução da Universidade Federal do Rio Grande do Norte, como requisitos para obtenção do título de Mestre em Sistemática e Evolução com ênfase em Padrões e Processos Evolutivos. Aprovado em 27 de Fevereiro de 2019. Comissão examinadora: Dr. Gideão Wagner Werneck Félix Costa – UFRN Dr. Clóvis Coutinho da Mota Neto – CPRM Dr. Paulo Augusto de Lima Filho - IFRN Dr. Wagner Franco Molina - UFRN “Dedico a Deus por tudo que faz por mim, aos meus pais, a minha querida Tia Hagar (in memorian) e a todos que contribuíram na minha formação” iv “Por isso, me alegro, por amor de cristo, com as fraquezas, com as ofensas, com as adversidades, com as perseguições, com as angústias. Porque quando sou fraco, então é que sou forte” (Apóstolo Paulo) “Não são nossas habilidades que mostram quem realmente somos, são nossas escolhas” (J. K. Rowling) v AGRADECIMENTOS Agradeço primeiramente a Deus, por tudo que tem feito na minha vida, pelas portas que ele abriu durante toda minha vida e no percurso desta pesquisa e por tudo que é para mim. Agradeço do fundo do meu coração aos meus pais Valdenir Dalvino da Silva e Antônia Eveline Soares da Silva, por todo apoio, paciência, carinho e auxilio que me proporcionou em toda minha vida. Agradeço a minha Vó Maria de Lourdes Mendonça Soares, pelas inúmeras orações e carinho, que foram de muita importância para eu chegar até aqui. Ao meu irmão por ter me ajudado várias vezes quando precisei e pelo apoio que tem me fornecido. Ao meu tio Saquel Pascoal de Mendonça Soares, por toda ajuda que me proporcionou e por sempre estar ao meu lado. Agradeço ao meu orientador Prof. Dr. Wagner Franco Molina, por ter acreditado nesse projeto e confiado na minha pessoa para realizar essa pesquisa, e por todo aprendizado que tive o prazer de ter recebido. Agradecimento especial ao meu co-orientador Prof. Dr. Paulo Augusto de Lima Filho, por ser uma pessoa excepcional, por ter acreditado e confiado em mim, por ter me proporcionado a oportunidade de desenvolver essa pesquisa, assim como, ter grade participação na minha formação. Agradeço ao meu tio ‘China’ e ao meu amigo ‘Pretinho’, por ter me ajudado em algumas coletas. Agradeço também a minha noiva Monike Paulino da Silva de Mendonça, por todo carinho prestado, apoio nos momentos de dificuldade, pelas coletas que fomos juntos que com a presença dela se tornaram especiais. Agradeço a Enildo e a Francisco pelas incontáveis ajudas que deram ao longo desses dois anos vindo de Guamaré para Natal. Agradeço ao meu amigo e parceiro de pesquisa Anderson, pelo apoio e dedicação no desenvolvimento do trabalho, como também, na coleta do material biológico. Agradeço ao meu amigo dos fungos Renan Oliveira, um amigo que ganhei nesse percurso sempre me ajudando nas disciplinas e na construção da dissertação. Agradeço também a todos os componentes do Laboratório de Genética de Recursos Marinho, Gideão, Karlla, Inaílson, Divana, Aparecida, Glaicon, Vanessa, Josy, Juliany, Erika, Alisson e os demais, por toda ajuda que me proporcionaram que foram essenciais para desenvolvimento deste trabalho, sem essa ajuda, nada disso seria possível. Agradeço ao pessoal que trabalha no CTA de Extremoz, pela grande ajuda que me proporcionaram. Agradeço também, a todos os professores desde o maternal até aqui, pois todos contribuíram para minha formação tanto pessoal como acadêmica. A professora Lidiane Guilhermino, pela grande ajuda que me deu, me substituindo em algumas aulas quando precisada ir para Natal. Agradeço aos componentes da “Melhor Turma” do IFRN campus Macau, pelo companheirismo, amizade e ajuda que ultrapassou os limites da graduação e tornou- se para vida. Resumo A ordem Gobiiformes, com mais de 2.200 espécies é um dos grupos mais diversificados dentre os teleósteos. Essa alta diversidade é acompanhada por significativa diversificação cariotípica, cujas causas biológicas não são completamente estabelecidas. Dentre seus grupos encontra-se a família Eleotridae, com 23 gêneros e 171 espécies, das quais seis habitam rios próximos à costa e regiões estuarinas em vastas áreas do litoral brasileiro. Com vistas a ampliar os dados citogenéticos para Eleotridae e revisar os padrões de diversificação cariotípica em Gobiiformes, aqui foram analisadas as espécies Dormitator maculatus, Eleotris pisonis, Erotelis smaragdus e Guavina guavina com ocorrência no Atlântico Sul e realizada uma ampla revisão citogenética e associação com os aspectos biológicos e ecológicos da Ordem Gobiiformes. As análises citogenéticas empregaram metodologias convencionais (coloração com Giemsa, impregnação por nitrato de prata, bandamento-C), coloração com fluorocromos base-específicos (MM/DAPI) e mapeamento por hibridação in situ fluorescente com sondas DNAr 18S, 5S e de sequências microssatélites (CA)15. As espécies E. smaragdus e E. pisonis apresentaram 2n=46 cromossomos (NF=46), D. maculatus exibiu um cariótipo estruturalmente diversificado, com 36sm+4st+6a (NF=86), enquanto G. guavina um cariótipo numericamente divergente com 2n=52 (NF=52). As regiões organizadoras nucleolares nas quatro espécies, estão localizadas em diferentes posições de um único par cromossômico, revelando-se em um efetivo marcador citotaxonômico entre as espécies. O mapeamento in situ do DNAr indicou uma significativa variação de arranjos estruturais dos genes 18S e 5S, confirmando a elevada dinâmica evolutiva dos cariótipos dessas espécies. O conjunto de dados citogenéticos confirma 2n=46 cromossomos acrocêntricos, como uma condição simplesiomórfica para Gobiiformes. O variado conjunto de sequências repetitivas nos cromossomos das espécies de Eleotridae, e em Gobiiformes em geral, pode atuar como fator promotor de diversificação cariotípica, associado a características biológicas, como hábitos bentônicos e ovos adesivos bentônicos. Os resultados encontrados contribuem para um melhor conhecimento da evolução cariotípica da família Eleotridae, indicando a participação de variados rearranjos (inversões, fusões e fissões) e amplia as informações citogenéticas para Gobiiformes. Os padrões estruturais heterogêneos dos cromossomos desta Ordem de peixes, aliado aos seus aspectos biológicos que favorecem estruturações populacionais elevando a dinâmica evolutiva dos cromossomos, tornando este grupo um dos modelos mais ilustrativos de megaevolução cariotípica em ambientes marinhos. Palavras-chave: Evolução cromossômica, diversificação cariotípica, rearranjos cromossômicos, DNAr, microssatélites. Abstract The Order Gobiiformes, with more than 2200 species is one of the most diversified groups among teleosts. This high diversity of species is accompanied by significant karyotypic diversification, whose the biological causes are not completely established. Among its groups there is the family Eleotridae, with 23 genera and 171 species, of which six inhabits rivers near the coast and estuarine regions in vast areas of the Brazilian coast. In order to extend the cytogenetic data for Eletridae and to review the patterns of karyotype diversification in Gobiiformes, the species Dormitator maculatus, Eleotris pisonis, Erotelis smaragdus and Guavina guavina with occurrence in the South Atlantic were analyzed and a broad review of cytogenetic, biological and ecological aspects of the Order Gobiiformes were performed. The cytogenetic analyses employed conventional techniques (Giemsa staining, silver nitrate impregnation, C-banding), base-specific fluorochromes (MM/DAPI) staining, and fluorescence in situ hybridization with probes of 18S rDNA, 5S rDNA and repeats (CA)15. The species E. smaragdus and E. pisonis presented 2n=46 chromosomes (NF=46). D. maculatus exhibited a structurally diverse karyotype
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