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DIVERSIFICAÇÃO CITOGENÔMICA EM SERRANIDAE (PERCIFORMES), UM MODELO DE ESTASE CARIOTÍPICA: ASPECTOS EVOLUTIVOS E APLICADOS KARLLA DANIELLE JORGE AMORIM ________________________________________________ Tese de Doutorado Natal/RN, março de 2020 KARLLA DANIELLE JORGE AMORIM DIVERSIFICAÇÃO CITOGENÔMICA EM SERRANIDAE (PERCIFORMES), UM MODELO DE ESTASE CARIOTÍPICA: ASPECTOS EVOLUTIVOS E APLICADOS Tese apresentada ao Programa de Pós- Graduação em Sistemática e Evolução/PPGSE, da Universidade Federal do Rio Grande do Norte, como parte dos requisitos para a obtenção do título de doutora em Sistemática e Evolução. Orientador: Prof. Dr. Wagner Franco Molina NATAL – RN 2020 Universidade Federal do Rio Grande do Norte - UFRN Sistema de Bibliotecas - SISBI Catalogação de Publicação na Fonte. UFRN - Biblioteca Setorial Prof. Leopoldo Nelson - •Centro de Biociências - CB Amorim, Karlla Danielle Jorge. Diversificação citogenômica em serranidae (Perciformes), um modelo de estase cariotípica: aspectos evolutivos e aplicados / Karlla Danielle Jorge Amorim. - Natal, 2020. 120 f.: il. Tese (Doutorado) - Universidade Federal do Rio Grande do Norte. Centro de Biociências. Programa de Pós-Graduação em Sistemática e Evolução. Orientador: Prof. Dr. Wagner Franco Molina. 1. Garoupas - Tese. 2. Evolução cariotípica - Tese. 3. Hibridização interespecífica - Tese. 4. DNA repetitivo - Tese. 5. Estase cariotípica - Tese. I. Molina, Wagner Franco. II. Universidade Federal do Rio Grande do Norte. III. Título. RN/UF/BSCB CDU 567 Elaborado por KATIA REJANE DA SILVA - CRB-15/351 KARLLA DANIELLE JORGE AMORIM DIVERSIFICAÇÃO CITOGENÔMICA EM SERRANIDAE (PERCIFORMES), UM MODELO DE ESTASE CARIOTÍPICA: ASPECTOS EVOLUTIVOS E APLICADOS Tese apresentada ao Programa de Pós- Graduação em Sistemática e Evolução/PPGSE da Universidade Federal do Rio Grande do Norte, como parte dos requisitos para a obtenção do título de doutora em Sistemática e Evolução. _________________________________________________ Dr. Clóvis Coutinho da Motta Neto (Serviço Geológico do Brasil) _________________________________________________ Dr. Luiz Antonio Carlos Bertollo (UFSCar) _________________________________________________ Dr. Marcelo de Bello Cioffi (UFSCar) _________________________________________________ Dr. Paulo Augusto de Lima Filho (IFRN) _________________________________________________ Dr. Wagner Franco Molina (Presidente da banca – UFRN) Aos meus avós, Francisco e Terezinha com todo o meu amor AGRADECIMENTOS Agradeço a minha família por todo apoio, por ser meu porto seguro, estender a mão quando tudo estava difícil, abrir mão de muitas coisas para que eu pudesse estudar e vibrar a cada degrau acadêmico que eu subo. Sem vocês o caminho teria sido mais difícil. Todas as minhas vitórias serão por e para vocês. Minha gratidão aos amigos que conquistei durante a vida e fizeram a diferença em momentos turbulentos. A vocês devo a minha sanidade e minha leve protuberância abdominal. Aos colegas que passaram ou estão no laboratório e contribuíram com minha formação e, adicionalmente, direta e/ou indiretamente para a realização deste trabalho: Allyson, Amanda, Aparecida, Cristiane, Clóvis, Dalvan, Divana, Éricka, Gideão, Glaicon, Inailson, Juliana, Josi, Leonardo Calado, Louise, Marcelo, Paulo, Rafael, Roberta, Rodrigo, Simião e Vanessa. Agradeço ao Professor Dr. Wagner Franco Molina por abrir as portas do laboratório para mim, pela orientação e por me fazer crescer pessoalmente e profissionalmente. Ao professor Marcelo de Belo Cioffi por disponibilizar as espécies coletadas na Tailândia e contribuir com o enriquecimento deste trabalho. Ao Programa de Pós-Graduação em Sistemática e Evolução e todo seu corpo docente pelo suporte oferecido durante o desenvolvimento deste trabalho. A CAPES (Coordenação de Aperfeiçoamento de Pessoal de Nível Superior) pela bolsa concedida. Nobody said it was easy. No one ever said it would be this hard. Coldplay – The Scientist RESUMO A família Serranidae, cujos representantes são conhecidos como garoupas, constitui um dos mais diversos grupos de peixes em ambientes recifais. São predadores eficientes que podem ocupar desde poças de maré a recifes profundos. Algumas espécies podem alcançar porte considerável e, em geral, significativo valor econômico. Com distribuição circunglobal, o grupo é composto por 577 espécies, distribuídas em 75 gêneros, dos quais Epinephelus é o mais representativo. Apesar do crescente conjunto de informações genéticas, menos de 10% das espécies apresentam aspectos citogenéticos conhecidos. Abordagens citogenéticas para a família se mostram dissociadas quanto aos seus aspectos filogenéticos, geográficos e taxonômicos, limitando a compreensão de seus aspectos evolutivos e aplicados ao manejo. No presente trabalho, análises citogenéticas convencionais e mapeamento por hibridização in situ fluorescente das sequências DNAr 18S, DNAr 5S, elementos transponíveis Tol2 e Rex3, microssatélites (CA)15, (GA)15, (CAA)10 e (CGG)10, foram realizadas em 11 espécies de Serranidae. O primeiro capítulo pretende comparar a distribuição de sequências repetitivas dentro do cenário filogenético e em diferentes regiões biogeográficas. Em virtude das informações citogenéticas estarem concentradas no gênero Epinephelus, o segundo capítulo traz informações citogenômicas para outras três espécies da família dos gêneros Cephalopholis e Rypticus. No terceiro capítulo análises citogenômicas de três espécies de garoupas de interesse comercial foram realizadas para gerar informações sobre a organização dos cromossomos e o nível de divergência cromossômica entre as espécies, além de aspectos adaptativos e potencialidades aplicáveis à práticas de hibridização interespecífica. Todas as espécies apresentaram 2n=48, das quais apenas três possuem cariótipos não compostos exclusivamente por elementos acrocêntricos. O conservadorismo cariotípico, apenas perturbado por inversões pericêntricas, se estende à organização do DNA repetitivo onde as sequências repetitivas aqui analisadas refletem um baixo dinamismo evolutivo. A divergência citogenética e genômica entre espécies decorrente de ancestralidade comum ocorrida em períodos entre 15 a 11 M.a aparentemente proporciona reduzido nível de bloqueio reprodutivo pós-zigótico, com valiosas implicações para a hibridização induzida na aquicultura. Sob perspectiva biogeográfica e filogenética, os grupos mais basais do Atlântico apontam 2n=48a como o padrão ancestral para a família, indica que a especiação no grupo não foi seguida por modificações cariotípicas significantes e a despeito de uma relativa frequência de cariótipos divergentes, esse grupo de peixes se adequa a um padrão evolutivo de estase cariotípica. Palavras-chave: Garoupas, evolução cariotípica, hibridização interespecífica, DNA repetitivo, estase cariotípica. ABSTRACT The Serranidae family, whose representatives are known as groupers, is one of the most diverse groups of fish in reef environments. They are efficient predators that can occupy everywhere from tide pools to deep reefs. Some species can reach considerable size and, in general, significant economic value. With a circunglobal distribution, the group consists of 577 species, distributed in 75 genera, of which Epinephelus is the most representative. Despite the growing set of genetic information, less than 10% of species have known cytogenetic aspects. Cytogenetic approaches to the family are dissociated in terms of their phylogenetic, geographical and taxonomic aspects, limiting the understanding of their evolutionary aspects and applied to management. Conventional cytogenetic analyzes and fluorescent in situ hybridization mapping of the 18S rDNA, 5S rDNA sequences, transposable elements Tol2 and Rex3, microsatellites (CA)15, (GA)15, (CAA)10 and (CGG)10, were performed in 11 species of Serranidae. The first chapter aims to compare the distribution of repetitive sequences within the phylogenetic scenario and in different biogeographic regions. Because the cytogenetic information is concentrated in the genus Epinephelus, the second chapter provides cytogenomic information for three other species in the family of the genera Cephalopholis and Rypticus. In the third chapter, cytogenomic analyzes of three species of groupers of commercial interest were performed to generate information on the organization of chromosomes and the level of chromosomal divergence between species, in addition to adaptive aspects and potentialities applicable to interspecific hybridization practices. All species presented 2n = 48, of which only three have karyotypes not composed exclusively of acrocentric elements. Karyotype conservatism, only disturbed by pericentric inversions, extends to the organization of repetitive DNA where the repetitive sequences analyzed here reflect a low evolutionary dynamism. The cytogenetic and genomic divergence between species due to common ancestry that occurred in periods from 15 to 11 M.a apparently provides a reduced level of post-zygotic reproductive block, with valuable implications for the hybridization induced in aquaculture. From a biogeographic and phylogenetic perspective, the most basal groups in the Atlantic Ocean demonstrate 2n = 48a as the ancestral pattern for the family, indicating that speciation in the group was not followed by significant karyotype changes and despite a relative frequency of divergent karyotypes, this group of fish is adapted to an evolutionary pattern of karyotype stasis. Keywords: Groupers, karyotype evolution, interspecific hybridization, repetitive DNA, karyotype
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