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Dissertação-Leandro-Araujo-Argôlo UNIVERSIDADE ESTADUAL DO SUDOESTE DA BAHIA PROGRAMA DE PÓS-GRADUAÇÃO EM GENÉTICA, BIODIVERSIDADE E CONSERVAÇÃO ANÁLISES GENÉTICAS PARA RESOLUÇÃO DE INCERTEZAS TAXONÔMICAS E IDENTIFICAÇÃO DE UNIDADES EVOLUTIVAS EM PEIXES PLEURONECTIFORMES DE ESTUÁRIOS DO BRASIL LEANDRO ARAUJO ARGÔLO PPGGBC Jequié-BA 2015 LEANDRO ARAUJO ARGÔLO ANÁLISES GENÉTICAS PARA RESOLUÇÃO DE INCERTEZAS TAXONÔMICAS E IDENTIFICAÇÃO DE UNIDADES EVOLUTIVAS EM PEIXES PLEURONECTIFORMES DE ESTUÁRIOS DO BRASIL Dissertação de mestrado apresentada ao Programa de Pós-graduação em Genética, Biodiversidade e Conservação da Universidade Estadual do Sudoeste da Bahia, para obtenção do Título de Mestre em Genética, Biodiversidade e Conservação. Orientador: Prof. Dr. Paulo Roberto Antunes de Mello Affonso Co-orientadora: Profa. Dra. Iracilda Sampaio PPGGBC Jequié-BA 2015 3 LEANDRO ARAUJO ARGÔLO ANÁLISES GENÉTICAS PARA RESOLUÇÃO DE INCERTEZAS TAXONÔMICAS E IDENTIFICAÇÃO DE UNIDADES EVOLUTIVAS EM PEIXES PLEURONECTIFORMES DE ESTUÁRIOS DO BRASIL Aprovada em ____/____/____ Banca Examinadora Dr. Paulo Roberto Antunes de Mello Affonso (orientador) Universidade Estadual do Sudoeste da Bahia - UESB Dra. Jamille de Araújo Bitencourt Universidade Estadual do Sudoeste da Bahia - UESB PPGGBC Dr. Henrique Batalha Filho Universidade Federal da Bahia - UFBA Programa de Pós-Graduação em Genética, Biodiversidade e Conservação Universidade Estadual do Sudoeste da Bahia – Jequié, BA 4 PPGGBC Dedico à Silvia Britto, por todo o esforço e auxílio neste trabalho, e à minha família, pelo apoio e liberdade para fazer o que amo. 5 AGRADECIMENTOS Agradeço à Universidade Estadual do Sudoeste da Bahia (UESB) e ao Programa de Pós- Graduação em Genética, Biodiversidade e Conservação (PPGGBC) pelo apoio e oportunidade de desenvolver este trabalho. À Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) pela concessão da bolsa. Ao meu orientador Dr. Paulo Roberto Antunes de Mello Affonso, pela inestimável amizade e orientação sempre impecável. À Dra. Iracilda Sampaio pela disponibilização do laboratório para realização do sequenciamento e à Luciana Watanabe pelo grande auxílio no estágio. Agradeço também a todos os amigos em Bragança pela imensa hospitalidade e disposição, especialmente à Adriane Freitas e Priscila Santos. Ao Dr. Robson Tamar da Costa Ramos pela imprescindível contribuição ao trabalho. À Dra. Jamille de Araújo Bitencourt, pela amizade, por ser minha “segunda orientadora”, me ensinando grande parte do que sei na área de pesquisa hoje, e por toda a ajuda. Obrigado! À minha melhor amiga e namorada Silvia Britto Barreto por estar sempre ao meu lado, por me auxiliar tanto e por toda a contribuição aos trabalhos. Você foi a pessoa mais importante nesta jornada. Aos amigos de curso ou laboratório Lídia, Henrique, Leo Aragão, Lorena, Fabilene, Aline, Juliani, Renata, Alexandre Falcão, Josivanda, Fernando, Josi, Arlete, Zaline, Ana Paula, Nathanna, Márcia, Lands, Mirian e todos os outros que tornaram esta etapa sempre divertida e prazerosa. A todos os professores que contribuíram com minha formação e crescimento durante estes anos, MarceloPPGGBC Cervini, Caroline Garcia, Ana Maria, Eduardo Lazzarin, Luiza Soares, Christine Steiner, Paulo Carneiro, Juvenal Cordeiro, Sérgio Siqueira e Débora Diniz. A meus pais e meus irmãos, pela educação que me deram, pelas influências, amizade, e por me fazer quem sou hoje. Vocês são tudo pra mim! 6 BIOGRAFIA Leandro Araujo Argôlo, filho de Evandro Argôlo e Antônia Araujo Argôlo, nasceu em 06 de fevereiro de 1992, em Jequié, Bahia. Cursou o Bacharelado em Ciências Biológicas com ênfase em Genética pela Universidade Estadual do Sudoeste da Bahia (UESB) entre os anos de 2009 e 2013. Durante a graduação, foi estagiário voluntário e, posteriormente, bolsista de iniciação científica pela FAPESB (2011-2012) no Laboratório de Citogenética Animal, sob orientação do Prof. Dr. Paulo Roberto Antunes de Mello Affonso. Ao fim da graduação, em abril de 2013, apresentou sua monografia intitulada: “Análise citogenética em Cichlasoma sanctifranciscense (Perciformes, Cichlidae)”. Imediatamente após a conclusão da sua graduação, ingressou no Programa de Pós-Graduação em Genética, Biodiversidade e Conservação (PPGGBC), da UESB, permanecendo sob orientação do Prof. Dr. Paulo Roberto Antunes de Mello Affonso e sendo bolsista CAPES. Ao longo do curso, desenvolveu pesquisas na área de Citogenética e Biologia Molecular em Pleuronectiformes da costa do Brasil. PPGGBC 7 PPGGBC “A Ciência é mais do que um conjunto de conhecimentos, é uma forma de pensar” (Carl Sagan) 8 RESUMO A ordem Pleuronectiformes constitui um grupo peculiar de peixes conhecidos como linguados ou solhas e caracterizados por sua anatomia diferenciada. Na fase adulta, o corpo destes organismos é altamente comprimido, bilateralmente assimétrico e com os dois olhos localizados no mesmo lado. Entretanto, a taxonomia dos representantes dessa ordem é bastante controversa em razão da sobreposição de caracteres morfológicos e variação intraespecífica, causando problemas de identificação pelos métodos tradicionais e subestimando a riqueza do grupo. Tais incertezas quanto à diversidade dos Pleuronectiformes são agravadas pela associação frequente destes animais com ecossistemas costeiros altamente impactados, como os estuários e pela exploração pesqueira. Mesmo diante desses fatores, poucos estudos que utilizem ferramentas variadas de análise da diversidade e variabilidade dos Pleuronectiformes estão disponíveis. Por outro lado, em conjunto com os dados morfológicos, a análise das espécies por ferramentas citogenéticas e moleculares representa uma abordagem informativa para resolução taxonômica em peixes, incluindo grupos historicamente controversos. Assim, este trabalho objetivou estabelecer as unidades taxonômicas e/ou evolutivas em representantes de diferentes famílias de Pleuronectiformes da costa brasileira, com ênfase em ecossistemas estuarinos por meio do uso do DNA barcoding, análises morfológicas e citogenéticas. Os resultados aqui apresentados sobre dados morfológicos e moleculares confirmam a eficiência do DNA barcoding em Pleuronectiformes e reforçam o poder da taxonomia integrativa na identificação de unidades evolutivas e resolução de inconsistências de dados com base em um único tipo de análise. No caso do gênero Bothus, os dados cromossômicos revelaram uma homogeneidade para cariótipos de Bothus ocellatus (2n=32) ao longo da costa brasileira, além de um padrão pouco usual em Bothus sp. (2n=44), ampliando a variação do número diplóide para o gênero. Da mesma forma, as análises morfológicas e moleculares indicam a presença de uma nova espécie desse o gênero com maior similaridade morfológica à B. lunatus e íntima relação genética com B. pantherinus proveninente dos oceanos Índico e Pacífico. Juntamente com cacterísticas larvais e biogeográficas, os dados genéticos em Bothus apoiam um cenário de especiação em anel com origem e dispersão do ancestral de Bothus sp. pela corrente de Benguela. Os resultados deste trabalho demonstram a necessidade de ampliação de estudos na ordem para o litoral brasileiro e reforçam a necessidade de revisões taxonômicas detalhadas para o grupo. Palavras-chave: Barcoding, citotaxonomia, DNA, Pleuronectiformes, sistemática. PPGGBC 9 ABSTRACT The order Pleuronectiformes encompasses a unique fish group, known as flounder or flatfish, with differentiated anatomical features. When adults, these fish present highly compressed and asymmetrical body with both eyes located on the same body side. Nonetheless, the taxonomy of species in this order is controversial because of overlapping of morphological traits and intraspecific variation thus hindering a proper identification by traditional methods and underestimating their actual richness. Such uncertainties about the diversity in Pleuronectiformes are critical since they inhabit highly threatened coastal environments, such as estuaries, besides being exploited in fisheries. Even though, few studies based on distinct tools to access the diversity and variability of Pleuronectiformes are available. On the other hand, morphological studies associated with cytogenetic and molecular analyses provide an informative approach to resolve taxonomic issues in fish, including controversial groups. Therefore, in the present work, we attempted to establish the taxonomic/evolutionary units in representatives of distinct families in Pleuronectiformes from Brazilian coast with emphasis on estuarine ecosystems by DNA barcoding, morphological and cytogenetic analyses. The results based on morphological and molecular data confirmed the efficacy of DNA barcoding in flatfish and reinforce the utility of integrative taxonomy in the identification of evolutionary units and resolution of inconsistent data based on a single dataset. In the case of Bothus, the chromosomal data revealed a karyotypic homogeneity in Bothus ocellatus (2n=32) throughout the Brazilian coast besides an unusual in Bothus sp. (2n=44), thus increasing the variation in diploid number within this genus. Likewise, both morphological and molecular analyses indicated the presence of a new species in this genus, being morphologically similar to B. lunatus but genetically related to B. pantherinus from Indian and Pacific oceans. Along with larval traits and biogeographic features, the genetic data in Bothus support the hypothesis of ring speciation with origin and dispersal of Bothus sp. ancestor via Benguela current. The present results show that studies in Pleuronectiformes should be intensified
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