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Diversidade Cromossômica E Padrões Ecomorfológicos Em Gobiidae (Perciformes) No Litoral E Ilhas Oceânicas Do Brasil

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Diversidade Cromossômica E Padrões Ecomorfológicos Em Gobiidae (Perciformes) No Litoral E Ilhas Oceânicas Do Brasil UNIVERSIDADE FEDERAL DO RIO GRANDE DO NORTE CENTRO DE BIOCIENCIAS PROGRAMA DE PÓS-GRADUAÇÃO EM ECOLOGIA Diversidade cromossômica e padrões ecomorfológicos em Gobiidae (Perciformes) no litoral e ilhas oceânicas do Brasil Paulo Augusto de Lima Filho Natal - RN 2015 Diversidade cromossômica e padrões ecomorfológicos em Gobiidae (Perciformes) no litoral e ilhas oceânicas do Brasil Tese apresentada ao Programa de Pós- graduação em Ecologia/PPGE da Universidade Federal do Rio Grande do Norte, como parte dos requisitos para obtenção do título de Doutor em Ecologia. Paulo Augusto de Lima Filho Orientador: Prof. Dr. Wagner Franco Molina Universidade Federal do Rio Grande do Norte – UFRN Natal - RN 2015 Catalogação da Publicação na Fonte. UFRN / Biblioteca Setorial do Centro de Biociências Lima Filho, Paulo Augusto de. Diversidade cromossômica e padrões ecomorfológicos em Gobiidae (Perciformes) no litoral e ilhas oceânicas do Brasil / Paulo Augusto de Lima Filho. – Natal, RN, 2015. 155 f.: il. Orientador: Prof. Dr. Wagner Franco Molina. Tese (Doutorado) – Universidade Federal do Rio Grande do Norte. Centro de Biociências. Programa de Pós-Graduação em Ecologia. 1. Evolução cromossômica – Tese. 2. Estratificação populacional. – Tese. 3. Dimorfismo sexual. – Tese. I. Molina, Wagner Franco. II. Universidade Federal do Rio Grande do Norte. III. Título. RN/UF/BSE-CB CDU 575 Diversidade cromossômica e padrões ecomorfológicos em Gobiidae (Perciformes) no litoral e ilhas oceânicas do Brasil Paulo Augusto de Lima Filho Tese apresentada ao Programa de Pós- graduação em Ecologia/PPGE da Universidade Federal do Rio Grande do Norte, como parte dos requisitos para obtenção do título de Doutor em Ecologia. _________________________________________________ Prof. Dr. Luiz Antônio Carlos Bertollo (Membro externo – UFSCar) _________________________________________________ Prof. Dr. Marcelo de Bello Cioffi (Membro externo – UFSCar) _________________________________________________ Prof. Dr. Vitor Oliveira Lunardi (Membro externo – UFERSA) _________________________________________________ Prof. Dr. José Garcia Junior (Membro externo – IFRN) _________________________________________________ Prof. Dr. Wagner Franco Molina (Presidente da Banca – UFRN) “Aprender é algo que nunca se cansa, nunca se tem medo e nunca se arrepende”. (Leonardo da Vinci) “A meu pai e mães, familiares, amigos e colegas, pelo apoio, confiança e paciência, dedico”. AGRADECIMENTOS Não só nesta trajetória, mas durante toda a vida em cuja presença eu estou agora, agradeço a Deus, pois me presenteia com a humildade, simplicidade, amor e felicidade. À presença familiar que me traz compreensão, carinho e paciência. Pais, Paulo Augusto de Lima, Maria da Conceição Dutra de Andrade e ―boadrasta‖ Geovana da Silva Nascimento. Às mães que sempre me apoiam e auxiliam: Dailva Bezerra, Joilza Cortez, Fátima (Tatinha), Rita Neide, Gracinete e Ivanoska de Lucena. Aos tios e tias, dedico gratidão adicional pelo apoio neste sonho: Hugo, Dedé, Garibadi, Nercy e Socorro, que me acolheram e auxiliaram durante toda esta trajetória no Rio Grande do Norte. Aos irmãos, Henrique, Ricardo, Isaura, Tatiane e José, que me dão forte base familiar. Sobrinhos, Felipe, Victor, Ane, Júlia, Isabela, Guilherme e aos muitos primos. Aos antigos e aos novos amigos que conquistei nesta jornada, desde o Cursinho da Poli, CEDAP, graduação e pós-graduação na Universidade Federal do Rio Grande do Norte. Aos colegas residentes de repúblicas, Rogerson Rodrigues, Carlos Eduardo e Helena, amigos que me apoiaram e incentivaram durante todo desenvolvimento acadêmico. Amigos e companheiros de trabalho do Instituto Federal Educação, Ciência e Tecnologia do Rio Grande do Norte, campi Macau e Pau dos Ferros. Acadêmicos que realmente me ajudaram no desenvolvimento deste trabalho, que agora os considero como amigos e colegas de ofício. Amigos de trabalho do Laboratório de Genética de Recursos Marinhos (UFRN), pelo ambiente que promoveram e pelos ensinamentos que melhoraram decisivamente nossa pesquisa: Allyson Souza, Amanda Borges, Aparecida, Clóvis Motta, Eurico, Gideão Wagner, Inailson Costa, Ingrid, Jeanne, Jose, Juliana Galvão, Karlla Danielle, Leonardo Calado, Roberta Godoy, Pablo Martinez, Rodrigo Xavier, Uedson Jacobina e Washington Candeia. Obrigado e parabéns a todos, verdadeiramente praticaram ciência, democratizando o conhecimento e trabalhando em equipe. Agradeço, especialmente, a orientação do professor Dr. Wagner Franco Molina, por acompanhar e contribuir com nosso amadurecimento acadêmico e por nos orientar com liberdade e exigência, permitindo nosso desenvolvimento científico e a realização deste trabalho. Sumário Resumo....................................................................................................................XIII 1. Introdução..............................................................................................................16 2.Objetivos..................................................................................................................21 3. Material e métodos..............................................................................................23 4. Referências............................................................................................................32 Capítulo 1: Chromosomal and morphological divergences in Atlantic populations of the frillfin goby Bathygobius soporator (Gobiidae: Perciformes) ......................................................................................................................................37 Capítulo 2: Multiabordagem evolutiva em espécies Atlânticas do gênero Bathygobius (Gobiidae): aspectos citogenéticos, morfológicos e sequências DNA barcode ......................................................................................................................................58 Capítulo 3: Influência latitudinal no dimorfismo sexual do peixe marinho Bathygobius soporator (Gobiidae: Perciformes) ......................................................................................................................................81 Capítulo 4: Karyotype divergence and spreading of 5S rDNA sequences between genomes of two species: darter and emerald Gobies (Ctenogobius, Gobiidae) .....................................................................................................................................105 Capítulo 5: Chromosomal mapping of repetitive DNAs in Gobionellus oceanicus and Gobionellus stomatus (Gobiidae: Perciformes): a shared XX/XY system and an unusual uneven distribution of 5S rDNA sites on the Y chromosome ...................................................................................................................................119 Capítulo 6: Aspectos carioevolutivos da família Gobiidae (Perciformes) e descrição citogenética de Coryphopterus glaucofraenum e Evorthodus lyricus .....................................................................................................................................135 Considerações finais:.............................................................................................154 LISTA DE FIGURAS INTRODUÇÃO Figura 1. Mapa indica os pontos do litoral e ilhas oceânicas brasileiras 24 utilizados para coleta de Gobiidae. Figura 2. Exemplares da família Gobiidae utilizados nas análises 25 citogenéticas. Bathygobius soporator (a), Bathygobius sp.1 (b), Bathygobius sp.2 (c), Ctenogobius smaragdus (d), Ctenogobius boleosoma (e), Coryphopterus glaucofraenum (f), Evorthodus lyricus (g), Gobionellus oceanicus (h) e Gobionellus stomatus (i). Barras= 1 cm. CAPÍTULO I Figura 1. Collection sites for B. soporator along the Brazilian coast and Rocas 40 Atoll sample (a); B. soporator specimen with the nine landmarks used for interpopulation morphological analysis (b). Schematic description of landmarks: 1. Distal extremity of the premaxillary bone; 2. Start of the first dorsal fin; 3. End of the first distal fin; 4. Start of the second dorsal fin; 5. End of the second dorsal fin; 6. End of the anal fin; 7. Start of the anal fin; 8. Start of the pelvic fin; 9. Medial position of the eye. Bar= 1,5 cm. Figura 2. Karyotypes of Bathygobius soporator populations MA, BA, RN and 43 RA arranged from sequentially Giemsa-stained (left) and C-banded chromosomes (right). Chromosomes bearing Ag-NOR, CMA3 + and DAPI− sites are boxed. Bar= 5 μm. Figura 3. Karyotypes of the Bathygobius soporator populations MA, BA, RN 44 and RA arranged from chromosomes probed with 5S rDNA (green), H1 hisDNA (red) counterstained with DAPI. Chromosomes carrying 18S rDNA (red) are boxed. Bar= 5 μm. Figura 4. Distribution of morphological data of B. soporator samples along 47 the first and second canonical variables. The largest highlighted symbols indicate the morphometric mean of each population; (a) all populations including Rocas Atoll individuals (CV1=63.5% and CV2=30.5%); (b) only continental populations (CV1=76.4% and CV2=23.5%). Schematic drawings at c, d, e and f represent comparisons between morphotypes of the populations analyzed generated by CV1. CAPÍTULO II Figura 1 Pontos de coleta de Bathygobius, Rio Grande do Norte (RN), Atol das 61 Rocas (RA) e Arquipélago de Fernando de Noronha (FNA). Figura 2. Espécies do gênero Bathygobius coletadas em áreas do litoral do 62 Brasil e ilhas oceânicas e analisadas quanto a padrões citogenéticos, morfológicos e das sequências mitocondriais COI. Barra= 1 cm. Figura 3. Cariótipos de B. soporator, Bathygobius sp.1, e Bathygobius sp.2 66 (Atol
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