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TESE Ralf Tarciso Silva Cordeiro.Pdf 0 llp UNIVERSIDADE FEDERAL DE PERNAMBUCO CENTRO DE CIÊNCIAS BIOLÓGICAS PROGRAMA DE PÓS-GRADUAÇÃO EM BIOLOGIA ANIMAL RALF TARCISO SILVA CORDEIRO REVISÃO TAXONÔMICA DO GÊNERO Plexaurella KÖLLIKER, 1865 (CNIDARIA: ANTHOZOA: OCTOCORALLIA) BASEADA EM DADOS MORFOLÓGICOS E MOLECULARES Recife 2018 1 RALF TARCISO SILVA CORDEIRO REVISÃO TAXONÔMICA DO GÊNERO Plexaurella KÖLLIKER, 1865 (CNIDARIA: ANTHOZOA: OCTOCORALLIA) BASEADA EM DADOS MORFOLÓGICOS E MOLECULARES Tese apresentada ao Programa de Pós- Graduação em Biologia Animal da Universidade Federal de Pernambuco, como requisito parcial para obtenção do título de Doutor em Biologia Animal. Área de Concentração: Zoologia Orientador: Prof. Dr. Carlos Daniel Pérez Co-orientador: Prof. Dr. Antonio M. Solé-Cava Recife 2018 2 Dados Internacionais de Catalogação na Publicação (CIP) de acordo com ISBD Cordeiro, Ralf Tarciso Silva Revisão taxonômica do gênero Plexaurella Kölliker, 1865 (Cnidaria: Anthozoa: Octorallia) baseada em dados morfológicos e moleculares/ Ralf Tarciso Silva Cordeiro- 2018. 156 folhas: il., fig., tab. Orientador: Carlos Daniel Perez Coorientador: Antonio M. Solé-Cava Tese (doutorado) – Universidade Federal de Pernambuco. Centro de Biociências. Programa de Pós-Graduação em Biologia Animal. Recife, 2018. Inclui referências 1. Octocorallia 2. Zoologia- classificação 3. Recifes e Ilhas de Coral I. Perez, Carlos Daniel (orient.) II. Solé-Cava, Antonio M. (coorient.) III. Título 593.6 CDD (22.ed.) UFPE/CB-2018-233 Elaborado por Elaine C. Barroso CRB4/1728 3 RALF TARCISO SILVA CORDEIRO REVISÃO TAXONÔMICA DO GÊNERO Plexaurella KÖLLIKER, 1865 (CNIDARIA: ANTHOZOA: OCTOCORALLIA) BASEADA EM DADOS MORFOLÓGICOS E MOLECULARES Tese apresentada ao Programa de Pós- Graduação em Biologia Animal da Universidade Federal de Pernambuco, como requisito parcial para obtenção do título de Doutor em Biologia Animal. Aprovada em: 27/02/2018 COMISSÃO EXAMINADORA ___________________________________________ Prof. Dr. Clovis Barreira e Castro – Titular Externo (MNRJ/ UFRJ) ___________________________________________ Prof. Jesser Fidelis de Souza Filho – Titular Externo (UFPE/ PPGO) __________________________________________ Prof. Dra. Paula Braga Gomes – Titular Interno (PPGBA) __________________________________________ Prof. Dr. Ulisses dos Santos Pinheiro – Titular Interno (PPGBA) __________________________________________ Prof. Dr. Leandro Manzoni Vieira – Titular Interno (PPGBA) 4 Às mulheres da minha vida 5 AGRADECIMENTOS A Deus, pelo modo como tem guiado minha vida, e às mulheres da minha vida (minha mãe, Ana Maria, e minha esposa, Karina Bezerra). A todos os amigos de fora da academia, principalmente os mais próximos, que me ajudam a viver o aqui e o agora, Caio Weimann, Mário Matos, Mary Lima, Augusto Albuquerque e Lidiane Macedo. Ao meu orientador, Carlos Pérez, pelo apoio, oportunidades e amizade de sempre. Aos amigos do Grupo de Pesquisas em Antozoários que convivi nesses últimos anos, David Oliveira, Felipe Campos, Gislaine Lima, Manuela Menezes, Erica Patrícia, Érika Crispim, Rafael Brandão, Alessandra Targino e a todos os que já se foram e aos que estão chegando. À Dra. Sigrid Neumam Leitão e Dr. Jesser Fidelis, por abrirem as portas do Museu de Oceanografia Petrônio Alves Coelho (UFPE). A todos do Laboratório de Zooplâncton que me acolheram nesses dois últimos anos e me trataram como família: Renata Campelo, Claudeilton Santana, Gabriela Figueiredo, Natália Lins, Igor Teixeira, Cynthia Lima, Morgana Lolaia e Lucas Figueredo. Em especial a Gleice Santos e Simone Lira, irmãs de longa data. Ao Dr. Stephen Cairns, pela amizade, tempo dedicado, conhecimentos transmitidos e acesso ao National Museum of Natural History – EUA. Ao Chad Walter, pela hospitalidade, paciência e aulas de inglês. Ao Dr. Herman Wirshing, pelas dicas nas análises moleculares e ajuda no LAB (NMNH). Também à toda família Cairns, pelo acolhimento e hospitalidade e aos amigos brasileiros que fiz durante a estadia nos EUA. Ao meu co-orientador Antonio Solé-Cava, pela amizade e conversas, assim como por abrir as portas de sua casa e laboratório. A todos que ajudaram, deram sua opinião ou apenas se mostraram solícitos durante os últimos quatro anos: Dr. Clovis Castro, Dr. Marcelo Kitahara, Dra. Liany Melo, Dra. Ghennie Rodríguez, Maria Eduarda Alves (Duda), Dr. Manfred Grasshoff, Dr. Juan Sánchez, Dr. Phil Alderslade, Dr. Ernesto Weil e Dra. Rosa Rodríguez-Martínez. Agradeço a todos os técnicos e curadores das seguintes coleções: Museu Nacional – Rio de Janeiro (Genivaldo Teixeira, em especial), National Museum of Natural History – Smithsonian Institution, Zoologisches Museum – Berlim, Peabody Museum of Natural History – Yalle, Museum of Comparative Zoology – Harvard. Sem eles, qualquer trabalho taxonômico seria inviável. 6 Sou grato também aos membros da banca, que se dispuseram de antemão a ler e criticar o presente texto. Ao ICMBio (Instituto Chico Mendes de Conservação da Biodiversidade), especificamente à Maurizélia Brito (Zelinha) e Fred Osório, cujo carinho e apoio não faltaram durante os 30 dias na Reserva Biológica do Atol das Rocas. À Universidade Federal de Pernambuco, pela estrutura e suporte fornecidos. À CAPES (Coordenação de Aperfeiçoamento de Pessoal de Nível Superior) e FACEPE (Fundação de Amparo a Ciência e Tecnologia de Pernambuco), pelas bolsas de estudo concedidas (PDSE 99999.006617/2015-08 e IBPG-0558-2.04/13, respectivamente). Evitando o risco de ser ainda mais injusto, se algum nome aqui foi esquecido, peço que me perdoe. Memória é uma das muitas coisas que perdi durante o doutorado, mas o amor permanece. Muito obrigado! Thank you ∞ Muchas gracias ∞ Vielen dank ∞ Grazie Mille ∞ Merci beaucoup 7 RESUMO Octocorais são engenheiros ecossistêmicos importantes em ambientes marinhos profundos e rasos. Gorgônias são octcorais particularmente diversificados e sempre presentes em ambientes recifais ao redor do mundo. Os representantes do gênero Plexaurella estão entre os gêneros mais conspícuos de gorgonias em recifes do Atlântico Ocidental, no entanto, o conhecimento atual sobre a diversidade do gênero é baseado em uma série de revisões sem consultas a espécimes-tipo. Além disso, mesmo níveis de classificação taxonômica superiores (subfamílias, famílias, subordens, etc.) são atualmente incertos, uma vez que reconstruções filogenéticas não dão suporte à classificação atual de Plexaurella dentro da família Plexauridae e o posicionam próximo à família Gorgoniidae, também polifilética. Assim, o presente estudo revisa o gênero, baseado no exame de espécimes-tipo disponíveis e testa sua monofilia, usando marcadores moleculares mitocondriais e um nuclear (COI+igr+mtMutS e 28S, respectivamente). Até o momento, até seis species eram consideradas válidas. Nossos resultados demonstram que o grupo é composto por pelo menos sete espécies válidas previamente descritas: P. dichotoma, P. nutans, P. grisea, P. teres, P. grandiflora, P. regia e P. obesa; e uma nova: Plexaurella rastrera sp. nov. A classificação atual do gênero é discutida e, baseado em reconstruções filogenéticas e distâncias genéticas, é proposta a elevação de Plexaurellinae, previamente uma subfamília de Plexauridae, ao nível de família. Embora Gorgoniidae tenha sido recuperado como polifilética, vários gêneros, incluindo Gorgonia, foram agrupados num clado monofilético, com o qual Plexaurellidae é relacionado. Palavras-chave: Alcyonacea. Gorgônias. Octocorais recifais. Plexauridae. Gorgoniidae. 8 ABSTRACT Octocorals are important ecosystem engineers in deep and shallow marine environments. Gorgonians are particularly diversified octocorals and ubiquitous in reef environments around the world. Representatives of the genus Plexaurella are amongst the most conspicuous genera of gorgonians in the western Atlantic reefs; however, current knowledge on the diversity of the genus is based on a series of revisions lacking examination of type-specimens. Furthermore, even higher classification levels (subfamilies, families, suborders, etc.) are currently uncertain, as phylogenetic reconstructions do not support the current classification of Plexaurella within the family Plexauridae and places it close to the polyphyletic family Gorgoniidae. Thus, this study reviews that genus based on examination of available types and test its monophyly using molecular markers, both mitochondrial and nuclear (COI+igr+mtMutS and 28S, respectively). Until now, up to six species were considered valid. Results show that the group is composed by at least seven previously described species: P. dichotoma, P. nutans, P. grisea, P. teres, P. grandiflora, P. regia and P. obesa; and one new: Plexaurella rastrera sp. nov. The current classification of the genus is discussed and, based on phylogenetic reconstructions and genetic distances, is proposed the elevation of Plexaurellinae, formerly a subfamily of Plexauridae, to the family level. Although Gorgoniidae has been recovered as polyphylethic, several genera, including Gorgonia, were grouped within a monophyletic clade, of which Plexaurellidae is closely related. Key words: Alcyonacea. Gorgonians. Reef octocorals. Plexauridae. Gorgoniidae. 9 SUMÁRIO 1 INTRODUÇÃO........................................................................................................10 1.2 OBJETIVOS............................................................................................................12 1.2.1 Objetivo Geral.....................................................................................................12 1.2.2 Objetivos específicos...........................................................................................12
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