Posicionamento Filogenético De Chaetognatha Baseado Em Dados Morfológicos

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Posicionamento Filogenético De Chaetognatha Baseado Em Dados Morfológicos Rudy Camilo Nunes Posicionamento filogenético de Chaetognatha baseado em dados morfológicos Dissertação apresentada como requisito final à obtenção do grau de Mestre em Ciências Biológicas, área de concentração em Zoologia. Curso de Pós-Graduação em Ciências Biológicas, Zoologia, Departamento de Sistemática e Ecologia da Universidade Federal da Paraíba. Orientador: Prof. Dr. Martin Lindsey Christoffersen. João Pessoa 2012 Rudy Camilo Nunes Posicionamento filogenético de Chaetognatha baseado em dados morfológicos Dissertação apresentada como requisito final à obtenção do grau de Mestre em Ciências Biológicas, área de concentração em Zoologia. Curso de Pós-Graduação em Ciências Biológicas, Zoologia, Departamento de Sistemática e Ecologia da Universidade Federal da Paraíba. Orientador: Prof. Dr. Martin Lindsey Christoffersen. João Pessoa 2012 i Agradecimentos Agradeço ao orientador Prof.º Dr. Martin Lindsey Christoffersen, pela infinita paciência e pelo apoio durante todo o processo. Pelo conhecimento acumulado em décadas de pesquisa e dedicação que, tão facilmente, me foi doado ao longo de nossas conversas formais e informais. E, sobretudo, pela relação de amizade e confiança construída durante o período em que trabalhamos juntos. A Prof.ª Dr. Maria Nazaré Stevaux (UFGO), que durante a graduação me apresentou a Biologia Evolutiva, fato que me proporcionou dar um sentido real ao estudo da história biológica do nosso planeta. A minha esposa Angélica Pereira Soares, pelo companheirismo e lealdade incondicionais. Por compartilhar de todas as dificuldades e conquistas vividas desde o início desta etapa. Pelo amor, carinho e dedicação empenhados todos os dias. Aos meus pais, irmãos e familiares que, em diferentes níveis e momentos, forneceram a ajuda necessária para que se rompessem as barreiras existentes até aqui. Aos grandes amigos, que hoje fazem parte da família, Carolina Caldas, Fabiano Nogueira, Filipe Eduardo Macedo Menezes, Lucas Aguiar Rosa e Marianne Krela por todos os dias em que vivemos juntos a mesma vida. Agradeço a eles pelas lindas tardes nas praias paraibanas, pela paz de espírito e sabedoria oferecidos em diversas situações do dia-a-dia e pelas inúmeras histórias vividas, e que sempre serão lembradas. Aos grandes amigos Danilo Rada, Flávia Moura, Hugo Fernandes, Luana Poliseli Ramos e Newton Mota Gurgel Filho pela parceria e pelos momentos de confraternização, pelas bebedeiras intelectualizadas ou não e por toda a diversão que proporcionaram. Foram vários os momentos em que a amizade de vocês foi decisiva pra resolver problemas pessoais, profissionais ou simplesmente pra tornar a vida mais rica e interessante. A José Eriberto de Assis, que sempre esteve pronto a ajudar em todas as situações, mesmo antes do meu ingresso no programa. Pelas boas discussões acerca da Sistemática Filogenética e dos aspectos evolutivos dos metazoários e pela participação decisiva em vários momentos importantes. A todos os colegas e amigos do Programa de Pós-Graduação em Ciências Biológicas (Zoologia) da Universidade Federal da Paraíba. A CAPES, pela bolsa de mestrado que pagou minha estadia na cidade de João Pessoa durante dois anos, e que deixou muita saudade após o seu término. ii Sumário Agradecimentos .............................................................................................................. ii Lista de tabelas ............................................................................................................... v Lista de figuras ............................................................................................................... vi Resumo ........................................................................................................................... 01 Abstract ........................................................................................................................... 02 1. Introdução ................................................................................................................... 03 1.1. Caracterização morfológica do grupo ..................................................................... 04 Caracteres externos .............................................................................................. 05 Parede corporal e musculatura ............................................................................ 06 Celoma e estruturas mesodérmicas ...................................................................... 07 Sistema nervoso e órgãos sensoriais .................................................................... 08 Sistema digestivo ................................................................................................... 10 Reprodução e desenvolvimento ............................................................................. 10 Registro fóssil ........................................................................................................ 12 1.2. Histórico do posicionamento filogenético e classificação do grupo ....................... 13 2. Metodologia ................................................................................................................ 15 2.1. Escolha dos grupos externos e composição do grupo interno ................................. 16 2.2. Fonte de caracteres .................................................................................................. 19 Codificação ........................................................................................................... 20 2.3. Polarização dos caracteres ....................................................................................... 20 2.4. Análise dos dados e reconstrução da árvore filogenética ........................................ 21 iii 3. Resultados ................................................................................................................... 22 3.1. Evidências de monofilia dos táxons terminais ........................................................ 28 3.2. Discussão dos caracteres ......................................................................................... 30 4. Discussão geral ........................................................................................................... 56 5. Conclusões .................................................................................................................. 73 6. Referências ................................................................................................................. 74 iv Lista de tabelas Tabela 1. Matriz gerada a partir de 25 caracteres morfológicos. Os caracteres foram codificados com “0-5” para os diferentes estados apresentados. Caracteres inaplicáveis foram codificados com “-” e caracteres duvidosos ou desconhecidos com “?”. Todos os caracteres foram tratados como não ordenados durante a análise. ................................. 25 Tabela 2. Lista de caracteres contendo 25 caracteres morfológicos (sendo 17 multiestado e 8 binários). Os caracteres foram codificados com “0-5” para os diferentes estados apresentados. ...................................................................................................... 25 Tabela 3. Matriz contendo os 25 caracteres morfológicos reanalisados e ordenados com base no cladograma da Fig. 1, gerado a partir da análise inicial dos caracteres não- ordenados. Os caracteres foram recodificados com “0-5”, onde “0” representa o estado plesiomórfico e “1-5” representam os estados apomórficos sucessivos de cada caráter. Caracteres inaplicáveis foram codificados com “-”. ...................................................... 27 Tabela 4. Lista de caracteres reordenados com base na árvore mais parcimoniosa obtida nesta análise (Fig. 1). Aqui, os caracteres foram codificados com “0” para o estado plesiomórfico e “1-5” para os diferentes estados apomórficos sucessivos. ................... 27 v Lista de figuras Fig. 1. Árvore mais parcimoniosa, gerada a partir de 25 caracteres morfológicos (tamanho: 59; CI: 0,91 e RI: 0,90). Os números abaixo de cada ramo indicam, respectivamente, os valores de suporte bootstrap e jacknife, obtidos de forma heurística, com 1000 pseudoréplicas e com as mesmas especificações utilizadas na obtenção do cladograma. Os números acima de cada ramo representam os caracteres que sustentam cada clado recuperado. ................................................................................................... 23 Fig. 2. Distribuição dos caracteres de Chaetognatha ao longo do sistema filogenético de Radialia + Pogonophora e Oweniida, demonstrando sua ligação com diferentes níveis hierárquicos dentro deste grupo. a) sistema nervoso intraepitelial [sni]; b) enterocelia [en]; c) clivagem radial [cr]; d) deuterostomia [de]; e) fotorreceptores ciliados [fc]; f) ausência de tentáculos mesossomais [atm], presença de células multiciliadas em tecidos ectodérmicos e endodérmicos [cm]; g) nadadeira caudal pós-anal [ncp], presença de apenas dois tagmas corporais [dt], intestino linear com ânus ventral pré-anal [avpa]; h) ausência de estágio larval e metamorfose [elm], fibras musculares transversalmente estriadas [me]; i) simetria bilateral perfeita [sb], sistema lateral de órgãos ciliares sensoriais [ocs], intestino muscularizado [im], predação ativa [pa], presença de epitélio multiestratificado [em]. .................................................................................................. 67 vi Resumo As relações evolutivas de Chaetognatha foram testadas com a utilização de métodos filogenéticos e a filogenia de Deuterostomia foi reconstruída.
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