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Bruno Cavalcanti Vila Nova De Albuquerque Universidade Federal de Pernambuco Bruno Cavalcanti Vila Nova de Albuquerque Preparação, re-descrição e posicionamento filogenético de Cearadactylus atrox Leonardi & Borgomanero, 1985 (Archosauria, Pterosauria) Orientador: Prof. Dr. Virgínio Henrique de Miranda Lopes Neumann Recife / 2010 Universidade Federal de Pernambuco Centro de Tecnologia e Geociências Departamento de Geologia Programa de Pós-Graduação em Geociências Preparação, re-descrição e posicionamento filogenético de Cearadactylus atrox Leonardi & Borgomanero, 1985 (Archosauria, Pterosauria) Dissertação apresentada ao programa de Pós-Graduação em Geociências da Universidade Federal de Pernambuco, como parte dos requisitos necessários para a obtenção do título de Mestre em Geociências. Aluno: Bruno Cavalcanti Vila Nova de Albuquerque Orientador: Virgínio Henrique de Miranda Lopes Neumann Recife, 2010 A345p Albuquerque, Bruno Cavalcanti Vila Nova de. Preparação, re-descrição e posicionamento filogenético de Cearadactylus atrox Leonardi & Borgomanero, 1985 (Archosauria, Pterosauria) / Bruno Cavalcanti Vila Nova de Albuquerque. - Recife: O Autor, 2010. xi, 91 folhas, il : figs. Dissertação ( Mestrado) – Universidade Federal de Pernambuco. CTG. Programa de Pós-Graduação em Geociências, 2010. Orientador: Profº. Virgínio Henrique de Miranda Lopes Neumann. Inclui Bibliografia e Anexos. 1. Geociências. 2.Pterosauria. 3.Formação Romualdo. 4.Análise Filogenética. 5.Cearadactylus. I. Título. UFPE 551 CDD (22. ed.) BCTG/2010-137 Dedicatória Para os pterossauros, que sempre tentaram alçar vôos cada vez mais altos, e inspiram aqueles que buscam novas terras. Agradecimentos Um trabalho como esse não teria sido realizado sem a ajuda de tantos que, de uma forma ou de outra, me incentivaram a continuar em frente, e com suas opiniões e personalidades, deram nova luz ao meu trabalho: - Ao Prof. Dr. Virgínio Neumann, por ter me aceitado como aluno, confiando na minha palavra, quando “caí de pára-quedas” no DGEO. Mais do que Orientador, um verdadeiro pai, que não só me guiou no meio acadêmico, como me abriu vários caminhos e mostrou diversas oportunidades. Para ele, agradecimentos nunca serão suficientes. - A Prof. Dra. Juliana Sayão, por me iniciar na Paleontologia, nos nossos queridos Pterossauros. Começamos como Orientadora e aluno, terminamos como bons amigos. Um dia eu irei retribuir tudo o que ela fez por mim. Até lá, espero que ela aceite minha eterna gratidão. - Ao Prof. Dr. Alexander Kellner, por toda a ajuda e orientação durante a minha estadia no Rio de Janeiro, ao longo da preparação do material no Museu Nacional/UFRJ. Suas palavras sempre dão um estímulo a mais, mesmo que seja a costumeira frase “Tem que trabalhar mais Bruno!”. - Aos amigos que, desde a época do colégio sempre me incentivaram a ir cada vez mais além – Thiago Leite, Vinícius Santiago, Alberto Lima, Frederico Rolim, Iris Tébeká, Marco Antônio Padilha, Mariana Arroxelas e Pedro Nícolas – sem vocês, toda essa aventura não teria tido o menor sentido. - Aos amigos da faculdade, por continuarem fazendo aquilo que sempre fizeram melhor: serem bons amigos. Bruno Rodrigues, Carolina Machado, Daniela Moura, Laura Leal, Mirela D’arc. Cada um, do seu jeito especial, tornou essas páginas possíveis de serem escritas. Meus sinceros agradecimentos. - Aos amigos da madrugada, que na etapa final, ao escrever a Dissertação me faziam companhia através da internet noite adentro. Gabriela Coutinho, Jully Iguchi, Rômulo Maciel e Veronica Slobodian, finalmente posso dormir cedo! - A todo o Setor de Paleovertebrados do Museu Nacional/UFRJ, pela ajuda, companheirismo e incentivo durante minha estadia no Rio de Janeiro e na preparação do material estudado. A Elaine Machado, Helder Silva, Kamila Bandeira, Rodrigo Giesta e Uiara Gomes, obrigado por tudo. Gostaria de agradecer especialmente a Gustavo Oliveira, por ter me recebido no MN, e me auxiliado sempre que eu precisei. A Jéssica Silva, por me ajudar com o passo- a-passo da preparação do material. Ao Pedro Romano, pela ajuda e conselhos com as matrizes e análises filogenéticas. E a Fabiana Rodrigues e Taíssa Rodrigues, pelas dicas e ajudas em relação aos Pterossauros em geral. - A secretária da Pós-Graduação em Geociências, Elizabeth Galindo, que por diversas vezes me tirou de vários problemas e sempre esteve pronta para me ajudar. - A minha querida família, que sempre esteve por perto com uma palavra de estímulo e carinho para que eu desse o melhor de mim em todos os momentos. - Agradeço a Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) pela bolsa de Mestrado, sem a qual eu não teria conseguido concluir o projeto. - E a cada um que, em maior ou menor grau, voluntariamente ou inconscientemente, me ajudou e estimulou a ir além, e completar objetivos cada vez maiores. A todos, eu serei eternamente grato. Resumo Pterossauros são um dos grupos de arcossauros fósseis melhor representados no registro fóssil brasileiro. Dentro desta diversidade, uma das primeiras espécies descritas, Cearadactylus atrox, tem causado divergências de interpretação entre pesquisadores desde a sua publicação. Devido ao fato de o holótipo e único espécime conhecido estar parcialmente preparado, as tentativas de classificações taxonômicas realizadas, posicionaram-no em grupos diferentes. Estas discordâncias geraram o colapso de certos ramos da filogenia de Pterosauria. O presente trabalho teve como objetivo a preparação do holótipo de Cearadactylus atrox, sua re-descrição e o posicionamento filogenético, utilizando-se matrizes filogenéticas disponíveis para o grupo. Evidências de adulterações foram observadas durante a preparação. Dentre as características exclusivas apresentadas por Cearadactylus atrox estão uma fenda no dentário bifurcada na extremidade rostral, órbita e narina em posição elevada relativo a fenestra nasoanterorbital e menos de 15 dentes em cada lado da maxila e mandíbula. Filogeneticamente Cearadactylus atrox se mostrou como grupo irmão do clado Anhangueridae, sendo uma espécie intermediária entre as formas européias (nominalmente “Ornithocheirus” compressirostris) e as formas brasileiras (Tropeognathus mesembrinus e as espécies que compõem o gênero Anhanguera). Palavras-chave: Pterosauria, Formação Romualdo, Análise filogenética, Anhangueridae, Cearadactylus. Abstract Pterosaurs are one of the best represented fossil archosaurs from the Brazilian register. Among this diversity, one of the first described species, Cearadactylus atrox, has caused divergences in scientists’ interpretations since its publication. Because the holotype and only known specimen been only partially prepared, the taxonomic classifications realized have positioned it in different groups. These discordances would create the collapse of some branches in the Pterosauria phylogeny. The present work had the objective the preparation of the Cearadactylus atrox holotype, its redescription and the phylogenetic position, using phylogenetic matrixes available for the group. Adulteration evidences were observed during the preparation. Among the exclusive characteristics presented by Cearadactylus atrox is a dentary groove with bifurcate rostral end, orbit and naris in high position related to the nasoanterorbital fenestra and less than 15 teeth on each side of maxilla and mandible. Phylogeneticaly Cearadactylus atrox was shown as sister-group of the Anhangueridae clade, being one intermediate species between the European (nominally “Ornithocheirus” compressirostris) and Brazilian (Tropeognathus mesembrinus and the species which compose the Anhanguera genus) forms. Keywords: Pterosauria, Romualdo Formation, Phylogenetic Analysis, Anhangueridae, Cearadactylus. Sumário Sumário ................................................................................................................... vii Lista de Abreviações ............................................................................................... viii Lista de Figuras ...................................................................................................... ix 1 - Introdução ......................................................................................................... 01 2 - Contexto Geológico .......................................................................................... 10 3 - Objetivos ........................................................................................................... 16 4 - Material e Métodos ........................................................................................... 17 5 – Resultados ....................................................................................................... 19 5.1 – Preparação ................................................................................................... 19 5.2 – Descrição do material ................................................................................... 22 5.3 – Análises filogenéticas ................................................................................... 33 6 - Discussão ......................................................................................................... 43 7 - Conclusão ......................................................................................................... 56 8 - Referências Bibliográficas ................................................................................ 57 9 - Anexos .............................................................................................................. 67 Anexo A – Estados de caracteres para Cearadactylus atrox
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