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Universidade Federal Do Rio Grande Do Sul Instituto De Geociências Programa De Pós-Graduação Em Geociências Contribuição UNIVERSIDADE FEDERAL DO RIO GRANDE DO SUL INSTITUTO DE GEOCIÊNCIAS PROGRAMA DE PÓS-GRADUAÇÃO EM GEOCIÊNCIAS CONTRIBUIÇÃO AO CONHECIMENTO DOS PTEROSSAUROS DO GRUPO SANTANA (CRETÁCEO INFERIOR) DA BACIA DO ARARIPE, NORDESTE DO BRASIL FELIPE LIMA PINHEIRO ORIENTADOR – Prof. Dr. Cesar Leandro Schultz Porto Alegre - 2014 UNIVERSIDADE FEDERAL DO RIO GRANDE DO SUL INSTITUTO DE GEOCIÊNCIAS PROGRAMA DE PÓS-GRADUAÇÃO EM GEOCIÊNCIAS CONTRIBUIÇÃO AO CONHECIMENTO DOS PTEROSSAUROS DO GRUPO SANTANA (CRETÁCEO INFERIOR) DA BACIA DO ARARIPE, NORDESTE DO BRASIL FELIPE LIMA PINHEIRO ORIENTADOR – Prof. Dr. Cesar Leandro Schultz BANCA EXAMINADORA Prof. Dr. Marco Brandalise de Andrade – Faculdade de Biociências, PUC, RS Profa. Dra. Marina Bento Soares – Departamento de Paleontologia e Estratigrafia, UFRGS Profa. Dra. Taissa Rodrigues – Departamento de Biologia, UFES, ES Tese de Doutorado apresentada ao Programa de Pós-Graduação em Geociências como requisito parcial para a obtenção do título de Doutor em Ciências. Porto Alegre – 2014 “Ao ser destampado pelo gigante, o cofre deixou escapar um hálito glacial. Dentro havia apenas um enorme bloco transparente, com infinitas agulhas internas nas quais se despedaçava em estrelas de cores a claridade do crepúsculo. Desconcertado, sabendo que os meninos esperavam uma explicação imediata, José Arcadio Buendía atreveu-se a murmurar: – É o maior diamante do mundo.” Gabriel García Marquez AGRADECIMENTOS Um trabalho como esse não é feito apenas a duas mãos. Durante o percurso de meu mestrado e doutorado, tive o privilégio de contar com o apoio (por vezes, praticamente incondicional) de diversas pessoas. Em primeiro lugar, pelo apoio irrestrito em todos os momentos, agradeço a minha família, em especial a meus pais, Sandra e Valmiro e a meus irmãos, Fernando e Sacha. A todos meus amigos de Fortaleza, em especial a Marina, Fernanda, Luana, Janaina, Gabriel, João Paulo, Heitor, Paulo e Lucas. Aos meus amigos e companheiros de Porto Alegre, por tornarem memorável o tempo em que vivi nessa cidade: Marcel, Leonardo, Tomaz, Igor, Elker, David, Marcos, Alexandre, Marquinho, Victor, Tiago, Bruno, Pola, Fábio, Heitor e Bibiana. Aos meus amigos de Munique, em especial ao Tiago, Raul, Luísa e Alexandra. À equipe do laboratório de paleontologia da UFRGS, por tornar minha rotina tão diverdida. Vários pesquisadores tiveram importância crucial em minha formação. Dentre eles, agradeço especialmente a Oliver Rauhut, Richard Butler, Taissa Rodrigues, John Maisey, Juliana Sayão (a qual também agradeço pela compreensão e tato em um começo turbulento do doutorado) e Bruno Vila Nova. Ao meu orientador, Schultz, pelo apoio irrestrito em todas as fases de meu mestrado e doutorado e a Marina Bento Soares, pelo apoio, disponibilidade e aconselhamento. Aos três membros da banca, Marco Brandalise, Marina Soares e Taissa Rodrigues por aceitarem avaliar este trabalho em um momento tão turbulento. Eu provavelmente nunca vou conseguir agradecer o suficiente aos meus amigos Daniel, Emilia e Paula, que com certeza sabem que sem eles eu não seria metade do profissional que sou hoje. Por último, um agradecimento especial a Adriana pelo amor, compreensão incondicional e companheirismo. RESUMO Neste trabalho, contribuímos com pesquisas originais sobre a anatomia e taxonomia de pterossauros provenientes do Grupo Santana (Cretáceo Inferior, Bacia do Araripe, Nordeste do Brasil). Além disso, revisamos aspectos taxonômicos e filogenéticos do clado e a geologia do Grupo Santana é descrita a partir de uma abrangente revisão bibliográfica. A origem de Pterosauria e suas relações filogenéticas com demais tetrápodes é abordada, explicitando a tendência atual de considerar estes arcossauros como grupo-irmão de Dinosauromorpha no clado Ornithodira. As relações filogenéticas internas de Pterosauria são discutidas através da apresentação de dados das principais análises cladísticas propostas para o grupo. As relações filogenéticas de táxons provenientes do Grupo Santana são tratadas em maior detalhe, principalmente no que tange o debate sobre a monofilia ou parafilia de Tapejaridae. É realizada, também, uma revisão taxonômica detalhada e atualizada, destacando todos os táxons válidos de pterossauros do Grupo Santana. Neste trabalho, reconhecemos 27 espécies como válidas. Três trabalhos científicos originais são apresentados. No primeiro, revisamos as atribuições taxonômicas do espécime descrito como “Pricesaurus megalodon”, concluindo que este possui afinidades a Anhangueridae, em especial, ao gênero Anhanguera. No segundo trabalho, descrevemos um espécime incomum, constituído por um palato fragmentário, tentativamente atribuído a Azhdarchoidea. Em adição, contribuímos na compreensão da anatomia e evolução do palato de Pterodactyloidea, reconhecendo tendências evolutivas até então não descritas para o grupo. No terceiro e último trabalho, reavaliamos um espécime de pterossauro da Formação Romualdo, anteriormente atribuído a Anhanguera santanae, concluindo que este, na verdade, representa um táxon ainda não descrito de Anhangueridae. A ecologia de pterossauros anhanguerídeos da Formação Romualdo é discutida, principalmente no que diz respeito à presença aparentemente simultânea de várias espécies que possivelmente compartilhariam o mesmo nicho ecológico. Palavras chave: Pterossauros; Grupo Santana; Bacia do Araripe; filogenia; taxonomia; anatomia. ABSTRACT We contribute, in this work, with original research on the anatomy and taxonomy of pterosaurs from the Santana Group (Lower Cretaceous, Araripe Basin, northeastern Brazil). Also, some taxonomic and phylogenetic aspects of pterosaurs are reassessed, and the geology of the Santana Group is described through a comprehensive literature revision. Pterosaur origins and its phylogenetic relationships with other tetrapods are discussed, detailing the current tendency to consider these archosaurs as the sister group of Dinosauromorpha, within the clade Ornithodira. Moreover, the internal phylogenetic relationships of Pterosauria are presented and discussed through the analysis of the main cladistic studies of this group. Phylogenetic relationships of Santana Group taxa are addressed in detail, especially regarding the debate on the monophyly or paraphyly of Tapejaridae. Furthermore, we conduct a detailed and updated taxonomic revision, depicting all the valid Santana Group pterosaur taxa. We recognize, on this work, 27 species as valid. Three original papers are presented. On the first one, we review the taxonomic affinities of “Pricesaurus megalodon”, concluding that this specimen has affinities with Anhangueridae, especially with the genus Anhanguera. On the second paper, we describe an unusual pterosaur palate, tentatively attributed to Azhdarchoidea. In addition, we contribute on the anatomy and evolution of the pterodactyloid palate, recognizing evolutionary trends that were never described for the clade. We reappraise, on the third paper, a pterosaur specimen from the Romualdo Formation that was formerly attributed to Anhanguera santanae, concluding that the material represents a yet undescribed anhanguerid. The ecology of anhanguerid pterosaurs from the Romualdo Formation is discussed, especially with respect to the apparent coexistence of several species that possibly shared the same ecological niche. Keywords: Pterosaurs; Santana Group; Araripe Basin; phylogeny; taxonomy; anatomy. SUMÁRIO TEXTO EXPLICATIVO DA ESTRUTURA DA TESE 9 Parte I 1. INTRODUÇÃO 11 2. OBJETIVOS 12 3. JUSTIFICATIVA 13 4. MATERIAIS E MÉTODOS 14 4.1. Materiais 14 4.2. Métodos 19 4.2.1. Preparação 19 4.2.2. Fotografia e desenhos 19 4.2.3. Comparação osteológica e anatômica com base em espécimes depositados em coleções 19 4.2.4. Descrição anatômica 23 5. ESTADO DA ARTE 24 5.1. Contexto geológico 24 5.1.1. Considerações gerais sobre a Bacia do Araripe 24 5.1.2. Geologia e paleontologia do Grupo Santana 26 5.2. Os Pterosauria 33 5.2.1. Aspectos gerais 33 5.2.2. Origem e relações de parentesco com outros tetrápodes 35 5.2.3. Relações filogenéticas internas de Pterosauria 44 5.2.4. Relações filogenéticas de pterossauros do Grupo Santana 55 5.2.4.1. Anhangueridae 55 5.2.4.2. Tapejaridae 55 5.2.5. Considerações sobre o registro espacial e temporal de Pterosauria 62 5.2.6. Taxonomia de pterossauros provenientes do Grupo Santana 65 6. REFERÊNCIAS 125 Parte II ARTIGOS 140 Parte III ANEXOS 190 9 Pinheiro, F. L. 2014 – Contribuição ao conhecimento dos pterossauros do Grupo Santana, Bacia do Araripe, Brasil TEXTO EXPLICATIVO DA ESTRUTURA DA TESE Esta tese de doutorado está estruturada em torno de artigos publicados ou submetidos para avaliação em periódicos científicos especializados. Assim, sua estrutura é organizada em três partes distintas, a saber: a) Introdução sobre o tema estudado, incluindo um estado da arte sobre o tema da pesquisa, objetivos do trabalho desenvolvido, seguidos de uma discussão integradora contendo um sumário dos principais resultados obtidos e interpretações deles derivadas. b) Artigos publicados ou submetidos a periódicos com corpo editorial permanente e revisores independentes, ou publicações equivalentes (capítulo de livro de publicação nacional ou internacional com corpo de revisores independente), escritos pelo autor durante o desenvolvimento de seu doutoramento. c) Anexos, incluindo artigos científicos nos quais o pós-graduando é co- autor; artigos científicos não diretamente relacionados ao tema principal da tese; resumos e artigos publicados em eventos, bem como documentação pertinente de natureza numérica (tabelas,
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