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Revisão Filogenética De Mesoeucrocodylia: Irradiação Basal E

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Revisão Filogenética De Mesoeucrocodylia: Irradiação Basal E UNIVERSIDADE DE SÃO PAULO FFCLRP - DEPARTAMENTO DE BIOLOGIA PROGRAMA DE PÓS-GRADUAÇÃO EM BIOLOGIA COMPARADA Revisão filogenética de Mesoeucrocodylia: irradiação basal e principais controvérsias Felipe Chinaglia Montefeltro Tese apresentada à Faculdade de Filosofia, Ciências e Letras de Ribeirão Preto da USP, como parte das exigências para a obtenção do título de Doutor em Ciências, Área: Biologia Comparada. RIBEIRÃO PRETO - SP 2013 UNIVERSIDADE DE SÃO PAULO FFCLRP - DEPARTAMENTO DE BIOLOGIA PROGRAMA DE PÓS-GRADUAÇÃO EM BIOLOGIA COMPARADA Revisão filogenética de Mesoeucrocodylia: irradiação basal e principais controvérsias Felipe Chinaglia Montefeltro Orientador: Max Cardoso Langer Tese apresentada à Faculdade de Filosofia, Ciências e Letras de Ribeirão Preto da USP, como parte das exigências para a obtenção do título de Doutor em Ciências, Área: Biologia Comparada. RIBEIRÃO PRETO - SP 2013 FICHA CATALOGRÁFICA Montefeltro, Felipe Chinaglia Revisão filogenética de Mesoeucrocodylia: irradiação basal e principais controvérsias. Ribeirão Preto, 2013. 285 p. : il. ; 30 cm Tese de Doutorado, apresentada à Faculdade de Filosofia, Ciências e Letras de Ribeirão Preto/USP. Área de concentração: Biologia Comparada. Orientador: Langer, Max Cardoso. 1. Crocodyliformes. 2. Mesoeucrocodylia 3. Metasuchia. 4. Notosuchia. 5. Pissarrachampsa . 6. Filogenia AGRADECIMENTOS Agradeço ao orientador Max Cardoso Langer pelo auxilio e oportunidade de desenvolver o projeto de doutorado sob sua tutela no Laboratório de Paleontologia da FFCLRP. Agradeço também ao orientador Hans C. E. Larsson pela oportunidade e auxilio durante o tempo desenvolvido no Redpath Museum da McGill University. Agradeço o suporte financeiro deste projeto às instituições: Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP), Programa de Pós- Graduação em Biologia Comparada da FFCLRP e Laboratório de Paleontologia da FFCLRP. Agradeço aos seguintes curadores, e pesquisadores das instituições visitadas pela autorização de análises de material sob sua responsabilidade: Mark Norell e Carl Mehling (AMNH), Bernhard Zipfel e Bruce Rubdge (BPI), Juliana Sayão e Edison Vicente Oliveira (DG-CTG-UFPE), Rodrigo da Rocha Machado (DGM), Marco Brandalise de Andrade (UFRGS), Norbert Micklich (HLMD), Hans C. E. Larsson e Louise Marie Meunier (Redpath Museum), Miguel Ramalho (SGP), Karl Rauscher (IPUW), Max Langer (LPRP), Alejandro Kramarz, Martin Ezcurra e Fernando Novas (MACN), Gladis Retamal (MAU), Diego Pol e Eduardo Ruigomez (MEF), Arturo Muñoz Saraiva e Ricardo Cespedes Paz (MHCN), Lucas Pomi e Marcelo Reguero (MLP), Alexander Kellner, Deise Henriques e Rodrigo Giesta Figueiredo (Museu Nacional), Alberto Garrido (MOZ), Antonio Celso Arruda-Campos, Fabiano Iori e Sandra Tavares (MPMA), Domenica Santos, Flávia Coelho, Jorge Calvo e Juan Porfiri (MUCP), Alberto Barbosa de Carvalho, Hussan Zaher e Paulo Miranda Nascimento (MZUSP), David Evans (ROM), Sheena Kaal (IZIKO), Paul Sereno (UCRC), Felipe Vasconcellos, Ismar Carvalho e Thiago S. Marinho (UFRJ). Agradeço coletivamente a todos os meus amigos de longa data e recentes, no Brasil e no exterior. Sem dúvidas, eles fizeram esse caminho árduo dos últimos 4 anos mais proveitoso e interessante. Desculpem-me por não citar nominalmente todos, porém não posso me abster de citar o imenso companheirismo provido pelos amigos Túlio Nunes, Estevan Eltink e Mario Bronzati. Agradeço também aos amigos de laboratório por promover um ambiente agradável durante as horas de trabalhos diários. Finalmente, agradeço ao apoio da minha família, em particular, meus pais Valéria, Antônio e minha irmã Suiane, dos quais nunca ouvi uma única palavra que não fosse de incentivo. Chegar ao final deste Doutorado não seria possível sem cada pequeno ato de encorajamento, carinho, suporte e compreensão durante toda minha vida. RESUMO MONTEFELTRO, F. C. Revisão filogenética de Mesoeucrocodylia: irradiação basal e principais controvérsias. 2013. 285 p. Tese (Doutorado) – Faculdade de Ciências e Letras de Ribeirão Preto, Universidade de São Paulo, Ribeirão Preto, 2013. Mesoeucrocodylia é um clado bem estabelecido, que circunscreve a maior parte da diversidade taxonômica e morfológica de Crocodyliformes. No entanto, questões amplas relacionadas à sua irradiação basal permanecem controversas. Trabalhos de cunho filogenético tem estabelecido hipóteses altamente contrastantes, em especial relacionadas à validade e inter-relações dos grupos Thalattosuchia, Sebecosuchia, Notosuchia, Peirosauridae e Trematochampsidae. Uma revisão filogenética ampla foi elaborada com o intuito de reavaliar a filogenia de Mesoeucrocodylia de maneira crítica. Para tal, baseou-se no estudo dos táxons envolvidos neste contexto, em especial os possivelmente atribuídos aos cinco subgrupos supracitados, mas também membros das outras principais linhagens de Mesoeucrocodylia, além de grupos externos imediatos ao clado. Os caracteres previamente propostos para este contexto também foram reavaliados, e novos caracteres propostos, visando abarcar a diversidade morfológica total dos diferentes táxons analisados. A matiz táxon-caráter resultante da revisão é composta por 122 táxons e 494 caracteres, que foi submetida à análise de parcimônia através do programa TNT (5000 RAS, TBR, Hold 20). O consenso estrito das 3575 árvores mais parcimoniosas resultantes (MPT) não concorda em sua totalidade com nenhuma hipótese prévia. A nova hipótese filogenética apresenta uma combinação de clados anteriormente propostos e relações pela primeira vez recuperadas. Thalattosuchia foi encontrado como o grupo de Mesoeucrocodylia mais basal, assim, grupo irmão de Metasuchia. Metasuchia, por sua vez, é formado por uma grande dicotomia com Neosuchia em um ramo e Gondwanasuchia no outro. Neosuchia é formado por Stomatosuchidae como ramo mais basal, seguido de Atoposauridae como o grupo irmão de uma politomia. Esta politomia envolve Tethysuchia, Goniopholididae, Bernissartia , e um clado congregando os Neosuchia derivados (Paralligatoridae, Susisuchus e Eusuchia). Gondwanasuchia circunscreve a maior diversidade dos táxons na presente análise sendo formado por uma politomia que envolve Doratodon , Peirosauridae + Trematochampsidae, Sebecidae + Mahajangasuchidae e Notosuchia. Assim, Sebecosuchia foi recuperado parafilético, com Baurusuchidae como um Notosuchia derivado. Adicionalmente, Sebecia, como originalmente proposto, não foi recuperado como uma hipótese mais parcimoniosa em todas as árvores, porém, encontra-se entre os possíveis arranjos dentre as MPT. Palavras-chave: Crocodyliformes, Mesoeucrocodylia, Metasuchia, Notosuchia, Pissarrachampsa e Filogenia. ABSTRACT MONTEFELTRO, F. C. The phylogenetic revision of Mesoeucrocodylia: basal radiation and major issues . 2013. 285 p. Tese (Doutorado) – Faculdade de Ciências e Letras de Ribeirão Preto, Universidade de São Paulo, Ribeirão Preto, 2010. Mesoeucrocodylia is a well established clade encompassing the greatest taxonomic and morphological diversity within Crocodyliformes. However, its basal phylogeny remains poorly understood. Previous works highly disagree on the relationships and validity of an array of basal mesoeucrocodylian groups, particularly Thalattosuchia, Sebecosuchia, Notosuchia, Peirosauridae and Trematochampsidae. The Mesoeucrocodylia basal phylogeny was critically revaluated based on a broad revision. Taxa belonging to the aforementioned subgroups, as well as taxa assigned to other Mesoeucrocodylia major lineages and its immediate outgroups were reanalyzed. Characters previously proposed within this context were also revised and new characters added to cover the morphological diversity within the selected taxa. The final matrix is 122 taxa and 494 character and the parsimony analyses was performed in TNT (5000 RAS, TBR and Hold 20). The strict consensus of the 3575 most parsimonious trees (MPTs) does not fully agree to any of the previous phylogenies. The new analysis depicts a mix of known clades and relationships proposed for the first time. Thalattosuchia is the basal most Mesoeucrocodylia and sister group of Metasuchia. Metasuchia is formed by a dichotomy including Neosuchia on one branch and Gondwanasuchia on the other. Stomatosuchidae is the basal most Neosuchia, followed by Atoposauridae as the sister group of a polytomy encompassing Tethysuchia, Goniopholididae, Bernissartia and a clade of derived neosuchians (Paralligatoridae, Susisuchus and eusuchians). Gondwanasuchia is formed by a basal polytomy including Doratodon , Peirosauridae + Trematochampsidae, Sebecidae + Mahajangasuchidae and Notosuchia. In this sense, Sebecosuchia was not depicted as a monophyletic group and Baurusuchidae was placed derived in the Notosuchia clade. In addition, Sebecia, as original, conceived was not recovered in all of the MPTs however it is a possible result among the MPTs. Keywords: Crocodyliformes, Mesoeucrocodylia, Metasuchia, Notosuchia, Pissarrachampsa e Phylogeny. SUMÁRIO PREFÁCIO ........................................................................ 01 I. INTRODUÇÃO ........................................................................ 03 I.I. Principais Controvérsias na Filogenia de Meoseucrocodylia 05 I.I.I. Posicionamento de Thalattosuchia ........................................ 05 I.I.II. Monofilia de Notosuchia .......................................................... 07 I.I.III. Monofilia de Sebecosuchia ...................................................... 09 I.I.IV. Posição filogenética de Peirosauridae, Mahajangasuchus e Stolokrosuchus ....................................................................
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