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Universidade De São Paulo Ffclrp UNIVERSIDADE DE SÃO PAULO FFCLRP – DEPARTAMENTO DE BIOLOGIA PROGRAMA DE PÓS-GRADUAÇÃO EM BIOLOGIA COMPARADA Dinossauromorfos triássicos do Sul do Brasil e padrões biogeográficos da irradiação dos dinossauros Júlio Cesar de Almeida Marsola 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 2018 UNIVERSIDADE DE SÃO PAULO FFCLRP – DEPARTAMENTO DE BIOLOGIA PROGRAMA DE PÓS-GRADUAÇÃO EM BIOLOGIA COMPARADA Dinossauromorfos triássicos do Sul do Brasil e padrões biogeográficos da irradiação dos dinossauros Júlio Cesar de Almeida Marsola Orientação: 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 2018 Autorizo a reprodução e divulgação total ou parcial deste trabalho, por qualquer meio convencional ou eletrônico, para fins de estudo e pesquisa, desde que citada a fonte. FICHA CATALOGRÁFICA Marsola, Júlio Cesar de Almeida Dinossauromorfos triássicos do Sul do Brasil e padrões biogeográficos da irradiação dos dinossauros, 2018. 199 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. Dinosauromorpha. 2. Dinosauriformes. 3. Dinosauria. 4. Saurischia. 5. Triássico. 6. Bioestratigrafia. 7. Biogeografia. 8. Gondwana iii AGRADECIMENTOS Agradeço ao Max Langer por me receber e concordar em me orientar não apenas durante o doutorado, mas por todos os 10 ou mais anos de jornada acadêmica, e pela amizade. É gratificante poder opinar agora. Agradeço ao Richard Butler por ter me recebido em Birmingham, pela orientação e pelo suporte sempre além do esperado de um orientador. Esta Bolsa de Doutorado foi concedida no âmbito do Convênio FAPESP/CAPES para a concessão de bolsas, para as quais expresso meus mais sinceros agradecimentos: processo nº 2013/23114-1, Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP). De igual modo, agradeço a FAPESP pela concessão da Bolsa Estágio de Pesquisa no Exterior: processo nº 2016/02473-1, Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP). Também sou grato ao Programa de Pós- Graduação em Biologia Comparada (FFCLRP-USP) pelo amparo institucional durante o doutorado. Sou grato aos seguintes curadores por permitirem que eu analisasse os espécimes sob seus cuidados: Alan Turner (Stony Brook University, Estados Unidos), Alejandro Kramarz (Museo Argentino de Ciencias Naturales Bernardino Rivadavia, Argentina), Ana Maria Ribeiro e Jorge Ferigolo (Fundação Zoobotânica do Rio Grande do Sul, RS), Átila Da-Rosa (Universidade Federal de Santa Maria, RS), Carl Mehling (American Museum of Natural History, Estados Unidos), Caroline Buttler (National Museum of Wales, País de Gales), César Schultz (Universidade Federal do Rio Grande do Sul, RS), Claudia Hildebrandt (University of Bristol, Inglaterra), Deborah Hutchinson (Bristol Museum and Art Gallery, Inglaterra), Gabriela Cisterna (Universidad Nacional de La Rioja, Argentina), Ingmar Werneburg (Eberhard Karls Universität Tübingen, iv Alemanha), Jaime Powell (Fundación Miguel Lillo, Argentina), Jessica Cundiff (Museum of Comparative Zoology, Estados Unidos), Marco Brandalise de Andrade (Museu de Ciências e Tecnologia da PUC, RS), Oliver Rauhut (Ludwig-Maximilians- Universität, Alemanha), Rainer Schoch (Staatliches Museum für Naturkunde, Alemanha), Ricardo Martínez (Museo de Ciencias Naturales, Argentina), Sandra Chapman (Natural History Museum, Inglaterra), Sérgio Cabreira (Museu de Ciências da Naturais da ULBRA, RS), Sifelani Jirah (Evolutionary Studies Institute, África do Sul), Thomaz Schossleitner (Museum für Naturkunde, Alemanha), Tomasz Sulej e Mateusz Talanda (Institute of Paleobiology, Polish Academy of Sciences, Polônia), e Zaituna Erasmus (Iziko South African Museum, África do Sul). Agradeço aos amigos que sempre me ajudaram das mais diversas maneiras ao longo do doutorado: Átila Da-Rosa, Estevan Eltink (Tevinho), Felipe Montefeltro (Fezão), Gabriel Ferreira (Fumaça), Marco França, Marcos Bissaro, Mariela Castro, Mario Bronzati (Roquinho), Jonathas Bittencourt, Pedro Godoy (Tomate) e Thiago Fachini (Schumi). Agradeço aos amigos do PaleoLab em geral, pelo fantástico ambiente de trabalho que sempre foi proporcionado. Também agradeço aos amigos de Birmingham, principalmente ao pessoal da 22 Roman Way. Agradeço todo apoio, suporte e confiança da minha família, Paulo, Lázara e Majú, que fizeram deste doutorado uma jornada muito mais agradável. Por fim, e mais importante, agradeço a minha esposa e companheira Dalila, por estar e por acreditar em mim em todas as partes que este doutorado nos levou. v RESUMO Marsola, J. C. A. Dinossauromorfos triássicos do Sul do Brasil e padrões biogeográficos da irradiação dos dinossauros. 2018. 199p. Tese (Doutorado) – Faculdade de Filosofia, Ciências e Letras de Ribeirão Preto, Universidade de São Paulo, Ribeirão Preto, 2018. Os depósitos triássicos continentais do sul do Brasil abrigam uma grande diversidade de tetrápodes terrestres, incluindo terápsidos, rincossauros, rincocefálios e arcossauros, como pseudosúquios e dinossauromorfos. Inserida neste contexto, a Formação Santa Maria, de porção superior datada do Carniano superior, tem papel fundamental no entendimento da origem e irradiação inicial dos dinossauromorfos, pois abriga alguns dos mais antigos registros do grupo em todo mundo, incluindo vários fósseis de dinossauros. Atualmente, a fauna de dinossauromorfos desta unidade é representada por Ixalerpeton polesinensis, Teyuwasu barberenai, Staurikosaurus pricei, Saturnalia tupiniquim, Pampadromaeus barberenai, Buriolestes schultzi e Bagualosaurus agudoensis, enquanto para o Noriano da Formação Caturrita são conhecidos Guaibasaurus candelariensis, Unaysaurus tolentinoi e Sacisaurus agudoensis. Visando o melhor entendimento da diversidade de dinossauromorfos oriundos destes depósitos, foram descritos, no contexto dessa tese, diversos novos fósseis do grupo: ULBRA-PVT 059, 280, LPRP/USP 0651, MCN PV 10007-8, 10026, 10027 e 10049. Adicionalmente, foi considerado o recente histórico de pesquisas sobre a origens dos dinossauros para examinar o impacto de novas descobertas e das diferentes hipóteses filogenéticas no entendimento dos padrões biogeográficos da irradiação dos dinossauros. vi ▪ ULBRA-PVT059 e 280 representam os holótipos de duas espécies de dinossauromorfos: Ixalerpeton polesinensis e Buriolestes schultzi. I. polesinensis é o primeiro lagerpetídeo descrito para o Brasil e o único no mundo que preserva elementos do crânio e do membro escapular. O material revela que algumas características antes inferidas como sinapomórficas para Dinosauria já estavam presentes em outros dinossauromorfos. B. schultzi é um sauropodomorfo, provável grupo-irmão dos demais representantes do grupo. Além disso, sua anatomia dentária e relações filogenéticas sugerem que os primeiros dinossauros, incluindo os sauropodomorfos, eram adaptados a faunivoria. ▪ LPRP/USP 0651 é o holótipo de uma nova espécie de dinossauro, Nhandumirim waldsangae, da Formação Santa Maria. Apesar de incompleto, as partes preservadas mostram que este se tratava de um indivíduo juvenil, mas que difere em vários aspectos dos demais dinossauros do Carniano, em especial daqueles provenientes dos mesmos níveis estratigráficos. As relações filogenéticas de N. waldsangae indicam que o novo táxon se trata de um dinossauro saurísquio não- sauropodomorfo, possivelmente afim aos terópodos. ▪ MCN PV 10007-8, 10026, 10027 e 10049 se tratam de materiais de dinossauros provenientes da localidade tipo de Sacisaurus agudoensis. Estes representam um sauropodomorfo morfologicamente mais semelhante a membros mais recentes do grupo do que aqueles do Carniano. Assim, correlações bioestratigráficas sugeridas pela presença destes sauropodomorfos indicam uma idade mais nova para a localidade tipo de S. agudoensis do que a das biozonas carnianas. vii ▪ As análises biogeográficas consistentemente otimizaram a porção sul do Gondwana como a área ancestral de Dinosauria, o mesmo se dando para clados mais inclusivos. Estes resultados mostram que a hipótese em questão é robusta mesmo com maior amostragem taxonômica e geográfica, e independentemente das hipóteses filogenéticas. Desta forma, é demonstrado que não há suporte para a hipótese da Laurásia representar a área ancestral dos dinossauros. viii ABSTRACT Marsola, J. C. A. Triassic dinosauromorphs from southern Brazil and biogeographic patterns for the origin of dinosaurs. 2018. 199p. Thesis (Doctorate) – Faculdade de Filosofia, Ciências e Letras de Ribeirão Preto, Universidade de São Paulo, Ribeirão Preto, 2018. The Triassic deposits of southern Brazil harbor a great diversity of terrestrial tetrapods, including therapsids, rhynchocephalians, rhynchosaurs, and archosaurs like pseudosuchians and dinosauromorphs. In this context, the Carnian Santa Maria Formation is important for the understanding of the origins and early diversifications of Dinosauromorpha, as it bears one of the oldest records for the group worldwide, including some of the oldest dinosaurs. Its dinosauromorph fauna is currently represented by Ixalerpeton polesinensis, Staurikosaurus pricei, Saturnalia tupiniquim, Pampadromaeus barberenai, Buriolestes schultzi, Bagualosaurus agudoensis, and Teyuwasu
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