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C:\Documents and Settings\Edio-Ernst Kischlat.TEYUWASU\My Documents\Artigos\Tese\Ufrgs-00-1.Wp7 Universidade Federal do Rio Grande do Sul Instituto de Geociências Departamento de Paleontologia e Estratigrafia Pós-Graduação em Paleontologia Padrão muscular da coxa de arcossauromorfos fósseis. Aplicação do cladismo reverso e teste de hipóteses. Volume I (TEXTO) Edio-Ernst Kischlat Tese apresentada ao curso de pós-graduação em Paleontologia e Estratigrafia como pré-requisito para obtenção do título de Doutor em Ciências. Orientador: Prof. (D.Sc.) Mário Costa Barberena Co-Orientadora: Profª. (D.Sc.) Lígia Krause BANCA EXAMINADORA: Prof. (D.Sc.) César Leandro Schulz (UFRGS) Prof. (D.Sc.) Ulisses Caramaschi (UFRJ) Profª. (D.Sc.) Ana María Báez (UBA) 21 de Fevereiro de 2003 Kischlat, Edio-Ernst Padrão muscular da coxa de arcossauromorfos fósseis: Aplicação do cladismo reverso e teste de hipóteses. / Edio-Ernst Kischlat. - Porto Alegre: UFRGS, 2003. 3v., xii+451p., 90fig., 28tab. Tese (Doutorado) - Universidade Federal do Rio Grande do Sul. Instituto de Geociências. Programa de Pós-Graduação em Geociências, Porto Alegre, Rio Grande do Sul, Brasil, 2003. 1. Paleontologia. 2. Archosauria. 3. Paleovertebrados. I. Título. Catalogação na Publicação Biblioteca do Instituto de Geociências Renata Cristina Grün CRB10/1113 - E.-E. Kischlat - iii RESUMO A metodologia aqui denominada de “Cladismo Reverso” foi aplicada no estudo da musculatura da coxa dos tetrápodes coronais, objetivando o entendimento das origens e inserções musculares femorais e pélvicas de tecodôncios fósseis, mais especificamente em crocodilotársios e avemetatarsálios basais. A idéia central é de que um modelo plesiomórfico hipotético arcossauriano resulta da intersecção entre os modelos plesiomórficos hipotéticos aviário e crocodiliano, e a diferença encontrada entre ambos seria representativa de aquisições no intervalo Archosauria-Aves e Archosauria-Crocodylia. Da mesma forma, o modelo plesiomórfico hipotético sauriano é resultado da intersecção entre os modelos plesiomórficos hipotéticos lepidossauriano e arcossauriano, e as diferenças encontradas em ambos, representativa de aquisições nos intervalos Sauria-Lepidosauria e Sauria-Archosauria. As hipóteses propostas são então testadas em espécimens fósseis, procurando-se seguir seu padrão de diferenciação temporal através de hipóteses cladistas prévias. Estruturas topográficas femorais e pélvicas foram relacionadas às origens e inserções musculares, procurando-se um entendimento homológico para aplicação em futuras análises filogenéticas. Especial atenção foi dado às estruturas comumente denominadas como “trocânteres”. Propostas prévias de reconstruções musculares utilizando formas fósseis foram discutidas e comparadas com o modelo proposto. No que se refere à nomenclatura miológica, foi relacionada sua aplicação em cada grupo coronal, visando hipóteses de homologia e propostas de universalização nomenclatural. Grande parte da nomenclatura sistemática filogenética, principalmente dos intervalos Sauria-Aves e Sauria-Crocodylia, foi revista, com definições nodais e estemáticas, além de proposição de novos nomes. Regras nomenclaturais foram discutidas, também com novas propostas operacionais. É descrito novo material triássico sul-rio-grandense, incluindo táxons dinossaurianos e crocodilotarsianos (Karamuru vorax, tax.n.), com comentários sobre os táxons relacionados, em especial a Staurikosaurus, Guaibasaurus e Prestosuchus. O modelo plesiomórfico hipotético miológico proposto é aplicado e considerações biomecânicas são feitas. Análises filogenéticas prévias são discutidas e alguns resultados são falseados. - E.-E. Kischlat - iv ABSTRACT The method named “Extant Phylogenetic Brackets Approach” is used in study of tetrapod thigh musculature, trying to understand the femoral and pelvic muscle origins and insertions in fossil thecodontians, mainly in basal crocodylotarsians and avemetatarsalians. The central idea is that an archosaurian hypothetic plesiomorphic model is the result of the intersection between the avian and crocodylian hypothetic plesiomorphic models, and the differences in both are acquisitions in the interval Archosauria-Aves and Archosauria-Crocodylia. In the same way, the saurian hypothetic plesiomorphic model is the result of the intersection between the lepidosaurian and archosaurian hypothetic plesiomorphic models, and the differences in both are acquisitions in the interval Sauria-Lepidosauria and Sauria-Archosauria. The hypothesis proposed are tested in fossil specimens, following their temporal diferentiation pattern using previous cladistic hypothesis. Femoral and pelvic topographic structures were related to muscle origins and insertions, trying a homologic knowledge for application in future phylogenetic analyses. Special attention was dealed to structures commonly denominated as “trochanters”. Previous proposals in myological reconstructions are discussed and compared with the proposed model. In reference to myological nomenclature, its application on each crown group was related, proposing homological hypothesis and universalization of names. Great part of the systematic phylogenetic nomenclature, mainly from the interval Sauria-Aves and Sauria-Crocodylia, were reviewed, applying node and stem based definitions, beyond proposition of new names. Nomenclatural rules were also discussed, with new operational proposals. New material from the Triassic of the State of Rio Grande do Sul (Brazil) is described, including dinosaurian and crocodylotarsan (Karamuru vorax, tax.n.) taxa, with comments on other taxa related, specially Staurikosaurus, Guaibasaurus and Prestosuchus. The proposed myological hypothetic plesiomorphic model is applied with biomechanic considerations. Previous phylogenetic analyses were discussed and some results falsified. - E.-E. Kischlat - v AGRADECIMENTOS Muitos me ajudaram no desenvolvimento do presente texto em tempos de crises. Meus orientadores, Dr. Mário Costa Barberena e Drª. Lígia Krause, estiveram presentes em momentos críticos e, sem a ajuda e compreensão deles, estes não seriam superados. O Conselho Nacional de Desenvolvimento Científico e Tecnológico, que forneceu subsídios básicos durante quatro anos, assim como na minha estadia em New York, no American Museum of Natural History (AMNH). Dr. Eugene Gaffney, forneceu a oportunidade de iniciar o desenvolvimento das idéias aqui contidas, disponibilizando a biblioteca do AMNH, a copiadora, a coleção de vertebrados e uma ajuda de custo para minha viagem à Alemanha. Na Alemanha, terra do meu avô, meus estudos sobre material triássico brasileiro não seriam possíveis sem a ajuda de muitos profissionais: Dr. P. Wellnhofer (Ludwig-Maximilians-Universität, München, Bayern), Dr. Alexander Liebau (Eberhard-Karls-Universität, Tübingen, Baden-Württemberg) e Dr. R. Wild (Staatliches Museum für Naturkunde, Stuttgart, Baden-Württemberg). Nos Estados Unidos, o Dr. P.M. Galton (University of Bridgeport, Connecticut, Bridgeport) foi muito gentil em disponibilizar o holótipo de Staurikosaurus sob sua custódia, assim como Mr. C. Schaff (Harvard University, Cambridge, Massachusetts) no que se refere ao material ali depositado. Vários curadores e responsáveis por material, me auxiliaram de maneira incondicional, disponibilizando espécimens para estudo e consulta: C.L. Schultz (Universidade Federal do Rio Grande do Sul, Porto Alegre), D.J. Gower (British Museum, Natural History, London), J. Ferigolo (Fundação Zoobotânica, Porto Alegre), M.C. Malabarba e M. Richter (Pontifícia Universidade Católica, Porto Alegre), M.W. Maisch (Institut und Museum für Geologie und Paläontologie, Eberhard-Karls-Universität, Tübingen) e R. Schoch (Staatliches Museum für Naturkunde, Stuttgart). Colegas (alguns já doutores), se tornaram amigos nestes tempos de overdose paleontológica, tanto no Brasil quanto na Europa: J.C. Cisneros, J.E.F. Dornelles, F. Bonfim, E. Dias, S. Dias-da-Silva, D.J. Gower, M. Holz, M.C. Langer, L. Maciel, M.W. Maisch, R. Negri, J. Pereira, A.M. Ribeiro, C.M. Scherer, R. Schoch, M.B. Soares e L. Timm. Muito deste trabalho se deteve na consulta de artigos depositados em bibliotecas. Em quase todas as ocasiões a minha presença significava trabalho dobrado para “desencavar” inúmeros artigos “fossilizados”. Meus agradecimentos a seus funcionários. Alguns fizeram verdadeiras mágicas, sendo muito competentes na disponibilização destes artigos. A biblioteca do Museu Nacional da Universidade Federal do Rio de Janeiro foi especialmente “sugada” desde meus tempos de mestrado. As bibliotecas do Instituto de Geociências e do Instituto de Biociências da Universidade Federal do Rio Grande do Sul foram também exauridas e seus funcionários não mediram esforços para me ajudar. No meu impedimento, algumas pessoas foram especialmente importantes na gentileza de “desencavar” artigos em bibliotecas “longínquas”. O meu agradecimento a V.A. Blank, Dr. J.F. Bonaparte, J.C. Cisneros, M. de F. Ferreira, R. Hammel, M. Porto, A. Springer, L.M.G. Takche e S. Zetkus. Finalmente, agradeço à minha esposa, que me aturou nestes tempos de “elocubrações” variadas, e aos meus pais, que não puderam ver este trabalho finalizado. - E.-E. Kischlat - vi As idéias aqui contidas foram desenvolvidas em conseqüência da trilha sonora composta de quase 200 horas de gravações mp3 de Tangerine Dream, Klaus Schulze, Mike Oldfield, Vangelis, Constance Demby, Enigma, Era e qualquer outro compositor que me tirasse da realidade e me induzisse a trilhar caminhos alternativos. - E.-E. Kischlat - vii Um trabalho nunca chega ao final, mesmo que tenha sido exaustivamente questionado… Aos meu
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