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Plano De Trabalho ____________________________________________________________________________________________________________ Rio C la ro UNIVERSIDADE ESTADUAL PAULISTA INSTITUTO DE GEOCIÊNCIAS E CIÊNCIAS EXATAS Curso de Pós - Graduação em GEOCIÊNCIAS Área de Concentração em GEOLOGIA REGIONAL ANÁLISES ESTATÍSTICA MULTIVARIADA E FILOGENÉTICA DOS DIPNOIFORMES BRASILEIROS.COMPARAÇÕES BIÓTICAS COM O GONDWANA OCIDENTAL CARLOS EDUARDO VIEIRA TOLEDO Rio Claro (SP) 2006 ____________________________________________________________________________________________________________ Rio C la ro UNIVERSIDADE ESTADUAL PAULISTA INSTITUTO DE GEOCIÊNCIAS E CIÊNCIAS EXATAS Curso de Pós - Graduação em GEOCIÊNCIAS Área de Concentração em GEOLOGIA REGIONAL ANÁLISES ESTATÍSTICA MULTIVARIADA E FILOGENÉTICA DOS DIPNOIFORMES BRASILEIROS.COMPARAÇÕES BIÓTICAS COM O GONDWANA OCIDENTAL CARLOS EDUARDO VIEIRA TOLEDO ORIENTADOR: REINALDO J. BERTINI CO-ORIENTADORA: MARTHA RICHTER Tese de Doutoramento elaborda junto ao Curso de Pós-Graduação em Geociências Área de Concentração em Geologia Regional, para obtenção do Título de Doutor em Geociências. Rio Claro (SP) 2006 ____________________________________________________________________________________________________________ 560 Toledo, Carlos Eduardo Vieira T649a Análises estatística multivariada e filogenética dos Dipnoiformes brasileiros. Comparações bióticas com o Gondwana ocidental / Carlos Eduardo Vieira Toledo. – Rio Claro : [s.n.], 2006 212 f. : il., tabs., fots., mapas Tese (doutorado) – Universidade Estadual Paulista, Instituto de Geociências e Ciências Exatas Orientador: Reinaldo José Bertini Co-orientador: Martha Richter 1. Paleontologia. 2. Análise estatística multivariada. 3. Histologia. 4. Análise filogenética. I. Título. Ficha Catalográfica elaborada pela STATI – Biblioteca da UNESP Campus de Rio Claro/SP ____________________________________________________________________________________________________________ Comissão Examinadora Prof. Dr. Reinaldo J. Bertini Prof. Dr. Paulo Milton Barbosa Landim Prof. Dr. Marcello Simões Profa Dra. Valéria Gallo da Silva Prof. Dr. Paulo Marques Machado Brito Carlos Eduardo Vieira Toledo Rio Claro, 10 de Março de 2006. Resultado: ____________________________________________________________________________________________________________ “Pensamos demasiadamente e sentimos muito pouco. Necessitamos mais de humildade que de máquinas. Mais de bondade e ternura que de inteligência. Sem isso, a vida se tornará violenta e tudo se perderá.“ Charles Chaplin ____________________________________________________________________________________________________________ A DENISE, LUÍSA E GABRIEL ____________________________________________________________________________________________________________ AGRADECIMENTOS Ao Prof. Dr. Reinaldo J. Bertini, pela orientação e amizade nestes anos. A Lília Bertini pelos auxílios técnicos e de laboratório. A Denise pelo apoio incondicional, pela paciência e dedicação e amor. Aos meus pais por sua compreensão e apoio. Aos professores Paulo M. B. Landim, Paulo Brito, Joel C. de Castro, Antenor Zanardo, Solange A. C. Monteiro, Valéria Gallo, Marcello Simões e Maurício Bacci, pelas sugestões, discussões e correções. Ao Prof. Dr. Manuel Alfredo de Medeiros pelos espécimens do Estado do Maranhão e pela agradável experiência de campo na Ilha do Cajual. Ao Prof. Dr. Mario Cozzuol pela notável saída de campo no Estado do Acre. Ao Prof. Dr. Ismar de Souza Carvalho pelas imagens das escavações de dipnoiformes da Formação Ponta Grossa. A Drª Marise Sardenberg Salgado de Carvalho e Drª Rita de Cássia Tardin Cassab pelos espécimens de Lepidosiren megalos. A Prof ª Drª Maria Claudia Malabarba pelos espécimens de Posto Queimado. Ao Prof. Edson Guilherme pelos espécimens de Lepidosiren megalos. Ao Prof. Dr. Jorge Ferigolo pelas imagens de dipnoiformes da Formação Santa Maria. A Profª. Drª. Martha Richter (Natural History Museum – Londres), Profª Drª. Moya Meredith Smith (King’s College - Londres), Prof. Dr. Daniel Goujet (Muséum National ____________________________________________________________________________________________________________ d’Histoire Naturelle - Paris), Profª. Drª. Magdalena Borsuk-Bialynicka (Institute of Paleobiology da Polish Academy of Sciences - Varsóvia), Prof. Dr. Stanislav Stamberg (Natural History Museum of Eastern Bohemia - Hradec Králové – República Tcheca), Profª. Drª. Gloria Arratia (Museum fur Naturkunde der Humboldt Universitat - Berlin), que possibilitaram a comparação dos espécimens e que muito me auxiliaram na execução deste trabalho. Aos meus amigos, Rubens, Egaz, José Vicente, Luis Felipe, Gil, Paulo, Dênis, Tadeu, Neto. Ao Leandro, que muito me auxilio durante meu estágio em Londres. Não tenho palavras para expressar minha gratidão. Aos técnicos Júnior, Adilson, Alan, Claudio, Vladimir, Cezário, Susi e Moema (Biblioteca), Jaime (Desenhista - I.B.). A Foto Santa Cruz pelo auxílio técnico na preparação das fotos utilizadas neste trabalho. À Fundação de Amaparo à Pesquisa do Estado de São Paulo pela bolsa de estudos (01/12570-0) e pelos recursos concedidos. Aos colegas do Núcleo de Evolução e Paleobiologia de Vertebrados, Rodrigo, Marco, Eliane, Darciléia, Felipe, André, Marco e Mírian por todo o auxílio e companheirismo. Aos Colegas da Pós-graduação e a todos aqueles que contribuíram para a realização desta Tese de Doutoramento. Carlos Eduardo Vieira Toledo i Análises Estatística Multivariada e Filogenética dos Dipnoiformes Brasileiros. Comparações Bióticas com o Gondwana Ocidental SUMÁRIO RESUMO _____________________________________________________________________xv ABSTRACT __________________________________________________________________ xvi Introdução _____________________________________________________________________1 Materiais e Métodos ______________________________________________________________6 Dipnoiformes __________________________________________________________________28 Dipnoiformes do Gondwana ______________________________________________________47 Ocorrências de Dipnoiformes no Brasil _____________________________________________61 Análise Estatística _____________________________________________________________100 Análise Histológica_____________________________________________________________136 Análise Filogenética____________________________________________________________158 Discussões e Conclusões ________________________________________________________174 Referências Bibliográficas_______________________________________________________179 Carlos Eduardo Vieira Toledo ii Análises Estatística Multivariada e Filogenética dos Dipnoiformes Brasileiros. Comparações Bióticas com o Gondwana Ocidental ÍNDICE Introdução _________________________________________________________ 1 Materiais e Métodos__________________________________________________ 6 Amostragem para preparação ___________________________________________________9 Preparação Química __________________________________________________________10 Preparação Mecânica _________________________________________________________12 Confecção de Lâminas Delgadas ________________________________________________12 Análise Estatística ____________________________________________________________13 Análise Estatística Multivariada ___________________________________________________________ 13 Análise de Agrupamento _____________________________________________________________ 14 Análise Fatorial_____________________________________________________________________ 15 Análise das Componentes Principais ___________________________________________________ 16 Análise de Correspondências__________________________________________________________ 17 Análise Filogenética___________________________________________________________18 Ordenação e polarização de caracteres___________________________________________20 Métodos de consenso e índices __________________________________________________20 Consenso Estrito _______________________________________________________________________ 21 Consenso Semi-Estrito __________________________________________________________________ 21 Consenso de Adams ____________________________________________________________________ 21 Consenso de Maioria ____________________________________________________________________ 22 Índice de Consistência___________________________________________________________________ 23 Índice de Retenção _____________________________________________________________________ 24 Índice de Consistência Reescalonado _______________________________________________________ 25 Análise Histológica ___________________________________________________________25 Dipnoiformes ______________________________________________________ 28 Dipnoiformes do Gondwana __________________________________________ 47 Austrália____________________________________________________________________47 Índia _______________________________________________________________________53 Carlos Eduardo Vieira Toledo iii Análises Estatística Multivariada e Filogenética dos Dipnoiformes Brasileiros. Comparações Bióticas com o Gondwana Ocidental África e Madagascar __________________________________________________________54 América do Sul ______________________________________________________________58 Antártica ___________________________________________________________________59 Ocorrências de Dipnoiformes no Brasil _________________________________ 61 Bacia do Paraná______________________________________________________________63 Formação Ponta Grossa__________________________________________________________________
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