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A Evolução Dos Metatheria: Sistemática, Paleobiogeografia, Paleoecologia E Implicações Paleoambientais

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A Evolução Dos Metatheria: Sistemática, Paleobiogeografia, Paleoecologia E Implicações Paleoambientais UNIVERSIDADE FEDERAL DE PERNAMBUCO CENTRO DE TECNOLOGIA E GEOCIÊNCIAS PROGRAMA DE PÓS-GRADUAÇÃO EM GEOCIÊNCIAS ESPECIALIZAÇÃO EM GEOLOGIA SEDIMENTAR E AMBIENTAL LEONARDO DE MELO CARNEIRO A EVOLUÇÃO DOS METATHERIA: SISTEMÁTICA, PALEOBIOGEOGRAFIA, PALEOECOLOGIA E IMPLICAÇÕES PALEOAMBIENTAIS RECIFE 2017 LEONARDO DE MELO CARNEIRO A EVOLUÇÃO DOS METATHERIA: SISTEMÁTICA, PALEOBIOGEOGRAFIA, PALEOECOLOGIA E IMPLICAÇÕES PALEOAMBIENTAIS Dissertação de Mestrado apresentado à coordenação do Programa de Pós-graduação em Geociências, da Universidade Federal de Pernambuco, como parte dos requisitos à obtenção do grau de Mestre em Geociências Orientador: Prof. Dr. Édison Vicente Oliveira RECIFE 2017 Catalogação na fonte Bibliotecária: Rosineide Mesquita Gonçalves Luz / CRB4-1361 (BCTG) C289e Carneiro, Leonardo de Melo. A evolução dos Metatheria: sistemática, paleobiogeografia, paleoecologia e implicações paleoambientais / Leonardo de Melo Carn eiro . – Recife: 2017. 243f., il., figs., gráfs., tabs. Orientador: Prof. Dr. Édison Vicente Oliveira. Dissertação (Mestrado) – Universidade Federal de Pernambuco. CTG. Programa de Pós-Graduação em Geociências, 2017. Inclui Referências. 1. Geociêcias. 2. Metatheria . 3. Paleobiogeografia. 4. Paleoecologia. 5. Sistemática. I. Édison Vicente Oliveira (Orientador). II. Título. 551 CDD (22.ed) UFPE/BCTG-2017/119 LEONARDO DE MELO CARNEIRO A EVOLUÇÃO DOS METATHERIA: SISTEMÁTICA, PALEOBIOGEOGRAFIA, PALEOECOLOGIA E IMPLICAÇÕES PALEOAMBIENTAIS Dissertação de Mestrado apresentado à coordenação do Programa de Pós-graduação em Geociências, da Universidade Federal de Pernambuco, como parte dos requisitos à obtenção do grau de Mestre em Geociências. Orientador: Prof. Dr. Édison Vicente Oliveira Aprovado em: 06/03/2017 BANCA EXAMINADORA Prof. Dr. Édison Vicente Oliveira (Orientador – 1º Titular) Departamento de Geologia – UFPE Prof. Dr. Francisco Javier Goin (Membro Externo – 2º Titular) Museu de La Plata – Argentina Prof. Dr. Kleberson Porpino (Membro Externo – 3º Titular) Departamento de Geologia – UFPE AGRADECIMENTOS Gostaria de agradecer a todos aqueles que de certa forma contribuíram para este momento da minha vida. Gostaria de expressar minha gratidão a Universidade Federal de Pernambuco (UFPE), ao Centro de Tecnologias e Geociências (CTG) e a todos os funcionários responsáveis pela organização e estruturação dessa instituição. E porque não, agradecer também a todas as experiências (positivas e negativas) que vivi nesse espaço e que hoje se mostram tão importantes. Gostaria de expressar minha gratidão a Fundação de Amparo à Ciência e Tecnologia do Estado de Pernambuco (FACEPE) pelo financiamento do projeto de Mestrado que terminou por transformar-se nessa dissertação. Sou eternamente grato ao professor Édison Vicente Oliveira por ter acreditado em mim quando me aceitou para estagiar em seu laboratório, por ter confiado esse projeto de mestrado que lhes é apresentado em forma de dissertação e pela ajuda e apoio prestados durante os mais de dois anos em que trabalhei no Laboratório de Paleontologia (PaleoLab). A todos os docentes que dedicaram o seu tempo à minha formação profissional e a todos os meus colegas e amigos de curso, com quem convivi, aprendi e cresci durante esses três anos e meio de UFPE! Agradeço em especial a Luana Andrade e Mariana Sena por terem se mostrado amigas de grande importância para a continuidade desse projeto de mestrado. Gostaria de agradecer especialmente aos professores Kleberson Porpino por ter me ajudado bastante nas discussões filogenéticas que me levaram a completar as filogenias aqui presentes. Também a professora Lílian Paglarelli Bergqvist por ter me indicado à gaveta mais importante de minha carreira e também pelas discussões e informações importantíssimas que me foram prestadas. A Luciana Carvalho, Rodrigo Machado e Flávia Figueiredo por terem em ajudado com as coleções de Itaboraí. Aos professores Cristian de Muizon e Pierre-Olivier por terem me ajudado com materiais do Peru e Europa. Aos Professores Francisco Javier Goin e Analia Forasiepi que me ajudaram com artigos, discussões e fotografias dos metatérios Argentinos. Janete Gillett e professor Gary Alpert por ter me enviado uma das fotografias mais importantes da minha carreira. Ao professor Richard Cifelli e seu aluno Joshua Cohen por terem me ajudado bastante com fotografias e materiais dos EUA, incluindo os moldes incríveis que recebi. Ao professor Leonardo Avilla e seus alunos por terem me ajudado enquanto estive no Rio de Janeiro. A minha família, as pessoas mais importantes de minha vida! Pessoas que sempre serviram de exemplo para mim e a quem devo a minha vida, educação, saúde e tudo o que tenho. Pessoas essas que me prestaram apoio nos momentos mais difíceis, que se desdobraram em mil para me ajudar, que sempre estiveram presentes... E, acima de tudo, que me mostraram o verdadeiro significado de dedicação, comprometimento, afeto e confiança! Dizem que a família é o pilar mais importante de nossas vidas e posso garantir que nada é mais certo! Aos meus irmãos, amigos e filhos de quatro patas, asas, nadadeiras e etc. Que me mostraram que a maior beleza da vida está na simplicidade, no companheirismo, na sinceridade, na dedicação e no amor verdadeiro! Esses que nunca desistiram de mim e com quem sempre pude contar! E porque não agradecer a toda minha família, porque família é quem ama, quem cuida e quem nos ama! Por último, mas não menos importante, gostaria de agradecer a todos aqueles que me guiam; que nunca me abandonam e que sempre acreditam em mim! Aqueles que já não estão mais entre nós, mas que vivem eternamente ao meu lado! Gostaria de deixar um agradecimento a minha pessoa... Apenas por nunca ter desistido e sempre ter acreditado que um dia a "vitória" chega para aqueles que a merecem... Não sei se sou merecedor de alguma coisa, mas creio (e espero) que um dia, a vida me responda essa pergunta! Se sou ou não, apenas sei que já conquistei muita coisa e espero continuar conquistando! Essa dissertação será, para sempre, uma eterna lembrança do que ganhamos com amor, dedicação, respeito, paciência e, talvez, merecimento! Para que eu sempre possa lembrar quem sou: "Somos todos visitantes deste tempo, deste lugar. Estamos só de passagem. O nosso objetivo é observar, crescer, amar... Depois vamos para casa" – Provérbio Aborígene. “Existem três coisas que não podem ser escondidas por muito tempo: o sol, a lua e a verdade” – Buda. A todos vocês, meu mais sincero e verdadeiro obrigado! "Australia has long been known as the Land of Marsupials, but South America has also been a land of marsupials (…). In South America, (…) marsupials (…) had an adaptative radiation that led into many ways of life occupied by placentals on the northern continents and Africa.” (George G. Simpson) RESUMO Os marsupiais sul-americanos, dentre os quais se incluem os atuais “gambás” e “cuícas”, consistiram em dos grupos de mamíferos de maior diversidade no Paleogeno sul- americano. A literatura apresenta diversos estudos que enfocam na sistemática e paleobiogeografia dos taxa sul-americanos, entretanto, poucos apresentam resultados bem suportados. Essa incerteza sistemática resultou na análise de quase todas as famílias já descritas para a linhagem dos marsupiais (excluindo as quatro ordens endêmicas do continente Australiano) com o intuito de validar as características consideradas diagnósticas para as mesmas. Os resultados mostram que os Metatheria sul-americanos originaram-se durante o Cretáceo Inicial na América do Norte. A filogenia demonstrou que as grandes linhagens de Metatheria já haviam se diversificado na América do Norte antes da chegada à América do Sul, entretanto foi neste continente que a linhagem atingiu a sua maior diversidade. A provável rota de chegada a América do Sul envolveu a região do Caribe e América Central, constituindo os arcos-de-ilha e as regressões marinhas importantes fatores ambientais para a evolução do grupo. A paleoecologia do grupo demonstrou que os Metatheria sul-americanos evoluíram para ocupar diferentes nichos tróficos durante todo o Paleogeno-Neogeno, estando sua maior diversidade relacionada ao Máximo Termal do Paleoceno-Eoceno (MTPE). A redução das temperaturas globais durante o início do Oligoceno foi provavelmente o principal evento de extinção para a maioria das linhagens no Hemisfério Sul. PALAVRAS-CHAVE: Metatheria. Paleobiogeografia. Paleoecologia. Sistemática. ABSTRACT The South American marsupials, which include the extant opossums, represented a diverse group in South America during the Paleogene. Most published studies focus on systematic and paleobiogeography of South American taxa, however, few present consistent results. This problematic resulted on the analysis of almost all described families of marsupials (excluding Australidelphia four endemic orders), in order to validate the characters considered as diagnostic for them. The results demonstrate that South American metatherians evolved in North America since Early Cretaceous, with almost all major lineages present in North America prior to the arrival in South America; nevertheless, the lineage reached its major diversity in the last continent. The probable dispersal rout was the Caribbean Plate, with ‘aves ridge’ and sea-level regressions during the Late Cretaceous representing important environmental events for the evolution of this group. The paleoecology demonstrated that South American taxa evolved to occupy different trophic niches during Paleogene and Neogene, with the greater diversity being recorded during the Paleocene-Eocene
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