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Microvertebrates of the Lourinhã Formation (Late Jurassic, Portugal)

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Microvertebrates of the Lourinhã Formation (Late Jurassic, Portugal) Alexandre Renaud Daniel Guillaume Licenciatura em Biologia celular Mestrado em Sistemática, Evolução, e Paleobiodiversidade Microvertebrates of the Lourinhã Formation (Late Jurassic, Portugal) Dissertação para obtenção do Grau de Mestre em Paleontologia Orientador: Miguel Moreno-Azanza, Faculdade de Ciências e Tecnologia da Universidade Nova de Lisboa Co-orientador: Octávio Mateus, Faculdade de Ciências e Tecnologia da Universidade Nova de Lisboa Júri: Presidente: Prof. Doutor Paulo Alexandre Rodrigues Roque Legoinha (FCT-UNL) Arguente: Doutor Hughes-Alexandres Blain (IPHES) Vogal: Doutor Miguel Moreno-Azanza (FCT-UNL) Júri: Dezembro 2018 MICROVERTEBRATES OF THE LOURINHÃ FORMATION (LATE JURASSIC, PORTUGAL) © Alexandre Renaud Daniel Guillaume, FCT/UNL e UNL A Faculdade de Ciências e Tecnologia e a Universidade Nova de Lisboa tem o direito, perpétuo e sem limites geográficos, de arquivar e publicar esta dissertação através de exemplares impressos reproduzidos em papel ou de forma digital, ou por qualquer outro meio conhecido ou que venha a ser inventado, e de a divulgar através de repositórios científicos e de admitir a sua cópia e distribuição com objetivos educacionais ou de investigação, não comerciais, desde que seja dado crédito ao autor e editor. ACKNOWLEDGMENTS First of all, I would like to dedicate this thesis to my late grandfather “Papi Joël”, who wanted to tie me to a tree when I first start my journey to paleontology six years ago, in Paris. And yet, he never failed to support me at any cost, even if he did not always understand what I was doing and why I was doing it. He is always in my mind. Merci papi ! This master thesis has been one-year long project during which one there were highs and lows. It also comes to conclude three years of working in Portugal and starts new beginnings in paleontology fields. Because I could not achieve this journey alone, I have some people to thank for helping me through this adventure. I would like to thank my family, who supports me since the very beginning, and especially both my parents, without whose nothing could have been enable, who let their son go to a foreign country without assurance of what he will find. More than anything else, they both have been, beyond all expectations, understanding and supportive since the time this little boy was saying “I want to study dinosaurs!”, even if it was sometimes scary and uncharted. My family was present for me, even if it was only to make jokes about the size of my croc teeth, and for that I am grateful. I would like to thank my supervisor Dr. Miguel Moreno-Azanza, who introduce me to the world of microvertebrates, help me in each stage of this thesis, support and motivate me during the low times, and share all the advises and knowledge I needed to complete this work. I would like to thank also my co-supervisor Prof. Octávio Mateus, who welcomed me in his team three years ago and provide the tools and environment to broaden my background and prepare my future career in paleontology. I would like to thank the Museu da Lourinhã for letting me work and use its facilities, and more particularly: Carla Alexandra Tomás in her help to store and prepare the new sediment samples, Alexandre Audigane, Bruno Pereira, and Carla Abreu in their help to organize and facilitate my work at the museum. I would like to thank the Departamento de Ciências da Terra of the Faculdade de Ciências e Tecnologia da Universidade Nova de Lisboa for allowing to use its equipment and facilities, and more particularly: Maria Eduarda Ferreira in her help to process the bulk samples collected on the field, and Prof. Paulo Legoinha for giving me the opportunity I to follow this master program and this master thesis. I would like to thank Prof. José Mirão and Luis Dias to enable the access to the scanning electron microscope in HERCULES facilities. I would like to express all my love and gratitude to Céline Viquesnel, who jumped on the bandwagon few months ago and decided to stay longer for bearing me, giving me the strength and motivation to complete this thesis; and to Alexandra Fernandes, who always believed in me and was inspiring and understanding when I needed courage and could not find the light in the darkest times, and for her amazing help in picking. For them, I will be always grateful. I would like to thank Eduardo Puértolas-Pascual, in his help of the description and analysis of the croc teeth, giving me thus the training to describe the other material studied during this thesis; Filippo Rotatori, my partner for so many things, for his friendship and all the motivation we mutually give to each other; André Saleiro, for having screen washed alone all the bulk sample from Valmitão used for this thesis, but also for his unique friendship and all the fun we have together; and to Francisco Costa, for his friendship and understanding as roommate during this year. I would like to thank my other classmate of the master, namely Darío Estraviz López, Tiago Pereira, and João Velez, for the special friendship we will always share. I would like to thank the students of the year 6 of the Master of Paleontology of FCT: André Bonito, Vincent Cheng, Miguel Marx, Helena Oliveira, Roberto Silva, Cátia Ribeiro, and Victor Takev, for their help in picking Valmitão sediments and their respective friendships. I would like to thank Carl Melhing, from the American Museum of Natural History, for the amazing revision on the English he has done in an impossible deadline and all his support. I would like to thank all my friends of the Lourinhã Paleo team that have not been named yet: Hugo Campos, João Marinheiro, Marco Marzola, Simão Mateus, João Muchagata, João Pratas, João Russo, and Sara Sneltorp. I would like to thank my friends back to France, particularly Adrien Aimard, Aurore Fino, Laurent Henry, Maxime LeCesne, Charlène Selva, Benoît Thevenot, and Thibaut Totis, for whom going back is always a pleasure. I would like to thank the “Héros de Papier Froissé” internet community, and particularly “Norya”, “Crystallina”, “Bibi”, and “princesse”, to have been this safety valve that allowed to clear my mind when I needed it. My final thank is for Steven Spielberg, who first gave me the dream to become a paleontologist and keeps making me dream 23 years later. Thank you all! II ABSTRACT The Upper Jurassic of Portugal has been globally known for its microfossil vertebrate fauna thanks to the Konzentrat-Lagerstätte of the Guimarota mine, which provided thousands of bone fragments, isolated teeth, and even complete specimens. Other vertebrate microfossil assemblages have been studied around the world. Besides Guimarota, no other Portuguese Jurassic assemblage has been extensively studied. Hereby is presented a revision of the state of the art on Portuguese microvertebrate record, and the first microvertebrate studies on three localities from the Lourinhã Formation (Late Jurassic) hosted by a Portuguese institution; Porto das Barcas, Zimbral, and Valmitão has provided 2,497 microvertebrates skeletal remains and teeth, from which 824 specimens have been identified, described and assessed to the conservative-most taxa. The stratigraphy and sedimentology of the localities suggest that Porto das Barcas and Zimbral were floodplain mud deposits, and Valmitão was an oxbow lake mud deposit, with a slow rate of sedimentation. The remains have been attributed to fishes, amphibians, squamates, crocodylomorphs, and dinosaurs; but unfortunately, no mammaliaform material has been collected. Paleoecological analyses suggest Zimbral and Valmitão were dominated by a terrestrial fauna and more diverse than Porto das Barcas, dominated by an amphibious fauna. The Lourinhã Formation appears to have been closer to the shoreline than American localities in the Morrison and Cloverly Formations were, but more continental than Buenache and Las Hoyas localities (Spain) with swamp to lacustrine paleoenvironments. A detailed study on 125 crocodylomorph teeth from Valmitão support the presence of Goniopholididae, at least two Atoposauridae taxa, and Bernissartiidae in the Late Jurassic of Portugal, with a fauna either dominated by relative small individuals, either juveniles or adults or small taxa. Keywords: Crocodylomorpha, Albanerpetontidae, Dinosauria, Lusitanian Basin, Kimmeridgian-Tithonian, Paleoecology RESUMO O Jurássico Superior de Portugal tem sido mundialmente conhecido pela sua fauna de vertebrados microfósseis graças à Konzentrat-Lagerstätte da mina de Guimarota, que forneceu milhares de fragmentos ósseos, dentes isolados e até espécimes completos. Outros cortejos de III microfósseis de vertebrados foram estudadas noutras partes do globo. Além da Guimarota, nenhuma localidade de microvertebrados jurássicos tem sido extensivamente estudada. É apresentada uma revisão do registo dos microvertebrados portugueses, e os primeiros estudos de microvertebrados de três localidades da Formação da Lourinhã (Jurássico Superior). Porto das Barcas, Zimbral e Valmitão forneceram 2.797 restos esqueléticos e dentes de microvertebrados, dos quais 824 espécimes foram identificados os táxons de forma conservadora. A estratigrafia e sedimentologia das localidades sugerem que Porto das Barcas e Zimbral eram depósitos de planície de inundação, e Valmitão era um depósito de planície de inundação, com baixa taxa de sedimentação. Os restos mortais foram atribuídos a peixes, anfíbios, escamas, crocodilomorfos e dinossauros; mas, infelizmente, nenhum mamífero foi identificado. Análises paleoecológicas sugerem que Zimbral e Valmitão eram dominados por uma fauna terrestre e mais diversa do que Porto das Barcas, dominada por uma fauna anfíbia. A Formação da Lourinhã parece ter estado mais perto da costa do que as localidades americanas nas Formações Morrison e Cloverly, mas mais continentais do que as localidades Buenache e Las Hoyas (Espahna), com paleoambientes pantanosos a lacustres. Um estudo detalhado em 125 dentes de crocodilomorfos de Valmitão suportam a presença de Goniopholidae, pelo menos dois táxones de Atoposauridae, e Bernissartiidae no Jurássico Superior de Portugal, com uma fauna dominada por animais de pequeno porte, possivelmente juvenis a jovens adultos ou formas pequenas.
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