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Ornithopod Dinosaurs from the Late Jurassic of Portugal

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Ornithopod Dinosaurs from the Late Jurassic of Portugal Filippo Maria Rotatori B.Sc. Natural Sciences Orientador: Octávio Mateus, Prof. Associado FCT-UNL Co-orientador: Miguel Moreno-Azanza, Pós-Doc, FCT-UNL Ornithopod dinosaurs from the Late Jurassic of Portugal Dissertação para obtenção do Grau de Mestre em Paleontologia September 2018 September 2018 Filippo Maria Rotatori B.Sc. Natural Sciences Ornithopod dinosaurs from the Late Jurassic of Portugal Dissertação para obtenção do Grau de Mestre em Paleontologia Orientador: Octávio Mateus, Prof. Associado FCT-UNL Co-orientador: Miguel Moreno-Azanza, Pós-Doc, FCT-UNL i Acknowledgements This is the end of a journey, as a student, as a scientist and most importantly as a person. As any other achievement, this thesis is not just the fruit of myself, but there are so many to thank. Since no one is a student if there is no one who teaches, I need to thank my main advisor, Prof. Octávio Mateus and my co-advisor Dr. Miguel Moreno-Azanza. Thanks for their patience and enthusiasm in teaching and leading this thesis project. Also thanks to the unknown arguent, whose work will surely improve this small piece of research. Thanks to Dr. Hübner for sharing with me his work and for being so enthusiastic about basal iguanodontians (which I was not at the time) and to Dr. Madzia for giving me advises about neornithischians and phylogenetics. Thanks to Prof. Will Harcourt-Smith for helping me out with Principal Component Analysis, morphometrics and multivariate statistics. Thanks also to Dr. Federico Fanti, my former advisor, for always giving me suggestions and sharing cool ideas. For the same reasons as wel, thanksl to Dr. Emanuel Tschopp, for sharing his experience in the field and as a research. Thanks to Dr. Bruno Camilo Silva of Sociedade de Historia Natural in Torres Vedras for giving me access to the holotye of Eousdryosaurus nanohallucis, and to Joana Ferreira for her assistance. The data taken there, were greatly useful for the realization of this thesis. Thanks also to my family, who gave me the possibility to live and study abroad, and kept sustaining me during these years: Mom, Dad, my two sisters: Anna and Rebecca. Also thanks to uncle Andrea, Tommaso, Renato and all the cousins. Thanks to my Grandma and all the aunties who actually fed me during this year of research and writing. Here in Lourinhã I need to thank all the members of Lourinhã Paleoteam: Alex, Darìo, Andrè, for they close supports, laughs and reciprocal help. Thanks also to Francisco, Catia, an indefinite number of Joãos, Edu, Xanna, Alex, Carla, Marco, Hugo. A special thank goes to Carla-Alexandra for her patience in helping with (and teaching me about) preparation. Also thanks to her for the affection she showed me in all this time. Another one goes to Micael, for sharing his work with me, since most of the material here described was introduced to me by him! Thanks also to Vincent for his kindess and joy shared with us this year. Thanks to Miggi for the fun, the honest talks about life, paleontology and many other things. I have also so many beautiful relationships back home, which I always bring with me: friends from college, especially Carlotta who shares with me the passion for old bones and kept an ear on me all time. Thanks also to Tiz, who is always there to cheer for me when the odds look overwhelming. And for the same reason, thanks to Fil. And thanks to all the others in Bologna I miss so much: Andrea, Silvia, Andrea, Mati, Gaspa, Leti, Giulia and Vicki. ii Back home, I have people who I have known for almost twenty years, who are almost siblings rather than friends: thanks to Barto, Jaco and Manu for hearing for too long about this thesis, PhD problems and how good Gin Tonic is at international level (and as international, I mean Spain and Portugal). Thanks also to Angie, Cento, Nichi, Riba, Michi, Panga, Luca, Marianna, Lella and Pié. Thanks to Don Baldo, for always listening and helping me out. Also thanks to Matti, Marco, Brilla, Pietro for the fun while I was writing the manuscript. Thanks to Mirco (and to the so many I could not fit here) for encouraging me to give my best. iii Abstract Ornithopods are the most successful group of herbivorous dinosaurs, ranging from the Middle Jurassic to the Cretaceous, becoming extinct at the Cretaceous Paleogene boundary. However, most of the attention has been given to derived forms (Hadrosaurids). Herein, ornithopod material coming from Upper Jurassic of Lourinhã Fm. and housed at Museu da Lourinhã (ML) is described and discussed. Specimens studied include cranial and dental material, including cranial remains of a dryosaurid, axial skeleton elements, pectoral girdle elements and limb bones. Comparison with reference literature and phylogenetic analyses has allowed the attribution of the material either to Dryosauridae or and to Ankylopollexia. Since the lack of autapomorphic characters, it was not possible to assign the material to the two valid ornithopod Portuguese taxa, Draconyx loureiroi and Eousdryosaurus nanohallucis, although phylogenetic analyses hint a close relationship between the Lourinha dryosaurid material and E. nanohallucis. Principal Component Analysis (PCA) plotting limb bones proportions indicates a not fully mature ontogenetic stage for the Portuguese specimens. Comparing the Portuguese ornithopod fauna with the one in Morrison Formation and Kimmeridge Clay Formation, it is remarked the key-role of Portugal to understand biogeographic patterns in the distribution of iguanodontians. Keywords: Late Jurassic, Lourinhã Formation, Ornithopoda, Dinosauria, Systematics. iv Resumo Os dinossauros ornitópodes são o grupo mais bem sucedido de dinossauros herbívoros, tendo aparecido no Jurássico Médio extinguindo-se no limite Cretácico/Paleogénico. É aqui descrito e discutido material das coleções do Museu da Lourinhã (ML) de ornitópode do Jurássico Superior da Formação da Lourinhã. Os espécimes estudados incluem material craniano e dentário, incluindo restos cranianos de um dinossauro driossaurídeo, elementos do esqueleto axial, elementos da cintura escapular e ossos dos membros. A comparação e análise filogenética permitiu a atribuição do material tanto a Dryosauridae como a Ankylopollexia. Não é possível atribuir o material aos dois táxons portugueses válidos de ornitópodes, Draconyx loureiroi e Eousdryosaurus nanohallucis, embora análises filogenéticas indiquem uma estreita relação entre o material de driossaurídeo da Lourinhã e E. nanohallucis. Análise de Componentes Principais (PCA) representando as proporções dos ossos dos membros indica um estágio ontogenético não maduro para os espécimes portugueses. É debatida a importância dos espécimes portugueses na distribuição de Iguanodontia na Formação Morrison e na Formação Kimmeridge Clay. Palavras-chave: Jurássico Superior, Formação da Lourinhã, Ornithopoda, Dinosauria, Sistemática. v Graphical Abstract vi Table of contents Acknowledgements ......................................................................................................................... ii Abstract ......................................................................................................................................... iv Resumo .......................................................................................................................................... v Graphical Abstract ......................................................................................................................... vi 1.Introduction .................................................................................................................................. 5 1.1 Systematics of ornithopod dinosaurs .................................................................................... 6 1.2 Ornithopod anatomy: state of the art and main synapomorphies ............................................. 12 Basal ornithopods (non-iguanodontians) ................................................................................... 12 Iguanodontians .......................................................................................................................... 17 Hadrosaurids ............................................................................................................................. 24 1.3 Ornithopods in Portugal: history of discoveries ........................................................................ 30 1.4 Geological framework: Late Jurassic of Portugal ..................................................................... 31 1.5 Objectives ............................................................................................................................... 36 2. Material and methods ................................................................................................................ 36 3.Results ....................................................................................................................................... 38 3.1 Systematic Palaeontology ................................................................................................... 38 Dryosauridae ............................................................................................................................. 38 Cranial skeleton ..................................................................................................................... 38 Axial skeleton ......................................................................................................................... 44 Appendicular skeleton ...........................................................................................................
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