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Pterosaurs from the Late Cretaceous of Angola

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Pterosaurs from the Late Cretaceous of Angola Alexandra Eugénio Fernandes B. A. Anthropology Pterosaurs from the Late Cretaceous of Angola Dissertação para obtenção do Grau de Mestre em Paleontologia Orientador: Octávio Mateus, PhD, Prof. Associado FCT-UNL Co-orientador: Brian Andres, PhD, Associado University of Sheffield Março 2020 Alexandra Eugénio Fernandes Eugénio Alexandra Pterosaurs from the Late Cretaceous of Angola of Cretaceous Late the from Pterosaurs 2020 Pterosaurs from the Late Cretaceous of Angola Alexandra Eugénio Fernandes B.A. Anthropology Dissertação para obtenção do Grau de Mestre em Paleontologia Março 2020 Orientador: Octávio Mateus, PhD, Prof. Associado FCT-UNL Co-orientador: Brian Andres, PhD, Associado University of Sheffield i Pterosaurs from the Late Cretaceous of Angola Copyright © Alexandra Eugénio Fernandes, da FCT/UNL e da UNL A Faculdade de Ciencias e Tecnologia e a Universidade Nova de Lisboa tém 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 atraves de repositôrios cientificos e de admitir a sua côpia e distnibuiçao com objectivos educacionais ou de investigacao, não comerciais, desde que seja dado crédito ao autor e editor’. Tambérn, de acordo corn os Regulamentos dos Cursos de 2.°, 3.° ciclos e Mestrados Integrados, e o Despacho 41/2010 de 21 de Dezembro de 2010, as teses sujeitas a periodo de ernbargo so são divulgadas quando este periodo terminar. Urn periodo de embargo da divulgação tambem pode ser solicitado para as dissertaçoes elaboradas com base em artigos previamente publicados por outros editores, sempre que tal seja necessário para respeitar os direitos de copia desses editores. ii Dedicated to Tia Teresa, who, with every bowl of soup, convinced me that I could. Acknowledgements Heartfelt thanks are extended to Maria, Alvaro, Nicki, and Joe Fernandes, for their encouragements and tolerance of my pipe dream of pursuing this course. Thank you to my aunt and godmother Ana Fontoura: you are the best personal librarian anyone could ever dream of. Thank you to Mike Eklund, the most entertaining fossil preparation mentor and cheese connoisseur I’ve ever met. Thank you to Jack Conrad, who planted this seed (when many others would have just pulled it out by the root). Thank you Călin Șuteu, digitization wizard and life running partner. Thank you to Elizabeth Eugenio, Kimberley Aparisio, Carl Mehling, Fiona Brady, Ana Balcarcel, Eileen Westwig, and all of my beloved family and friends (who, to name individually, might just exceed the length of this thesis). I would like to deeply thank the members of the PaleoAngola Project, to Louis Jacobs, Mike Polcyn, Octavio Mateus, Bill Johnson, and my brother Miguel Marx (meet me at “The Old Monk” when this is all over), Euridice Pacheco (for her naming creativity) and the rest of the Dream Team. Sincerest gratitude is extended to Carla Alexandra Tomás for her impeccable preparation and guidance, the staff of the Museu da Lourinhã, to the Lourinhã PaleoTeam for countless laughs and incomparable brute strength, and the SMU and FCT-UNL staff. Thank you to Brian Andres for so many superb insights and such generosity in sharing profuse pterosaurian knowledge, to Miguel Moreno-Azanza (for teaching me almost everything I learned throughout this degree), to Eduardo Puertolas-Pascual (for teaching me whatever Miguel didn’t), and to Filippo Rotatori (pro auxilio sunt Latin). Many thanks to the participants of Flugsaurier 2018 for extending me a warm welcome into their club, and an especial round of gratitude to Oliver Rauhut, Matyas Vremir, Mark Norell, Mike Habib, Chris Bennett, Nick Longrich, Dave Martill, Olaf Dulfer, Pia Schucht, and Elsa Marlene Vorderwuelbecke. iii Abstract Fourteen fossil bones from the Lower Maastrichtian marine locality of Bentiaba in the Namibe Basin of Angola, Africa, identified as pterosaurs, are here described and attributed to Pteranodontia. One articulated set of bones from within the fossil assemblage can be further attributed to the Nyctosauridae, gen. et sp. nov. This is the first record of pterosaur material from the central west coast of Africa, contributing new information about the paleoecological environment of Gondwana during this time frame. Ontogenetic deductions are also drawn from preliminary histological work. This fossil assemblage provides a first glimpse of Angolan pterosaur paleobiodiversity, giving further insight into the ecosystems of the Late Cretaceous. Keywords: Pterosauria, Lower Maastrichtian, Angola, Namibe Basin Resumo Quatorze ossos fósseis da localidade marinha de Bentiaba, no Maastrichtiano Inferior, na Bacia do Namibe, em Angola, África, identificados como pterossauros, são aqui descritos e atribuídos a Pteranodontia. Um conjunto articulado de ossos da assembléia fóssil pode ser ainda atribuído a Nyctosauridae, gen. et sp. nov. Este é o primeiro registo de material de pterossauro da costa oeste central da África, contribuindo com novas informações sobre o ambiente paleoecológico de Gondwana durante esse período. Informações ontogenéticos também são extraídas de trabalhos histológicos preliminares. Este conjunto fóssil fornece um primeiro vislumbre da paleobiodiversidade Angolano dos pterossauros, fornecendo uma visão mais detalhada dos ecossistemas do final do Cretáceo. Palavras-chave: Pterosauria, Maastrichtiano Inferior, Angola, Bacia do Namibe iv Table of Contents Title Page ...................................................................................................................................... i Copyright Information .................................................................................................................. ii Dedication and Acknowledgements ............................................................................................. iii Abstract ...................................................................................................................................... iv Table of Contents ......................................................................................................................... v Introduction ................................................................................................................................ 1 History and Record of Pterosaur Discoveries on the African Continent ............................................. 2 Materials and Methods ....................................................................................................................... 5 Geographical and Geological Setting .................................................................................................. 7 Results ....................................................................................................................................... 12 Systematic Paleontology ................................................................................................................... 12 Descriptions ...................................................................................................................................... 12 Phylogenetic Analysis ........................................................................................................................ 41 Wingspan .......................................................................................................................................... 43 Bone Histology .................................................................................................................................. 44 Discussion .................................................................................................................................. 47 Conclusion ................................................................................................................................. 53 Bibliography .............................................................................................................................. 54 Appendix ................................................................................................................................... 63 Data Matrix ....................................................................................................................................... 63 Figures Figure 1. José Pereira do Nascimento and his campaign in Angola, circa 1894 ..................................... 4 Figure 2. Location and Region of Bentiaba ............................................................................................. 7 Figure 3. Geological Map of Angola ........................................................................................................ 8 Figure 4. Vertebrate and Invertebrate Fauna of Bentiaba ..................................................................... 9 Figure 5. Map of Angolan Pterosaur Localities ..................................................................................... 10 Figure 6. Stratigraphic Section of Bentiaba .......................................................................................... 11 Figure 7. Humerus MGUAN-PA650 ....................................................................................................... 13 Figure 8. Ulna MGUAN-PA651 .............................................................................................................. 16 Figure 9. Ulna MGUAN-PA653 .............................................................................................................
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