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A New Specimen of Cardiocorax Mukulu with a Nearly Complete Cranium from Angola

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A New Specimen of Cardiocorax Mukulu with a Nearly Complete Cranium from Angola Miguel Patrick Marx BS Geologia A new specimen of Cardiocorax mukulu with a nearly complete cranium from Angola Dissertação para obtenção do Grau de Mestre em Paleontologia Orientador: Octávio Mateus, Prof. Associado com Agregação FCT-UNL 19 May 2020 May 2020 Miguel Patrick Marx BS Geologia A new specimen of Cardiocorax mukulu with a nearly complete cranium from Angola Dissertação para obtenção do Grau de Mestre em Paleontologia Orientador: Octávio Mateus, Prof. Associado com Agregação FCT-UNL i Acknowledgments Firstly, I would like to thank my advisor, Octávio Mateus, for allowing me to work on this incredible specimen, guiding me throughout the process of this work, and being an incredible mentor and friend. I would like to thank Mike Polcyn and Louis Jacobs for their generous advice, mentorship, and for introducing me to the two rules. I want to also thank Ricardo Araújo for the advice on how to measure plesiosaur vertebrae. I am immensely grateful to Projecto PaleoAngola for funding my stay in Angola for the 2019 summer excavation, and the Institute of Study for Earth and Man at SMU for funding my trip to Dallas in Dec. of 2017, providing financial aid for my visit in Nov. of 2019, and funding the CT scan of the cranium from MGUAN PA278. I also thank Chris Beard and Megan Sims for organizing my collection visit to the University of Kansas and Ross Seacord, George Corner, and Shane Tucker for accommodating my visit to the University of Nebraska- Lincoln in April of 2019. I would also like to thank Elliot Smith for providing a photograph of the skull of Thalassomedon haningtoni (UNSM 50132). My friends and colleagues in Portugal are the best. Starting with Lourinhã, a big thanks to everyone at the Museu da Lourinhã for being so kind to me the past five years. I have learned a lot from my master’s, PhD, and postdoc colleagues during my time in Portugal and I want to thank all of them for helping me develop as a scientist. I want to also thank Alex Guillaume, Andre Saleiro, João Russo, João Pratas, Hugo Campos, Darío Estraviz-Lopez, João Muchagato, Vincent Cheng, Francisco Costa, Cátia Ribeiro, Miguel Moreno-Azanza, Eduardo Puértolas and Filippo Rotatori for all the laughs and memories. Alexandra Fernandes (gurl!) for the amazing time spent together in the classroom and the field across three continents, and Roberto Pereira, Helena Oliveira, Andre Bonito, and Pedro Fialho for the great times we shared in class and during fieldtrips. A big thanks to Miguel Moreno-Azanza and Eduardo Puértolas for helping with the systematics portion of this work. Another big thanks to the Dream Team, Isabel, Celso, Euridice, and Simão for an amazing time excavating in Angola. I also want to thank Maciej Ruciński and Bruno Costa for making the Caparica PaleoHouse a great place to live and for the late nights playing Catan and Street Fighter. João Russo, Alexandra Fernandes, and Maciej Ruciński all helped prepare the flip jacket for the skull block, Bruno Costa helped mechanically prepare the mandible block, and Melanie Lex helped prepare the propodial block. Lastly, I need to thank my parents for their love and support, making this thesis possible at all. ii Abstract We report a plesiosaur specimen that includes the most well complete skull of an Elasmoaurid plesiosaurian from Sub-Saharan Africa. The new specimen, MGUAN PA278 from the Maastrichtian outcrops of Bentiaba, Angola includes fully preserved semicircular canals, both hyoids, a nearly complete mandible, seventeen cervical vertebrae, a rib and rib fragment, a propodial, and paddle elements. A CT scan of the cranium was performed at UTCT in Austin, TX to visualize all aspects of the cranium and to produce a digital model of the skull and casts of the endosseous labyrinths. The cranium was segmented at SMU using AMIRA software and compared to other well preserved elasmosaurid plesiosaurian skulls from collection visits and from the literature. MGUAN PA278 is diagnosed as Cardiocorax mukulu based on the morphology of the middle cervical neural spines. C. mukulu is returned from a phylogenetic analysis as a basal elasmosaurid outside of the Elasmosaurinae-Aristonectinae clade and the sister taxon of Libonectes morgani. The compact morphology of the endosseous labyrinth in MGUAN PA278 may be an adaptation to living in a pelagic environment. Keywords: Africa, Angola, Maastrichtian, cranium, endosseous labyrinth iii Resumo Reportamos aqui o crânio mais completo dos plesiossauros de África Subsaariana. O novo espécime, MGUAN PA278, dos afloramentos maastrichtianos de Bentiaba, Angola inclui um crânio quase completo, incluindo canais semicirculares totalmente preservados, ambos hióides, uma mandíbula quase completa, dezessete vertebras cervicais, um fragment de costela e uma costela, um fêmur e elementos das barbatanase. Uma tomografia computadorizada do crânio foi realizada na UTCT em Austin, Texas, para visualizar todos os aspectos do crânio e produzir um model digital do crânio e moldes dos labirintos interósseos. O crânio foi segmentado na SMU usando o software AMIRA e comparado a outros crânios dos plesiossauros elasmosaurídeos bem preservados, através de visitas a coleções e da literatura. MGUAN PA278 é diagnosticado como Cardiocorax mukulu com base na morfologia dos espinhos neurais cervicais. C. mukulu é retornado de uma análise filogenética como um elasmosaurídeo basal fora do clado Elasmosaurinae-Aristonectinae e o táxon irmão de Libonectes morgani. A morfologia compacta do labirinto endósseo em MGUAN PA278 pode ser uma adaptação à vida em um ambiente pelágico. Palavras-chave: África, Angola, Maastrichtian, crânio, labirinto interósseo iv Graphical Abstract v Table of Contents Acknowledgements ...................................................................................................................................... ii Abstract ........................................................................................................................................................ iii Resumo ........................................................................................................................................................ iv Graphical abstract ......................................................................................................................................... v Anatomical and institutional abbreviations .................................................................................................. 5 1. Introduction ............................................................................................................................................... 6 1.1 A history of Plesiosauria in paleontology ....................................................................................... 6 1.2 Angolan and southern hemisphere elasmosaurids .......................................................................... 9 1.3 Geological setting of Bentiaba, Angola ........................................................................................ 17 1.4 Classification and Systematics of Plesiosauria ............................................................................. 22 1.5 A review of the systematics of Elasmosauridae............................................................................ 33 2. Methods .................................................................................................................................................. 41 3. Stratigraphic provenance of MGUAN PA278 ........................................................................................ 44 4. Taphonomy ............................................................................................................................................. 46 5. Results ..................................................................................................................................................... 54 5.1 Systematic paleontology, material, revised diagnosis of Cardiocorax mukulu ........................... 54 5.2 Craniofacial skeleton description .................................................................................................. 64 5.3 Palate description .......................................................................................................................... 86 5.4 Braincase and basicranium description ......................................................................................... 89 5.5 Endosseous labyrinth digital cast description ............................................................................... 98 5.6 Mandible description................................................................................................................... 100 5.7 Dentition description ................................................................................................................... 110 5.8 Axial skeleton description ........................................................................................................... 114 5.9 Appendicular skeleton description .............................................................................................. 132 6. Phylogenetic analysis ............................................................................................................................ 137 7. Discussion ............................................................................................................................................. 150 7.1 Amended diagnosis of Cardiocorax mukulu with comments on MGAUN PA103 .................... 150 7.2
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