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Revision of Basal Sauropods from the Middle Jurassic of Patagonia and the Early Evolution of Eusauropods

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Revision of Basal Sauropods from the Middle Jurassic of Patagonia and the Early Evolution of Eusauropods Revision of basal sauropods from the Middle Jurassic of Patagonia and the early evolution of eusauropods Dissertation zur Erlangung des Doktorgrades an der Fakultät für Geowissenschaften der Ludwig- Maximilians-Universität München Vorgelegt von Femke Marleen Holwerda München, 23. Januar 2019 Erstgutachter: PD Dr. Oliver Walter Mischa Rauhut Zweitgutachter: Dr. Diego Pol Tag der mündlichen Prüfung: 05.06.2019 II Statutory declaration and statement I hereby confirm that my thesis entitled “Revision of basal sauropods from the Middle Jurassic of Patagonia and the early evolution of eusauropods”, is the result of my own original work. Furthermore, I certify that this work contains no material which has been accepted for the award of any other degree or diploma in my name, in any university and, to the best of my knowledge and belief, contains no material previously published or written by another person, except where due reference has been made in the text. In addition, I certify that no part of this work will, in the future, be used in a submission in my name, for any other degree or diploma in any university or other tertiary institution without the prior approval of the Ludwig-Maximilians-University Munich. Munich, 23.01.2019 Femke Holwerda III Contents Statutory declaration and statement III Acknowledgements VII Abstract VIII Chapter 1: Introduction 1.1 Sauropods 1 1.2 Evolution of early sauropods 3 1.3 The world during the Jurassic: a sauropod perspective 5 1.4 Systematic overview of important Jurassic taxa 6 1.5 The importance of Patagosaurus fariasi 13 1.6 Overview of Chapters 2 – 5 14 1.7 References 16 Chapter 2: Osteological revision of the holotype of Patagosaurus fariasi BONAPARTE 1979 2.1 Abstract 35 2.2 Introduction 36 2.3 Material and methods 39 2.3.1 Anatomical abbreviations 39 2.3.2 Terminology 40 2.3.3 Institutional abbreviations 40 2.3.4 Systematic palaeontology 41 2.3.5 Holotype 41 2.3.6 Original diagnosis 42 2.3.7 Emended diagnosis 42 2.3.8 Horizon, locality and age 43 2.3.9 Geological setting 44 2.4 Description 48 2.4.1 Axial skeleton 48 2.4.2 Appendicular skeleton 156 2.5 Discussion and conclusions 167 2.5.1 Diagnostic characters of Patagosaurus fariasi 167 2.5.2 Other characteristics and comparison with other eusauropods 173 2.6 Acknowledgements 179 2.7 References 180 IV Chapter 3: Revising associated material of Patagosaurus fariasi BONAPARTE 1979, with notes on the species diversity of the Cerro Condor bonebeds 3.1 Abstract 193 3.2 Introduction 194 3.3 Horizon, locality and material 195 3.3.1 Material 196 3.3.2 Anatomical abbreviations 200 3.3.3 Geological setting 201 3.4 Description Material Cerro Condor Norte (Patagosaurus) 206 3.5 Description Material Cerro Condor Sur 281 3.5.1 Cerro Condor Sur Patagosaurus referable 281 3.5.2 Cerro Condor Sur Specimens that are not Patagosaurus 306 3.6 Discussion 338 3.6.1 Species diversity overview from Cerro Condor Norte and Sur 338 3.6.2 New elements to be added to the taxon Patagosaurus 343 3.6.3 Ontogenetic features on vertebrae of Patagosaurus 345 3.6.4 Ontogenetic features on axial and appendicular skeleton of Patagosaurus 347 3.7 Conclusions 349 3.8 Acknowledgements 350 3.9 References 351 Chapter 4: The phylogeny of Patagosaurus fariasi and implications for evolutionary relationships of Early-Middle Jurassic sauropods and their biogeography 4.1 Abstract 365 4.2 Introduction 367 4.2.1 Sauropod diversification in the Late Triassic-Early Jurassic 367 4.2.2 Historical perspective and previous analyses of Patagosaurus phylogeny 368 4.2.3 Phylogenetic importance of Patagosaurus 372 4.3 Methods 373 4.3.1 Coding 373 4.3.2 Software 374 4.3.3 New characters 374 4.4 Results 380 4.5 Discussion and conclusions 395 4.5.1 Phylogenetic signal 395 4.5.2 Biogeographic implications 399 4.5.3 Phylogenetic implications for the Cañadón Asfálto Formation sauropods 403 4.6 Acknowledgements 406 4.7 References 406 Chapter 5: Geometric morphometrics on Patagosaurus fariasi and Jurassic sauropods: implications for diversity and ontogeny 5.1 Abstract 424 5.2 Introduction 425 V 5.3 Methods 426 5.4 Results 433 5.5 Discussion 441 5.6 Conclusions 444 5.7 References 445 Chapter 6: General conclusions and future perspectives 6.1 Conclusions 453 6.2 Future perspectives 459 Appendix 1.1 Appendix Chapter 3: Patagosaurus Material list updated i 1.2 Appendix Chapter 4: Phylogenetic Character list and PCA analysis iv 1.3 Appendix Chapter 5: Procrustes analysis and PCA analysis xlv VI Acknowledgements I would never have made it this far but for the support of Diego Pol, who took me on as a MSc student and then kept supporting me throughout. And we still work together! Thanks to Oliver Rauhut for providing the DFG grant so that I could work on this great group, the sauropods! #sauropodsrule #gosauropodsgo Thanks also to the members of the workgroup Mesozoic Vertebrates. And thanks to: Munich: Gertrud, Nicola, Martin&Camila, Mike, Warren, Bill, and the awesome student gang Tom, Lukardis, Sara Nederland: Sofie, Eva, De Bioloogjes; Petra, Madelon, Tryntsje, Linda, Tammo, Anne Schulp, Jelle Reumer, Wilma Wessels, en mijn familie. Argentina: Trelew: Diego, Jose, Nacho, Juli&Juli, Marcos, Andrea, Marcelo, Pablo, Dudu, Leandro&Lorena Buenos Aires: Alejandro Kramarz, Stella Alvarez, Juli Desojo, Chorno, Agus, Tucuman: Jaime Powell, Rodrigo, Graci …..and a huge thanks to The Randomers Becky and Santi!! Portugal: João, João, Nury (just counting you there fofinha) Alex, Christophe, Emanuel, Alexandra UK: Paul&Paul (the sauropauls), Mark Evans, Hillary Ketchum, Sandra Chapman, Mike Taylor, Matt Wedel, Steve Brusatte, Stig Walsh, Elsa Panciroli, David Button, Phil Mannion Germany: Daniela Schwarz, Nils Knötschke, Martin Sander, Heinrich Mallison, Ulrich Joger Paris: Ronan Allain, Claire PdF, Emilie Läng My Sauroporacles: a word of my own invention (sauropod + oracle) which stands for amazing people who've helped me with every stupid question imaginable concerning sauropods, and that on a regular (sometimes daily) basis. Without (at least audibly) getting annoyed with me. Without complaining. A magnificent thank you goes to Emanuel Tschopp, Jose Luis Carballido, Phil Mannion and Veronica Díez Díaz. Moral support (very important); listening to me when I needed someone to listen, giving sound advice and swearing along with my cursing: Madelon, Liz, Nicola, Franzi, Roland, the EAVP gossip group, and Jon and Daenerys for a right proper swooning. And last, but certainly not least: My husband Jeff. I’m looking forward to a long life together and becoming Fat like Pat J Lastly, a big thank you to Fat Pat and the Patsies (#FatPat, #gosauropodsgo, #sauropodsrule). I may go on to date Cetio and Rut and Moos, but you will always be My Pat. And finally, this thesis is dedicated to the memory of Jaime Powell. VII Abstract The Early and Middle Jurassic are widely regarded as the critical time for sauropod radiation and worldwide dispersal. The accepted theory is that the common ancestor of Early Jurassic sauropods had its provenance in South Africa, South Gondwana at the time. The major sauropod diversification and dispersal out of South Africa happened between the Early and Middle Jurassic (±180-160 Ma), with major clades being firmly established by the Late Jurassic (±150 Ma). The Early and Middle Jurassic, however, have a generally poor sauropod body fossil record with the exception of a few taxa from Gondwana (e.g. Patagosaurus, Volkheimeria, Amygdalodon, Vulcanodon, Tazoudasaurus, Spinophorosaurus, Barapasaurus, Bothriospondylus, Lapparentosaurus) and even fewer from Laurasia (e.g. Cetiosaurus, Cetiosauriscus, Shunosaurus, Klamelisaurus, Mamenchisaurus, Omeisaurus). The Gondwanan sauropod with the most material preserved is Patagosaurus fariasi, from the Cañadón Asfálto Formation of west-central Chubut province, Patagonia, Argentina. It has a holotype consisting of axial and appendicular elements, and associated material consisting of cranial, axial, appendicular and dermal bones. The completeness of the material makes it valuable for Middle Jurassic sauropod research. Moreover, recent dating of the sediments belonging to the Cañadón Asfálto Formation resulted in a much older age range for the latter, redating it and all vertebrates found there from a Callovian to a much older Aalenian- Bajocian age, and thus placing Patagosaurus in a critical time for sauropod evolution; the latest Early to the early Middle Jurassic. Patagosaurus has been found in two bonebeds, not far apart; the holotype and several other associated specimens from Cerro Condor Norte, north of the village of Cerro Cóndor by the Chubut river, and more associated material, as VIII well as another sauropod taxon, Volkheimeria chubutensis, from Cerro Condor Sur, a site close to the Chubut river. Both beds are lacustrine deposits. However, the original 1986 description of Patagosaurus consisted of a blend of the holotype and associated material, without clearly distinguishing the former from the latter. Moreover, some material from Cerro Condor Sur, specimen MACN-CH 934, has since been considered to be a taxon other than Patagosaurus or Volkheimeria. Thus, a revision of all material has been done. The holotype has been revised and the alpha taxonomy of Patagosaurus fariasi has been established, confirming or discarding old diagnostic characters as appropriate, and generating new diagnostic characters for the new assignment of associated material to Patagosaurus fariasi, Volkheimeria, sauropod indet., or a new taxon. After the revision of all material, the Cerro Condor Norte bonebed was found to be monospecific, and includes an ontogenetic series of a small juvenile Patagosaurus, an intermediate juvenile, a subadult, the holotype (being a subadult to adult specimen) and another adult specimen. Cerro Condor Sur proved to be more problematic; yielding Patagosaurus material of an adult and a fully grown large adult, material that can only be assigned to an indeterminate sauropod, due to the fragmentary nature, and finally, two potential new taxa, one of which is more likely a valid new taxon than the other: MACN-CH 934 and MACN-CH 230.
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