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Junggarsuchus Sloani, an Early Late Jurassic Crocodylomorph with Crocodyliform Affinities

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Junggarsuchus Sloani, an Early Late Jurassic Crocodylomorph with Crocodyliform Affinities Junggarsuchus sloani, an Early Late Jurassic Crocodylomorph with Crocodyliform Affinities by Alexander Altieri Ruebenstahl B.S. in Biology, May 2018, The George Washington University B.S. in Geology, May 2018, The George Washington University A Thesis submitted to The Faculty of The Columbian College of Arts and Sciences of The George Washington University in partial fulfillment of the requirements for the degree of Master of Science August 31st, 2019 Thesis directed by James M Clark Professor novitas of Biology © Copyright 2019 by Alexander Altieri Ruebenstahl All rights reserved ii Acknowledgments I would like to give my sincere thanks and appreciation to James Clark, Catherine Forster, Andrew Moore, Joseph Stiegler for their help in this project and my education. I would also like to thank the George Washington University, the Institute of Vertebrate Paleontology and Anthropology, the University of Berkeley at California, the Carnegie Museum of Natural History and the American Museum of Natural History for access to specimens. Thanks to the Hennig Society for providing TNT. The skull of Junggarsuchus was prepared by Wang Hai-jun. The skulls of Dibothrosuchus and Junggarsuchus were scanned by Yi Hongyu. This work was supported by NSF grant EAR 1636753. iii Abstract of Thesis Junggarsuchus sloani, an Early Late Jurassic Crocodylomorph with Crocodyliform Affinities The holotype of Junggarsuchus sloani, from the early Late Jurassic of Xinjiang, China, consists of a nearly complete skull and the anterior half of an articulated skeleton, including the pectoral girdles, nearly complete forelimbs, vertebral column, and ribs. This taxon shares many features with a cursorial group of crocodylomorphs, known as ‘sphenosuchians’ whose relationships are poorly understood. However, it also displays several derived crocodyliform features that are not found among members of this group. A phylogenetic analysis corroborates the hypothesis that Junggarsuchus is closer to Crocodyliformes than Dibothrosuchus and Sphenosuchus, but not as close to crocodyliforms as Almadasuchus and Macelognathus, which includes extant crocodylians, and that the “Sphenosuchia” are a paraphyletic assemblage. Two other species of “sphenosuchians,” Dibothrosuchus elaphros and Sphenosuchus acutus, are hypothesized to be more closely related to Crocodyliformes while the rest of the ‘Sphenosuchia’ form several smaller groups and are largely unresolved. We find that Dibothrosuchus is not as closely related to crocodyliforms as Junggarsuchus, but also possesses several unique autapomorphies. We also report an elongate blade-like coronoid in Junggarsuchus that is also widely present in ‘sphenosuchians’. iv Table of Contents Acknowledgements……………………………………………………………………...iii Abstract of Thesis……………………………………………………………………….iv List of Figures……………………………………………………………………………vi List of Tables……………………………………………………………………………vii Institutional Abbreviations…………………………………………………………....viii Chapter 1: Introduction……………………………………………………………………1 Chapter 2: Methods and Materials……………………………………………………….10 Chapter 3: Systematic Paleontology…………………………………………………......16 Chapter 4: Description of Junggarsuchus and comparison with Dibothrosuchus………21 Chapter 5: Phylogenetic Results……………………………………………………......105 Chapter 6: Discussion…………………………………………………………………..117 Chapter 7: Conclusion……………………………………………………………….....125 Literature Cited:………………………………………………………………………..127 Appendices……………………………………………………………………………..142 v List of Figures Figure 1: Material of Junggarsuchus sloani holotype IVPP 14010……………………218 Figure 2: Generalized tree of crocodylomorph relationships…………………………..219 Figure 3: Life reconstruction of Junggarsuchus sloani………………………………...220 Figure 4: Skulls of Junggarsuchus and Dibothrosuchus in left lateral view…………...221 Figure 5: Skulls of Junggarsuchus and Dibothrosuchus in dorsal view……………….223 Figure 6: Skulls of Junggarsuchus and Dibothrosuchus in ventral view……………...224 Figure 7: Skulls of Junggarsuchus and Dibothrosuchus in occipital view……………..225 Figure 8: Braincase of Junggarsuchus …………………………………………………227 Figure 9: Ear region of Junggarsuchus…………………………………………………229 Figure 10: Braincase of Dibothrosuchus……………………………………………….231 Figure 11: Mandible of Junggarsuchus……………………………………………..….232 Figure 12: Mandible of Dibothrosuchus………………………………………………..234 Figure 13: Shoulder bones of Junggarsuchus and Dibothrosuchus…………………....235 Figure 14: Left forelimb of Junggarsuchus and Dibothrosuchus………………………236 Figure 15: Left wrist bones of Junggarsuchus and Dibothrosuchus…………….……..237 Figure 16: Left manus of Junggarsuchus and Dibothrosuchus………...………………238 Figure 17: Axis and atlas of Junggarsuchus………………………………………..….239 Figure 18: Cervical vertebrae of Junggarsuchus and Dibothrosuchus ………………...240 Figure 19: Dorsal vertebrae and ribs of Junggarsuchus………………………………..241 Figure 20: Phylogenetic tree with Tennant’s characters and equal weights……………242 Figure 21: Phylogenetic tree with Tennant’s characters and implied weights…………243 Figure 22: Phylogenetic tree without Tennant’s characters and with implied weights...244 vi List of Tables Table 1: Table for CI, RI, and Steps from Phylogenetic analyses without Tennant’s characters……………………………………………………………………………….245 Table 2: Table for CI, RI and steps from Phylogenetic analyses with Tennant’s characters……………………………………………………………………………….245 Table 3: Relationships and node support for groups in analyses without Tennant’s characters……………………………………………………………………………….246 Table 4: Relationships and node support for groups in analyses with Tennant’s characters……………………………………………………………………………….248 Table 5: Table of Synapomorphies for group found……………………………………249 vii Institutional Abbreviations AMNH: American Museum of Natural History (Fossil Reptiles), New York, United States BP: Evolutionary Studies Institute (formerly Bernard Price Institute for Palaeontological Research), University of the Witwatersrand, Johannesburg, South Africa CM: Carnegie Museum of Natural History, Pittsburg, PA United States CUP: Fujen Catholic University of Peking (Beijing) collection in Field Museum of Natural History IGM: Institute of Geology, Mongolian Academy of Sciences IVPP: Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing, China UCMP: University of California Museum of Paleontology, Berkeley, United Stat viii INTRODUCTION The Shishugou Formation of Xinjiang, China is a continuous series of sediments spanning the poorly known late Middle to early Late Jurassic (Eberth et al., 2001; Clark et al., 2006). The lower part of the formation has yielded a variety of turtles, brachyopoid amphibians, a mammaliaform, and theropod and sauropod dinosaurs while the upper contains a more diverse fauna of dinosaurs and non-dinosaurian amniotes. An expedition in 2001 recovered the skull and part of the postcranial skeleton of an unknown crocodylomorph, briefly described and named Junggarsuchus sloani by Clark et al. (2004), from the lower part of the Shishugou, now dated to the early Late Jurassic (Choiniere et al, 2010; see below). This taxon shares many similarities—such as adaptations toward cursoriality and an elongate, rod-like coracoid—with basal crocodylomorph archosaurs known as ‘sphenosuchians.’ However, it also shares features—such as modifications to the braincase that solidify the skull—with Crocodyliformes, the group that includes modern crocodylians. Furthermore, Junggarsuchus possesses autapomorphies not seen in any other crocodylomorph, such as a surangular foramen and a massively dorso-ventrally expanded and pneumatic basisphenoid, which is farther expanded than that in Sphenosuchus (Walker, 1990). The ‘Sphenosuchia’ (Bonaparte 1971, 1982) are archosaurs known from the Late Triassic to the Late Jurassic (Clark et al. 2000, Göhlich et al. 2005; Leardi et al, 2017) that fall within the Crocodylomorpha (Hay 1930 emend Walker 1970) but outside the Crocodyliformes. The Crocodyliformes (includes modern crocodylian species and their fossil relatives that possess specializations that solidify the skull); ‘sphenosuchians’ are 1 the most basal crocodylomorphs. At least 13 valid monotypic genera are considered potential sphenosuchians: Sphenosuchus (Haughton 1915; Walker 1990), Saltoposuchus (Heune 1921; Sereno and Wild, 1992), Hallopus (Marsh 1877; Walker 1970), Terrestrisuchus (Crush 1984), Dibothrosuchus (Simmons 1965; Wu and Chatterjee 1993), Hesperosuchus (Colbert 1952), Pseudhesperosuchus (Bonaparte 1971), Litargosuchus (Clark and Sues 2002; Pedeticosaurus of Gow and Kitching 1988), Kayentasuchus (Clark and Sues 2002), Dromicosuchus (Sues et al. 2003), Macelognathus (Marsh 1884; Göhlich et al. 2005), Almadasuchus Pol et al. (2013) and Junggarsuchus Clark et al. (2004). Phyllodontosuchus (Harris et al., 2000), Trialestes (Nesbitt et al., 2013); and Redondavenator (Nesbitt et al., 2005), are ‘sphenosuchians’ but known from incomplete or poorly preserved material and their affinities are not well understood. Terrestrisuchus has been considered a junior synonym of Saltoposuchus (e.g., Benton and Clark, 1988), or as distinct taxa (e.g., Sereno and Wild, 1992); Allen (2003) considered Terrestrisuchus material to be juvenile individuals of Saltoposuchus. Nesbitt (2011) tentatively recognized Terrestrisuchus, but considered the specimen assigned to Hesperosuchus by Clark et al. (2000), Carnegie Museum 29894, to potentially belong to a different taxon due to its younger age within the Chinle Formation and the lack of autapomorphies shared by this specimen and the holotype
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