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Osteology of a Near-Complete Skeleton of Tenontosaurus Tilletti (Dinosauria: Ornithopoda) from the Cloverly Formation, Montana, USA

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Osteology of a Near-Complete Skeleton of Tenontosaurus Tilletti (Dinosauria: Ornithopoda) from the Cloverly Formation, Montana, USA Osteology of a Near-Complete Skeleton of Tenontosaurus tilletti (Dinosauria: Ornithopoda) from the Cloverly Formation, Montana, USA A report submitted in partial fulfilment of the requirements for the degree of MEarthSci Geology (Hons.), University of Manchester Jonathan Tennant 70406912 MEarthSci Geology (Hons.) School of Earth, Atmospheric and Environmental Sciences University of Manchester Williamson Building Oxford Road Manchester M13 9PL Osteology of Tenontosaurus tilletti Contents Figures and Tables..............................................................................................................................................3 1.0 Abstract .............................................................................................................................................................6 2.0 Introduction ......................................................................................................................................................7 2.1 Aims and Objectives....................................................................................................................................7 2.2 Institutional Abbreviations ........................................................................................................................9 3.0 Previous Work ............................................................................................................................................... 11 3.1 The Cloverly Formation ........................................................................................................................... 11 3.2 Geological Setting..................................................................................................................................... 13 3.3 Palaeoecology ........................................................................................................................................... 17 3.4 Biomechanics ............................................................................................................................................ 23 3.5 Palaeobiogeography ................................................................................................................................ 25 3.6 Phylogeny................................................................................................................................................... 36 4.0 Systematic Palaeontology ........................................................................................................................... 48 4.1 Revised Diagnosis ..................................................................................................................................... 48 4.2 Material Studied ....................................................................................................................................... 49 4.31 Skull ...................................................................................................................................................... 53 4.32 Vertebral Column .............................................................................................................................. 60 4.33 Pectoral Girdle.................................................................................................................................... 67 4.44 Forelimb .............................................................................................................................................. 71 4.35 Pelvic Girdle ........................................................................................................................................ 80 4.36 Hindlimb .............................................................................................................................................. 87 5.0 Discussion....................................................................................................................................................... 92 6.0 Conclusions .................................................................................................................................................... 99 7.0 Acknowledgements .................................................................................................................................... 102 8.0 References.................................................................................................................................................... 103 9.0 Appendices................................................................................................................................................... 117 9.1 Appendix 1: Morphological Description of LL.12275 (Tenontosaurus tilletti) ............................. 117 9.11 Skull .................................................................................................................................................... 117 9.12 Pectoral Girdle.................................................................................................................................. 126 9.13 Forelimb and Manus ....................................................................................................................... 130 9.14 Pelvic Girdle ...................................................................................................................................... 139 9.15 Hindlimb and Pes ............................................................................................................................. 145 9.2 Appendix 2: Comparative Morphology I – Hypsilophodon foxii..................................................... 152 9.21 Skull .................................................................................................................................................... 152 9.22 Pectoral Girdle.................................................................................................................................. 161 1 Osteology of Tenontosaurus tilletti 9.23 Forelimb ............................................................................................................................................ 164 9.24 Pelvic Girdle ...................................................................................................................................... 168 9.25 Hindlimb ............................................................................................................................................ 173 9.3 Appendix 3: Comparative Morphology II - Thescelosaurus neglectus .......................................... 177 9.4 Appendix 4: Comparative Morphology III – Valdosaurus canaliculatus ...................................... 181 9.5 Appendix 5: Comparative Morphology IV – Lesothosaurus diagnosticus .................................... 187 9.6 Appendix 6: Character description and recoded data matrix ........................................................ 192 2 Osteology of Tenontosaurus tilletti Figures and Tables Figure 1 – Fully articulated display specimens of Tenontosaurus tilletti, including LL.12275, in bipedal and quadrupedal stances (page 10) Figure 2 – Typical outcrop of the Cloverly Formation, Montana (page 13) Figure 3 – Hypaxial and epaxial ossified tendons dorsal and ventral to caudal series in (page 23) Figure 4 – Cladograms from Barrett and Han (2009), Butler et al. (2009), and Sereno (1999) (page 38) Figure 5 – Cladograms from Weishampel and Heinrich (1992), Winkler et al. (1997), and Weishampel et al. (2003) (page 40) Figure 6 – Cladograms from Pisani et al. (2002), Galton (2009), Varrichio et al. (2007) and Boyd et al. (2009) (page 41) Figure 7 – Various reconstructed skulls of ornithischian dinosaurs (including T. tilletti), (Coria and Salgado, 1996) (page 53) Figure 8 – Reconstructed sketch of Tenontosaurus skull, (Maidment and Porro, 2010) (page 54) Figure 9 – Sketches of LL.12275 cranium in lateral, rostral and dorsal aspects (page 56) Figure 10 – Photographs of cranium in lateral, rostral and dorsal aspects (page 57) Figure 11 – Photograph and sketch of lower jaw in lateral aspect (page 58) Figure 12 – Photographs of articulated and mounted dorsal and cervical series, and atlas and axis (page 62) Figure 13 – Photographs of articulated sacrum (with ilia) in lateral aspects (page 63) Figure 14 – Graphs illustrating vertebral dimensions recorded in Table 4 (page 65) Figure 15 – Sketch and photograph of paired sternal plates (page 68) Figure 16 – Sketches of sinistral and dextral scapulae and coracoids in lateral aspect (page 69) Figure 17 – Sinistral proximal humerus sketch and photo, and dextral humerus sketch in lateral aspect (page 71) Figure 18 – Photographs of dextral humerus in lateral, cranial and caudal aspects (page 72) Figure 19 – Sketches of dextral humerus in cranial and lateral aspects (page 73) Figure 20 – Sketches of dextral ulna in cranial, caudal (proximal half), distal and proximal aspects (page 75) 3 Osteology of Tenontosaurus tilletti Figure 21 – Sketch and photograph of dextral ulna in lateral aspect (page 76) Figure 22 – Sketches of dextral and sinistral ulnae in lateral aspect (page 77) Figure 23 – Photograph of artificially articulated, mounted complete dextral forelimb (page 79) Figure 24 – Sketches of sinistral and dextral ischia in lateral aspect (page 81) Figure 25 – Photographs of sinistral and dextral ilia in lateral aspect (page 83) Figure 26 – Sketches of dextral ilium in lateral and proximal medial and ventral aspects (page 84) Figure 27 – Sketch of sinistral ilium in
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