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A Revision of Temnodontosaurus Crassimanus (Reptilia: Ichthyosauria) From A revision of Temnodontosaurus crassimanus (Reptilia: Ichthyosauria) from the Lower Jurassic (Toarcian) of Whitby, Yorkshire, UK A thesis submitted to The University of Manchester for the degree of Master of Philosophy (MPhil) in the Faculty of Science and Engineering 2019 Emily J. Swaby School of Earth and Environmental Sciences TABLE OF CONTENTS TABLE OF CONTENTS ......................................................................................................................... 2 LIST OF PRIMARY FIGURES ................................................................................................................ 4 LIST OF PRIMARY TABLES ................................................................................................................. 8 APPENDIX A TABLES ........................................................................................................................... 9 APPENDIX B TABLES ........................................................................................................................... 9 ABSTRACT........................................................................................................................................... 10 DECLARATION .................................................................................................................................... 11 COPYRIGHT STATEMENT .................................................................................................................. 11 DEDICATIONS ..................................................................................................................................... 12 INSTITUTIONAL ABBREVIATIONS .................................................................................................... 13 1. ACKNOWLEDGEMENTS................................................................................................................. 14 2. INTRODUCTION ............................................................................................................................... 15 3. MATERIALS AND METHODS ......................................................................................................... 17 4. GEOLOGICAL BACKGROUND ...................................................................................................... 23 5. LITERATURE REVIEW .................................................................................................................... 26 5.1 A HISTORY OF ICHTHYOSAURS FROM THE YORKSHIRE COAST ........................... 26 5.2 ICHTHYOSAUR SPECIES FROM YORKSHIRE ............................................................. 29 5.2.1 Eurhinosaurus longirostris ................................................................................... 29 5.2.2 ?Temnodontosaurus acutirostris ......................................................................... 30 5.2.3 Temnodontosaurus crassimanus ......................................................................... 32 5.2.4 Ichthyosaurus communis ...................................................................................... 32 5.2.5 Stenopterygius sp. ................................................................................................. 34 5.2.6 Other Ichthyosaurs ................................................................................................. 35 6. COMPARATIVE MATERIAL ............................................................................................................ 37 6.1 SPECIMEN PB 1 ............................................................................................................... 38 6.2 SPECIMEN SMNS 15950 .................................................................................................. 39 6.3 SPECIMEN SMNS 50000 .................................................................................................. 41 6.4 SPECIMEN SMNS 17560 .................................................................................................. 43 7. THE HISTORY OF TEMNODONTOSAURUS CRASSIMANUS...................................................... 45 2 8. REDESCRIPTION OF THE HOLOTYPE OF TEMNODONTOSAURUS CRASSIMANUS AND COMPARISON WITH TEMNODONTOSAURUS TRIGONODON ....................................................... 48 8.1 COMPARATIVE DESCRIPTION ....................................................................................... 49 9. ASSESSMENT OF POSSIBLE ADDITIONAL SPECIMENS OF TEMNODONTOSAURUS CRASSIMANUS ................................................................................................................................... 80 9.1 SPECIMEN WHITM: SIM2546.S ....................................................................................... 80 9.2 SPECIMEN WHITM: SIM5 ................................................................................................ 87 9.3 SPECIMEN MANCH: LL. 16096 ....................................................................................... 90 9.4 SPECIMEN MANCH: L. 1688 ........................................................................................... 92 10. PHYLOGENETIC ANALYSIS ........................................................................................................ 99 10.1 RESULTS ...................................................................................................................... 100 11. DISCUSSION ................................................................................................................................ 101 11.1 COMPARISON, TAXONOMIC IDENTIFICATION AND INTERPRETATION ............... 101 11.2 LIMITATIONS ................................................................................................................ 103 12. CONCLUSION .............................................................................................................................. 104 13. FUTURE RESEARCH .................................................................................................................. 106 14. REFERENCES .............................................................................................................................. 107 15. APPENDICES ............................................................................................................................... 116 Appendix A: Measurements for Temnodontosaurus trigonodon specimens ............... 116 Appendix B: Measurements for additional Temnodontosaurus specimens ................. 124 Appendix C: List of phylogenetic characters for cladistic analysis ............................... 128 Appendix D: Results of second cladistic analysis ........................................................... 133 Appendix E: Individual species character coding used with the cladistic analysis ..... 134 Final word count, excluding references, appendices and figure explanations: 27,114 3 LIST OF PRIMARY FIGURES Figure 1. An embryo-bearing specimen of Stenopterygius (SMNS 6293) from the Toarcian of Germany, highlighting how ichthyosaur young were typically born tail-first. Photograph courtesy of Dr Erin Maxwell. Figure 2. Digital restoration of the large Temnodontosaurus alongside the smaller Stenopterygius, in comparison to a scuba diver (~ 2 m) as a scale by Fabio Manucci (Insacco et al., 2014). Figure 3. Map of the UK and of the Yorkshire coast, highlighting the location of Whitby and several other coastal localities, the majority of which are important ichthyosaur-bearing locations. Grey shaded areas represent Lower Jurassic rocks, including foreshore exposures. Modified from Benton and Taylor (1984) and Howarth (2002). Figure 4. A stratigraphic column of the lower three divisions of the Whitby Mudstone Formation, which yield many of the marine reptiles from Whitby (Swaby, 2018). The term ‘Liassic’ was formerly used within older literature to refer to the Lower Jurassic, however the word is now technically abolished and ‘Lias’ is used instead. Figure 5. YORYM: 2016.316; a cut and polished boulder (Block A and B) containing between six and eight ichthyosaur embryos, accompanied by illustrations of the identifiable sections of embryos (Block A1 and B1). Scale bar equals 10 cm (Boyd and Lomax, 2018, fig. 1). Figure 6. The practically complete specimen of Eurhinosaurus longirostris (SMNS 14931) from the Upper Lias of Germany. Photograph courtesy of Dr Erin Maxwell. Figure 7. A nearly complete skull in dorsal view of Temnodontosaurus acutirostris (NHMUK PV OR 15500a) from the Upper Lias of Whitby, Yorkshire. Scale bar equals 10 cm. Figure 8. Skull of Ichthyosaurus communis (SMNS 13111) from the Lower Lias of Yorkshire. Scale bar equals 5 cm. Modified from Massare and Lomax (2017, fig. 5C). Figure 9. Practically complete skeleton of Stenopterygius (SMNS 17500) from the Toarcian Posidonienschiefer, Lower Jurassic, Upper Lias. Photograph courtesy of Dr Dean Lomax. Figure 10. The nearly complete skull of the holotype specimen of Temnodontosaurus trigonodon (PB 1), on display at the Banz Castle Museum. Photograph courtesy of Dr Erin Maxwell. Figure 11. An early illustration of the skull of the holotype specimen of Temnodontosaurus trigonodon (PB 1) (Theodori, 1854). Figure 12. A nearly complete skeleton of Temnodontosaurus trigonodon (SMNS 15950). Scale bar equals ~ 160 cm. Figure 13. The nearly complete skeleton of Temnodontosaurus trigonodon (SMNS 50000). Scale bar equals ~ 170 cm. 4 Figure 14. A three-dimensional mounted skeleton of Temnodontosaurus trigonodon (SMNS 17560). Scale bar equals ~ 70 cm. Figure 15. The three-dimensional mounted
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