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Vertebrate Ichnology Behavioural, Sedimentological and Palaeoecological Aspects of Vertebrate Ichnology

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Vertebrate Ichnology Behavioural, Sedimentological and Palaeoecological Aspects of Vertebrate Ichnology Vertebrate Ichnology Behavioural, Sedimentological and Palaeoecological Aspects of Vertebrate Ichnology Jesper Milàn Ph.D. Thesis Department of Geography and Geology, Geology Section Faculty of Science University of Copenhagen April 2007 Jesper Milàn – Vertebrate Ichnology Title ________________________________________________________________________________ Behavioural, Sedimentological and Palaeoecological Aspects of Vertebrate Ichnology Ph.D. thesis Jesper Milàn Department of Geography and Geology Faculty of Science University of Copenhagen April 2007 Superviser: Docent Nanna Noe-Nygaard Department of Geography and Geology University of Copenhagen i Jesper Milàn – Vertebrate Ichnology Title ________________________________________________________________________________ “A footprint is not an organism but the by-product of dynamic contact between an organism and its environment” - Donald Baird 1957. ii Jesper Milàn – Vertebrate Ichnology Preface ________________________________________________________________________________ Preface This thesis is the result of a three-year Ph.D. programme supported by the Faculty of Science, University of Copenhagen and conducted at the Department of Geography and Geology – Geology Section, University of Copenhagen, under supervision of Docent Nanna Noe-Nygaard, Institute of Geography and Geology – Geology Section, University of Copenhagen. In accordance with the guidelines from the Faculty of Science, University of Copenhagen, the thesis comprises an introduction, overview and discussion and the following 12 papers. 1. Milàn, J. & Gierlinski, G. 2004. A probable thyreophoran (Dinosauria, Ornithischia) footprint from the Upper Triassic of southern Sweden. Bulletin of the Geological Society of Denmark, v. 51, p. 71–75. 2. Milàn, J., Clemmensen, L.B. & Bonde, N. 2004. Vertical sections through dinosaur tracks (Late Triassic lake deposits, East Greenland) – undertracks and other subsurface deformation structures revealed. Lethaia, v. 37, p. 285–296. 3. Milàn, J., Avanzini, M., Clemmensen, L.B., Garciá-Ramos, J.C. & Piñuela, L. 2006. Theropod foot movement recorded from Late Triassic, Early Jurassic and Late Jurassic fossil footprints. In Haris et al. (eds.). The Triassic-Jurassic Terrestrial Transition. New Mexico Museum of Natural History and Science Bulletin, v. 37, p 352–364. 4. Milàn, J., Loope, D.B. & Bromley, R.G. (submitted). Dinosaur tracks showing evidence of individual behaviour from the Lower Jurassic Navaho Sandstone, Coyotes Buttes locality, Utah, USA. Acta Palaeontologia Polonica. 5. Milàn, J. & Loope, D.B. 2007. Preservation and erosion of theropod tracks in eolian deposits; examples from the Middle Jurassic Entrada Sandstone, Utah, USA. The Journal of Geology, v. 115, p. 375–386. 6. Milàn, J. & Bromley, R.G. 2005. Dinosaur footprints from the Middle Jurassic Bagå Formation, Bornholm, Denmark. Bulletin of the Geological Society of Denmark, v. 52, p. 7–15. 7. Mateus, O. & Milàn, J. (Submitted). Ichnological evidence for giant ornithopod dinosaurs in the Late Jurassic Lourinhã Formation, Portugal. Oryctos. iii Jesper Milàn – Vertebrate Ichnology Preface ________________________________________________________________________________ 8. Hurum, J.H., Milàn, J., Hammer, Ø., Midtkandal, I., Amundsen, H. & Sæther, B. 2006. Tracking polar dinosaurs - new finds from the Lower Cretaceous of Svalbard. Norwegian Journal of Geology, v. 83, p. 397–402. 9. Milàn, J., Bromley, R.G., Titschack, J. & Theodouru, G. (in press). A diverse vertebrate ichnofauna from a Quaternary eolian oolite from Rhodes, Greece. In: Bromley, R.G., Buatois, L.A., Márango, M.G., Genise, J.F. & Melchor, R.N. (eds.), Sediment-organism interactions: A multifaceted ichnology. SEPM Special Publications, v. 88. 10. Milàn, J., Clemmensen, L.B., Buchardt, B. & Noe-Nygaard, N. (in prep). A Late Holocene tracksite in the Lodbjerg dune system, northwest Jylland, Denmark. New Mexico Museum of Natural History and Science Bulletin. 11. Bromley, R.G., Uchman, A., Milàn, J. & Hansen, K.S. (in press). Rheotactic Macaronichnus, and human and cattle trackways in Holocene beachrock, Greece: reconstruction of palaeoshoreline orientation. Ichnos. 12. Milàn, J. & Bromley, R.G. (in press). Do shod humans leave true tracks?. Ichnos. iv Jesper Milàn – Vertebrate Ichnology Acknowledgements ________________________________________________________________________________ Acknowledgements During my work on this thesis I have had the fortune to work together with a number of different researchers, each internationally recognized expert in their respective fields, and who in many cases appear as co-authors on my works. I thank my superviser Nanna Noe-Nygaard, Department of Geography and Geology, University of Copenhagen, for her patient supervision and her open mindedness to my many wild ideas. Richard G. Bromley, University of Copenhagen, has been a constant source of support and inspiration to new projects and his efforts in proof reading and improving my English has been endless. Octávio Mateus and all the staff and various volunteers at Museu da Lourinhã have given me numerous unforgettable excavation seasons and fruitful co-operations during the years. David B. Loope, University of Lincoln, Nebraska, are thanked for his hospitality and excellent co-operations on the fieldwork with tracks from the Jurassic sandstones of Utah. Andrew S. Gale, University of Portsmouth, is thanked for a very enjoyable track hunt around the Isle of Wight. I thank Jørn H. Hurum, Natural History Museum, Oslo for his hospitality and great co-operations. Marco Avanzini, Museo Tridentino di Scienze Naturali, Italy; Gerard Gierlinski, Polish Geological Institue, Poland; Jürgen Titschack, University of Erlangen, Georgios Theodorou, University of Athens, Alfred Uchman, Institute of Geological Sciences, Poland; Lars B. Clemmensen, Niels Bonde and Bjørn Buchardt all from Department of Geography and Geology, University of Copenhagen, have all been important partners in the various papers and projects. Eckart Håkansson, Department of Geography and Geology, University of Copenhagen, has always been helpful with good advises, proof reading and coffe drinking. I would like to thank the staff and students of the Department of Geology and Geography, Section for Geology, and the Geological Museum, University of Copenhagen for making my time here as enjoyable and memorable as it has been. If anybody feels that I have forgotten them in the previous listing please enter your name here _________________. The music of Lumsk, Tiamat, Rammstein and Theatre of Tragedy has been a great relief during the long hours of writing. And finally I sincerely thank my wonderful wife, Inken Mueller-Töwe for her support and patience with me, and my parents and family who always supported my choice to become a palaeontologist instead of getting a real job. v Jesper Milàn – Vertebrate Ichnology ________________________________________________________________________________ vi Jesper Milàn – Vertebrate Ichnology Contents ________________________________________________________________________________ Contents Title…………………………..………………..…………….……………….…………….………..i Preface………………………………………..…………………………………….………………iii Acknowledgements………………………...…………………….……………….…………………v Contents……………………………………..…………………………………………..…………viii Abstract……………………………………………………..……...……………..………………….1 Introduction………………………………………..…………………………………………………3 Summary of papers………………………………………………………………….……………….7 Material and methods…………………………………………………………….…………………21 Discussion…………………………………………………………………………………………..23 Conclusions…………………………………………………………………………………………30 References………………………………………………………………..…………………………32 Appendix: Paper 1: A probable thyreophoran (Dinosauria, Ornithischia) footprint from the Upper Triassic of southern Sweden Paper 2: Vertical sections through dinosaur tracks (Late Triassic lake deposits, East Greenland) – undertracks and other subsurface deformation structures revealed. Paper 3: Theropod foot movement recorded from Late Triassic, Early Jurassic and Late Jurassic fossil footprints. Paper 4: Dinosaur tracks showing evidence of individual behaviour from the Lower Jurassic Navaho Sandstone, Coyotes Buttes locality, Utah, USA. Paper 5: Preservation and erosion of theropod tracks in eolian deposits; examples from the Middle Jurassic Entrada Sandstone, Utah, USA. Paper 6: Dinosaur footprints from the Middle Jurassic Bagå Formation, Bornholm, Denmark. Paper 7: Ichnological evidence for giant ornithopod dinosaurs in the Late Jurassic Lourinhã Formation, Portugal. Paper 8: Tracking polar dinosaurs - new finds from the Lower Cretaceous of Svalbard. vii Jesper Milàn – Vertebrate Ichnology Contents ________________________________________________________________________________ Paper 9: A diverse vertebrate ichnofauna from a Quaternary eolian oolite from Rhodes, Greece. Paper 10: A Late Holocene tracksite in the Lodbjerg dune system, northwest Jylland, Denmark. Paper 11: Rheotactic Macaronichnus, and human and cattle trackways in Holocene beachrock, Greece: reconstruction of palaeoshoreline orientation. Paper 12: Do shod humans leave true tracks?. viii Jesper Milàn – Vertebrate Ichnology Abstract ________________________________________________________________________________ Abstract Twelve case studies, each representing different aspects of vertebrate ichnology, are published in order to examine the applications of vertebrate ichnology in reconstructing behaviour, sedimentological properties, and palaeoecology. Tracks of sauropods and thyrophorean
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