Of Mice and (Neanderthal) Men: The small mammal record of the Middle to Upper Paleolithic transition in the Swabian Jura, Germany Dissertation der Mathematisch-Naturwissenschaftlichen Fakultät der Eberhard Karls Universität Tübingen zur Erlangung des Grades eines Doktors der Naturwissenschaften (Dr. rer. nat.) vorgelegt von Sara E. Rhodes, M.Sc. aus Toronto (Kanada) Tübingen 2019 Gedruckt mit Genehmigung der Mathematisch-Naturwissenschaftlichen Fakultät der Eberhard Karls Universität Tübingen. Tag der mündlichen Qualifikation: 8. Juli 2019 Dekan: Prof. Dr. Wolfgang Rosenstiel 1. Berichterstatter: Prof. Nicholas J. Conard, PhD 2. Berichterstatter: PD Dr. Britt M. Starkovich Table of Contents I ABBREVIATIONS ................................................................................................................ III II ACKNOWLEDGEMENTS .................................................................................................. IV III SUMMARY ......................................................................................................................... VI IV LIST OF PUBLICATIONS ............................................................................................... XIV V PERSONAL CONTRIBUTION .......................................................................................... XIV 1.0 INTRODUCTION ............................................................................................................... 1 1.1 NEANDERTHALS IN THE EUROPEAN CONTEXT ..................................................................... 1 1.2 THE MIDDLE TO UPPER PALEOLITHIC TRANSITION ............................................................. 3 1.3 THEORIES OF NEANDERTHAL EXTINCTION .......................................................................... 6 1.4 THE MIDDLE AND UPPER PALEOLITHIC ARCHAEOLOGY OF THE SWABIAN JURA .............. 17 1.4.1 THE SWABIAN JURA ................................................................................................................. 17 1.4.2 HOHLE FELS CAVE ................................................................................................................... 18 1.4.3 GEIßENKLÖSTERLE CAVE ........................................................................................................ 20 1.4.4 THE UPPER PALEOLITHIC CULTURAL FLUORESCENCE ............................................................ 22 1.5 THE MIDDLE AND UPPER PALEOLITHIC ACH VALLEY PALEOECOLOGICAL RECORD ......... 24 1.5.1 THE MIDDLE PALEOLITHIC PALEOENVIRONMENTAL RECORD ................................................. 24 1.5.2 THE AURIGNACIAN PALEOENVIRONMENTAL RECORD ............................................................ 26 2.0 OBJECTIVES AND EXPECTED OUTPUT OF RESEARCH ........................................ 29 3.0 METHODS ........................................................................................................................ 31 3.1 SAMPLE ............................................................................................................................... 31 3.2 TAXONOMIC IDENTIFICATIONS ........................................................................................... 32 3.2.1 MORPHOLOGICAL IDENTIFICATION ......................................................................................... 32 3.3 TAPHONOMIC IDENTIFICATIONS ......................................................................................... 43 3.4 DIVERSITY MEASURES ........................................................................................................ 44 3.5 PALEOENVIRONMENTAL RECONSTRUCTIVE METHODS ....................................................... 45 3.5.1 MODIFIED INDICATOR SPECIES METHOD ................................................................................ 45 3.5.2 HABITAT WEIGHING METHOD ................................................................................................. 46 4.0 RESULTS .......................................................................................................................... 46 4.1 TAPHONOMIC RESULTS ....................................................................................................... 47 4.2 TAXONOMIC RESULTS ......................................................................................................... 49 4.3 PALEOENVIRONMENTAL RESULTS ...................................................................................... 50 4.3.1 PREDATOR HUNTING BEHAVIOURS AND HABITAT PREFERENCES ............................................ 50 I 4.3.2 MODIFIED INDICATOR SPECIES METHOD .................................................................................. 52 4.3.3 HABITAT WEIGHTING METHOD ................................................................................................ 54 5.0 DISCUSSION .................................................................................................................... 55 5.1 CORRELATIONS WITH THE ACH VALLEY RECORD .............................................................. 55 5.2 CORRELATIONS WITH THE GREENLAND STADIAL/INTERSTADIALS .................................... 56 5.3 CORRELATION WITH OTHER FAUNAL PALEOECOLOGICAL SIGNALS ................................... 58 5.4 CORRELATION WITH OTHER PALEOECOLOGICAL ARCHIVES ............................................. 60 6.0 CONCLUSIONS ................................................................................................................ 62 REFERENCES ........................................................................................................................... 67 APPENDIX ................................................................................................................................. 90 II I Abbreviations NISP – Number of Identified Specimens MNI – Minimum Number of Individuals NTAXA – Number of Taxonomic Designations at the genera and/or species level. AMH – Anatomically Modern Humans OIS – Oxygen Isotope Stage GS – Greenland Stadial GIS – Greenland Interstadial D-O – Dansgaard-Oeschger event (numbered) H – Heinrich event (numbered) ACC – anterioconid complex AL – anterior lobe PL – posterior lobe BRA – buccal re-entrant angle LRA – lingual re-entrant angle BSA – buccal salient angle/edge LSA – lingual salient angle/edge T – triangle (numbered) Qu. – quadrant SDQ – Schmelz-Dicken-Quotient LE – leading edge TE – trailing edge LUF – length of upper condylar facet LLF – length of lower condylar facet HC – height of the condyle TRW – trogonid width TAW – talonid width W – width L - length III II Acknowledgements By any measure of the word, the undertaking of a thesis project is a community effort. I consider myself exceptionally lucky to have had a true community of colleagues, friends, and family from which I drew guidance, strength, and motivation over the past four and a half years. First and foremost, I would like to express my sincere gratitude to my two supervisors, Nicholas Conard and Britt Starkovich, for their intellectual and personal support throughout the entire process of this thesis. Together, we weathered every storm and celebrated every victory, and I cannot thank you both enough. Nick, thank you for guiding me through the history of research on the German Paleolithic, and for teaching me the value in recognizing the long record of scientific endeavor of which I can now proudly consider myself a part. Our season together at Sibudu and our long discussions over scotch and cigarettes (or in your case, a pipe) about life and academia are fond memories I will keep for years. Britt, thank you for your trust in my scientific abilities, for providing the space to work independently, and for your messages of encouragement when they were most needed. I am honored to have been your first doctoral student and I hope this thesis lives up to that role. I would also like to extend sincere gratitude to Dr. Simone Riehl, Prof. Michael Bolus, and Prof. Hervé Bocherens for agreeing to sit on my doctoral committee and for their faith in my abilities as an academic. Further thanks go to the various collaborators with whom I have had the pleasure of working throughout this project. My sincerest thanks go to Dr. Reinhard Ziegler for teaching me all that I know about small mammal dental morphology and evolution. This thesis would not have been possible without your patience and generosity. To Dr. Susanne Münzel, for putting me in contact with Reinhard initially, for countless hours of discussion and advice, and your continued and valued friendship. Drs. Chris Miller, Simone Riehl, and Michael Bolus, thank you for providing your time and expertise. Mima Fatima Batalovic and Monika Doll, thank you for helping me navigate the German academic system. Dr. Lutz Maul, thank you for inviting me to be a part of the INQUA Ground Squirrels on the March Project, which provided the opportunity to meet the superstars of our field and taught me more than I had ever imagined I could know about squirrels. My additional thanks to you and Dr. Thijs van Kolfschoten for always being available to help with any questions or difficult ID’s which arose. IV I am indebted to hundreds of student excavators who volunteered their time to sort out tiny bones from the fine sediment of Hohle Fels and Geißenklösterle caves. Without their painstaking tweezing, this thesis would not have been possible. Furthermore, I must extend my sincere gratitude to Maria Malina, Alexander Janas, Sarah Rudolf, Marziyah Zare Khalili and Anthony Fernandes for maintaining