On the individual learning of primate material culture Dissertation der Mathematisch-Naturwissenschaftlichen Fakultät der Eberhard Karls Universität Tübingen zur Erlangung des Grades eines Doktors der Naturwissenschaften (Dr. rer. nat.) vorgelegt von Miss Elisa Bandini aus Boulogne Billancourt, Paris, France Tübingen 2018 Gedruckt mit Genehmigung der Mathematisch-Naturwissenschaftlichen Fakultät der Eberhard Karls Universität Tübingen. Tag der mündlichen Qualifikation: 16.07.2018 Dekan: Prof. Dr. Wolfgang Rosenstiel 1. Berichterstatter: Prof. Nicholas J. Conard PhD 2. Berichterstatter: Dr. Claudio Tennie To my parents, and theirs. “Anything which I have done which you may consider worthwhile, has been made possible by the genius of my mother and father” -Howard Hughes Chimpanzee algae scooping (drawing by William Daniel Snyder, 2018) Acknowledgments It is not often that we have the opportunity to formally thank the people in our lives. Although I will never have enough (or the right) words to properly thank everyone who has supported, understood and enriched my life over the last couple of years, I hope this clumsy attempt demonstrates how much I appreciate all that everyone has done for me over the years. I will never be able to fully repay the incredible debt I owe to my parents. Without you, none of this would have been possible. Everyday I feel grateful to have been raised by such supportive, open-minded and intelligent people who taught me all I that I know. Everything I do, I do in an attempt to make you proud, as that is the only way I can envision to try to repay my debt. This PhD is for my father, who never stopped encouraging me to push a bit harder in everything I do, leading by example, and always being there for me when I thought I couldn’t go any farther. My father introduced me to the things I love most in life, and some of my happiest memories are of hiking in the Dolomiti all together. I will be forever grateful that you opened this world up for me. This PhD is for my mother, my best friend and my role model. I hope someday to become half of the strong, independent, brilliant and beautiful woman that you are. I am incredibly grateful for everything my sister has done for me over the last 26 years. I couldn’t have wished for a better big sister. Marghe, everyday you amaze me with your strength, resilience and passion for everything you do. You are an inspiration, and I know I have so much more to learn from you. Of course, this PhD would not have been possible without my supervisors. I thank my new supervisor, Nick Conard, who welcomed me so warmly to Tübingen. I am also very grateful to Sarah Beck, who provided such valuable advice and support over the years, and who, despite being the busiest person I know, always had time to share a cup of (decaf) coffee and a word of wisdom. Sarah is one of my role models in academia, and I am very grateful to have been part of her lab. Lastly, I would like to thank Claudio Tennie for his immense generosity over the years, which spanned from invaluable financial to academic support. From Claudio I gained the courage to stand- up for my ideas, even when it resulted in going against the status quo, and inevitably ruffling some (pretty important) feathers. Claudio also demonstrated the importance of taking our work seriously and never cutting corners, whilst always staying humble enough to be open to new ideas. Yet, I also learnt that this attitude goes hand-in-hand with not taking everything too seriously. There is always time for Snoop Lion and (terrible) puns in the Tennie-lab, and (thus) I will forever be grateful that I laughed so much during my PhD. One of the best things I gained from this PhD has been the many friendships made over the years. Eva: thank you for everything. You are one of the best friends, sisters, daughters, and women I have had the pleasure of meeting. Your passion, drive, strength and kindness are an inspiration and I will never stop being in awe of you. You are right, we did create something special. Damien: despite being a silly goose, you are one of the wisest people I know. Your unwavering integrity, work ethic and generosity are astounding, and I so lucky to have you in my life. To all my other Brummie friends: the fifth-floor fitties, Abi, George, Rèmi, Daniella, Rachel. I will never say that I liked Birmingham, but all of you made being there worth it. I am so grateful to all my fellow inmates. Alba, thank you for sitting with me on the floor and talking me through all of my (many) crises of confidence, for reading (and re-reading) my work and providing such insightful comments, for putting up with Vivaldi’s four seasons on repeat, and for being the ambitious, funny and sweet person you are. Snorellana: thanks for making the last couple of months in the prison such fun. Will, thank you for not getting angry every time I make a joke about the US, and for creating the beautiful artwork used in my thesis, you are incredibly talented! Thank you to my new Tübingen family: Recha for being so welcoming when I first arrived, Mima for letting me convince you to go to boot-camps in nightclubs on a Sunday, and for everything you do for all of us. The Schloß would be in ruins without you. Michi, for being such an integral part of my life in Tübingen. I will never stop being amazed, and amused, by your never-ending peep-show references. And of course, Viola, for all the lunches, coffees and chats over the past year where you patiently answered all my questions and taught me so much about the ‘real archaeology’. You are so passionate about what you do, I have no doubt that you will succeed. Thank you to the rest of the Bandini & Gottardi gang, for always reminding me where home is. And to the Coyles, my English family, thank you for welcoming me into your family, for always asking how work was going (and for listening to the inevitable subsequent rant), for accepting me even if I don’t drink or like Coldplay, and for always having a home for us in Kent. I am so grateful to all the research assistants who I have had the pleasure of working with over the years: Flo Rocoque, Alice Coombes, Alice Cope, Polly Cowdell, David Boysen, you all became good friends and made testing so much more fun. To my wonderful Zambian dream-team: Linda (Twinnie), you made Chimfunshi the best experience for me. Thank you for helping me with testing, for our run-days-fun-days, our late-night giggles and for all your generous advice on life and work. You are the best Twinnie I could have asked for. Ruthie: thank you for showing me how important it is to stand-up for what you believe in, and for demonstrating what passion and hard-work can lead to. And for letting me call you Ruthie. Silvan: thanks for cooking for us, for making me laugh until my stomach hurt and for being a real- life example of why one shouldn’t walk around the bush without shoes. Thank you to all the inspiring researchers I met along the way, who put aside their differing views to help out a struggling PhD student. Thank you especially to Lydia Luncz & Michael Gumert, for their patience, support and invaluable advice on the macaque study, and to Katja Liebal & Cathy Crockford for adopting me in Zambia and providing such insightful comments on my studies. Cathy, I will never forget how you taught me chimpanzee vocalisations over dinner and stitched my (last pair!) of trousers. Thank you to all the testing institutions that allowed me to carry-out my research: Twycross Zoo, il Bioparco di Roma, Curraghs Wildlife Park, Shepreth Wildlife Park, Chimfunshi Wildlife Orphanage, Think-Tank and to all the keepers and research managers that helped so much during testing. And of course, thanks to all the primates that participated in my studies. I hope the peanut butter was worth it. My last thank you goes to Ben. I will never be able to express how grateful I am for having you with me throughout this whole process. This PhD would be a shadow of itself without you. But, more importantly, I would be a shadow of myself without you. I cannot wait to see what our next adventures will bring. Look at that, we did it! Abstract The cognition behind tool-use in primates is studied across a wide variety of fields, ranging from animal behaviour, biology, archaeology, anthropology, history, to psychology. This thesis encapsulates an interdisciplinary approach to studying and interpreting the phylogenetic origins of individual learning behind tool-use in non- human primates. The introduction chapter presents the background to this thesis, introducing concepts such as the Zone of Latent Solutions hypothesis (Tennie et al., 2009), Vygotsky’s (1978) ZAD and ZPD theories, and the current state of animal tool-use research. Chapter II provides a literature review of individual learning of tool-use across animal species, presenting a novel picture of animal tool-use: one in which individual learning, rather than social learning, can drive and sustain animal material culture. Chapter III then discusses the positive results of four independent experimental studies on the individual learning abilities of naïve, captive chimpanzees (Pan troglodytes) across wildlife parks in Europe and a sanctuary in Africa.