Université de Liège Institut Royal des Sciences Naturelles Département des Sciences et Gestion de Belgique de l’Environnement Operational Directorate Natural Environment Unite de Biologie du Comportement Conservation Biology Unit Eco-ethology of a population of bonobos (Pan paniscus) living in the western forest-savannah mosaics of the Democratic Republic of Congo Dissertation submitted for the degree of Doctor of Sciences Adeline Serckx April 2014 Collaborating institutions: Max Planck Institute for Evolutionary Anthropology – Department of Primatology Université de Kinshasa - Ecole Régionale post-universitaire d’Aménagement et de gestion Intégrés des Forêts et Territoires tropicaux WWF - Congo JURY COMPOSITION FOR THE THESIS DEFENSE Emmanuel Serusiaux President Université de Liège Jan Bogaert Reporter Gembloux AgroBiotech. - Université de Liège Zjef Pereboom Reporter Royal Zoological Society of Antwerp, Center of Research and Conservation University of Utrecht, the Netherlands Barbara Fruth Reporter Royal Zoological Society of Antwerp, Center of Research and Conservation Ludwig-Maximilian University, Germany Hjalmar Kühl Reporter Max Planck Institute for Evolutionary Anthropology, Germany German Centre for Integrative Biodiversity Research, Germany Alain Hambuckers Reporter Université de Liège Roseline C. Beudels-Jamar Co-supervisor Institut Royal des Sciences Naturelles de Belgique Marie-Claude Huyen Co-supervisor Université de Liège Pascal Poncin Supervisor Université de Liège i “A long time ago, Bonobo and the humans were living together in the villages. One day, the tax inspector visited the village and Bonobo had no money. For lack of two francs, Bonobo run away in the forest with his wife. Too shameful of its behaviour and while hairs were growing on his body, Bonobo did not dare to come back in the village and became the Human of the forest.” “A long time ago, the oil palm and the raphia palm were fighting to know which one was the most powerful. The oil palm said “my fruits provide oil”, the raphia palm retorted “with my branches, clothes are weaved”. While they could not find a common ground, they asked Bonobo to stop the quarrel. Bonobo agreed that oil palm was right. The raphia palm was angry and responded “since you gave right to the oil palm, I take back my branches with which your clothes are weaved”. He took back the clothes and Bonobo was naked. Shameful, Bonobo ran away in the forest and never came back in the village.” Teke tales ii ACKNOWLEDGEMENTS / REMERCIEMENTS When I started my PhD thesis, I never imagined how much this project would become a wonderful adventure. I learnt so much about sciences, I met so many interesting people. I also had the opportunity to experience the life in Congo. I never expected I should have to fight against ants to protect my house, to spend 36 hours on a plastic seat in a boat to reach the field or to attach my phone on a stake to be able to make phone calls! But it was so wonderful to be immersed in the Teke culture, I only keep good memories of all these months spent in the field. I must confess that I do not consider this thesis as my work but rather as a wonderful project that I had the chance to conduct. So many people help me by giving time, energy, advices and I really would like to thank you all for your contributions in my thesis! Let’s go back a few years ago to remember thanks to whom this project has been possible. November 2008 Roseline, I remember when I entered for the first time in your office. I did not have any idea of what I could do in my life but I was sure that I wanted to work in conservation biology. You asked me “if you had the choice, what would you really like to do?”. I was dreaming about bonobos and this wonderful adventure started. Thanks you so much for having opened the doors of my deepest dreams, thanks you for all your advices, for sharing your experience, for all the time you spent to correct my work even when you were travelling across the world. You really support me since the beginning and nothing would have been possible without you! December 2008 One month later, I arrived at the University of Liege with an outline of this PhD project. I really thank Pascal Poncin for welcoming me in his team, for having supported my project and for giving me the opportunity to work in the Primatology Research Group. The second person who gave me the opportunity to accomplish this project is without any doubt Marie-Claude. You gave me judicious advices, constant support, so many times for reading, correcting and commenting my writings. You always find the solution to overcome each obstacle in this PhD and I cannot thank you enough for what you have done. September 2009 My thesis really started after months of preparation and thanks to the funding of the FRIA (Fonds pour la Formation a la Recherche dans l’Industrie et l’Agriculture) and of the F.R.S- F.N.R.S (Fonds National pour la Recherche Scientifique). Other funding also helped me later, that I should already thank: the University of Liege, the Leopold III foundation and the ERAIFT (Ecole Régionale post-universitaire d’Aménagement et de gestion Intégrés des Forêts et Territoires tropicaux). iii Since this period, I am also benefiting from the help and the advices of Alain Hambuckers and Regine Vercauteren. Thanks you so much for having participated to the development of this work! I also would like to thank Jan Bogaert and Jean-Louis Doucet, who were always present to answer my questions. I warmly thank all my colleagues from the University and the Museum for those funny lunch time and for the nice atmosphere in our offices. A special thanks to Barbara, Fany and Chab for the great times at the congresses. March 2010 When I arrived for the first time in the field, I met Pupa (Valentin Mbenzo). You gave yourself the mission to make me meet everybody in the villages, to teach me everything about the place, the local culture, the bonobos and your work. I am really convinced this is thanks to you that all my work on the field was possible, even when you were not there anymore. Thanks you for continuing to take care of me and I really hope that we could have new opportunities to work together. I am grateful to Menard Mbende for all his help and his advices in the field, for his wonderful capability to resolve each problem, for all his stories about Congo and for his humour in each situation. Je dois aussi remercier Daniel Malalu, qui m’a conduit un peu partout sur le terrain, à répétitions de « sois calme » à chaque pente où j’avais peur que les freins de la voiture lâchent et en me mettant en tête ce refrain qui ne m’a toujours pas quitté (« Mon père avait raison, ma mère avait raison,… »). I also thank Jean Mobuli and Petra Lahaan, even if they arrived later in the field. Jean, tu as vraiment transformé cette base WWF en un endroit agréable à vivre et tu fais un travail formidable. Don’t give up ! Nous revenons bientôt avec un disque dur pleins de films pour occuper tes longues soirées d’hiver. Petra, thanks you so much for our collaboration, I really hope we will have new opportunities to meet each other. Je remercie également toutes les autres personnes du WWF-Congo pour leur aide dans la préparation de chaque mission de terrain, et tout particulièrement Raymond Luebuenamo qui m’a ouvert les portes de Malebo. And besides all the support of WWF-Congo, I warmly thank people from WWF-Belgique, who helped me numerous time! Je remercie également Mbou-Mon-Tour de m’avoir accueilli dans leur ferme, Paulin pour ton aide quotidienne, Jean-Christophe pour le travail formidable que vous accomplissez. Je dois aussi dire un tout grand merci aux chefs de terre de Nkala et Mpelu, merci de m’avoir accueilli dans vos villages, de m’avoir permis de travailler dans vos forêts, de m’avoir fait partager avec vous et avec les autres villageois de si bons moments, de si bonnes fêtes du 30 juin. Nazali kobosana bino te, nakozonga mpembeni ! iv September 2010 Mais l’ensemble de ce travail n’aurait vraiment pas été possible sans l’aide de tous mes assistants de terrain, Yaya Cicé, Petit Mowa, Mpesse, Mafa, Bula, Pala, Papa Fido, et tous ceux qui nous ont aidé à couper les transects chaque année. Cicéron, tu as vraiment accompli un travail formidable et je t’en suis énormément reconnaissante. Sans toi, cette thèse n’aurait pas été possible. Tu as su gérer nos équipes en mon absence, apprendre à chacun le travail avec une patience et un savoir-faire impressionnant. Travailler avec toi était une expérience magnifique, tu t’es investi, tu t’es intéressé à ce que nous faisions et tu m’as appris tellement de choses. J’espère que tu garderas cette curiosité et cette soif d’apprendre et qu’elles te permettront d’accomplir cette nouvelle aventure qu’est l’université. Je te souhaite énormément de courage, je suis toujours là si tu as besoin de moi et j’attends avec impatience de retravailler avec toi. Mowa, j’ai passé des moments formidables avec toi en forêt, merci pour ton dévouement dans notre travail. Je te souhaite pleins de succès à l’école et j’espère que nous pourrons retravailler ensemble bientôt. Je remercie également les pisteurs bonobos, qui m’ont aidé à localiser les sites de nids, à collecter les fèces, et qui ont partagé avec moi leurs connaissances des bonobos de la région.