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Marion N. C. Laporte Phd Thesis PANT-GRUNTS IN WILD CHIMPANZEES (PAN TROGLODYTES SCHWEINFURTHII): THE VOCAL DEVELOPMENT OF A SOCIAL SIGNAL Marion N. C. Laporte A Thesis Submitted for the Degree of PhD at the University of St Andrews 2011 Full metadata for this item is available in Research@StAndrews:FullText at: http://research-repository.st-andrews.ac.uk/ Please use this identifier to cite or link to this item: http://hdl.handle.net/10023/1973 This item is protected by original copyright This item is licensed under a Creative Commons License Pant-grunts in wild chimpanzees (Pan troglodytes schweinfurthii ): the vocal development of a social signal Marion N. C. Laporte Thesis submitted to the School of Psychology of the University of St. Andrews for the degree of Doctor of Philosophy August 2010 Candidate’s declarations: I, Marion Laporte, hereby certify that this thesis, which is approximately 45,000 words in length, has been written by me, that it is the record of work carried out by me and that it has not been submitted in any previous application for a higher degree. I was admitted as a research student in September, 2006 and as a candidate for the degree of Doctor of Philosophy in June 2007; the higher study for which this is a record was carried out in the University of St Andrews between 2006 and 2010. Date ……………………………. Signature of candidate ……………………………. Supervisor’s declaration: I hereby certify that the candidate has fulfilled the conditions of the Resolution and Regulations appropriate for the degree of Doctor of Philosophy in the University of St Andrews and that the candidate is qualified to submit this thesis in application for that degree. Date ……………………………. Signature of supervisor ……………………………. Permission for electronic publication: In submitting this thesis to the University of St Andrews I understand that I am giving permission for it to be made available for use in accordance with the regulations of the University Library for the time being in force, subject to any copyright vested in the work not being affected thereby. I also understand that the title and the abstract will be published, and that a copy of the work may be made and supplied to any bona fide library or research worker, that my thesis will be electronically accessible for personal or research use unless exempt by award of an embargo as requested below, and that the library has the right to migrate my thesis into new electronic forms as required to ensure continued access to the thesis. I have obtained any third-party copyright permissions that may be required in order to allow such access and migration, or have requested the appropriate embargo below. The following is an agreed request by candidate and supervisor regarding the electronic publication of this thesis: Access to all of printed copy but embargo of all electronic publication of thesis for a period of 2 years on the following ground: publication would preclude future publication. Date………… Signature of candidate……………… Signature of supervisor ……………… i Acknowledgements A great number of people have contributed to this thesis, all in their own ways. I would like to thank them all for having helped me through this “rite de passage”. First and foremost, I would like to thank my supervisor Pr. Klaus Zuberbühler for giving me the chance to work with his team, both in St Andrews and in Budongo. His constant support, guidance and encouragements at every step of the realisation of this thesis have been extremely valuable. I am grateful to the EU FP6 Pathfinder initiative “what it means to be human” for providing the core funding of this work and the Lucie Burgers Foundation for Comparative Behaviour Research, Arnhem, The Netherlands for additional support. For allowing me to conduct research in Uganda, I must thank the president’s office, the Uganda Wildlife Authority (UWA) and the Ugandan National Council for Science and Technology. I must also thank Dr. Fred Babweteera for opening the Budongo doors and for his wisdom and integrity. In St Andrews, thanks to Dick Byrne, Juan-Carlos Gómez, and Andy Whiten for the discussions, to Tecumseh Fitch and Vincent Janik for their support on acoustic analyses and their patience with my questions and Mike Oram for the statistical advice. I would also like to thank all the people of the BDG and the acoustic and cognitive journal clubs to keep these initiatives alive and lively. Brian, Jackie Ken and Helen are also to be thanked for their constant support whenever I encountered problems with my computer or my university records. Without other St Andrews PhD students, the life in St Andrews would not have been as wonderful as it was. My thanks are especially going to our ladies’ team including Anne, Erica, Simone, Emilie, Cris, Cat and Zanna who were like a second family and Simon, Verena, Elainie, Thomas, Thibaud, Volker, Katie, the Claudia(s), the Valentina(s), Bess and Caroline for all the great moments spent together. Special thanks go to Kate and Anne for all the great night long discussions! Andrea and Amy, thank you for your dynamism, your talents in everything you are doing, your friendship and for being fantastic flatmates. In Budongo, asante mkubwa sana to Geresomu Muhumuza, for his legendary rigour and knowledge of the forest. For all the good moments spent in the forest and the help, I thank him and also Monday and his memorable kubwa cheka, James, Stephen, Jackson, and Sam. Many thanks to Marie Plasse, Verena Kersken and Rebecca Schnittger for their help with the ii data collection and the great discussions. The life at camp would not have been so nice without Zephyr, Mary and Sarah and the Nyakafunjo women who are truly wonderful. Richard, a bit of Budongo’s heart has gone with you and a year later I cannot still believe you really left us. On a happier note, thanks to all Budongo students, it was just wonderful to share this experience with you, especially the A’s and Z’s: Anne, Agnès, Anna, Zinta, Zarin and Zanna. Thanks also to the other letters, Roman and Cathy, Cat and Tanja. A special thank to Daryl, my Masindi’s angel. Finally a soft grunt must go to the Sonso chimps, it was truly wonderful they let us spend so much time running around the forest and observing them. En France, un grand merci a tous les amis qui ne comprenaient pas toujours ce que je faisais ni pourquoi je le faisais mais qui ont été d’un immense soutien tout au long de ces années. Merci en tout premier a Anahita pour “tout” depuis plus de 10 ans. Merci à Élodie pour sa sagesse et ses cours de yoga et à Georgette Elgey pour son soutien et pour les chaussures indispensables à la forêt equatoriale. Merci aussi à Géraldine, Raphaëlle, Mathilde et Isabelle, Charlotte et Thomas, Gat et Flo, Till et Julie, Hélène et Daniel, Julie et Selma. Un immense merci à Danièle Courteil pour avoir survécu a la correction d’une thèse sur un sujet qu’elle ne connaissait pas. Fabien, tu as rendu cette thèse plus belle et plus bouleversante que je ne le pensais, le reste serait trop long à écrire et j’espère que nous écrirons encore longtemps ensemble. The last but not least, ma famille chérie, Marie-Françoise, Gérard et Jade, merci pour votre enthousiasme constant, votre foi inébranlable en moi et par tous les temps, votre amour et vos discussions sans fin. J’ai adoré vous faire découvrir ce pays que j’aime tant et partager ces moments privilégiés avec vous au milieu des chimpanzés. Jamais cette thèse n’aurait pu être réalisée sans avoir grandi au milieu de ton émerveillement au monde, maman et de tes questionnements philosophiques papa. Cette thèse vous est dédiée. iii Contents General Abstract 1 Chapter one: General introduction 2 Introduction 3 Animal communication systems in the light of human language 4 Human language and evolution 4 Studying natural vocalisations in animals and the problem of meaning 6 “Functionally referential” communication 8 Intentionality and flexibility in vocal communication 11 Audience effects 11 Vocal flexibility in adult animals 12 Social learning of vocalisations 14 Chapter two: Chimpanzee social structure and vocalisations 21 Social structure 22 Communication 23 The chimpanzee pant-grunt 24 Acoustics 24 The social function of pant-grunts 25 Development 28 Babies 29 Infants 30 Juveniles 31 Subadults 32 Maternal influence on offspring sociability 32 Thesis outline 34 Chapter three: General Methods 35 Study Site 36 Fauna 38 The Sonso community 38 Data collection 42 Sampling method 43 Behavioural and contextual description of vocalisations 46 Vocal data and Acoustic analysis 46 Statistical analyses 53 iv Chapter four: Vocal greeting behaviour in wild chimpanzee females 54 Introduction 56 Methods 59 Study Site 59 Data collection 59 Dominance hierarchy 60 Statistical analyses 61 Results 62 General patterns 62 Effects of target individual identity 63 Effects of target individual behaviour 64 Audience effects: alpha male and female 65 Audience effects: group size 67 Discussion 69 Chapter five: Acoustic structure of chimpanzee pant-grunts co-varies with the receiver’s social class and intended type of interaction 73 Introduction 75 Methods 78 Study Site 78 Data collection 78 Selection of pant-grunts 78 Acoustic analysis 79 Statistical analysis 81 Results 81 Individual identity 81 Receiver’s status 82 Context of emission 87 Discussion 92 Chapter six: The development of a greeting signal in wild chimpanzees 96 Introduction 98 Methods 101 Study site 101 Data collection 102 Vocalisations 104 Context analysis 104 Response to the type of vocalisations and the identity of the emitter 105 Response to
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