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The Acoustic Structure and Ontogeny of Vervet Monkey Vocalizations

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The Acoustic Structure and Ontogeny of Vervet Monkey Vocalizations University of Calgary PRISM: University of Calgary's Digital Repository Graduate Studies The Vault: Electronic Theses and Dissertations 2019-11 The Acoustic Structure and Ontogeny of Vervet Monkey Vocalizations Dubreuil, Colin Dubreuil, C. (2019). The Acoustic Structure and Ontogeny of Vervet Monkey Vocalizations (Unpublished doctoral thesis). University of Calgary, Calgary, AB. http://hdl.handle.net/1880/111238 doctoral thesis University of Calgary graduate students retain copyright ownership and moral rights for their thesis. You may use this material in any way that is permitted by the Copyright Act or through licensing that has been assigned to the document. For uses that are not allowable under copyright legislation or licensing, you are required to seek permission. Downloaded from PRISM: https://prism.ucalgary.ca UNIVERSITY OF CALGARY The Acoustic Structure and Ontogeny of Vervet Monkey Vocalizations by Colin James Dubreuil A THESIS SUBMITTED TO THE FACULTY OF GRADUATE STUDIES IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY GRADUATE PROGRAM IN ANTHROPOLOGY CALGARY, ALBERTA NOVEMBER, 2019 © Colin James Dubreuil 2019 Abstract Vervet monkey (Chlorocebus pygerythrus) vocalizations have been the subject of considerable research, most notably for their putative language-like qualities. While this focus has inspired a productive research effort investigating vocal communication in non-human primates, it has diverted attention away from other, non-linguistically inspired mechanisms by which vocal signals exert their effects on receivers. My research focuses on two vocal classes, grunts and alarm calls, and how their acoustic structures vary according to sender-specific attributes, including age, sex, body size, and identity - all of which have the potential to influence receiver response. I recorded calls from three wild groups of vervet monkeys over a 7-month period on the Samara Game Reserve, South Africa. I used random forest models to determine whether grunts varied in structure in relation to caller age, sex, identity and context, and whether alarms varied with sex and caller identity. I performed a cluster analysis to determine whether alarms segregated into different call types based on variation in acoustic structure. Finally, I used a series of mixed effects models to determine whether call structure in males and females correlated with overall body size (using body weight as a proxy). I found grunts varied in structure with age and sex, but not among individuals or contexts. Alarm calls varied with both sex and identity. My cluster analysis identified two qualitatively distinct alarm call types corresponding to the calls of males and females respectively. My analysis demonstrated that the relationship between body weight and the distribution of acoustic energy throughout alarm calls differed between the sexes. Relative to females, acoustic energy in the calls of males was concentrated at disproportionately lower frequencies. These results suggest that alarm calls may be under selection to exaggerate caller body size and/or extend the effective range of these signals in males. My results highlight that vocalizations can vary in structure along a number of dimensions simultaneously. While it is possible that in some instances, vocalizations convey specific information surrounding their context of production, it is likely that the mechanisms by which vocal signals exert their effects on receivers are more numerous and diverse. ii Acknowledgements Firstly, I would like to thank my supervisory team: Dr. Hugh Notman, Dr. Mary Pavelka, and Dr. Peter Henzi and Dr. Louise Barrett for their support, direction, and mentorship. I’d like to thank Dr. Kurt Hammerschmidt, and Dr. Julia Fischer for their hospitality and help with my acoustic analysis. Special thanks to Dr. Tyler Bonnell for his help and guidance with my statistical analysis, and his endless patience. Thanks to Dr. Jonathan Jarrett and Dr. Richard McFarland for their help with the weight data. I would also like to thank the field team: Dr. Christopher Young, Miri Minkner, Rose Blersch, Christina Nord, Stephen Morris, Lucy Twitcher, Tatiana Bortolato, and Pia Marlena for their help, patience, and perseverance in the field. None of this would have been possible without you. Special thanks to Kitty and Richard Viljoen for making the field feel like home. Financial support for this research was provided by the Natural Sciences and Engineering Research Council of Canada, the Killam Trust, the Faculty of Graduate Studies, and the Department of Anthropology at the University of Calgary. Special thanks to Mom and Dad, for constantly being there for me. Thanks to Alex, Veronica, Lily and Theodore for all the smiles over the last 5 years. Finally, I’d like to thank Lucy Millington for being there through this entire process, whether in person or through skype. Your support and patience have meant a lot to me. iii Contents Chapter 1 Introduction ................................................................................................................................. 1 1.1 Theoretical Framework ....................................................................................................................... 1 1.2 Research Goals, and Layout of this Thesis .......................................................................................... 4 1.3 Literature Review ................................................................................................................................ 7 Source Filter Theory and the Relationship Between Body Size and Acoustic Structure ...................... 7 Variation in Relation to Body Size ......................................................................................................... 9 Variation Related to Age and Sex........................................................................................................ 10 Variation Related to Individual Identity .............................................................................................. 14 Variation related to Motivation .......................................................................................................... 18 Variation related to Context ............................................................................................................... 20 Information and Influence .................................................................................................................. 24 1.4 Brief outline of subsequent chapters ............................................................................................... 27 Chapter 2 Sources of acoustic variation in the grunt vocalizations of wild vervet monkeys ..................... 28 2.1 Introduction ...................................................................................................................................... 28 Vervet Grunts ...................................................................................................................................... 30 2.2 Methods ............................................................................................................................................ 34 Study Species ...................................................................................................................................... 34 Study Site and Population ................................................................................................................... 36 Vocal data collection in the field ......................................................................................................... 37 2.3 Data Analysis ..................................................................................................................................... 39 Assigning Contexts to calls .................................................................................................................. 39 Selecting calls for analysis ................................................................................................................... 42 Acoustic analysis ................................................................................................................................. 44 Statistical Analysis ............................................................................................................................... 49 2.4 Results ............................................................................................................................................... 51 Classification of caller sex ................................................................................................................... 51 Classification by age class ................................................................................................................... 54 Classification of individuals using their grunt vocalizations ............................................................... 58 Effects of context on grunt variation .................................................................................................. 59 2.5 Discussion .......................................................................................................................................... 70 Age and Sex ......................................................................................................................................... 70 iv Individual Identity ..............................................................................................................................
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