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Individual Variation in Cooperative Behaviour in Meerkats Individual variation in cooperative behaviour in meerkats Sinéad English A dissertation submitted to the University of Cambridge in application for the degree of Doctor of Philosophy Clare College October 2009 i Preface This dissertation is my own work and contains nothing which is the outcome of work done in collaboration with others, except as specified in the text and acknowledgements. The total length of the text does not exceed 60,000 words, including the bibliography and appendices. No part of this dissertation has been submitted to any other university in application for a higher degree. Sinéad English 9 October 2009 ii Summary Individual variation in cooperation is a striking yet poorly understood feature of many animal societies, particularly in cooperative breeders where individuals assist in the care of young that are not their own. While previous research on these systems has emphasised the plasticity of helping and how it varies with current environmental and social conditions, in this dissertation I examine how individual variation is constrained and influenced by trade-offs with other behaviours and experiences in early life. I demonstrate that variation in cooperative pup care (babysitting and provisioning) is consistent within individuals over time (Chapter 3). Provisioning is more consistent than babysitting, although the two behaviours are highly correlated within individuals. I then focus on the variation in helping that remains once current factors, such as condition, group size and food availability, are taken into account. In Chapter 4, I explore the possibility that variation in helping can be explained by personality, or consistency in behavioural traits such as exploration or risk-taking. I find little evidence for consistent individual differences in field measures of personality traits, however, with such behaviours instead being group-specific. Early social experiences are known to have important and lasting effects on later fitness and behaviour: in Chapter 5, I demonstrate that, in female meerkats only, growing up in a group with more helpers is correlated with reduced cooperation later in life. This result suggests the importance of future fitness in influencing current cooperative behaviour, as females raised in larger groups are more likely to attain dominance. Finally, I examine the extent to which vocal communication between carers and young is influenced by variation in contributions to cooperation. Females are more sensitive to increased begging rate (Chapter 6), which reflects general sex differences in cooperative behaviour. Carers modify their vocalizations but not their foraging behaviour in the presence of pups, and the way in which they vocalize during provisioning events suggests these calls serve to increase efficiency of prey transfer (Chapter 7). iii Acknowledgements I am most grateful Tim Clutton-Brock and Joah Madden for supervising this thesis. My interest in Behavioural Ecology and field research is largely due to Tim and the opportunity he gave me to work in the Kalahari as an undergraduate. I thank him for his encouragement to embark on this PhD, providing access to the habituated population of meerkats and the long-term database, and his extensive support and advice both when initially formulating my ideas and over the course of writing up this dissertation. I am very grateful to Joah for his encouragement and advice on formulating hypotheses, executing experiments and preparing manuscripts. In the Kalahari, I was lucky to have two excellent field assistants. Biz Turnell, as well as being a wonderful help and calming influence, was inspirational in her determination and conscientiousness. Finnie (Mike) brought fun to fieldwork, helped with ideas and acted as a firm negotiator with film crews. As well as direct assistance from Biz and Finnie, my experiences of fieldwork all ran smoothly with managerial support from Tom Flower, Rob Sutcliffe and Dave Bell. Most of the work in this thesis would not have been possible without the hard work of the many volunteers who have collected data for the Meerkat Project over the past ten years. I was fortunate to overlap with Marta Manser during my trips to the Kalahari, who was always very helpful and encouraging in discussing ideas for experiments and generous in giving me access to her bank of meerkat vocal recordings for use in playback experiments. During my first field season, I benefited from working alongside both Joah and Hansjoerg Kunc, who both gave me the opportunity to participate in a series of playback experiments, and continued to involve me in subsequent publications from this period. My research on carer-offspring communication also benefited from the hard work of Lizzie Rowe, a Part II undergraduate student in Cambridge whose project I supervised from January to April 2009. Lizzie was an excellent student, hard-working and independent, who patiently transcribed hours of sound recordings without having the pleasure of seeing a real meerkat. I hope that she manages to go to the Meerkat Project one day. This dissertation has also greatly benefited from the assistance of Shinichi iv Nakagawa: starting with a polite email enquiring after R code, a fruitful collaboration has been born, and the analyses presented in the first data chapter would not have been possible without his guidance. I have been blessed with a team of self-appointed supervisors (and friends) in Cambridge and beyond over the past three years, without whom I would have floundered long ago. Alex Thornton might possibly be targeted as the culprit who set me on this course to begin with. As well as demonstrating by example what an amazing and fun PhD one could do on meerkats, he has helped me at every step of the way, in writing my application proposal to the final touches of this dissertation. His accomplice, Kelly Moyes, is almost as much to blame. Kelly helped me enormously not only in moving my stuff around, keeping my stress levels down (with occasional headaches as a side-effect) and taking time to help me deal with the quagmire of R. Also leading by example, Nikki Raihani has been an inspiration in her pragmatic approach both to science and life, as well as being a wonderful source of academic support and fun adventures, including encounters with jellyfish in France and bear-squirrels in Canada. Stu Sharp and Matt Bell have both also been important scientific advisors and life coaches. Stu has had the misfortune of sharing an office with me for the last two months of my PhD but his Liverpudlian sense of humour has weathered the experience considerably well, and has kept me mostly sane. Matt Bell has been a great source of encouragement and advice through all stages of this PhD, from tantalizing me with the wonders of banded mongooses to advising me on ideas, writing skills and the fineries of cheese and wine both in Cambridge and the Kalahari. Nikki, Stu, Matt and Alex in particular have been exceptionally helpful about reading through chapters and going far beyond the call of duty as friends or colleagues. As well as my academic support team, I have received moral support from the extended family of LARG. Julian Drewe has been my trail-blazer, not only on half-marathon courses but in following his steps during the writing up process. I thank Bonnie Metherell for trying to inject some kind of organisation into my scatty life, providing me with sweets and cake and being a great fellow Arts Picturehouse nerd. Raff · Mares, needless v to say, completely transformed my PhD experience, and gave me a sense of perspective both during fieldwork and back in Cambridge. Other fellow PhD students that have given me help or advice along the way include: Neil Jordan, Krys Golabek, Andrew Bateman (and Dr. Love), Lucy Browning, Martha (Maple) Nelson-Flower, Caroline Dingle and Jenny Oates. I have also received considerable help and guidance from many of the friendly and helpful post-docs in Cambridge, and beyond: I thank Dieter Lukas, Sarah Hodge, Kavita Isavaran, Andy Young, Camilla Hinde, Dan Nussey, Andy Radford and Simon Townsend. Particular gratitude is owed to Linda Wheatley and Penny Roth. Both Linda and Penny have always been extremely patient and approachable with my frequent, nagging requests. I am grateful to Nick Davies, Mike Cant and Becky Kilner for their advice on my yearly reports and insightful suggestions on my research, which I hope I managed to incorporate into this dissertation. Finally, I thank my parents and sisters for their patience with my long periods of absence, or, when present, my extreme levels of stress and putting up with constant anecdotes from the Kalahari. My mother was a pillar of emotional support, especially during the more trying times of my PhD. In particular, I am extremely grateful to my father for inspiring in me a motivation to work hard and enjoy the outdoors, as well as offering his professional services as an editor to proof-read this entire dissertation. This research was supported financially by the Natural Environment Research Council and the generosity of Clare College, for which I am most grateful. vi Table of contents Preface i Summary ii Acknowledgements iii Table of Contents vi Chapter One Introduction 1 1.1 Individual variation in cooperation 2 1.2 Quantifying individual variation: plasticity and consistency 6 1.2.1 Consistency across time 6 1.2.2 Personality types and behavioural syndromes 8 1.3 Environmental influences on development 10 1.4 Carer-offspring communication in a cooperative species 12 1.5
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