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Post-Fledgling Begging and Development in Southern Pied Babblers

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Post-Fledgling Begging and Development in Southern Pied Babblers Post-fledging begging and development in Southern Pied Babblers (Turdoides bicolor) Alex Thompson Thesis Presented for the Degree of DOCTOR OF PHILOSOPHY At the Percy FitzPatrick Institute of African Ornithology, Department of Biological Sciences UNIVERSITY OF CAPE TOWN May 2013 Supervised by Professor Phil Hockey (deceased Jan 2013) Co-supervised by Dr. Nichola Raihani, Associate Professor Peter Ryan (from Feb 2013) The copyright of this thesis vests in the author. No quotation from it or information derived from it is to be published without full acknowledgementTown of the source. The thesis is to be used for private study or non- commercial research purposes only. Cape Published by the University ofof Cape Town (UCT) in terms of the non-exclusive license granted to UCT by the author. University Abstract Abstract Elaborate solicitation displays are a common feature of care-giver-offspring interactions throughout the animal kingdom. These displays, termed begging, are seen to be the physical manifestation of parent-offspring conflict over provisioning. The environment that offspring find themselves in is likely to have consequences for the costs and benefits of begging behaviour, and thus the dynamics of begging. However, the overwhelming majority of empirical and theoretical studies have used nestling birds as their model system. Nestlings are stationary, in close proximity to their siblings, and broods tend to be provisioned by one adult at a time. Therefore, current understanding of begging dynamics has potentially been narrowed by studies of stationary offspring. There is however, a growing literature on the begging of offspring that are mobile, able to move between care-givers and provide constant begging signals. In this thesis I use fledgling Southern Pied Babblers (Turdoides bicolor) as a model system in which to investigate mobile offspring begging and the developmental consequences of post-fledging care. Firstly, I give a general background to the field and address the questions that can still be asked in Chapter 1. I then discuss the observational, experimental and analytical methods used throughout the study in Chapter 2. In Chapter 3, I investigate the structure of fledgling pied babbler begging calls. I show that like other mobile offspring, pied babblers have multiple begging call types that are used in different contexts. I also show that the structure and use of begging calls changes with hunger level, information that may be used by care-givers in provisioning decisions. Using a series of feeding and playback experiments, in Chapter 4, I explore the begging dynamics between fledgling pied babblers and care-givers. I show that fledgling begging is positively related to hunger, adult provisioning is positively related to begging and that fledgling begging is independent of the begging behaviour of siblings. I propose that the dispersed nature of the fledgling environment means that fledglings gain no provisioning benefits nor incur any provisioning costs from changes in their siblings begging. In Chapter 5, I use a novel approach to investigate Zahavi’s blackmail hypothesis. I show that young fledglings face a trade-off: staying in the safety of trees or moving to the profitable yet risky ground to solicit care. Adults change their provisioning behaviour based on the risk that fledglings face, related to their location, and fledglings are thus able to gain more food by moving to riskier locations. In Chapter 6, I consider the long term consequences of fledgling behaviour during i Abstract dependence. Using natural observations and feeding experiments I disentangle the effects of initial advantages, investment in foraging and investment in offspring-adult associations on foraging development. I show that individuals who invest more in behaviours to promote individual-learning, foraging when young, develop relatively higher foraging efficiencies than their siblings. Finally in Chapter 7, I synthesize the finding of my thesis and relate these to current understanding in the field. In addition, I propose future work that could build on the research conducted in this study. The mobility of fledgling Southern Pied Babblers has led to the evolution of a care-giver-offspring interactions where offspring solicitation appears to signalling reliable signal of hunger, with the information contained within the structure of begging calls and fledgling’s response to each other’s begging revealing that begging is a way for individuals to signal their hunger. However, fledglings can still manipulate adults into provisioning them by threatening their own destruction. ii Acknowledgments Acknowledgments This research was funded by the NRF Centre of Excellence at the Percy FitzPatrick Institute of African Ornithology and the British Ecological Society, under ethical clearance from the Science Faculty Animal Ethics Board, University of Cape Town (R2012/2006/V15/AR). Firstly, I would like to thank my family. My parents always told me to pursue a career that I enjoyed and only put pressure on me to excel at whatever I chose to undertake. Without their sacrifices, support and encouragement I would not be doing what I am doing today. They have given me so much and there is no way I will ever be able to repay them in full or thank them enough. Thanks also to my brother, Morgan, for always bringing me back down to earth and also to his beautiful daughter Bella. Secondly, thanks go to my supervisors. Phil Hockey and Nikki Raihani helped to guide me in my planning, preparations and write up. They helped me to hone my writing skills, and to restrict the number of tangents that I am prone to meander onto. They managed to do this whilst both going through major upheavals in their lives, and for the efforts during those trying times I will be forever grateful. I also thank Peter Ryan for stepping in after Phil’s sudden death, you helped to steer me through the final choppy waters of UCT bureaucracy and streamline my writing at the very end . Phil enabled me to pursue my dream of doing a PhD and of doing it on the study system that I really wanted to do it on, the babblers. His support has helped to change my life and I will be eternally grateful. His constant support has allowed me to follow my ideas, whilst reining me in when I become tangential. He was the heartbeat of the Fitz, its guiding light and its energy. He will be forever missed, but his legacy will live on. Thanks to Tim Clutton-Brock and Marta Manser for allowing me to use the facilities at the Kuruman River Reserve and for their past support of the pied babbler project. Thanks also to Jamie Samson and Nate Thavarajah, for keeping the meerkat volunteers in check on the reserve and for the amusing radio chat. Joe Fenton and Melissa Delport helped out by keeping the ‘friends’ and Earthwatchers sufficiently occupied and out of our way at iii Acknowledgments Gannavlakte (GV). And finally, thanks to the meerkat volunteers for all of their radio calls that helped us find the babblers that little bit faster. It’s also important to thank the De Bruins and Kotzes; I am sure they have no idea how much science is done on their land and how many papers they have been acknowledged in. Sound analysis presented a horrible challenge during this PhD but I had help along the way from various people. Thanks to Simon Townsend for his patients while teaching me the fun world of pDFAs, I’ll miss all of our silly Oxbridge jibes. Dave Humphries helped me out a lot during my brief stint at Macquarie, you introduced me to the scary world of Raven and helped out loads during field work at GV. Roger Mundry went above and beyond the call of duty by providing help to a poor PhD student drowning in R code on the other side of the world, many thanks. Thanks to Matt Bell; who knew that a lowly 2nd year Natsci from Girton would still be asking for your help in formulating his scientific ideas 7 years later? And remember, Jesus rides a panda. I would also like to thank Amotz Zahavi for his comments on one of my chapters, interacting with him helped me to understand his thought processes more fully. I have to thank my three field assistants: Fiona Finch, Adam Britton and John Forecast. Fiona, I am sorry that lugging that ladder around to all of those nests, in the middle of the Kalahari summer, has resulted in almost no data finishing up in this thesis. Legs, you were part of a legendary GV football team and provided good banter whilst in the field. Time in the field wouldn’t have been the same without the myriad of characters that traversed the poorly built and maintained stage of GV. We developed a strange pride for our corner of the reserve, climaxing in the unbeaten GV football team, with such esteemed members as Mike Finnie, Tom Flower, Matt Gribble, Dave Humphries, Mark Whiteside and Adam Britton. The girls offered a tonic to the comic machismo of the boys at GV, Lizzie and Tanya were always ready with quick one liners and a cup of tea/coffee and a piece of cake. Thanks to Boo for the Insanity. The Nelson-Flowers/Flower-Nelsons need a big thanks as well. Tom and Maple have given me advice throughout and helped keep the life history on the babblers ticking, and it’s been fun to see Audry grow up in the Kalahari. iv Acknowledgments There are some people at the Fitz, without which this research would never have gotten off the ground. Hilary Buchanan, you are a magician, thanks for registering me every year and knowing the UCT system backwards and so allowing everything to seemingly flow smoothly.
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