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Understanding conservation conflicts surrounding predation and game shooting interests Submitted by George Julius Fraser Swan to the University of Exeter as a thesis for the degree of Doctor of Philosophy in Biological Sciences, November 2017 This thesis is available for Library use on the understanding that it is copyright material and that no quotation from the thesis may be published without proper acknowledgement. I certify that all material in this thesis which is not my own work has been identified and that no material has previously been submitted and approved for the award of a degree by this or any other University. (Signature) …………………………… Hawk Roosting “I sit in the top of the wood, my eyes closed. Inaction, no falsifying dream Between my hooked head and hooked feet: Or in sleep rehearse perfect kills and eat. The convenience of the high trees! The air's buoyancy and the sun's ray Are of advantage to me; And the earth's face upward for my inspection. My feet are locked upon the rough bark. It took the whole of Creation To produce my foot, my each feather: Now I hold Creation in my foot Or fly up, and revolve it all slowly - I kill where I please because it is all mine. There is no sophistry in my body: My manners are tearing off heads - The allotment of death. For the one path of my flight is direct Through the bones of the living. No arguments assert my right: The sun is behind me. Nothing has changed since I began. My eye has permitted no change. I am going to keep things like this.” Ted Hughes, 1960 2 Abstract Many predatory species cause negative impacts on human interests by threatening game, livestock or human safety. These impacts can create conflicts where stakeholders differ over wildlife management and when one party is perceived to exert their interests at the expense of the other. Finding effective methods to mitigate conservation conflicts requires an interdisciplinary perspective that investigates (i) the reality of the apparent impacts, (ii) the efficacy of any methods intended to remedy them and (iii) the perceptions, motivations and objectives of key stakeholders. In this thesis, I investigated a conservation conflict in the U.K. surrounding predators and game management. I did so with specific reference to the common buzzard Buteo buteo, a species that, due to predation of released pheasants Phasianus colchicus, is both subject to illegal persecution and on- going controversy concerning the licenced selective removal of ‘problem individuals’. I first review the literature to assess the ecological evidence that certain ‘problem individuals’ can be both disproportionately responsible in impacts upon human interests and more likely to reoffend. I show that while there is evidence for these animals across many different taxa, the benefits of their removal can sometimes be short-lived. I highlight possible indirect impacts of selective management and identify it as a potential compromise between different stakeholder groups. Next, I evaluate the performance of Bayesian stable isotope mixing models (BSIMMs) in quantifying the diets of wild animals. By comparing indirect and direct observations of buzzard foraging, I demonstrate that, with the correct selection of trophic discrimination factors, stable isotope analyses can provide a reliable picture of dietary composition that mirrors direct observations. I then apply these mixing models to evaluate the ecological basis of selective removal of ‘problem buzzards’. The results suggest that the consumption by buzzards of released pheasants is not limited to release pens where 3 gamekeepers perceive buzzard predation to be a problem. However, I then show that stable isotope analysis of blood sampled from two of the four buzzards caught inside pens indicates frequent consumption of released pheasants, relative to the rest of the buzzard population. These results suggest that, while some pheasant consumption may go undetected, selecting only buzzards inside pens for removal is likely to target ‘problem birds’. I then investigate buzzard foraging and breeding ecology on land managed for pheasant shooting. I find that buzzards nest at higher density in areas with greater abundances of pheasants and rabbits Oryctolagus cuniculus. However, records of provisioning from nest cameras showed that only rabbits were caught in proportion to their abundance and only rabbit provisioning rate was associated with buzzard productivity. I suggest that the positive relationship between buzzard and pheasant abundance, although seemingly unconnected to pheasant predation, might influence how gamekeepers perceive buzzard impact. Next, I conduct semi-structured interviews on the subject of predator control with 20 gamekeepers across the south of England, to explore the underlying beliefs, norms and information sources that motivate their behaviour. From these interviews, I identify a number of separate, but interconnected, motivations that influence predator control including professional norms, potential penalties, and interpretations of what is ‘natural’. The influences of these motivations are discussed in detail and a conceptual model, incorporating the theory of planned behaviour, is developed. Finally, the key contributions of this thesis are drawn together and discussed in their wider context. Taken together, the results of this thesis illustrate how predator management occurs simultaneously within social and ecological contexts that incorporate the individual attributes of both predators and people. The results of this thesis have direct implications for the management of predators, the representation of stakeholder perspectives and the design of conflict mitigation measures. 4 Acknowledgements It’s a strange thing to be writing my thesis acknowledgments and thinking back over the last four years to all the people who have played a role in getting me to this point. Suffice to say that the list is long and whatever I write over the next few pages will not do it justice… First and foremost, this thesis would not have been possible without the support, advice and encouragement that I have received from my three supervisors. I am particularly grateful to Robbie McDonald, not only for the day-to-day guidance he provided, but also for the opportunity to come down to the University of Exeter and work on a subject that I am (still) totally fascinated by. I owe a great debt to Steve Redpath who got me hooked on studying conflicts and very kindly agreed to be my external supervisor, bringing indispensable knowledge and experience to this study. Lastly, my thanks go to Stuart Bearhop for providing big-picture perspective and making time for me when I needed it. It is a real honour to have worked with all three of you. Next, I would like to express my sincere gratitude to the landowners and tenants who granted me access to their estates. In particular I would like to thank the gamekeepers without whose help this thesis would not have been possible. This includes those individuals that gave up their time to share their experiences of predator management. The study sites used in my fieldwork and visited for the interviews represent some of the most beautiful countryside England has to offer and it was a real privilege to have been able to share it with them. Over the course of my fieldwork I received a considerable amount of support from various sources. David and Katie Anderson (Forestry Commission Scotland) provided all the training for handling and ringing raptors. Amy Campbell and Tom David (Naturally Green Ltd.) provided assistance with the tree climbing and safety support. Mike Nash (HandKam Ltd.) was a total wizard with the nest cameras. James Yule provided in-house technical support on 5 many different occasions. Dave Cruse took control of all the battery charging and made a very stressful time that little bit easier. My last four years in Cornwall have been an amazing time, made extra special by the people. This PhD has, at times, been tough, but it has been the support and friendship from my colleagues that has kept it enjoyable. The last few days of ‘end-game’ PhD illustrate this well with pizza, chilli, soup and beer delivered to my desk in displays of kindness I’m sure I don’t deserve. In particular I would like to thank my two partners in crime Cecily Goodwin and Sarah Crowley who have both provided immense practical and emotional support and shown saintly patience over the course of our studies. Others in the brilliant-friends- that-I-also-ocassionaly-bug-for-academic-advice category are Matt Silk, Rebecca Short, David Fisher, Sean Meaden, Cat McNicol, Rich Inger, Katie Shanks, Sarah Paul, James Duffy, Jared Wilson-Aggarwal, Helen Fielding and Daniel Padfield. You guys have been totally amazing. I also owe a huge debt of gratitude to my parents who have always encouraged me to pursue my interest in natural history and taught me to keep pushing myself. Finally, I would like to thank my amazing partner Paz Duran for her companionship and support on my PhD journey. 6 Contents CHAPTER 1: INTRODUCTION ........................................................................................ 16 People and predators ............................................................................................ 16 Predators, gamebirds and gamekeepers .............................................................. 21 The common buzzard ........................................................................................... 24 Thesis outline and aims .......................................................................................

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