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A Case Study Using the Tuatara-Fairy Prion Association

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A Case Study Using the Tuatara-Fairy Prion Association Interspecific Interactions: A Case Study using the Tuatara-Fairy Prion Association Ilse Corkery A thesis submitted to Victoria University of Wellington in fulfilment of the requirements for the degree of Doctor of Philosophy in Ecology and Biodiversity Victoria University of Wellington Te Whare Wānanga o te Ūpoko o te Ika a Māui 2012 All animals are equal but some animals are more equal than others. George Orwell Animal Farm. v Abstract Some of the key relationships in the life of an organism are interactions with individuals of other species within the community, for example, negative interactions such as predation and competition are well known to shape natural communities. Positive interactions also have well documented influences, such as intertidal seaweed canopies extending the distribution of many organisms to higher tidal heights, by reducing thermal and desiccation stresses. However, investigating interactions and measuring their significance for fitness is notoriously difficult. For example, several groups of fish are known to ‘clean’ other fish species by feeding on their ectoparasites, a mutually beneficial arrangement. However, foraging by cleaners can damage scales of their hosts and this interaction can become parasitic in times of low ectoparasite abundance. Using both field and laboratory data, I investigated factors that influenced the dynamics of an unusual vertebrate association, the cohabitation of tuatara and fairy prions in a burrow. The end goal was to contribute to the understanding of the classification of this association. The fairy prion is a seabird that comes to land only for the breeding season and the tuatara is a burrowing reptile, active primarily at night in a temperate climate. Specifically, I measured the effects that this association had on tuatara thermoregulation, and demonstrated the difficulty in applying that information to categorize a complex interaction. Investigations into the temporal and spatial habitat of the tuatara, and the degree to which this influenced thermal opportunities, revealed that mean tuatara body temperatures were always within mean environmental temperatures. Males and females did not differ in mean body temperature or effectiveness of thermoregulation. Body size did not predict body temperature or cooling rates, but heating rates were influenced, with larger animals heating faster than smaller individuals. The presence of a fairy prion in a burrow increased humidity within the burrow, and tuatara that occupied burrows containing a fairy prion were able to maintain up to 1.8°C higher body temperatures through the night during the austral summer months. Thus, burrow use behaviour and burrow selection had greater influences on tuatara body temperature than an individual’s sex or size. Experimental evidence revealed that tuatara are capable of adjusting their habitat selection behaviour in response to different humidity constraints. More time was spent outside the burrows and tuatara were more active under humid laboratory conditions. Use of the burrow by tuatara almost halved the time that fairy prions spent at the burrow with their chick, indicating that tuatara were having a negative effect on fairy prions’ use of their burrow. There v vi was no evidence to support the fact that fairy prions were gaining any fitness benefits from their association with tuatara. Thus, we cannot call this interaction a commensalism or a mutualism. In certain instances, it may be that this interaction is best classed as a parasitism with the tuatara benefitting from burrow use and easy predation opportunities, to the detriment of the lifetime reproductive success of the fairy prion. In other instances it may simply be a case of competition for a limited resource (a burrow) with the outcome varying depending on the individuals and the circumstances involved. Being able to categorize interactions between species of high conservation value or at least to have an understanding of the costs and benefits associated with the interaction is desirable for conservation purposes, as failure to consider the ecological network within which a threatened species is embedded, may lead to counterproductive management measures. Further, these results can be used to develop future research into how climatic changes in temperature and rainfall may interact with habitat availability to influence the full range of natural outcomes of the tuatara-fairy prion association. vi vii Acknowledgements I would like to express my deepest gratitude to my supervisors Nicky Nelson and Ben Bell. I am very grateful for their guidance and support throughout, and their encouragement was often what kept me going. Thank you for your enthusiasm and for your meticulous corrections of endless drafts! I am very thankful for the financial support of the following organizations: Victoria University Doctoral Research Scholarship, Faculty of Science at Victoria University, Centre for Biology and Restoration Ecology (CBRE), Allan Wilson Centre for Molecular Ecology and Evolution, Kathleen Stewart Award, Forest and Bird, Society for Research on Amphibians and Reptiles in New Zealand (SRARNZ), and New Zealand Ecological Society. Their contribution had a major part in the success of my numerous fieldtrips and laboratory work, and also allowed me to present my work at a number of international conferences. My work was also supported by Ngati Koata iwi with permission from DoC (permit NM-23620-RES and the VUW Animal Ethic Committee (permit #2006R12). A number of Department of Conservation staff were incredibly supportive of this project. They were always quick with permits, provided transport to Stephens Island and went above and beyond to provide assistance and support whenever and wherever I needed it. Thank you to Stuart Cockburn and Jim Hunnewell for assistance with building and acquiring my RFID loggers, scanners and batteries. Thanks to Peter Gaze, Mike Aviss and in particular to Karen Ismay and John McKewon, the rangers on Stephens Island who first introduced me to the wonders of that remarkable place, and to Margie Grant-Caplan and Tim Bacon who not only did everything to make my fieldtrips run as smoothly as possible, but also provided endless hours of fun. Fieldwork would also not have been possible without the help from my field assistants. Many thanks go to Jen Moore, Andy Douglas, Evan Bredeweg, Dawson Dunning, Heidy Kikillus, Monica Awasthy, John Ballinger and in particular Danielle Middleton. Danielle was my partner in crime throughout much of my fieldwork and many adventures and laughs were had. A big thank you goes to Holly Jones who first introduced me to the joys of working on offshore islands. I never forgot my first trip to Maud Island, having a rare bird the takehe, snatch a muesli vii viii wrapper from my hand and seeing my first giant weta, by far the largest insect I’d ever seen! I think I was hooked on New Zealand ecology after that trip! Thank you Monica Awasthy for giving me my first paid ecology job, introducing me to New Zealand birds and making me learn New Zealand trees! I am very thankful to the warm welcome that I received at SBS, which made it a great and relaxed place to study. My many office mates and friends were amazing, always ready for a laugh and ready to provide distraction when work was just a little bit too full on! Lastly, I would like to thank my partner John, my parents Mia and Frank, and my sister Maria. My love goes to my family, who have been incredibly understanding, and put up with my constant travels and decision to study and live on the other side of the world. I could not have made it through this PhD without the constant support and encouragement from both John and my family, to whom I am forever grateful. I would like to dedicate this thesis to my father Frank Corkery; I wish he could have seen me complete this and I miss him very much. viii ix Table of Contents Abstract ................................................................................................................................ v Acknowledgements ........................................................................................................... vii CHAPTER ONE. Interspecific interactions: A case study using the tuatara-fairy prion association ........................................................................................................................... 1 1.1 Introduction ............................................................................................................... 1 1.2 Categorizing interspecific interactions ..................................................................... 2 1.3 Retreat selection ........................................................................................................ 5 1.4 Physiological ecology ................................................................................................. 6 1.5 Tuatara and seabird ecology ..................................................................................... 7 1.6 Key questions, thesis outline and style .................................................................. 10 1.6.1 KEY QUESTIONS ............................................................................................. 11 1.6.2 THESIS OUTLINE AND STYLE .......................................................................... 11 1.7 References ....................................................................................................................... 13 CHAPTER TWO. Behaviour not morphology determines
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