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Factors Influencing Foraging Behaviour in the Australasian Gannet and for Seabirds in General, and Discusses Future Directions for Research

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Factors Influencing Foraging Behaviour in the Australasian Gannet and for Seabirds in General, and Discusses Future Directions for Research Factors influencing foraging behaviour in the Australasian gannet by Lauren Patricia Angel B.Sc (Marine Biology & Zoology) (Hons) Submitted in fulfilment of the requirements for the degree of Doctor of Philosophy Deakin University October, 2015 Abstract The foraging decisions made by an individual are reflected in their foraging success (i.e. prey capture), resulting in enough nutrition to support self-maintenance and chick growth. As such, foraging success is directly linked to reproductive success in breeding seabirds. However, only a small proportion of individuals contribute to future generations. Hence, foraging behaviour can greatly influence an individuals’ fitness and species demography. The Australasian gannet (Morus serrator) is a large pelagic seabird breeding in Australia and New Zealand. However, despite the accelerated warming of sea surface temperatures in south-eastern Australia, little is known about whether this species can buffer the effects of climate change. Furthermore, an understanding of natural variability in foraging behaviour is needed to estimate species’ adaptability. Two colonies, Point Danger (ca. 300 breeding pairs) and Pope’s Eye (ca. 180 breeding pairs), of opposing oceanographic regimes were studied to identify factors which may cause variation in foraging metrics, fine-scale behaviour, foraging effort and diet. These variables were obtained using GPS, tri-axial accelerometry and stable isotope analysis. The factors investigated as potential causes of variation were sex, age, colony size/habitat, year, breeding stage, inter- and intra-individual preferences. The present study found Australasian gannets are sexually dimorphic with a degree of sexual segregation in their foraging area use. Colony size/location influenced foraging trip distance and duration. The foraging ranges of the two colonies did not overlap indicating colony specific foraging ranges. Inter-annual variation was compared in years of differing prey availability at Pope’s Eye. However, birds appeared to be constrained, not increasing trip distance or duration in iii poor years. Rather, energy expenditure and the proportion of a foraging trip spent foraging increased. Furthermore, females were found to be more flexible in their foraging behaviour and strategies than males in both breeding stages. At Pope’s Eye males typically foraged closer to the colony with a high site fidelity. The sex difference in site fidelity may be due to females searching for high quality prey, reflecting the greater initial reproductive investment. The findings of the present study highlight potential drivers of variability in foraging behaviour. However, sexual segregation and colony-specific foraging ranges present potential limitations to foraging behaviour flexibility in the face of climate change. iv Preface The ethical guidelines of Deakin University Animal Ethics Committee and Animal Welfare Committee were followed during the present studies. The protocol was approved by Deakin University Animal Ethics Committee and Animal Welfare Committee (Permit No. A86/2010). The project was conducted in accordance with the regulations of the Department of Sustainability and Environment Victoria (Permit No. 0005745). Birds captured for the duration of the study were banded under ABBBS R-Class Banding Permit No. 3046. This project was supported financially from research grants provided by Holsworth Wildlife Research Endowment, Victorian Marine Science Consortium, Point Danger Management Committee and the Centre for Integrative Ecology, Deakin University. The core chapters of this thesis (Chapters 2-6) have been published or submitted for publication in peer-reviewed journals. I am the principal contributor to all chapters of this thesis and the primary author on all publications arising from this thesis. Associate Professor John Arnould is co-author on all publications for his contribution to the study design, fieldwork, and editorial advice throughout the project. Maud Berlincourt is co-author on Chapters 4-6 due to her assistance with initial analysis of GPS data. John Speakman is co-author on Chapter 2 and 3 for his contribution to processing blood samples. Emma Tew (Chapter 2 and 4), Melanie Wells (Chapter 2), Marlenne Rodríguez-Malagón (Chapter 2), and Sophie Barker (Chapter4) are co-authors for their contributions of additional data to the project. Tiphaine Jeanniard-du-Dot (Chapter 2) and Victoria Warwick Evans (Chapter 4) are co-authors for their contribution to the conception of data analysis methods used in these chapters. v Publications arising from thesis Chapter 2 Angel L.P., Wells M. R., Rodríguez-Malagón M.A., Tew E., Speakman J.R., Arnould J.P.Y (in review) Sexual size dimorphism and body condition in the Australasian gannet. PLoS ONE. Chapter 3 Angel L.P., Jeanniard-du-Dot, T., Speakman J.R., Arnould J.P.Y. (in review) Determining activity-specific energy expenditure in a large seabird using accelerometry. Royal Society Open Science. Chapter 4 Angel L.P., Berlincourt M., Barker S., Tew E., Warwick-Evans V., Arnould J.P.Y. (2015) Eating locally: Australasian gannets increase their effort in a restricted foraging range. Biology Open 4:1298-1305 (doi:10.1242/bio.013250). Chapter 5 Angel L.P., Berlincourt M., Arnould J.P.Y. (in review) Foraging ecology of Australasian gannets: The influence of oceanographic regimes. Marine Ecology Progress Series. Chapter 6 Angel L.P., Berlincourt M., Arnould J.P.Y. (in review) Inter- and intra-individual variation in foraging behaviour in Australasian gannets. Marine Biology. vi Acknowledgements First and foremost, I would like to thank my supervisor, Associate Professor John Arnould. Without your encouragement and support in all aspects of my candidature I would never have been able to see the light. Thank-you for the opportunity to work on such a graceful, yet aggressive species, it has truly been an amazing experience. The amount of field work this project required could not have been done without my field team. Thank-you for making the days waiting for gannets to come home enjoyable. In particular, thank-you Marion Fourquez and Emma Tew for the great times out in the brutal wind. I would like to thank my Point Danger family: Andy and Tilly Govanstone for the warm welcome into their home each field season, Ewen Lovell for his insight and guidance, and the Point Danger Management Committee who completely understand the beauty of gannets and the need to protect them. Thank-you also to my Pope’s Eye family: Robbie and Jenny Main for providing a home to stay in, Sea All Dolphin Swims for the trips out to the colony on weekends (and getting to swim with seals and dolphins each time), the Victorian Marine Science Consortium and Liz McGrath. In particular, I’d like to thank Rod Watson, you made every trip out to Pope’s Eye an absolute joy. I’ll see you at Napona! Thank-you to the Arnould lab group for making my time in the BA building enjoyable – in particular, Andrew Hoskins, Maud Berlincourt, Laëtitia Kernaléguen, Travis Knox, Emma Tew, Melanie Wells and Grace Sutton. Thank- you also to Evelyn Chia, Michelle Bassett, Kasun Ekanayake, John Patykowsi, and Carolyn Ewers for being instrumental in my procrastination attempts. vii Thank-you to my all my friends for putting up with me complaining about data analysis and stress about writing up. Andrew Hoskins and Holly Lourie, thank-you for providing me with almost daily updates on Benson and Nelson. Maud Berlincourt and Melanie Wells, you have truly ignited a spark of obsession in me with your passion for seabirds. To my ever loving family (dad, mum, Karl, Emma, Hannah, Lily, Mia, Kieran, Elise, Greg, and Elizabeth!), thank-you so much for coming to see me when I was too stressed to visit you. Things are always easier when you know family has your back. Thank-you to my meower, Basil, for being a constant source of cuddles and an anti-depressant to this experience. I met Tom at the very beginning. Sometimes you don’t know what you’re missing until it walks in. Who knew you were exactly what I needed to get through this stage of my life. “I’m a seagull. I like to fly” “Don’t you forget that the reason you fly is to eat” – Jonathan Livingstone Seagull viii Table of Contents Abstract ................................................................................................................... iii Preface .......................................................................................................................v Publications arising from thesis ............................................................................ vi Acknowledgements................................................................................................ vii General overview....................................................................................................13 Factors influencing the foraging behaviour of seabirds ......................................14 Changes in the environment and the Australasian gannet...................................16 Research aims and thesis overview .....................................................................20 Sexual dimorphism and body condition in the Australasian gannet.................23 Abstract................................................................................................................24 Introduction..........................................................................................................25 Methods ...............................................................................................................28
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