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Anthornis Melanura) Lincoln University Digital Thesis Copyright Statement The digital copy of this thesis is protected by the Copyright Act 1994 (New Zealand). This thesis may be consulted by you, provided you comply with the provisions of the Act and the following conditions of use: you will use the copy only for the purposes of research or private study you will recognise the author's right to be identified as the author of the thesis and due acknowledgement will be made to the author where appropriate you will obtain the author's permission before publishing any material from the thesis. Information use during foraging by New Zealand bellbirds (Anthornis melanura) A thesis submitted in partial fulfilment of the requirements for the Degree of Doctor of Philosophy (PhD) at Lincoln University by Jennifer May Dent Lincoln University 2019 Abstract of a thesis submitted in partial fulfilment of the requirements for the Degree of Doctor of Philosophy (PhD) Abstract Information use during foraging by New Zealand bellbirds (Anthornis melanura) by Jennifer May Dent Nectivorous foragers rely on patchily distributed, static resources of highly variable quality. The ability to learn and remember information concerning the spatial distribution of available resources, and to associate this with variation in resource quality, is an adaptive trait for many nectivorous species. I examine informed foraging in regard to the New Zealand bellbird (Anthornis melanura). Bellbirds are facultative nectarivores; although they demonstrate an affinity for nectar, they readily switch to alternate food sources when it is unavailable. Informed foraging by nectarivores has largely been examined with respect to more highly specialised species (e.g. Trochilidae). Focus on a facultative nectarivore allows for examination of cognitive constraints in a less specialised system, where the availability of nectar poses fewer restrictions on forager behaviour. The first part of this thesis explored information use in natural environments by assessing engagement in resource tracking by bellbird populations at two spatial scales (Chapter 2-3). At a local scale, the nectivorous activity of resident bellbirds within 1 ha plots of native bush was correlated with spatial patterns of flowering for four key nectar species. The strength of the tracking response observed was linked to the pollination syndrome of the focal plants. Availability of ornithophilous plant species elicited stronger and more consistent tracking responses than the availability of entomophilous species. At a larger, landscape scale the movements of seasonally transient birds ii were quantified using a dialectal song type matching technique. Dispersal was found to be more distance limited than previously assumed. This suggests that large scale resource tracking may operate at an intermediary patch scale, rather than at a landscape scale. The second part of this thesis examined the cognitive mechanisms that may underlie informed foraging behaviours. I performed a series of experimental feeder trials on free-living male bellbirds (Chapter 4-6). Trials examined the ability of individuals to return to the location of a highly rewarding feeder within an array of less rewarding feeders. Treatments varied with respect to the reward concentration, the duration of the withholding period, the presence of visual cues, and the size of the experimental array. Bellbirds were found to have accurate spatial memory. Individuals were able to relocate rewards across a range of concentration treatments, retain spatial information for periods of up to 20 days, learn to associate visual cues with reward characteristics, and use spatial cues in a scale dependent manner. Overall, my research indicates that bellbirds were capable of engaging in memory informed foraging on nectar resources. Targeted exploitation of resources across multiple spatial scales likely translates to increased foraging efficiency. Low dietary dependence on nectar is associated with flexible usage of informed foraging, rather than a lack of cognitive ability in this species. Keywords: facultative nectarivores, memory informed foraging, resource tracking, resource value, spatial memory, visual cues iii Acknowledgements I am extremely grateful for the many people who have assisted me during my PhD journey. Firstly, I would like to thank my original primary supervisor, Laura Molles. Working with Laura was one of the main reasons I decided to embark upon my doctoral studies at Lincoln University as I have much admiration for her approach to both research and life. Thank you for helping me to develop a project that I still find exciting 4 years down the track, and for encouraging me to pursue all the weird and exciting research ideas that caught my fancy. Thank you also for all your help with bird banding, setting up feeder trials, sorting through acoustic data, and for your continued help in developing manuscripts. I would also like to express my gratitude to Adrian Paterson, who has been my primary supervisor since Laura departed the university. Thank you for helping me to rally for the final leg of my PhD journey – your assistance with preparation of my thesis and manuscripts has been invaluable. Thank you also to Jon Sullivan for your help with development of my project, assistance with collection of acoustic data, and in interpretation of landscape scale data trends, your perspective has helped me to understand the bigger picture of bellbird behaviour. Special thanks to Gill and Trevor Bedford who allowed me to conduct experimental trials at their property and went above and beyond to help me maintain those trials. My trips to Takamatua were some of the highlights of my PhD - I doubt I would have tried to accomplish so many feeder trials if it hadn’t been for your warm hospitality (and cups of tea)! I would also like to thank Michael Fake and Vanessa Mander for your help with fieldwork and for providing company on some otherwise long and tiring days in the field. This research would not have been possible without the assistance of a Lincoln University Doctoral Scholarship, and I am also grateful to have received a Shirtcliffe Fellowship and the LW McCaskill award which assisted me during this research. All activities described in this thesis were reviewed iv and authorized by the Lincoln University Animal Ethics committee (AEC 606 and AEC 2016-20) and were performed with permission of the Department of Conservation (Wildlife Act Authorisation 43202-FAU). Banding associated with this project was performed under supervision of Laura Molles (License 2013/085) v George Weasley, the bellbird Photo: Jennifer Dent vi Table of Contents Abstract ....................................................................................................................................... ii Acknowledgements ..................................................................................................................... iv Table of Contents ....................................................................................................................... vii List of Tables ................................................................................................................................ x List of Figures .............................................................................................................................. xi Chapter 1 Introduction ................................................................................................................. 1 1.1 Information use in foraging ........................................................................................................... 1 1.2 Value of information ...................................................................................................................... 2 1.3 Adaptive specialisation .................................................................................................................. 4 1.4 Nectivory and information use ...................................................................................................... 5 1.5 Avian nectarivores ......................................................................................................................... 7 1.6 Avian nectivory in New Zealand..................................................................................................... 9 1.7 Bellbirds ......................................................................................................................................... 9 1.8 Thesis structure ............................................................................................................................ 12 1.8.1 Part One .......................................................................................................................... 13 1.8.2 Part Two .......................................................................................................................... 13 Chapter 2 Spatiotemporal resource tracking at a local scale by New Zealand bellbirds (Anthornis melanura) .................................................................................................................................. 15 2.1 Abstract ........................................................................................................................................ 15 2.2 Introduction ................................................................................................................................. 16 2.3 Methodology ...............................................................................................................................
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