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Complex Interactions Between Intrinsic and Extrinsic Factors Govern the Diets of Herbivores

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Complex Interactions Between Intrinsic and Extrinsic Factors Govern the Diets of Herbivores Complex interactions between intrinsic and extrinsic factors govern the diets of herbivores. Keryn Frances Bain This thesis is in fulfilment of the requirements for the degree of Doctor of Philosophy School of Biological, Earth and Environmental Sciences Faculty of Science April 2018 THE UNIVERSI1YOF NEW SOUTH WALES Thesis/Dissertation Sheet Surname or Family name: Bain First name: Keryn Other name/s: Frances Abbreviation fordegree as given in the University calendar: PhD School: BEES Faculty: Science Title: Complex interactions between intrinsic and extrinsic factors govern the diets of herbivores. Thesis Abstract Consumer prey interactions are crucial for the transfer of energy through food webs and variations in those interactions have important consequences on the structure and composition of ecological communities. In the field, consumer diets vary widely both among and within consumer species. This variability can be attributed to intrinsic factors acting on individuals that alter preferences and extrinsic factors that alter their ability to obtain resources. As a consequence, what is consumed by an individual at a given time often does not reflectthe total dietary niche of that species or even the life span of that individual. In this thesis, I use a common generalist herbivore, the marine gastropod Luoella torquatus, as a model consumer to examine the intrinsic and extrinsic constraints on diet choice. First, I establish a method using near infrared reflectance spectroscopy (NIRS) to measure the diets of herbivores in the field and then use a combination of field surveys and experiments to identify how extrinsic and intrinsic factors impact the diets of marine herbivores. Given the slow mobility of chis herbivore relative to others, the local availability of macroalgae across temperate and spatial scales was the primary extrinsic factor considered and as there was some evidence for diet mixing in this herbivore, the primary intrinsic factor considered was recent dietary history. With careful calibrations and feeding assays, NIRS provided a non-destructive method to quantify the realised diets of free-living consumers. I identified that the local availability of algae, and its variation through time, can inhibit individuals from fully expressing their preference. Consequently, the diets of individuals in the field closely track the availability of resources. Nevertheless, at small scales, individuals will display preference for favoured algal species, if they are available, and are capable of altering diet choices depending on recent consumption, potentially compensating for poor diets through diet mi.xing. I hereby grant to the University of New South Wales or its agents the right to archive and to make available my thesis or dissertation in whole or in part in the University libraries in all forms of media, now or here after known, subject to the provisions of the Copyright Act 1968. I retain all property rights, such as patent rights. I also retain the right to use in future works (such as articles or books) all or part of this thesis or dissertation. I also authorise University Microfilmsto use the 350 word abstract of my thesis in Dissertation Abstracts Intern · his is applicable to doctoral theses only). • 11/04/2018 The University recognises that there may be exceptional circumstances requiring restrictions on copying or conditionson use. Requests for restriction for a period of up to 2 years must be made in writing. Requests for a longer period of restriction may be considered in exceptional circumstances and re uire the a roval of the Dean of Graduate Research. FOR OFFICE USE ONLY Date of completion of requirements for Award: --- -------------------------- ORIGINALITY STATEMENT ‘I hereby declare that this submission is my own work and to the best of my knowledge it contains no materials previously published or written by another person, or substantial proportions of material which have been accepted for the award of any other degree or diploma at UNSW or any other educational institution, except where due acknowledgement is made in the thesis. Any contribution made to the research by others, with whom I have worked at UNSW or elsewhere, is explicitly acknowledged in the thesis. I also declare that the intellectual content of this thesis is the product of my own work, except to the extent that assistance from others in the project's design and conception or in style, presentation and linguistic expression is acknowledged.’ Signed …………………………………………….............. Date …………………………………………….............. COPYRIGHT STATEMENT ‘I hereby grant the University of New South Wales or its agents the right to archive and to make available my thesis or dissertation in whole or part in the University libraries in all forms of media, now or here after known, subject to the provisions of the Copyright Act 1968. I retain all proprietary rights, such as patent rights. I also retain the right to use in future works (such as articles or books) all or part of this thesis or dissertation. I also authorise University Microfilms to use the 350 word abstract of my thesis in Dissertation Abstract International (this is applicable to doctoral theses only). I have either used no substantial portions of copyright material in my thesis or I have obtained permission to use copyright material; where permission has not been granted I have applied/will apply for a partial restriction of the digital copy of my thesis or dissertation.' Signed ……………………………………………........................... Date ……………………………………………........................... AUTHENTICITY STATEMENT ‘I certify that the Library deposit digital copy is a direct equivalent of the final officially approved version of my thesis. No emendation of content has occurred and if there are any minor variations in formatting, they are the result of the conversion to digital format.’ Signed ……………………………………………........................... Date ……………………………………………........................... Acknowledgements I would like to specifically acknowledge the following people and organizations that have made this thesis possible. Firstly, I would like to express my special appreciation and thanks to my supervisor Professor Alistair Poore. Without his guidance, constant feedback and mentoring this PhD would not have been achievable. I would not be the scientist I am today without Alistair’s brilliant supervision. I would also like to thank my committee members Professor Peter Steinberge, Professor Angela Moles, Dr Adriana Vergès and Professor Rob Brooks for providing much needed advice on strategies for writing, publication and research direction. A special thanks to Gordana Popovic for providing advice on statistical methods and analysis of data collected for Chapter 5. Chapter 2 was improved by the comments provided by Professor Aaron Galloway and Professor Jennifer Sorenson-Forbey. To my fellow Ecology lab mates, thanks for providing the fun, Friday drinks and field help. In particular, Damon Bolton, Brendan Lanham, Kingsley Griffin and Janine Ledet, who were always available and happy (felt obliged) to help out in the field. A special thanks to my parents for their encouragement throughout my studies and to my husband Wes, for his continued support, love, understanding and keeping me sane(ish). This accomplishment would not have been possible without my family. Contents page List of abbreviations and symbols i List of Figures iii List of Tables v List of Supplementary Figures and Tables vii Thesis abstract viii Chapter 1. General Introduction Intrinsic and extrinsic constraints on diet 1 Quantifying the diets of free-living consumers 4 Scope and objectives of research 6 Chapter 2. Use of near infrared reflectance spectroscopy to quantify diet mixing in a generalist marine herbivore Abstract 10 Introduction 11 Methods 16 Study organisms and sites 16 Discrimination among singles species diets 17 Calibration and validation of prediction models 19 Discrimination among components in a mixed species diet 22 Results 26 Discrimination among singles species diets 26 Discrimination among components in a mixed species diet 28 Discussion 40 Supplementary material 47 Chapter 3. Spatial and temporal patterns in individual diets of a marine herbivore relative to local availability of food sources Abstract 54 Introduction 55 Methods 59 Species description and study sites 59 Spatial and temporal patterns of algal availability 62 Spatial and temporal patterns of L. torquatus diets 64 Linking diets to locally available algal species 66 Results 68 Spatial and temporal patterns of algal availability 68 Spatial and temporal patterns of L. torquatus diets 69 Linking diets to locally available algal species 70 Discussion 83 Spatial and temporal variation in food availability 83 Evidence for selection among available algae on small scales 87 Using near infrared to assess spatial and temporal patterns in field diets 88 Chapter 4. Field diets of a generalist herbivore are determined first by availability then by preference. Abstract 91 Introduction 92 Methods 97 Study organisms and collections 97 Feeding rates and preferences among species of algae 98 Expanding NIRS predictive models to multiple species 100 Temporal and spatial patterns in the consumption of three algal species 103 Results 105 Feeding rates and preferences among species of algae 105 Expanding NIRS predictive models to multiple species 107 Temporal and spatial patterns in the consumption of three algal species 108 Discussion 120 Predicting field consumption from laboratory
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