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Nutritional Ecology of Brushtail Possums (Trichosurus Vulpecula Kerr)

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Nutritional Ecology of Brushtail Possums (Trichosurus Vulpecula Kerr) Nutritional Ecology of Brushtail Possums (Trichosurus vulpecula Kerr) in New Zealand Jenifer Eve Morley Thesis submitted for the degree of Doctor of Philosophy University of East Anglia School of Environmental Sciences April 2018 "This copy of the thesis has been supplied on condition that anyone who consults it is understood to recognise that its copyright rests with the author and that use of any information derived there from must be in accordance with current UK Copyright Law. In addition, any quotation or extract must include full attribution.” Abstract In this thesis, a combination of field observations and laboratory experiments are used to address the question: how do the nutritional strategies of brushtail possums enable this alien pest species to obtain a balance of nutrients at native forest-pasture margins in New Zealand? Possums had a polyphagous diet with a core of leaves from three species, supplemented by flowers in spring and summer, fruits in autumn and invertebrates in winter. Graphical models confirmed that possums were polyphagous in all seasons, although diet width and the extent to which individual species of food dominated the diet varied between seasons. Four nutrient characteristics of potential foods were analysed: total nitrogen, available nitrogen, soluble organic matter and total digestible dry matter. They varied significantly between food species and between seasons but were not significantly associated with either proportion by mass, or frequency of occurrence in the diet. In laboratory experiments relative preferences between leaves of three woody species changed when the basal leaf diet was supplemented by flowers or by pasture species. Intake rates of nutrients also varied according to diet. The target nutrient intake ratio of available nitrogen to digestible dry matter was calculated. Predictions from optimal foraging, nitrogen limitation and mixed diet theories were compared but mixed diet theory was supported most strongly. Home ranges were delineated using data from GPS telemetry. Nocturnal foraging pathways were characterised and divided to distinguish between searching within patches and travelling between them. Possums travelled further per night and travelled faster between search patches in winter than they did in autumn. Selection ratios showed that native forest and native shrub/scrubland were preferred habitats for both travelling and searching. The implications of applying mixed diet theory to possum nutritional ecology, for improving the efficiency of control programmes and reducing interactions between possums and cattle are explored. 2 Contents Title Abstract .................................................................................................................2 Contents ................................................................................................................3 List of tables and figures .....................................................................................9 Acknowledgements ............................................................................................14 1 Introduction ..............................................................................................16 1.1 Philosophical background .....................................................16 1.2 Nutritional ecology.................................................................16 1.3 Foraging strategies................................................................17 1.4 Plant defence........................................................................19 1.5 Foraging theory.....................................................................20 1.6 A multifaceted approach........................................................22 1.7 Chapter summaries...............................................................22 1.7.1 Chapter 2: The study species and study sites............22 1.7.2 Chapter 3: Seasonal and sex variation in the foraging strategies of an alien invader: the brushtail possum in New Zealand..................................................................................22 1.7.3 Chapter 4: Seasonal variation in nutritional content of plant foods available to brushtail possums at forest-pasture margins in New Zealand........................................................23 1.7.4 Chapter 5: Diet composition and nutrient intake rates of brushtail possums fed mixed diets.........................................24 1.7.5 Chapter 6: Foraging pathway characteristics and habitat selection of the brushtail possum at forest-pasture margins in New Zealand.........................................................24 1.7.6 Chapter 7: Discussion.................................................25 2. The study species and study sites .........................................................26 2.1 The study species: Trichosurus vulpecula .......................................26 3 2.1.1 Possums as alien pest species..............................................26 2.1.2 Possum biology and ecology.................................................27 2.1.3 Possum diet and foraging strategies.....................................28 2.1.4 Possum impacts and control in New Zealand........................33 2.1.5 Possums as a model species for increasing our understanding of nutritional ecology.........................................................................34 2.1.6 Opportunities for future research into the nutritional ecology of T. vulpecula in New Zealand............................................................35 2.2 Study sites .......................................................................................37 2.2.1 The wider landscape..............................................................37 2.2.2 Stomach and vegetation sampling sites – 300 and 353 Mount Cargill Road......................................................................................37 2.2.3 GPS telemetry site – Pigeon Flat..........................................38 3. Seasonal and sex variation in the foraging strategies of an alien invader: the brushtail possum in New Zealand......................................42 3.1 Abstract ...........................................................................................42 3.2 Introduction ......................................................................................44 3.2.1 Resource use and foraging strategies...................................44 3.2.2 Seasonal and sex variation in foraging strategies.................44 3.2.3 Measuring diet and the Amundsen approach........................45 3.2.4 Variation in possum foraging strategies.................................46 3.2.5 Specific ‘gaps in the science’.................................................47 3.2.6 Aims, objectives and hypotheses..........................................48 3.3 Methods ...........................................................................................49 3.3.1 Possum trapping and stomach sampling...............................49 3.3.2 Identification of species eaten...............................................50 3.3.3 Calculations of diet composition descriptors..........................52 3.3.4 Hypothesis testing.................................................................53 3.3.5 Statistical analyses................................................................53 4 3.3.7 Graphical analyses................................................................53 3.4 Results ............................................................................................56 3.4.1 Possum diet and availability of foods....................................56 3.4.2 Influence of season and sex on foraging patterns.................59 3.4.3 Seasonal variation in species composition of diet.................66 3.4.4 Graphical approach results....................................................69 3.5 Discussion .......................................................................................71 3.5.1 Possum diet and availability of foods....................................71 3.5.2 Use of foliar and non-foliar foods..........................................72 3.5.3 Influence of season and sex on diet composition..................74 3.5.4 Seasonal changes in diet width, food importance, feeding strategy and niche breadth....................................................75 4. Seasonal variation in nutritional content of plant foods available to brushtail possums at forest-pasture margins in New Zealand............ 78 4.1 Abstract ...........................................................................................78 4.2 Introduction .....................................................................................80 4.2.1 Determinants of food quality for herbivores..........................80 4.2.2 Possum-specific background.................................................81 4.2.3 Aims, objectives and hypotheses..........................................81 4.3 Methods ..........................................................................................83 4.3.1 Sampling design................ ..................................................83 4.3.2 Collection and preparation of plant tissues...........................85 4.3.3 Measures of food chemistry..................................................85 4.3.3.1 The availN assay.............................................85 4.3.3.2 Total nitrogen determination............................87 4.3.4 Statistical analyses................................................................88
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