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Swan-Plant Interactions in a Chalk River Catchment

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Swan-Plant Interactions in a Chalk River Catchment Swan-plant interactions in a chalk river catchment Kevin Anthony Wood This thesis has been submitted in partial fulfilment of the requirements of the degree of Doctor of Philosophy. Bournemouth University in collaboration with the Centre for Ecology & Hydrology (Natural Environment Research Council) May 2012 1 This copy of the thesis has been supplied on condition that anyone who consults it is understood to recognise that its copyright rests with its author and due acknowledgement must always be made of the use of any material contained in, or derived from, this thesis. 2 Abstract “Swan-plant interactions in a chalk river catchment”. Kevin A. Wood Plants are of fundamental importance to the structure, functioning and service provision of many ecosystems. However, herbivores can have negative ecological and socioeconomic effects on plant communities through consumption, trampling and alteration of nutrient cycles. In this thesis I address a particular herbivore-plant interaction: the grazing of plants in chalk river catchments, principally the submerged macrophyte water crowfoot (Ranunculus penicillatus ssp. pseudofluitans (Syne) S.D. Webster) and terrestrial pasture grass species, by flocks of non-breeding mute swans (Cygnus olor Gmelin, 1789). This research was carried out over two years in the River Frome catchment (Dorset, UK). Based on a meta-analysis of previous waterfowl grazing studies I show that waterfowl biomass density (kg ha-1) rather than individual density (ind. ha-1) is a better predictor of reductions in plant standing crop. Most studies to date have analysed such reductions using only individual densities, despite large between-taxa variation in waterfowl body mass, diet and intake rates. I quantified the abundance, species richness, evenness, flowering and dominance of the chalk river aquatic plant community in relation to biotic and abiotic factors during the growth-, peak-, and recession-phases of the growth cycle. The relative importance of herbivory, riparian shading, water temperature and distance downstream varied between different phases of the plant growth cycle, highlighting the importance of seasonal patterns in regulation of plant community structure. The River Frome swan population varied seasonally, being highest in the winter. The population was dominated by non-breeding adults and juveniles that lived in flocks. These flocks exhibited strong seasonal habitat switches between terrestrial pasture in winter and spring, and river in summer and autumn. I provided evidence that this switch was linked to the seasonal decrease in water velocity between spring and summer, which reduced the metabolic costs of river feeding and increased the relative profitability of aquatic food resources. I used a mathematical population model and an individual-based behavioural model respectively to explore two management options for the alleviation of the swan grazing conflict in chalk rivers: population control and habitat alterations. Population control measures, such as clutch manipulations, fertility control, culling or translocations, were predicted to be unsuccessful except at impractically high levels of management effort, due to the effects of immigration and high survival rates in offsetting removed eggs or individuals. Habitat alterations, in particular the narrowing of river channels to cause a local increase in water velocity and thus swan foraging costs, are more promising management options as they require lower management effort, are less ethically controversial, and address the fundamental reason why swans select their food resources, the rate of net energy gain (‘profitability’). 3 List of contents Copyright statement............................................................................................... p 2 Abstract.................................................................................................................. p 3 List of contents....................................................................................................... p 4 List of figures.......................................................................................................... p 8 List of tables........................................................................................................... p 12 Acknowledgements............................................................................................... p 14 Declaration............................................................................................................. p 15 1: ‘Swan-plant interactions in a chalk river catchment: an introduction’................. p 16 1.1: The roles of plants in ecosystems.............................................................. p 16 1.2: Herbivore effects on plants: individuals to communities............................. p 17 1.3: Herbivorous waterfowl................................................................................ p 21 1.4: Grazing conflicts......................................................................................... p 24 1.5: The management of herbivores and their effects....................................... p 25 1.6: The role of modelling in plant-herbivore interactions.................................. p 26 1.7: The chalk river ecosystem.......................................................................... p 27 1.8: Mute swan ecology..................................................................................... p 30 1.9: The study system: The River Frome (Dorset, UK)..................................... p 33 1.10: Thesis overview........................................................................................ p 37 2: ‘The impact of waterfowl herbivory on plant standing crop: a meta-analysis’..... p 39 2.1: Abstract...................................................................................................... p 39 2.2: Introduction................................................................................................. p 39 2.3: Methods...................................................................................................... p 41 2.3.1: Study species..................................................................................... p 41 2.3.2: Waterfowl densities and reductions in plant standing crop................ p 42 2.3.3: Differences between waterfowl taxa: implications of body mass....... p 43 2.3.4: Waterfowl size and rates of food consumption................................... p 43 2.3.5: Waterfowl size and herbivorous diet.................................................. p 43 2.3.6: Statistical analyses............................................................................. p 44 2.4: Results........................................................................................................ p 44 2.4.1: Waterfowl densities and reductions in plant standing crop................ p 44 2.4.2: Differences between waterfowl taxa: implications of body mass....... p 45 2.4.3: Waterfowl size and rates of food consumption.................................. p 46 2.4.4: Waterfowl size and herbivorous diet.................................................. p 47 2.5: Discussion.................................................................................................. p 48 3: ‘Measuring submerged macrophyte standing crop in shallow rivers: a test of methodology’.......................................................................................................... p 51 3.1: Abstract...................................................................................................... p 51 3.2: Introduction................................................................................................ p 51 3.3: Methods...................................................................................................... p 52 4 3.3.1: Study sites.......................................................................................... p 52 3.3.2: Measuring macrophyte cover, volume, and stand height................... p 53 3.3.3: Estimating biomass from stand height............................................... p 54 3.3.4: Statistical analyses............................................................................. p 54 3.4: Results........................................................................................................ p 55 3.4.1: Cover, volume, and biomass measures............................................. p 55 3.4.2: Biomass measures and estimates from stand height......................... p 56 3.5: Discussion.................................................................................................. p 60 4: ‘Responses of a shallow river plant community to four variables vary across the plant growth cycle’............................................................................................ p 63 4.1: Abstract...................................................................................................... p 63 4.2: Introduction................................................................................................. p 63 4.3: Methods...................................................................................................... p 65 4.3.1: Study sites.......................................................................................... p 65 4.3.2: Estimating required sample size........................................................ p 66 4.3.3: Plant abundance...............................................................................
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