Go with the flow: Water velocity regulates herbivore foraging decisions in river catchments

Foragers typically attempt to consume food resources that offer the greatest energy gain for the least cost, switching between habitats as the most profitable food resource changes over time. Optimal foraging models require accurate data on the gains and costs associated with each food resource to successfully predict temporal shifts. Whilst previous studies have shown that seasonal changes in food quantity and quality can drive habitat shifts, few studies have shown the effects on habitat choice of seasonal changes in metabolic foraging costs. In this study we combined field and literature data to construct an optimal foraging model to examine the effect of seasonal changes in food quantity, food quality and foraging costs on the timing of a switch from terrestrial to aquatic habitat by non-breeding mute swans Cygnus olor in a shallow river catchment. Feeding experiments were used to quantify the functional response of swans to changes in aquatic plant biomasses. By sequentially testing alternative models with fixed or variable values for food quantity, food quality and foraging cost, we found that we needed to include seasonal variance in foraging costs in the model to accurately predict the observed habitat switch date. However, we did not need to include seasonal variance in food quantity and food quality, as accurate predictions could be obtained with fixed values for these two parameters. Therefore, the seasonal changes in foraging costs were the key factor influencing the behavioural decision to switch feeding habitats. These seasonal changes in foraging costs were driven by changes in water velocity; the profitability of aquatic foraging was negatively related to water velocity, as faster water required more energy to be expended in swimming. Our results demonstrate the importance of incorporating seasonal variation in foraging costs into our understanding of the foraging decisions of animals.