Non-lethal effects of predation in birds

Predators can affect individual fitness and population and community processes through lethal effects (direct consumption or 'density' effects), where prey is consumed, or through non-lethal effects (trait-mediated effects or interactions), where behavioural compensation to predation risk occurs, such as animals avoiding areas of high predation risk. Studies of invertebrates, fish and amphibians have shown that non-lethal effects may be larger than lethal effects in determining the behaviour, condition, density and distribution of animals over a range of trophic levels. Although non-lethal effects have been well described in the behavioural ecology of birds (and also mammals) within the context of anti-predation behaviour, their role relative to lethal effects is probably underestimated. Birds show many behavioural and physiological changes to reduce direct mortality from predation and these are likely to have negative effects on other aspects of their fitness and population dynamics, as well as affecting the ecology of their own prey and their predators. As a consequence, the effects of predation in birds are best measured by trade-offs between maximizing instantaneous survival in the presence of predators and acquiring or maintaining resources for long-term survival or reproduction. Because avoiding predation imposes foraging costs, and foraging behaviour is relatively easy to measure in birds, the foraging-predation risk trade-off is probably an effective framework for understanding the importance of non-lethal effects, and so the population and community effects of predation risk in birds and other animals. Using a trade-off approach allows us to predict better how changes in predator density will impact on population and community dynamics, and how animals perceive and respond to predation risk, when non-lethal effects decouple the relationship between predator density and direct mortality rate. The trade-off approach also allows us to identify where predation risk is structuring communities because of avoidance of predators, even when this results in no observable direct mortality rate.