Papers focus on population ecology, plant-animal interactions, ecosystem ecology, community ecology, global change ecology, conservation ecology, behavioral ecology and physiological ecology.
His main areas of research are evolutionary biology, behavioural and community ecology, vertebrate conservation, and feeding and defence in lizards and snakes.
His research in community ecology has been in two broad areas: the role of disturbance in structuring natural communities and the ecology of host-parasite interactions.
Community ecology studies those characteristics of communities that either are not apparent or which are much less apparent if a community consists of only a single population.