The evolutionarily stable strategy, animal contests, parasitoids, pest control and sociality

The evolutionarily stable strategy, ESS, concept was first used in biology to understand sex ratio bias and, shortly afterwards, to explore the logic of contests over essential and indivisible resources. ESS models formed the basis of much subsequent research on animal behaviour and placed game-theoretic thinking firmly within the behavioural ecology approach. Among behavioural ecologists studying parasitoids, it was those asking questions about the evolution of sex ratios who first made extensive use of the game-theoretic approach. A later growth of interest in parasitoid host defence and fighting behaviour made use of these tractable study species to explore contests and their connections to further aspects of life-history evolution plus some pest control applications. Our aims are to (i) introduce the topic of contests, which are engaged in by a very wide array of animal taxa, and the importance, both historical and conceptual, of the game-theoretic approach to their study, and (ii) review recent studies of parasitoid contests, including those that have considered the context of social evolution and the performance of parasitoids as agents of biological control. We consider that game-theoretic models are eminently testable and applicable and will likely endure as valuable tools in studies of parasitoid biology. This article is part of the theme issue 'Half a century of evolutionary games: a synthesis of theory, application and future directions'.