Maynard Smith & Parker's (1976) rule book for animal contests, mostly

Abstract

Maynard Smith & Parker (1976 , Animal Behaviour , 24 , 159–175) offered animal behaviourists and behavioural ecologists a theoretical framework/guide to understanding animal behaviour in competitive contexts. In this essay I trace the influence of this ‘contest rule book’ from the factors that led the two researchers to develop a treatise on the logic of the asymmetric game to empirical tests of the contest rules and theoretical additions made to the basic model and its underlying assumptions. Over a thousand studies cite this paper directly and thousands more cite work spurred by the original paper. The vast majority of these studies confirm the evolutionarily stable strategy (ESS) predictions made by Maynard Smith & Parker. Theoretical and empirical deviations from EES can largely be explained by the need for further structuring of the analyses into subgames and investigation of less obvious asymmetries than apparent size and resource value. To date, much progress has been made in three areas of interest to behaviourists: (1) understanding of the strategic nature of contests between conspecifics over limited resources; (2) modelling developments that deal with how information about potential asymmetries is gained; and (3) evaluation of the question of honest signalling with specific reference to threat displays. Here, I propose suggestions for future work, much of which will either require collaboration with mathematicians, or require that students interested in animal behaviour obtain a strong foundation in biomathematics. My preference is for the latter strategy. • I describe contributions of Maynard Smith & Parker's (1976) simple asymmetric game to our understanding of animal contests. • Their hawk–dove game, while simple, provided insight into the issue of honest signalling during assessment. • Nonobvious assessments appear to underlie apparent cases of honest signalling. • I discuss recent theoretical contributions in two areas: analysis and assessment. • Formal ESS analyses of the subgame type may explain deviations from ESS predictions in the case of repeated contests.