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'.Peer reviewe

Defection on the bounty? Kinship and cooperative exploitation of a rich, essential but dangerous resource

We develop a game-theoretic model to explore the question of whether two animals should cooperate in the dangerous activity of obtaining a rich and essential resource. We consider variation in the risks incurred to individuals and in how the activities of the two animals interact to influence the probability of success. We also consider that the animals may be relatives and thus share evolutionary interests. The model is general and can, for instance, be applied to mammalian predators attempting to capture and subdue large and dangerous prey or to female parasitoid wasps that attack and, if successful, paralyse aggressive hosts that then provide the only feeding resource for their offspring. This minimal model of cooperation contains three dimensionless parameters: vulnerability (the ratio between the average time for a lone attacker to subdue the defending resource and the average time for the defender to fatally strike the attacker), the dilution ratio (the extent to which attack by animals acting in tandem reduces a defender's ability to kill its attackers) and the relatedness between the potential attackers. The model predicts that higher values of all three parameters favour cooperation and that for small values cooperation is not evolutionarily stable. Cooperation can arise from an ancestral state of non-cooperation if values of all parameters are sufficiently high but cannot arise among non-relatives, irrespective of other parameter values. Once cooperation has emerged in a population, it can be maintained among nonrelatives at modest values of dilution ratio and vulnerability. We discuss these general predictions in particular relation to the parasitoid genus Sclerodermus, in which multiple females may attack unusually large and aggressive hosts and in which host attack behaviour is mediated by kinship. (c) 2021 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.Peer reviewe

Volatile chemical emission as a weapon of rearguard action: a game-theoretic model of contest behavior

We use a game-theoretic model to explore whether volatile chemical (spiroacetal) emissions can serve as a weapon of rearguard action. Our basic model explores whether such emissions serve as a means of temporary withdrawal, preventing the winner of the current round of a contest from translating its victory into permanent possession of a contested resource. A variant of this model explores an alternative possibility, namely, that such emissions serve as a means of permanent retreat, attempting to prevent a winner from inflicting costs on a fleeing loser. Our results confirm that the underlying logic of either interpretation of weapons of rearguard action is sound; however, empirical observations on parasitoid wasp contests suggest that the more likely function of chemical weapons is to serve as a means of temporary withdrawal. While our work is centered around the particular biology of contest behavior in parasitoid wasps, it also provides the first contest model to explicitly consider self-inflicted damage costs, and thus responds to a recent call by empiricists for theory in this area

The impact of competition on elephant musth strategies: a game–theoretic model

Mature male African Savannah elephants are known to periodically enter a temporary state of heightened aggression called “musth,” often linked with increased androgens, particularly testosterone. Sexually mature males are capable of entering musth at any time of year, and will often travel long distances to find estrous females. When two musth bulls or two non-musth bulls encounter one another, the agonistic interaction is usually won by the larger male. However, When a smaller musth bull encounters a larger non-musth bull, the smaller musth male can win. The relative mating success of musth males is due partly to this fighting advantage, and partly to estrous females’ general preference for musth males. Though musth behavior has long been observed and documented, the evolutionary advantages of musth remain poorly understood. Here we develop a game–theoretic model of male musth behavior which assumes musth duration as a parameter, and distributions of small, medium and large musth males are predicted in both time and space. The predicted results are similar to the musth timing behavior observed in the Amboseli National Park elephant population, and further results are generated with relevance to Samburu National Park. We discuss small male musth behavior, the effects of estrous female spatial heterogeneity on musth timing, conservation applications, and the assumptions underpinning the model

An introduction to game-theoretic modelling

This book introduces game theory and its applications from an applied mathematician's perspective, systematically developing tools and concepts for game-theoretic modelling in the life and social sciences. Filled with down-to-earth examples of strategic behavior in humans and other animals, the book presents a unified account of the central ideas of both classical and evolutionary game theory. Unlike many books on game theory, which focus on mathematical and recreational aspects of the subject, this book emphasizes using games to answer questions of current scientific interest. In the present third edition, the author has added substantial new material on evolutionarily stable strategies and their use in behavioral ecology. The only prerequisites are calculus and some exposure to matrix algebra, probability, and differential equations

An Introduction to Game-Theoretic Modelling

This is an introduction to game theory and applications with an emphasis on self-discovery from the perspective of a mathematical modeller. The book deals in a unified manner with the central concepts of both classical and evolutionary game theory. The key ideas are illustrated throughout by a wide variety of well-chosen examples of both human and non-human behavior, including car pooling, price fixing, food sharing, sex allocation and competition for territories or oviposition sites. There are numerous exercises with solutions

Variation between Self- and Mutual Assessment in Animal Contests

Limited resources lead animals into conflicts of interest, which are resolved when an individual withdraws from a direct contest. Current theory suggests that the decision to withdraw can be based on a threshold derived from an individual’s own state (self-assessment) or on a comparison between their own state and their opponent’s (mutual assessment). The observed variation between these assessment strategies in nature does not conform to theory. Thus, we require theoretical developments that explain the functional significance of different assessment strategies. We consider a hawk-dove game with two discrete classes that differ in fighting ability, in which the players strategically decide on their investment toward mutual assessment. Analysis of the model indicates that there are simultaneous trade-offs relating to assessment strategies. First, weak individuals in a population must decide on whether to acquire information about their opponents at the cost of providing opponents with information about themselves. Secondly, all individuals must decide between investing in mutual assessment and being persistent in contests. Our analysis suggests that the potential for individuals to make errors during contests and differences in the consequences of sharing information within a population may serve as fundamental concepts for explaining variation in assessment strategy.peerReviewe

Victory displays: a game-theoretic analysis

Two rationales have been proposed verbally for the function of victory displays, which are performed by the winners of contests but not by the losers. The "advertising" rationale is that victory displays are attempts to communicate victory to other members of a social group that do not pay attention to contests or cannot otherwise identify the winner. The "browbeating" rationale is that victory displays are attempts to decrease the probability that the loser of a contest will initiate a future contest with the same individual. We formally explore the logic of these rationales with game-theoretic models. The models show that both rationales are logically sound; however, all other things being equal, the intensity of victory displays will be highest through advertising in groups where the reproductive advantage of dominance is low and highest through browbeating in groups where the reproductive advantage of dominance is high. Copyright 2006.bystander effects; game theory; signaling; victory displays

Modeling the evolution of winner and loser effects: A survey and prospectus

International audienc