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

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

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

Escaping the evolutionary trap: Can size-related contest advantage compensate for juvenile mortality disadvantage when parasitoids develop in unnatural invasive hosts?

The quality of hosts for a parasitoid wasp may be influenced by attributes such as host size or species, with high quality for successful development usually coincident with high quality for larger offspring. This is not always the case: for the Scelionid wasp Trissolcus basalis, oviposition in eggs of the Brown Marmorated Stink Bug, Halyomorpha halys, rather than of the normal host, the Southern Green Stink Bug, Nezara viridula, leads to lower offspring survival, but survivors can be unusually large. Adult female T. basalis engage in contests for host access. As larger contestants are typically favoured in contests between parasitoids, the larger size of surviving offspring may compensate for the mortality of others. We construct a general game-theoretic model to explore whether size advantage can sustain a maternal preference to utilize a more deadly host species. We find that size advantage alone is unlikely to sustain a shift in host preference, yet such an outcome is possible when size asymmetries act simultaneously with advantages in host possession (ownership effect). Halyomorpha halys is an invasive pest of major agro-economic importance in Europe and the Americas, and use of its eggs as hosts by native parasitoids such as T. basalis has been seen as an evolutionary trap due to their high developmental mortality. Our model suggests that the recently discovered effect of host choice on offspring size may provide an escape from the trap via effects on contest biology of T. basalis which could foster a more stable association with H. halys. An evolutionary shift in the reproductive value of H. halys could increase the efficiency of T. basalis as a biological control agent of this invasive stink bug pest

Exploring the cooperative regimes in a model of agents without memory or "tags": indirect reciprocity vs. selfish incentives

The self-organization in cooperative regimes in a simple mean-field version of a model based on "selfish" agents which play the Prisoner's Dilemma (PD) game is studied. The agents have no memory and use strategies not based on direct reciprocity nor 'tags'. Two variables are assigned to each agent $i$ at time $t$, measuring its capital $C(i;t)$ and its probability of cooperation $p(i;t)$. At each time step $t$ a pair of agents interact by playing the PD game. These 2 agents update their probability of cooperation $p(i)$ as follows: they compare the profits they made in this interaction $\delta C(i;t)$ with an estimator $\epsilon(i;t)$ and, if $\delta C(i;t) \ge \epsilon(i;t)$, agent $i$ increases its $p(i;t)$ while if $\delta C(i;t) < \epsilon(i;t)$ the agent decreases $p(i;t)$. The 4!=24 different cases produced by permuting the four Prisoner's Dilemma canonical payoffs 3, 0, 1, and 5 - corresponding,respectively, to $R$ (reward), $S$ (sucker's payoff), $T$ (temptation to defect) and $P$ (punishment) - are analyzed. It turns out that for all these 24 possibilities, after a transient,the system self-organizes into a stationary state with average equilibrium probability of cooperation $\bar{p}_\infty$ = constant $> 0$.Depending on the payoff matrix, there are different equilibrium states characterized by their average probability of cooperation and average equilibrium per-capita-income ($\bar{p}_\infty,\bar{\delta C}_\infty$).Comment: 11 pages, 5 figure

Coveting thy neighbors fitness as a means to resolve social dilemmas

In spatial evolutionary games the fitness of each individual is traditionally determined by the payoffs it obtains upon playing the game with its neighbors. Since defection yields the highest individual benefits, the outlook for cooperators is gloomy. While network reciprocity promotes collaborative efforts, chances of averting the impending social decline are slim if the temptation to defect is strong. It is therefore of interest to identify viable mechanisms that provide additional support for the evolution of cooperation. Inspired by the fact that the environment may be just as important as inheritance for individual development, we introduce a simple switch that allows a player to either keep its original payoff or use the average payoff of all its neighbors. Depending on which payoff is higher, the influence of either option can be tuned by means of a single parameter. We show that, in general, taking into account the environment promotes cooperation. Yet coveting the fitness of one's neighbors too strongly is not optimal. In fact, cooperation thrives best only if the influence of payoffs obtained in the traditional way is equal to that of the average payoff of the neighborhood. We present results for the prisoner's dilemma and the snowdrift game, for different levels of uncertainty governing the strategy adoption process, and for different neighborhood sizes. Our approach outlines a viable route to increased levels of cooperative behavior in structured populations, but one that requires a thoughtful implementation.Comment: 10 two-column pages, 5 figures; accepted for publication in Journal of Theoretical Biolog

Kleptoparasitic Interactions under Asymmetric Resource Valuation

We introduce a game theoretical model of stealing interactions. We model the situation as an extensive form game when one individual may attempt to steal a valuable item from another who may in turn defend it. The population is not homogeneous, but rather each individual has a different Resource Holding Potential (RHP). We assume that RHP not only influences the outcome of the potential aggressive contest (the individual with the larger RHP is more likely to win), but that it also influences how an individual values a particular resource. We investigate several valuation scenarios and study the prevalence of aggressive behaviour. We conclude that the relationship between RHP and resource value is crucial, where some cases lead to fights predominantly between pairs of strong individuals, and some between pairs of weak individuals. Other cases lead to no fights with one individual conceding, and the order of strategy selection is crucial, where the individual which picks its strategy first often has an advantage

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

Cooperation and Self-Regulation in a Model of Agents Playing Different Games

A simple model for cooperation between "selfish" agents, which play an extended version of the Prisoner's Dilemma(PD) game, in which they use arbitrary payoffs, is presented and studied. A continuous variable, representing the probability of cooperation, $p_k(t) \in$ [0,1], is assigned to each agent $k$ at time $t$. At each time step $t$ a pair of agents, chosen at random, interact by playing the game. The players update their $p_k(t)$ using a criteria based on the comparison of their utilities with the simplest estimate for expected income. The agents have no memory and use strategies not based on direct reciprocity nor 'tags'. Depending on the payoff matrix, the systems self-organizes - after a transient - into stationary states characterized by their average probability of cooperation $\bar{p}_{eq}$ and average equilibrium per-capita-income $\bar{p}_{eq},\bar{U}_\infty$. It turns out that the model exhibit some results that contradict the intuition. In particular, some games which - {\it a priory}- seems to favor defection most, may produce a relatively high degree of cooperation. Conversely, other games, which one would bet that lead to maximum cooperation, indeed are not the optimal for producing cooperation.Comment: 11 pages, 3 figures, keybords: Complex adaptive systems, Agent-based models, Social system