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

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

Facilitators on networks reveal optimal interplay between information exchange and reciprocity

Reciprocity is firmly established as an important mechanism that promotes cooperation. An efficient information exchange is likewise important, especially on structured populations, where interactions between players are limited. Motivated by these two facts, we explore the role of facilitators in social dilemmas on networks. Facilitators are here mirrors to their neighbors—they cooperate with cooperators and defect with defectors—but they do not participate in the exchange of strategies. As such, in addition to introducing direct reciprocity, they also obstruct information exchange. In well-mixed populations, facilitators favor the replacement and invasion of defection by cooperation as long as their number exceeds a critical value. In structured populations, on the other hand, there exists a delicate balance between the benefits of reciprocity and the deterioration of information exchange. Extensive Monte Carlo simulations of social dilemmas on various interaction networks reveal that there exists an optimal interplay between reciprocity and information exchange, which sets in only when a small number of facilitators occupy the main hubs of the scale-free network. The drawbacks of missing cooperative hubs are more than compensated for by reciprocity and, at the same time, the compromised information exchange is routed via the auxiliary hubs with only marginal losses in effectivity. These results indicate that it is not always optimal for the main hubs to become leaders of the masses, but rather to exploit their highly connected state to promote tit-for-tat-like behavior

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

Overestimating Resource Value and Its Effects on Fighting Decisions

Much work in behavioral ecology has shown that animals fight over resources such as food, and that they make strategic decisions about when to engage in such fights. Here, we examine the evolution of one, heretofore unexamined, component of that strategic decision about whether to fight for a resource. We present the results of a computer simulation that examined the evolution of over- or underestimating the value of a resource (food) as a function of an individual's current hunger level. In our model, animals fought for food when they perceived their current food level to be below the mean for the environment. We considered seven strategies for estimating food value: 1) always underestimate food value, 2) always overestimate food value, 3) never over- or underestimate food value, 4) overestimate food value when hungry, 5) underestimate food value when hungry, 6) overestimate food value when relatively satiated, and 7) underestimate food value when relatively satiated. We first competed all seven strategies against each other when they began at approximately equal frequencies. In such a competition, two strategies–“always overestimate food value,” and “overestimate food value when hungry”–were very successful. We next competed each of these strategies against the default strategy of “never over- or underestimate,” when the default strategy was set at 99% of the population. Again, the strategies of “always overestimate food value” and “overestimate food value when hungry” fared well. Our results suggest that overestimating food value when deciding whether to fight should be favored by natural selection

Rich Pickings Near Large Communal Roosts Favor ‘Gang’ Foraging by Juvenile Common Ravens, Corvus corax

Ravens (Corvus corax) feed primarily on rich but ephemeral carcasses of large animals, which are usually defended by territorial pairs of adults. Non-breeding juveniles forage socially and aggregate in communal winter roosts, and these appear to function as ‘information centers’ regarding the location of the rare food bonanzas: individuals search independently of one another and pool their effort by recruiting each other at roosts. However, at a large raven roost in Newborough on Anglesey, North Wales, some juveniles have been observed recently to forage in ‘gangs’ and to roost separately from other birds. Here we adapt a general model of juvenile common raven foraging behavior where, in addition to the typical co-operative foraging strategy, such gang foraging behavior could be evolutionarily stable near winter raven roosts. We refocus the model on the conditions under which this newly documented, yet theoretically anticipated, gang-based foraging has been observed. In the process, we show formally how the trade off between search efficiency and social opportunity can account for the existence of the alternative social foraging tactics that have been observed in this species. This work serves to highlight a number of fruitful avenues for future research, both from a theoretical and empirical perspective

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

Biology and evolutionary games

This chapter surveys some evolutionary games used in biological sciences. These include the Hawk-Dove game, the Prisoner’s Dilemma, Rock–Paper–Scissors, the war of attrition, the Habitat Selection game, predatorprey games, and signalling games