The use of multiplayer game theory in the modeling of biological populations

The use of game theory in modeling the natural world is widespread. However, this modeling mainly involves two player games only, or "playing the field" games where an individual plays against an entire (infinite) population. Game-theoretic models are common in economics as well, but in this case the use of multiplayer games has not been neglected. This article outlines where multiplayer games have been used in evolutionary modeling and the merits and limitations of these games. Finally, we discuss why there has been so little use of multiplayer games in the biological setting and what developments might be useful

Balancing risks and rewards: the logic of violence

Violence is widespread throughout the natural world, prominent examples being predatory violence between species, seasonal violent competition for mating rights and territories within species and food competition both within and between species. These interactions are generally between unrelated individuals with no social connection. There are, however, examples of violent behaviour which occurs within groups of individuals who otherwise cooperate to live, have significant social bonds and may also be related, and that is the primary focus of this paper. Examples are in the establishment and maintenance of dominance hierarchies, or in infanticide, where (usually) incoming males attempt to kill existing infants in a group. Such violence can seem paradoxical, but in fact is often perfectly logical for the individual perpetrating the violence, as distinct from the group as a whole. We discuss such situations from the perspective of evolutionary game theory, and also consider wider questions of interspecific violence

A game theoretical model of kleptoparasitism with incomplete information

Kleptoparasitism, the stealing of food from one animal by another, is a common natural phenomenon that has been modelled mathematically in a number of ways. The handling process of food items can take some time and the value of such items can vary depending upon how much handling an item has received. Furthermore this information may be known to the handler but not the potential challenger, so there is an asymmetry between the information possessed by the two competitors. We use game-theoretic methods to investigate the consequences of this asymmetry for continuously consumed food items, depending upon various natural parameters. A variety of solutions are found, and there are complex situations where three possible solutions can occur for the same set of parameters. It is also possible to have situations which involve members of the population exhibiting different behaviours from each other. We find that the asymmetry of information often appears to favour the challenger, despite the fact that it possesses less information than the challenged individual

A framework for modelling and analysing conspecific brood parasitism

Recently several papers that model parasitic egg-laying by birds in the nests of others of their own species have been published. Whilst these papers are concerned with answering different questions, they approach the problem in a similar way and have a lot of common features. In this paper a framework is developed which unifies these models, in the sense that they all become special cases of a more general model. This is useful for two main reasons; firstly in order to aid clarity, in that the assumptions and conclusions of each of the models are easier to compare. Secondly it provides a base for further similar models to start from. The basic assumptions for this framework are outlined and a method for finding the ESSs of such models is introduced. Some mathematical results for the general, and more specific, models are considered and their implications discussed. In addition we explore the biological consequences of the results that we have obtained and suggest possible questions which could be investigated using models within or very closely related to our framework

Evolutionary games on graphs and the speed of the evolutionary process

In this paper, we investigate evolutionary games with the invasion process updating rules on three simple non-directed graphs: the star, the circle and the complete graph. Here, we present an analytical approach and derive the exact solutions of the stochastic evolutionary game dynamics. We present formulae for the fixation probability and also for the speed of the evolutionary process, namely for the mean time to absorption (either mutant fixation or extinction) and then the mean time to mutant fixation. Through numerical examples, we compare the different impact of the population size and the fitness of each type of individual on the above quantities on the three different structures. We do this comparison in two specific cases. Firstly, we consider the case where mutants have fixed fitness r and resident individuals have fitness 1. Then, we consider the case where the fitness is not constant but depends on games played among the individuals, and we introduce a ‘hawk–dove’ game as an example

A Hawk-Dove game in kleptoparasitic populations

Kleptoparasitism, the parasitism by theft, is a widespread biological phenomenon. In this paper we extend earlier models to investigate a population of conspecifics involved in foraging and, potentially, kleptoparasitism. We assume that the population is composed of two types of individuals, Hawks and Doves. The types differ according to their strategic choices when faced with an opportunity to steal and to resist a challenge. Hawks use every opportunity to steal and they resist all challenges. Doves never resist and never steal. The fitness of each type of individual depends upon various natural parameters, for example food density, the handling time of a food item, density of the population, as well as the duration of potential fights over the food. We find the Evolutionarily Stable States (ESSs) for all arameter combinations and show that there are three possible ESSs, pure Hawks, pure Doves, and a mixed population of Hawks and Doves. We show that for any set of parameter values there is exactly one ESS. We further investigate the relationship between our findings and the classical Hawk-Dove game as defined in Maynard Smith 1982. We also show how our model extends the classical on

Models of kleptoparasitism on networks: the effect of population structure on food stealing behaviour

The behaviour of populations consisting of animals that interact with each other for their survival and reproduction is usually investigated assuming homogeneity amongst the animals. However, real populations are non-homogeneous. We focus on an established model of kleptoparasitism and investigate whether and how much population heterogeneities can affect the behaviour of kleptoparasitic populations. We consider a situation where animals can either discover food items by themselves or attempt to steal the food already discovered by other animals through aggressive interactions. Representing the likely interactions between animals by a network, we develop pairwise and individual-based models to describe heterogeneities in both the population structure and other individual characteristics, including searching and fighting abilities. For each of the models developed we derive analytic solutions at the steady state. The high accuracy of the solutions is shown in various examples of populations with different degrees of heterogeneity. We observe that highly heterogeneous structures can significantly affect the food intake rate and therefore the fitness of animals. In particular, the more highly connected animals engage in more conflicts, and have a reduced food consumption rate compared to poorly connected animals. Further, for equivalent average level of connectedness, the average consumption rate of a population with heterogeneous structure can be higher

Chris Cannings: A Life in Games

Chris Cannings was one of the pioneers of evolutionary game theory. His early work was inspired by the formulations of John Maynard Smith, Geoff Parker and Geoff Price; Chris recognized the need for a strong mathematical foundation both to validate stated results and to give a basis for extensions of the models. He was responsible for fundamental results on matrix games, as well as much of the theory of the important war of attrition game, patterns of evolutionarily stable strategies, multiplayer games and games on networks. In this paper we describe his work, key insights and their influence on research by others in this increasingly important field. Chris made substantial contributions to other areas such as population genetics and segregation analysis, but it was to games that he always returned. This review is written by three of his students from different stages of his career

An evolutionarily stable joining policy for group foragers

For foragers that exploit patchily distributed resources that are challenging to locate, detecting discoveries made by others with a view to joining them and sharing the patch may often be an attractive tactic, and such behavior has been observed across many taxa. If, as will commonly be true, the time taken to join another individual on a patch increases with the distance to that patch, then we would expect foragers to be selective in accepting joining opportunities: preferentially joining nearby discoveries. If competition occurs on patches, then the profitability of joining (and of not joining) will be influenced by the strategies adopted by others. Here we present a series of models designed to illuminate the evolutionarily stable joining strategy. We confirm rigorously the previous suggestion that there should be a critical joining distance, with all joining opportunities within that distance being accepted and all others being declined. Further, we predict that this distance should be unaffected by the total availability of food in the environment, but should increase with decreasing density of other foragers, increasing speed of movement towards joining opportunities, increased difficulty in finding undiscovered food patches, and decreasing speed with which discovered patches can be harvested. We are further able to make predictions as to how fully discovered patches should be exploited before being abandoned as unprofitable, with discovered patches being more heavily exploited when patches are hard to find: patches can be searched for remaining food more quickly, forager density is low, and foragers are relatively slow in traveling to discovered patches