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

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

The effects of land transport on animal welfare

Animal welfare during and as a result of transport can be assessed by using a range of behavioural, physiological, pathological and carcass-quality indicators that are described in this paper. Measures of the extent of any disease, injury or mortality resulting from, or exacerbated by, transport are important because health is an important part of welfare. Many of the indicators are measures of stress as they involve long-term adverse effects on the individual. Factors affecting the welfare of animals before, during and after transport which are discussed are: definition of the responsibilities and competence, attitudes to animals and need for training of staff; planning of journeys and methods of payment of staff; laws and retailers’ codes; genetics, especially selection for high productivity; rearing conditions and experience; the mixing of animals from different social groups; handling and loading procedures; driving methods; space allowance; increased susceptibility to disease and efforts to minimise the spread of disease

A game-theoretic model of kleptoparasitic behavior in polymorphic populations

Kleptoparasitism, the stealing of food by one animal from another, is a widespread biological phenomenon. In this paper we build upon earlier models to investigate a population of conspecifics involved in foraging and, potentially, kleptoparasitism. We assume that the population is composed of four types of individuals, according to their strategic choices when faced with an opportunity to steal and to resist an attack. The fitness of each type of individual depends upon various natural parameters, for example food density, the handling time of a food item and the probability of mounting a successful attack against resistance, as well as the choices that they make. We find the evolutionarily stable strategies (ESSs) for all parameter combinations and show that there are six possible ESSs, four pure and two mixtures of two strategies, that can occur. We show that there is always at least one ESS, and sometimes two or three. We further investigate the influence of the different parameters on when each type of solution occurs

Kleptoparasitic melees--modelling food stealing featuring contests with multiple individuals

Kleptoparasitism is the stealing of food by one animal from another. This has been modelled in various ways before, but all previous models have only allowed contests between two individuals. We investigate a model of kleptoparasitism where individuals are allowed to fight in groups of more than two, as often occurs in real populations. We find the equilibrium distribution of the population amongst various behavioural states, conditional upon the strategies played and environmental parameters, and then find evolutionarily stable challenging strategies. We find that there is always at least one ESS, but sometimes there are two or more, and discuss the circumstances when particular ESSs occur, and when there are likely to be multiple ESSs

Evolutionarily stable sexual allocation by both stressed and unstressed potentially simultaneous hermaphrodites within the same population.

Factors influencing allocation of resources to male and female offspring continue to be of great interest to evolutionary biologists. A simultaneous hermaphrodite is capable of functioning in both male and female mode at the same time, and such a life-history strategy is adopted by most flowering plants and by many sessile aquatic animals. In this paper, we focus on hermaphrodites that nourish post-zygotic stages, e.g. flowering plants and internally fertilising invertebrates, and consider how their sex allocation should respond to an environmental stress that reduces prospects of survival but does not affect all individuals equally, rather acting only on a subset of the population. Whereas dissemination of pollen and sperm can begin at sexual maturation, release of seeds and larvae is delayed by embryonic development. We find that the evolutionarily stable strategy for allocation between male and female functions will be critically dependent on the effect of stress on the trade-off between the costs of male and female reproduction, (i.e. of sperm and embryos). Thus, we identify evaluation of this factor as an important challenge to empiricists interested in the effects of stress on sex allocation. When only a small fraction of the population is stressed, we predict that stressed individuals will allocate their resources entirely to male function and unstressed individuals will increase their allocation to female function. Conversely, when the fraction of stress-affected individuals is high, stressed individuals should respond to this stressor by increasing investment in sperm and unstressed individuals should invest solely in embryos. A further prediction of the model is that we would not expect to find populations in the natural world where both stressed and unstressed individuals are both hermaphrodite

A theory for investment across defences triggered at different stages of a predator-prey encounter

We introduce a general theoretical description of a combination of defences acting sequentially at different stages in the predatory sequence in order to make predictions about how animal prey should best allocate investment across different defensive stages. We predict that defensive investment will often be concentrated at stages early in the interaction between a predator individual and the prey (especially if investment is concentrated in only one defence, then it will be in the first defence). Key to making this prediction is the assumption that there is a cost to a prey when it has a defence tested by an enemy, for example because this incurs costs of deployment or tested costs as a defence is exposed to the enemies; and the assumption that the investment functions are the same among defences. But if investment functions are different across defences (e.g. the investment efficiency in making resources into defences is higher in later defences than in earlier defences), then the contrary could happen. The framework we propose can be applied to other victim-exploiter systems, such as insect herbivores feeding on plant tissues. This leads us to propose a novel explanation for the observation that herbivory damage is often not well explained by variation in concentrations of toxic plant secondary metabolites. We compare our general theoretical structure with related examples in the literature, and conclude that coevolutionary approaches will be profitable in future work

Signal verification can promote reliable signalling.

The central question in communication theory is whether communication is reliable, and if so, which mechanisms select for reliability. The primary approach in the past has been to attribute reliability to strategic costs associated with signalling as predicted by the handicap principle. Yet, reliability can arise through other mechanisms, such as signal verification; but the theoretical understanding of such mechanisms has received relatively little attention. Here, we model whether verification can lead to reliability in repeated interactions that typically characterize mutualisms. Specifically, we model whether fruit consumers that discriminate among poor- and good-quality fruits within a population can select for reliable fruit signals. In our model, plants either signal or they do not; costs associated with signalling are fixed and independent of plant quality. We find parameter combinations where discriminating fruit consumers can select for signal reliability by abandoning unprofitable plants more quickly. This self-serving behaviour imposes costs upon plants as a by-product, rendering it unprofitable for unrewarding plants to signal. Thus, strategic costs to signalling are not a prerequisite for reliable communication. We expect verification to more generally explain signal reliability in repeated consumer-resource interactions that typify mutualisms but also in antagonistic interactions such as mimicry and aposematism

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