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

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

Modeling scan and interscan durations in antipredator vigilance

Many prey species alternate between bouts of foraging and bouts of antipredator vigilance. Models of vigilance typically predict how much total time prey animals should allocate to vigilance but do not specify how that time should be scheduled throughout foraging. Here, we examine how the scheduling of vigilance pays off in terms of food intake and predator detection. Specifically, we study how changes in ecological factors affect the expected duration of scans to look out for predators and the duration of interscan intervals dedicated to foraging. Our framework includes factors like the risk of attack, how difficult it is to locate food and predators, and the distance to protective cover. Our individual-based model makes several predictions about scan and interscan durations, which are discussed in relation to the available empirical evidence in birds and mammals. This model of antipredator vigilance is a first step in incorporating constraints related to food gathering and the detection of predators. Adding such constraints adds a novel dimension to vigilance models and produces a variety of predictions that await empirical scrutiny. (C) 2015 Elsevier Ltd. All rights reserved.PostprintPeer reviewe

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

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

Developing a methodology for social network sampling (abstract)

Researchers are increasingly turning to network theory to describe and understand the social nature of animal populations. To make use of the statistical tools of network theory, ecologists need to gather relational data, typically by sampling the social relations of a population of animals over a given time-period. Due to effort constraints and the practical difficulty involved in tracking animals, these sampled relational data are almost always a subset of the actual network. Measurements of the sample – such as average path length, clustering, and assortativity – are assumed to be informative as to the structure of the real-world social network. However, this assumption is problematic. Due to artefacts of the sampling process, the various network measures taken on the sample may be biased estimators of the true values. For example, just as we would get a biased estimate of mean human height by selecting for a sample those people who stood out in a crowd, we will get a biased estimate of a measure like mean connectivity if we sample individuals who are socially prominent. This problem can only be solved by developing a qualitative theory of network sampling, answering questions such as what proportion of the whole network needs to be sampled before a given level of accuracy is achieved, and what sampling procedures are least biased? To develop such a theory, we need to be able to generate networks from which to sample. Ideally, we need to perform a systematic study of sampling protocols on different known network structures. But currently available data on animal social networks are unsuitable as these networks were themselves sampled. The simulation methods of artificial life provide the way forward. We have developed a computational tool for generating artificial social networks that have user-defined distributions for network properties (such as the number of nodes, and the density) and for key the measures of interest to ecologists (such as the average degree, average path length, clustering, betweenness, and assortativity). This tool allows us to perform the required systematic analyses of the biases inherent in different sampling regimes (e.g., snowball sampling) applied to different network structures. We will present details of this system, and show we are using it to develop robust sampling methods for social network data. We see the system as the first in a series of works that will allow us to develop a qualitative theory of social network sampling to aid ecologists, and eventually social scientists, in their social network data collection

Temporally fluctuating prey and coexistence among unequal conspecific interferers

Coexistence among unequal conspecific interferers should be unlikely to persist if stronger interferers always experience a relative fitness increment from their higher foraging rates. In this study, we suggest that decreased relative costs to weaker interferers with increasing temporal fluctuations in prey availability may be a mechanism enhancing coexistence of unequal conspecific interferers. Previous work on fluctuation and coexistence has dealt with oscillations over a time-scale measured in generations of competitor species and their resources, while our work shows that fluctuations in prey availability facilitate coexistence of different phenotypic strategies within species and generations, and over short time-scales. With increasing amplitude of temporal fluctuation about an average prey density, cumulative intakes for differently strong interferers are affected differently. Because of the prey-dependent effect of interference, high amplitudes of fluctuation allow for relatively lower foraging-rate costs in weaker interferers, which decreases the difference in foraging success between strong and weaker interferers. This decreased difference in foraging success could thus significantly relax the conditions allowing for unequal interferer coexistence

The impact of detoxification costs and predation risk on foraging : implications for mimicry dynamics

This work was supported by the European Research Council (Advanced Grant 250209 to Alasdair Houston), a Natural Environment Research Council Independent Research Fellowship (NE/L011921/1) awarded to A.D.H., a BBSRC-NERC project grant (BB/G00188X/1) awarded to J.S., C.R. and G.D.R. and a faculty fellowship awarded to C.G.H. (Medical Sciences, Newcastle University) with strategic support funding from the Wellcome Trust.Prey often evolve defences to deter predators, such as noxious chemicals including toxins. Toxic species often advertise their defence to potential predators by distinctive sensory signals. Predators learn to associate toxicity with the signals of these so-called aposematic prey, and may avoid them in future. In turn, this selects for mildly toxic prey to mimic the appearance of more toxic prey. Empirical evidence shows that mimicry could be either beneficial (‘Mullerian’) or detrimental (‘quasi-Batesian’) to the highly toxic prey, but the factors determining which are unknown. Here, we use state-dependent models to explore how tri-trophic interactions could influence the evolution of prey defences. We consider how predation risk affects predators’ optimal foraging strategies on aposematic prey, and explore the resultant impact this has on mimicry dynamics between unequally defended species. In addition, we also investigate how the potential energetic cost of metabolising a toxin can alter the benefits to eating toxic prey and thus impact on predators’ foraging decisions. Our model predicts that both how predators perceive their own predation risk, and the cost of detoxification, can have significant, sometimes counterintuitive, effects on the foraging decisions of predators. For example, in some conditions predators should: (i) avoid prey they know to be undefended, (ii) eat more mildly toxic prey as detoxification costs increase, (iii) increase their intake of highly toxic prey as the abundance of undefended prey increases. These effects mean that the relationship between a mimic and its model can qualitatively depend on the density of alternative prey and the cost of metabolising toxins. In addition, these effects are mediated by the predators’ own predation risk, which demonstrates that, higher trophic levels than previously considered can have fundamental impacts on interactions among aposematic prey species.Publisher PDFPeer reviewe