Is Food Worth Fighting for? ESS’s in Mixed Populations of Kleptoparasites and Foragers

We extend the game theoretic model of kleptoparasitism discussed in Broom et al. (2004), by considering a population of foragers consisting of two groups with different behaviours—those who forage and steal from other feeders, and those who only forage. We a sume that those who do not steal have a better foraging rate than those who are also looking out for opportunities to steal. We also allow either type to resist an attack or not resist. We look for Evolutionary Stable States, of either a mixture of the two behaviours, or where the whole population has just one of these behaviours. We find nine such ESS’s, dependent on the environmental parameters, although in fact only five of these are distinguishable. In general, we find that if the overall population density is low, food-stealing becomes less viable, and there is an ESS consisting of only foragers. Conversely, when there are many animals looking for, and finding, food, there is an ESS consisting of just kleptoparasites (which are also foraging). In between, an ESS will contain both pure-foragers and stealers. There is some empirical evidence of such behaviours. We find that when there is a mixture of the two types, they must both have the same resistive behaviour. We can thus have some individuals challenging for food but not resisting challenges, and others not challenging and not resisting. This shows how aggressive behaviour may be context-dependent, as seen in practice.     

Rapid convergence to an equilibrium state in kleptoparasitic populations

Previous papers have modelled the behaviour of populations which are subject to kleptoparasitism, and found those ecological situations in which kleptoparasitism should occur. Individuals were considered to be in one of several states, and an equilibrium distribution for the population was found. It was then assumed, for analytical purposes but without proof, that the population was actually in that equilibrium. In this paper, we show that the equilibrium is a stable one, and that it is reached in a relatively short time for all reasonable values of the ecological parameters. Thus, a population may be expected to spend most of the time in equilibrium, and this assumption of these previous works is justified.Research of the first author supported by EPSRC.The authors are also members of The Centre for the Study of Evolution, at the University of Sussex.     

Initial predation and parasitism by muricid whelks demonstrated by the correspondence between drilled holes and their apparent enveloper

The morphologies of drilled holes enveloped by three coexisting muricid whelks, Morula musiva, Cronia margariticola and Ergalatax contractus, were compared. The results were used to demonstrate the replacement of feeders using drilled holes, and to estimate the frequency of this occurrence in the field. In the laboratory, M. musiva made circular internal holes, whereas C. margariticola and E. contractus made oval holes, when they preyed upon the mussel Hormomya mutabilis. The maximum inner diameter of the hole correlated with the driller's shell height in all three muricid species. In the field, mussels enveloped by M. musiva had circular drilled holes, as was found in the laboratory, and the correlation between maximum diameter and enveloper's shell height was not significantly different from that found in the laboratory. In contrast, holes enveloped by C. margariticola in the field were more circular than those observed in the laboratory, and there was no correlation between maximum diameter and enveloper's shell height. Statistical analyses of these results indicated that more than half of the mussels being ingested by C. margariticola had been drilled by M. musiva, i.e. C. margariticola would often kleptoparasitize or scavenge M. musiva's prey by taking over the drilled hole. Few M. musiva would take over the holes drilled by C. margariticola or E. contractus. Furthermore, probably few M. musiva would take over the hole made by a conspecific initial drilling predator. It appears that E. contractus rarely initiates predation by drilling.     

黑眉苇莺存在对东方大苇莺巢材的盗窃行为

筑巢是大多数鸟类繁殖成功的必要条件,筑巢花费精力较少的鸟类可以将更多的时间投入到后期繁殖中。因此,对于筑巢繁殖的鸟类来说,如何快速有效地建筑一个巢尤为重要。然而,对雀形目鸟类快速和投机筑巢的行为报道较少。2021年6至8月,在黑龙江扎龙国家级自然保护区,通过录像对繁殖期东方大苇莺（Acrocephalus orientalis）和黑眉苇莺（A. bistrigiceps）的筑巢行为进行研究,记录到黑眉苇莺盗取东方大苇莺巢材料的行为。根据视频内容和现场观察,本研究初步描述了黑眉苇莺偷窃行为的全过程。最后,通过文献查阅,对报道过巢材盗窃行为的部分雀形目鸟类进行了统计。本文研究结果为进一步确定雀形目鸟类的巢材盗窃行为的流行程度提供了基础信息。     

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.     

The Evolution of a Kleptoparasitic System under Adaptive Dynamics

Kleptoparasitism, the stealing of food items, is a common biological phenomenon which has been modelled mathematically in a series of recent papers. A common assumption, following early work, was that mixed strategy solutions were not possible. In this paper we consider the evolution of mixed strategies under adaptive dynamics and show that such mixed strategies can be stable solutions under certain assumptions. In particular we revisit the recent paper of Broom et al. (Bull math Biol 66, 1645–1658, 2004) which assumed pure solutions only, and reanalyze the model under this new formulation. The second author was supported by the UNCG New Faculty Grant.     

Habitat utilisation, feeding tactics and age related feeding efficiency in the Caspian Gull Larus cachinnans

The feeding behaviour of the Caspian Gull Larus cachinnans was analysed in southern Poland in 2001. During the pre-breeding period, most birds foraged on a refuse dump and some foraged in a river valley. During incubation, similar numbers of birds foraged on fishponds, gravel pits and the refuse dump. During the chick-rearing period, fishponds were the most important foraging grounds. The foraging success of three main foraging tactics was analysed: digging on refuse, fishing and kleptoparasitism. We found that digging success was higher in juveniles than in immature or adult birds. However, older birds moved and ate more items per unit of time than juveniles, which indicates that older birds improved their energy gain simply by a higher speed of searching. The opposite was found for fishing success. As juvenile birds made fewer attempts than immature or adult birds, fishing success was higher in adults. Adults and immature birds interrupted more attacks than juveniles, which indicates that older birds were better able to assess the probability of fish catching than juveniles. Kleptoparasitism was observed almost exclusively on the refuse dump during the pre-breeding period. Young birds kleptoparasitised more frequently than adults, but they had a lower rate of success. However, the lower success in young birds was due to victim choice, rather than differences in flight skills. Young birds kleptoparasitised Black-headed Gulls Larus ridibundus and Jackdaws Corvus monedula more frequently than adults, but none of the attacks towards these species was successful. Generally, Caspian Gulls kleptoparasitised conspecifics more often than expected from species frequency. Only attacks towards conspecifics yielded any success.