Evidence that tufted puffins Fratercula cirrhata use colony overflights to reduce kleptoparasitism risk

Predation, foraging and mating costs are critical factors shaping life histories. Among colonial seabirds, colony overflights may enhance foraging or mating success, or diminish the risk of predation and kleptoparasitism. The latter possibility is difficult to test because low predation or kleptoparasitism rates could be due either to low danger or to effective counter-tactics by prey. Tufted puffins Fratercula cirrhata breeding at a large colony in British Columbia, Canada, deliver several loads of fish each day to their nestlings and are targets for kleptoparasitism by glaucous-winged gulls Larus glaucescens . In the present study, we documented the ecological conditions under which overflights occurred in order to assess when overflights were made and to statistically isolate the effect of overflights on kleptoparasitism risk at this site. Load-carrying puffins engaged in overflights under ecological conditions associated with relatively high rates of kleptoparasitism. Further, when ecological factors determining risk were statistically controlled, overflights were correlated with marginally lower chances of kleptoparasitism than when the risk factors were ignored. The results suggest that breeding puffins at this site use overflights for kleptoparasite avoidance. This tactic is used sparingly, suggesting it is costly. Costs of overflight behaviour might contribute to the impact of kleptoparasitism on the breeding success of tufted puffins.     

Decision time modulates social foraging success in wild common ravens,Corvus corax

Social foraging provides several benefits for individuals but also bears the potential costs of higher competition. In some species, such competition arises through kleptoparasitism, that is when an animal takes food which was caught or collected by a member of its social group. Except in the context of caching, few studies have investigated how individuals avoid kleptoparasitism, which could be based on physical strength/dominance but also cognitive skills. Here, we investigated the foraging success of wild common ravens, Corvus corax, experiencing high levels of kleptoparasitism from conspecifics when snatching food from the daily feedings of captive wild boars in a game park in the Austrian Alps. Success in keeping the food depended mainly on the individuals’ age class and was positively correlated with the time to make a decision in whether to fly off with food or consume it on site. While the effect of age class suggests that dominant and/or experienced individuals are better in avoiding kleptoparasitism, the effect of decision time indicates that individuals benefit from applying cognition to such decision-making, independently of age class. We discuss our findings in the context of the ecological and social intelligence hypotheses referring to the development of cognitive abilities. We conclude that investigating which factors underline kleptoparasitism avoidance is a promising scenario to test specific predictions derived from these hypotheses.     

Ideal free distributions when individuals differ in competitive ability: phenotype-limited ideal free models

A series of prospective models is developed to investigate ideal free distributions in populations where individuals differ in competitive ability. The models are of three types. In the continuous-input models, there is continuous arrival of food or mates into each habitat patch, and competitors scramble to obtain as large a share as possible. In the interference models, the prey density in a particular patch stays constant but the presence of competitors slows down the rate at which prey are captured. In the kleptoparasitism model, individuals have food or females stolen from them by competitors higher in the dominance hierarchy, and in turn steal items from subordinates. A general result of the continuous-input and interference models is that the population of competitors can be truncated between patches so that the individuals with the highest competitive ability occur in the best patches, or in the patches where competitive differences are greatest. Individuals of lowest competitive ability occur in the poorest patches or where competitive differences are least, and intermediate phenotypes are ranked between these two extremes. Thus the ideal free prediction that all individuals will achieve equal fitness will not apply. However, in continuous-input cases where competitive differences between phenotypes remain constant across patches, this solution is only neutrally stable, and forms only one element of a set of equilibrium distributions. The fact that many empirical studies of continuous-input have found approximately equal mean fitness across patches may relate to this finding. Most interference studies contradict the simple ideal free solution by having different mean intake rates across patches; this may relate to the predicted positive correlation of competitive ability with patch quality. The kleptoparasitism model usually generated continuous cycling of individuals between habitat patches, though some correlation could be found between competitive ability and patch quality.     

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.     

The effect of kleptoparasite and host numbers on the risk of food‐stealing in an avian assemblage

Kleptoparasitism involves the theft of resources such as food items from one individual by another. Such food‐stealing behaviour can have important consequences for birds, in terms of individual fitness and population sizes. In order to understand avian host–kleptoparasite interactions, studies are needed which identify the factors which modulate the risk of kleptoparasitism. In temperate European intertidal areas, Eurasian oystercatchers Haematopus ostralegus feed primarily on bivalve molluscs, which may be stolen by kleptoparasitic species such as carrion crows Corvus corone and herring gulls Larus argentatus. In this study we combined overwinter foraging observations of oystercatchers and their kleptoparasites on the Exe Estuary, UK, with statistical modelling to identify the factors that influence the likelihood of successful food stealing behaviour occurring. Across the winter, 16.4% of oystercatcher foraging attempts ended in successful kleptoparasitism; the risk of theft was lowest in February (10.8%) and highest in December (36.3%). Using an information theoretic approach to compare multiple logistic regression models we present evidence that the outcome of host foraging attempts varied with the number of kleptoparasites per host within the foraging patch for two out of five individual months, and for all months grouped. Successful, kleptoparasitism was more likely to occur when the total number of all kleptoparasites per host was greater. Across the entire winter study period, oystercatcher foraging attempts that resulted in kleptoparasitism were associated with a mean number of kleptoparasites per host that was more than double that for foraging attempts that ended in the oystercatcher successfully consuming the mussel. Conversely, the stage of the tidal cycle within the estuary did not affect the outcome of oystercatcher foraging attempts. Our study provides evidence that bird numbers influence the risk of kleptoparasitism within avian assemblages.     

Stochastic models of kleptoparasitism

In this paper, we consider a model of kleptoparasitism amongst a small group of individuals, where the state of the population is described by the distribution of its individuals over three specific types of behaviour (handling, searching for or fighting over, food). The model used is based upon earlier work which considered an equivalent deterministic model relating to large, effectively infinite, populations. We find explicit equations for the probability of the population being in each state. For any reasonably sized population, the number of possible states, and hence the number of equations, is large. These equations are used to find a set of equations for the means, variances, covariances and higher moments for the number of individuals performing each type of behaviour. Given the fixed population size, there are five moments of order one or two (two means, two variances and a covariance). A normal approximation is used to find a set of equations for these five principal moments. The results of our model are then analysed numerically, with the exact solutions, the normal approximation and the deterministic infinite population model compared. It is found that the original deterministic models approximate the stochastic model well in most situations, but that the normal approximations are better, proving to be good approximations to the exact distribution, which can greatly reduce computing time.     

Evolution of kleptoparasitism as a war of attrition

Previous models of kleptoparasitism (resource stealing) assume that contests over resource items are of fixed duration. Here we suggest that such contests will often be well represented as a war of attrition, with the winner being the individual who is prepared to contest for the longer time. Given that time spent in contests cannot be used to search for other resource items, we provide an analytical expression for the evolutionarily stable distribution of contest times. This can be used to investigate the circumstances under which we would expect kleptoparasitism to evolve. In particular, we focus on situations where searching for conspecifics to kleptoparasitize can only be achieved at a cost of reduced resource discovery by other means; under such circumstances we show that kleptoparasitism is not evolutionarily stable.     

Advantage of storage in a fluctuating environment

We will elaborate the evolutionary course of an ecosystem consisting of a population in a chemostat environment with periodically fluctuating nutrient supply. The organisms that make up the population consist of structural biomass and energy storage compartments. In a constant chemostat environment a species without energy storage always out-competes a species with energy reserves. This hinders evolution of species with storage from those without storage. Using the adaptive dynamics approach for non-equilibrium ecological systems we will show that in a fluctuating environment there are multiple stable evolutionary singular strategies (ss's): one for a species without, and one for a species with energy storage. The evolutionary end-point depends on the initial evolutionary state. We will formulate the invasion fitness in terms of Floquet multipliers for the oscillating non-autonomous system. Bifurcation theory is used to study points where due to evolutionary development by mutational steps, the long-term dynamics of the ecological system changes qualitatively. To that end, at the ecological time scale, the trait value at which invasion of a mutant into a resident population becomes possible can be calculated using numerical bifurcation analysis where the trait is used as the free parameter, because it is just a bifurcation point. In a constant environment there is a unique stable equilibrium for one species following the “competitive exclusion” principle. In contrast, due to the oscillatory dynamics on the ecological time scale two species may coexist. That is, non-equilibrium dynamics enhances biodiversity. However, we will show that this coexistence is not stable on the evolutionary time scale and always one single species survives.     

Evolutionarily stable stealing: game theory applied to kleptoparasitism

We present an individual-based model of a group of foraginganimals. Individuals can obtain food either by discovering itthemselves or by stealing it from others (kleptoparasitism).Given that challenging another individual for a discovered fooditem costs time (which could otherwise be spent searching foran undiscovered item), attempting to steal from another maynot always be efficient We show that there is generally a uniquestrategy that maximizes uptake rate—always or never challengingothers. For any combination of parameter values, we can identifywhich strategy is appropraite. As a corollary to this, we predictthat small changes in ecolgical conditions can, under some circumstances,cause a dramatic change in the aggressive behavior of individuals.Further, we investigate situations where searching for undiscoveredfood and searching for potential opportunities for stealingare mutually exclusive activities (i.e., success at one canonly be improved at the expense of the other). Using game theory,we are able to find the evolutionarily stable strategy for investmentin these two activities in terms of the ecological parametersof the model.     

Do kleptoparasites reduce their own foraging effort in order to detect kleptoparasitic opportunities?: An empirical test of a key assumption of kleptoparasitic models

Determining if, or when, individuals trade off time spent personally feeding against time spent monitoring others for kleptoparasitism opportunities is essential to an understanding of the evolution of scrounging and usurpation behaviours. We provide a first field test of whether kleptoparasites reduce their personal foraging effort in situations where the frequency and rewards of kleptoparasitism increase. We provided experimental food patches for wild European blackbirds that varied in the distribution of prey and that had a potentially high rate of kleptoparasitism within pairs of blackbirds feeding in them. Although individuals differed in their rate of kleptoparasitism, they did not vary in the size of the reward that they gained from kleptoparasitism. As prey became more clumped, kleptoparasitism rate and its reward per incident increased on average. There was, however, no evidence that individuals that were kleptoparasitising more quickly and/or at a higher frequency had lower personal foraging effort. In contrast, foraging effort increased in both birds compared to when they were foraging alone, independent of dominance, kleptoparasitic opportunity or reward. Our evidence suggests that in some circumstances a kleptoparasite can detect kleptoparasitic opportunities without compromising its own personal foraging rate.     

Advantage of storage in a fluctuating environment

We will elaborate the evolutionary course of an ecosystem consisting of a population in a chemostat environment with periodically fluctuating nutrient supply. The organisms that make up the population consist of structural biomass and energy storage compartments. In a constant chemostat environment a species without energy storage always out-competes a species with energy reserves. This hinders evolution of species with storage from those without storage. Using the adaptive dynamics approach for non-equilibrium ecological systems we will show that in a fluctuating environment there are multiple stable evolutionary singular strategies (ss's): one for a species without, and one for a species with energy storage. The evolutionary end-point depends on the initial evolutionary state. We will formulate the invasion fitness in terms of Floquet multipliers for the oscillating non-autonomous system. Bifurcation theory is used to study points where due to evolutionary development by mutational steps, the long-term dynamics of the ecological system changes qualitatively. To that end, at the ecological time scale, the trait value at which invasion of a mutant into a resident population becomes possible can be calculated using numerical bifurcation analysis where the trait is used as the free parameter, because it is just a bifurcation point. In a constant environment there is a unique stable equilibrium for one species following the "competitive exclusion" principle. In contrast, due to the oscillatory dynamics on the ecological time scale two species may coexist. That is, non-equilibrium dynamics enhances biodiversity. However, we will show that this coexistence is not stable on the evolutionary time scale and always one single species survives.     

Prey abundance and the strength of interference in a foraging shorebird

Interference is an important component of food competition but is often difficult to detect and measure in natural animal populations. Although interference has been shown to occur between oystercatchers Haematopus ostralegus L. feeding on mussels Mytilus edulis L., four previous studies have not detected interference between oystercatchers feeding on cockles Cerastoderma edule L. In contrast, this study detected interference between cockle-feeding oystercatchers in the Baie de Somme, France. Prey stealing (kleptoparasitism), one of the main causes of interference between mussel-feeders, also occurred between oystercatchers in the Baie de Somme. The kleptoparasitism rate was related to the natural variation in the food supply, tending to be higher when cockles were rare. Feeding rate was negatively related to competitor density, so providing evidence for interference, but, as in mussel-feeders, only above a threshold density of about 50–100 birds ha⁻¹. The strength of interference at a fixed competitor density was related to the cockle food supply, usually being greater when cockles were rare. Previous studies probably failed to detect interference between cockle-feeders because competitor densities were too low, or cockles were too abundant, or because they were not conducted during late winter when interference is most intense. The study shows that natural variation in the food supply can influence the strength of interference within an animal population and provides support for those behaviour-based interference models which predict that the strength of interference will be greatest when competitor densities are high and prey scarce.     

Floquet theory: a useful tool for understanding nonequilibrium dynamics

Many ecological systems experience periodic variability. Theoretical investigation of population and community dynamics in periodic environments has been hampered by the lack of mathematical tools relative to equilibrium systems. Here, I describe one such mathematical tool that has been rarely used in the ecological literature but has widespread use: Floquet theory. Floquet theory is the study of the stability of linear periodic systems in continuous time. Floquet exponents/multipliers are analogous to the eigenvalues of Jacobian matrices of equilibrium points. In this paper, I describe the general theory, then give examples to illustrate some of its uses: it defines fitness of structured populations, it can be used for invasion criteria in models of competition, and it can test the stability of limit cycle solutions. I also provide computer code to calculate Floquet exponents and multipliers. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

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.     

具有年龄结构的食蚜蝇-蚜虫捕食模型

现有的具有年龄结构的捕食-食饵模型总是假设只有成年捕食者捕食猎物,这与实际情况不符。建立了一个幼年捕食者捕食食饵的具有年龄结构的食蚜蝇-蚜虫模型,应用微分方程定性理论,讨论了系统平衡点及其稳定性:其中平衡点E1(0,0,0)为不稳定的;满足一定条件时,边界平衡点E2(K,0,0)及正平衡点E3(x*,y1*,y2*)为局部渐近稳定的;且应用一致持续生存理论得到了系统永久持续生存的条件,为有害生物综合治理提供了理论依据。     

Stealing of dunked food in Carib grackles (Quiscalus lugubris)

The use of tool or tool-like food processing behaviours can render animals vulnerable to theft (kleptoparasitism) because (1) large, nutritious items are usually involved, (2) value is added to the food due to long and/or complex handling, and (3) physical control of items is often temporarily lost during handling. In Barbados, Carib grackles (Quiscalus lugubris) immersing items in water before consumption (a behaviour known as food dunking) lose a larger proportion of items to conspecific food thieves than grackles that do not dunk. In this paper, we first show that dunking in Carib grackles functions as a proto-tool food-processing technique that speeds up ingestion. We then examine five potential predictors of kleptoparasitism: only conspecific density and loss of physical control on food were found to influence the probability that birds would be attacked and successfully robbed of food by conspecifics. Grackles could reduce the probability of kleptoparasitism by holding items in the bill while dunking and engaging in head-up displays. These behaviours were used flexibly depending on variation in the risk of kleptoparasitism. We suggest that costs like the ones incurred from theft might limit the profitability and frequency of tool and proto-tool food processing behaviours, creating a context where counter-strategies might be selected.     

Kleptoparasitism and complexity in a multi-trophic web

In this work the effects of kleptoparitism in a multi-trophic food-web, in which an omnivorous species scavenges the kills of a top predator, are investigated. Scavengers are assumed to be able to steal predators’ kills by direct interference and aggression. The amount of prey shared depends on the relative competitive abilities of omnivores and predators and on their abundances. To make the proposed model consistent, scavenging is accompanied by other features. First, the predator can prey only on the juvenile life stage of scavengers (i.e. not on adult individuals) as well as on a different species of herbivores. Hence, an age structure is enforced within scavengers, so that they belong to the guild of preys when they are young and vulnerable. Second, predators can switch between prey species selecting the most abundant one. This condition grants the competitive exclusion of the two vegetation eaters, allowing them to co-exist: the omnivore and the herbivore compete for the same vegetation pool but pure herbivores are indeed assumed to be more efficient in its exploitation. The omnivorous species is not able to kill the herbivores, but nonetheless it may exploit their carcasses after they have been killed by predators. Finally, the juvenile/adult ratio of omnivores varies depending on the available resources and predators’ amount. In such a scenario it is shown that kleptoparasitism can modify to a large extent the stability of the system, leading either to a regularization or to chaos, depending both on the scavenger's and the predator's functional response. Moreover, an excess of kleptoparasitism is shown to compromise the whole trophic web, making extinct both predators and scavengers. As far as the authors are aware of, this paper is the first one to explicitly introduce kleptoparasitism in trophic web models.     

Evidence of nest material kleptoparasitism in Worm‐eating Warblers (Helmitheros vermivorum) in east‐central Arkansas,USA

Nest material kleptoparasitism likely evolved in birds to reduce the cost of searching for and collecting material themselves. Although nest material kleptoparasitism has been reported commonly in colonially nesting species, reports for solitary breeding species are infrequent, especially for neotropical migratory species. Here, we report potential and actual nest material kleptoparasitism in the Worm‐eating Warbler (Helmitheros vermivorum). We deployed video camera systems at passerine nests (n = 81) in east‐central Arkansas during summers 2011–2012. In one video, we observed a Worm‐eating Warbler stealing nesting material from a Hooded Warbler (Setophaga citrina) nest. One day later, we later observed a Worm‐eating Warbler landing within 0.5 m of the same warbler nest when the female was incubating, which possibly deterred a second theft of nesting material. In a third video recording, we observed another Worm‐eating Warbler landing within 1 m of an Indigo Bunting (Passerina cyanea) nest. As far as we could determine, neither of these latter two nest visits resulted in nest material kleptoparasitism. Potential benefits of nest material kleptoparasitism include reduced competition for limited nest materials, easy access to suitable material, reduced travel distance, and reduction of nest predation risk; however, costs include risk of attack by host or introducing parasites to one''s nest. Importantly, this behavior could ultimately affect the behavioral and life history evolution of a species. We suggest further work should be conducted to determine the prevalence of nest material kleptoparasitism in Worm‐eating Warblers and other solitary breeding passerines, including efforts to quantify the benefits and costs of this behavior.     

Competitive or weak cooperative stochastic Lotka–Volterra systems conditioned on non-extinction

We are interested in the long time behavior of a two-type density-dependent biological population conditioned on non-extinction, in both cases of competition or weak cooperation between the two species. This population is described by a stochastic Lotka–Volterra system, obtained as limit of renormalized interacting birth and death processes. The weak cooperation assumption allows the system not to blow up. We study the existence and uniqueness of a quasi-stationary distribution, that is convergence to equilibrium conditioned on non-extinction. To this aim we generalize in two-dimensions spectral tools developed for one-dimensional generalized Feller diffusion processes. The existence proof of a quasi-stationary distribution is reduced to the one for a d-dimensional Kolmogorov diffusion process under a symmetry assumption. The symmetry we need is satisfied under a local balance condition relying the ecological rates. A novelty is the outlined relation between the uniqueness of the quasi-stationary distribution and the ultracontractivity of the killed semi-group. By a comparison between the killing rates for the populations of each type and the one of the global population, we show that the quasi-stationary distribution can be either supported by individuals of one (the strongest one) type or supported by individuals of the two types. We thus highlight two different long time behaviors depending on the parameters of the model: either the model exhibits an intermediary time scale for which only one type (the dominant trait) is surviving, or there is a positive probability to have coexistence of the two species.