Density‐Dependent Foraging and Interference Competition by Common Gartersnakes are Temperature Dependent

The ideal free distribution (IFD) predicts that optimal foragers will select foraging patches to maximize food rewards and that groups of foragers should thus be distributed between food patches in proportion to the availability of food in those patches. Because many of the underlying mechanisms of foraging are temperature dependent in ectotherms, the distribution of ectothermic foragers between food patches may similarly depend on temperature because the difference in fitness rewards between these patches may change with temperature. We tested the hypothesis that the distribution of Common Gartersnakes (Thamnophis sirtalis) between food patches can be explained by an IFD, but that conformance to an IFD weakens as temperature departs from the optimal temperature because fitness rewards, interference competition and the number of individuals foraging are highest at the optimal temperature. First, we determined the optimal temperature for foraging. Second, we examined group foraging at three temperatures and three density treatments. Search time was optimized at 27°C, handling time at 29°C and digestion time at 32°C. Gartersnakes did not match an IFD at any temperature, but their distribution did change with temperature: snakes at 20°C and at 30°C selected both food patches equally, while snakes at 25°C selected the low food patch more at low density and the high food patch more at high density. Food consumption and competition increased with temperature, and handling time decreased with temperature. Temperature therefore had a strong impact on foraging, but did not affect the IFD. Future work should examine temperature‐dependent foraging in ectotherms that are known to match an IFD.     

The effect of travel costs on the ideal free distribution in stickleback

The ideal free distribution (IFD) theory, which predicts that a population of individuals will match the distribution of a patchily distributed resource, is widely used in ecology to describe the spatial distribution of animals. While many studies have shown general support of its habitat matching prediction, others have described a systematic pattern of undermatching, where too many animals feed at patches with fewer resources, and too few animals feed in richer patches. These results have been attributed to deviations from several of the assumptions of the IFD. One possible variable, the cost of travelling between patches, has received little attention. Here, we investigated the impact on resource matching when travel costs were manipulated in a simple laboratory experiment involving two continuous input patches. This experiment allowed us to control for extraneous variables and decouple time costs from energetic costs of travel. Two experiments examined the impact of varying travel costs on movement rates between foraging patches and how these travel costs impact conformity to the IFD. Our data demonstrated that there was less movement between patches and greater discrepancies from the IFD predictions as the cost of travel increased.     

Foraging in nature: Contrasting responses to resource heterogeneity at small‐ and large‐spatial scales

A key problem faced by foragers is how to forage when resources are distributed heterogeneously in space. This heterogeneity and associated trade‐offs may change with spatial scale. Furthermore, foragers may also have to optimize acquiring multiple resources. Such complexity of decision‐making while foraging is poorly understood. We studied the butterfly Ypthima huebneri to examine how foraging decisions of adults are influenced by spatial scale and multiple resources. We predicted that, at a small‐spatial scale, the time spent foraging in a patch should be proportional to resources in the patch, but at large‐spatial scales, due to limitations arising from large travel costs, this relationship should turn negative. We also predicted that both adult and larval resources should jointly affect foraging butterflies. To test these predictions, we laid eleven plots and sub‐divided them into patches. We mapped nectar and larval resources and measured butterfly behavior in these patches and plots. We found that adult foraging behavior showed contrasting relationships with adult resource density at small versus large‐spatial scales. At the smaller‐spatial scale, butterflies spent more time feeding in resource‐rich patches, whereas at the large‐scale, butterflies spent more time feeding in resource‐poor plots. Furthermore, both adult and larval resources appeared to affect foraging decisions, suggesting that individuals may optimize search costs for different resources. Overall, our findings suggest that the variation in foraging behavior seen in foragers might result from animals responding to complex ecological conditions, such as resource heterogeneity at multiple spatial scales and the challenges of tracking multiple resources.     

Titrating the cost of plant toxins against predators: determining the tipping point for foraging herbivores

1. Foraging herbivores must deal with plant characteristics that inhibit feeding and they must avoid being eaten. Principally, toxins limit food intake, while predation risk alters how long animals are prepared to harvest resources. Each of these factors strongly affects how herbivores use food patches, and both constraints can pose immediate proximate costs and long-term consequences to fitness. 2. Using a generalist mammalian herbivore, the common brushtail possum (Trichosurus vulpecula), our aim was to quantitatively compare the influence of plant toxin and predation risk on foraging decisions. 3. We performed a titration experiment by offering animals a choice between non-toxic food at a risky patch paired with food with one of five toxin concentrations at a safe patch. This allowed us to identify the tipping point, where the cost of toxin in the safe food patch was equivalent to the perceived predation risk in the alternative patch. 4. At low toxin concentration, animals ate more from the safe than the risky patch. As toxin concentration increased at the safe patch, intake shifted until animals ate mainly from the risky patch. This shift was associated with behavioural changes: animals spent more time and fed longer at the risky patch, while vigilance increased at both risky and safe patches. 5. Our results demonstrate that the variation in toxin concentration, which occurs intraspecifically among plants, can critically influence the relative cost of predation risk on foraging. We show that herbivores quantify, compare and balance these two different but proximate costs, altering their foraging patterns in the process. This has potential ecological and evolutionary implications for the production of plant defence compounds in relation to spatial variation in predation risk to herbivores.     

Foraging behaviour and dispersion of eastern grey kangaroos (Macropus giganteus) in an ideal free framework

Ideal free distribution (IFD) theory predicts that animals in competitive situations should distribute themselves among available habitat patches according to the density of conspecifics and its regulatory effect on resources. To investigate the applicability of IFD models to free-ranging herbivores, we quantified the dispersion and foraging behaviour of eastern grey kangaroos Macropus giganteus among habitat patches of differing suitability, within and outside a reservoir catchment in southern Victoria, Australia. Kangaroo densities within the catchment had a regulatory effect on resource density, while surrounding farmland maintained a higher standing crop despite higher densities of competitors. This difference was slight in autumn, however, when the system was apparently close to equilibrium. Gross bite rates of individuals foraging in farmland were lower than for individuals foraging within the catchment, and vigilance behaviour occurred more frequently in farmland habitat than any other, decreasing time devoted to feeding. Interference competition occurred in only 1.9% of focal samples, although competitive differences based on phenotype were observed. Although resource gains by individual kangaroos are likely to be influenced by other factors, including resource dynamics, predation risk and phenotypic differences, IFD theory provides a valuable analytical framework for this herbivore foraging system.     

The Little Miss Muffet Effect: Quantifying the Effect of Predation Risk on Foraging Patch Choice by Houseflies (<Emphasis Type="Italic">Musca domestica</Emphasis>)

The ability of the ideal free distribution (IFD) to predict patch choice of female houseflies (Musca domestica) was determined by examining their distribution between two patches containing unequal amounts of food. The effect of predation risk was then quantified in energetic terms by examining fly distribution between patches of equal food, with one containing spiders. Results were used to predict how much extra food must be added to the risky patch to offset the risk of predation. Flies were found to conform fairly closely to the IFD. Predation risk had a major effect on their distribution, with fewer flies feeding in the presence of predators as risk increased. Addition of extra food to the risky patch was successful in offsetting the risk of predation. These results suggest that the effect of risk on housefly foraging behavior can be quantified in energy terms, providing a common currency for predicting the effects of resources and predation risk on habitat use.     

Taking the rough with the smooth: foraging for particulate food in continuous time

This paper describes the development of the general dynamical model of foraging developed by Ollason (1980, Theoret. Popul. Biol. 18, 44-65) to predict foraging for particulate food in three different types of environment. In an environment containing particles of different types of food, the model predicts the selection of an approximately optimal diet; in an environment in which the particles occur in patches, the model predicts a time budget of patch occupancy that approximates to the optimal time budget; and in an environment containing patches of particles that regenerate by the addition of particles of food at constant rates, the model predicts that animals will dispose themselves among the patches approximately as predicted by the ideal free distribution. Where the predictions of the model depart from the predictions of optimal foraging theory, they are qualitatively similar to the observed departures of the behaviour of real animals from the predictions of optimal foraging theory. The model provides a general representation of the foraging decisions of animals whether they feed strictly continuously or discontinuously on particles of food, and does so without explicit reference to optimization processes.     

Distribution of a naturally fluctuating ungulate population among heterogeneous plant communities: ideal and free?

1. Herbivore distribution is often assumed to follow the ideal free distribution (IFD) model. This assumes that organisms are omniscient about forage quality and availability within the area available to them and are free to move, with negligible cost, throughout this environment. If this were the case we would expect that, at lowest densities, all animals would be found in the best habitat patches, with less desirable habitats being occupied stepwise as population density increases. We test this using data from a naturally fluctuating population of feral Soay sheep. 2. We show that, although the distribution of individuals is correlated positively with food quality, in line with patterns reported for hill sheep in Scotland, their distribution does not conform to the predictions of the IFD model. We argue that it is the dynamic nature of their food resource that causes this departure from the predictions of the IFD model and make the case that the IFD model, in its unmodified form, is inappropriate for use in modelling distribution among patches containing dynamic resources.     

State-dependent ideal free distributions

Summary The standard ideal free distribution (IFD) states how animals should distribute themselves at a stable competitive equilibrium. The equilibrium is stable because no animal can increase its fitness by changing its location. In applying the IFD to choice between patches of food, fitness has been identified with the net rate of energetic gain. In this paper we assess fitness in terms of survival during a non-reproductive period, where the animal may die as a result of starvation or predation. We find the IFD when there is a large population that can distribute itself between two patches of food. The IFD in this case is state-dependent, so that an animal's choice of patch depends on its energy reserves. Animals switch between patches as their reserves change and so the resulting IFD is a dynamic equilibrium. We look at two cases. In one there is no predation and the patches differ in their variability. In the other, patches differ in their predation risk. In contrast to previous IFDs, it is not necessarily true that anything is equalized over the two patches.     

Can spatial and temporal food variability explain the winter foraging movements of a threatened saltmarsh insectivore?

The white‐fronted chat (Epthianura albifrons) is a small, insectivorous passerine that is threatened with extinction in the north‐eastern part of its range, partially due to loss and degradation of its saltmarsh habitat. Food availability is a potential limiting factor for the disjunct populations that survive in saltmarsh refugia, surrounded by urbanized land, because the anthropogenic matrix reduces the capacity of birds to commute to alternative grassland habitat to exploit temporary insect outbreaks. This limitation is likely to be exacerbated during the winter months when local arthropod abundance in saltmarsh is reduced. This study measured temporal and spatial variation in the abundance of saltmarsh arthropods to determine whether patch switching by foraging flocks can be explained by variation in food availability. Arthropods in the size range known to be important in the diet of white‐fronted chats were vacuum‐sampled from six patches within a continuous area of Sarcocornia‐dominated saltmarsh over a four‐month period. The location of foraging birds was recorded during the same period. Despite superficial similarity in vegetation composition and structure, there was significant variation in arthropod biomass among sites through time, such that the patches with the highest food abundance changed from month to month. There was little evidence, however, to suggest that white‐fronted chats foraged in saltmarsh patches with the highest overall food abundance. During the course of the study, birds were discovered flying 2 km from the saltmarsh to a development site where they foraged in weedy grassland. Arthropod samples collected from this site contained an extremely high abundance of Hemiptera and Neuroptera larvae, supporting previous research indicating that white‐fronted chats forage on irruptions of particular arthropod taxa. These findings indicate that food abundance is unlikely to be the main determinant of foraging site selection within saltmarsh, but highlights the potential importance of alternative foraging habitat types for this species.     

Foraging on spatially distributed resources with sub‐optimal movement,imperfect information,and travelling costs: departures from the ideal free distribution

Ideal free distribution (IFD) theory offers an important baseline for predicting the distribution of foragers across resource patches. Yet it is well known that IFD theory relies on several over‐simplifying assumptions that are unlikely to be met in reality. Here we relax three of the most critical assumptions: (1) optimal foraging moves among patches, (2) omniscience about the utility of resource patches, and (3) cost‐free travelling between patches. Based on these generalizations, we investigate the distributions of a constant number of foragers in models with explicit resource dynamics of logistic type. We find that, first, when foragers do not always move to the patch offering maximum intake rate (optimal foraging), but instead move probabilistically according to differences in resource intake rates between patches (sub‐optimal foraging), the distribution of foragers becomes less skewed than the IFD, so that high‐quality patches attract fewer foragers. Second, this homogenization is strengthened when foragers have less than perfect knowledge about the utility of resource patches. Third, and perhaps most surprisingly, the introduction of travelling costs causes departures in the opposite direction: the distribution of sub‐optimal foragers approaches the IFD as travelling costs increase. We demonstrate that these three findings are robust when considering patches that differ in the resource's carrying capacity or intrinsic growth rate, and when considering simple two‐patch and more complex multiple‐patch models. By overcoming three major over‐simplifications of IFD theory, our analyses contribute to the systematic investigation of ecological factors influencing the spatial distribution of foragers, and thus help in deriving new hypotheses that are testable in empirical systems. A confluence of theoretical and empirical studies that go beyond classical IFD theory is essential for improving insights into how animal distributions across resource patches are determined in nature.     

Foraging patterns of voles at heterogeneous avian and uniform mustelid predation risk

Temporal variation of antipredatory behavior and a uniform distribution of predation risk over refuges and foraging sites may create foraging patterns different from those anticipated from risk in heterogenous habitats. We studied the temporal variation in foraging behavior of voles exposed to uniform mustelid predation risk and heterogeneous avian predation risk of different levels induced by vegetation types in eight outdoor enclosures (0.25 ha). We manipulated mustelid predation risk with weasel presence or absence and avian predation risk by reducing or providing local cover at experimental food patches. Foraging at food patches was monitored by collecting giving-up densities at artificial food patches, overall activity was automatically monitored, and mortality of voles was monitored by live-trapping and radiotracking. Voles depleted the food to lower levels in the sheltered patches than in the exposed ones. In enclosures with higher avian predation risk caused by lower vegetation height, trays were depleted to lower levels. Unexpectedly, voles foraged in more trays and depleted trays to lower levels in the presence of weasels than in the absence. Weasels match their prey's body size and locomotive abilities and therefore increase predation risk uniformly over both foraging sites and refuge sites that can both be entered by the predator. This reduces the costs of missing opportunities other than foraging. Voles changed their foraging strategy accordingly by specializing on the experimental food patches with predictable returns and probably reduced their foraging in the matrix of natural food source with unpredictable returns and high risk to encounter the weasel. Moreover, after 1 day of weasel presence, voles shifted their main foraging activities to avoid the diurnal weasel. This behavior facilitated bird predation, probably by nocturnal owls, and more voles were killed by birds than by weasels. Food patch use of voles in weasel enclosures increased with time. Voles had to balance the previously missed feeding opportunities by progressively concentrating on artificial food patches.     

Ant foraging strategies vary along a natural resource gradient

Food selection by foragers is sensitive to the availability of resources, which may vary along geographical gradients. Hence, selectivity of food types by foragers is expected to track these resource gradients. Here we addressed this hypothesis and asked if foraging decisions of seed-eating ants differ along a geographic gradient of habitat productivity. The study was carried out for two years at five sites along a natural climatic gradient, ranging from arid to Mediterranean, where plant productivity varies six-fold across a short geographic distance of 250 km. We found that in ant colonies of the genus Messor, collective foraging decisions differed along the gradient. Specifically, at the high-productivity sites, a stronger association was found between plant seed availability and selectivity, suggesting that colonies respond more accurately to within-patch variation in food amounts. In contrast, colonies in low-productivity sites foraged in patches with higher concentration of seeds, suggesting that they respond more accurately to among-patch variation in food amounts. Moreover, at the high-productivity sites, colonies were more discriminating in their choice of food and preferred bigger seeds, while in the low-productivity sites, where smaller seeds were relatively more abundant, food collection depended mostly on seed availability. An experiment with artificial seed patches performed along the same climatic gradient, revealed no difference in food selectivity across sites when food type and availability were similar, and a general preference for bigger over medium-sized seeds. Overall, our findings suggest that resource availability is an important factor explaining food choice along a climatic gradient and imply that in low-productivity regions small-seeded species incur high predation pressure, whereas in high-productivity regions, large-seeded species suffer higher predation. This could have important consequences for plant species composition, particularly at the face of climate change, which could dramatically alter the foraging decisions of granivores.     

Indirect Evidence that Flying Foxes Track Food Resources Among Islands in a Pacific Archipelago

Although flying foxes (fruit bats in the genus Pteropus ) in continental forests often fly between scattered resources, little is known about their ranging behavior among islands. The inhospitable water matrix that surrounds the food patches (islands) in archipelagos may prevent flying foxes from tracking resources as efficiently as their counterparts on larger landmasses do. Our aim in this study was to determine whether the abundance of foraging flying foxes ( Pteropus tonganus ) reflected food availability on islands in the Vava'u archipelago of Tonga, regardless of island size and isolation. Overall, food availability was the strongest determinant of flying fox abundance, and spatial aspects of the islands (land area within 10 km) had only a small influence. Food availability appears to regulate flying fox abundance only when food resources are low, but when food sources are plentiful, flying fox abundance may be high or low. These results provide indirect evidence that flying foxes are able to track food resources efficiently in an archipelago, and the water matrix that surrounds the food patches (islands) is not a strong deterrent for foraging animals.     

Foraging by <Emphasis Type="Italic">Drosophila melanogaster</Emphasis> Larvae in a Patchy Environment

The food resources of Drosophila comprise decaying vegetable matter distributed in patches, yet foraging behavior has not been examined in larvae reared continuously in a patchy environment. Here, the extent and rate of inter-patch movement was studied in larvae of four wild strains of D. melanogaster inhabiting an experimental arena from the egg stage to the third larval instar. The hypotheses were that larvae would forage primarily in the third instar, that larvae would move from low-protein patches at higher rates than from high-protein patches, and that foraging rates would be higher on an agar substrate than on sand. Larvae hatching on a nutrient-poor food patch switched to a nutrient-rich patch during the first instar. The rate of interpatch switching increased with larval age, as did the number of larvae roving on the substrate between food patches. Inter-patch distance affected switching speed---the closer the patches, the faster the switching. High protein patches were preferred over low-protein patches, but there was a bias towards staying on the natal patch. Significant variation among strains in latency to forage, in proportion of larvae that switched patches, and in the rate of roving between patches suggests that there is natural genetic variation for these traits. Larvae switched food patches on a substrate of moist sand as quickly as on an agar substrate.     

Using habitat selection theories to predict the spatiotemporal distribution of migratory birds during stopover – a case study of pink‐footed geese Anser brachyrhynchus

Understanding how animals select for habitat and foraging resources therein is a crucial component of basic and applied ecology. The selection process is typically influenced by a variety of environmental conditions including the spatial and temporal variation in the quantity and quality of food resources, predation or disturbance risks, and inter‐ and intraspecific competition. Indeed, some of the most commonly employed ecological theories used to describe how animals choose foraging sites are: nutrient intake maximisation, density‐dependent habitat selection, central‐place foraging, and predation risk effects. Even though these theories are not mutually exclusive, rarely are multiple theoretical models considered concomitantly to assess which theory, or combination thereof, best predicts observed changes in habitat selection over space and time. Here, we tested which of the above theories best‐predicted habitat selection of Svalbard‐breeding pink‐footed geese at their main spring migration stopover site in mid‐Norway by computing a series of resource selection functions (RSFs) and their predictive ability (k‐fold cross validation scores). At this stopover site geese fuel intensively as a preparation for breeding and further migration. We found that the predation risk model and a combination of the density‐dependent and central‐place foraging models best‐predicted habitat selection during stopover as geese selected for larger fields where predation risk is typically lower and selection for foraging sites changed as a function of both distance to the roost site (i.e. central‐place) and changes in local density. In contrast to many other studies, the nutritional value of the available food resources did not appear to be a major limiting factor as geese used different food resources proportional to their availability. Our study shows that in an agricultural landscape where nutritional value of food resources is homogeneously high and resource availability changes rapidly; foraging behaviour of geese is largely a tradeoff between fast refuelling and disturbance/predator avoidance.     

Energy benefits and emergent space use patterns of an empirically parameterized model of memory‐based patch selection

Many species frequently return to previously visited foraging sites. This bias towards familiar areas suggests that remembering information from past experience is beneficial. Such a memory‐based foraging strategy has also been hypothesized to give rise to restricted space use (i.e. a home range). Nonetheless, the benefits of empirically derived memory‐based foraging tactics and the extent to which they give rise to restricted space use patterns are still relatively unknown. Using a combination of stochastic agent‐based simulations and deterministic integro‐difference equations, we developed an adaptive link (based on energy gains as a foraging currency) between memory‐based patch selection and its resulting spatial distribution. We used a memory‐based foraging model developed and parameterized with patch selection data of free‐ranging bison Bison bison in Prince Albert National Park, Canada. Relative to random use of food patches, simulated foragers using both spatial and attribute memory are more efficient, particularly in landscapes with clumped resources. However, a certain amount of random patch use is necessary to avoid frequent returns to relatively poor‐quality patches, or avoid being caught in a relatively poor quality area of the landscape. Notably, in landscapes with clumped resources, simulated foragers that kept a reference point of the quality of recently visited patches, and returned to previously visited patches when local patch quality was poorer than the reference point, experienced higher energy gains compared to random patch use. Furthermore, the model of memory‐based foraging resulted in restricted space use in simulated landscapes and replicated the restricted space use observed in free‐ranging bison reasonably well. Our work demonstrates the adaptive value of spatial and attribute memory in heterogeneous landscapes, and how home ranges can be a byproduct of non‐omniscient foragers using past experience to minimize temporal variation in energy gains.     

Preference in patchy landscapes: the influence of scale-specific intake rates and variance in reward

Understanding the responses of foragers to patchy distributionsof resources has formed a fundamental challenge in behavioralecology. Two currencies have been used to assess the patch preferencesof herbivores—intake rate maximization and risk sensitivity.We wished to understand if small mammalian foragers, collaredlemmings (Dichrostonyx groenlandicus), choose patches to maximizefood intake rate or to reduce risk of starvation in "variable"environments. Moreover, we examined the possibility that maximizingintake rate depends on the spatial scale of patchiness. We designedan experiment offering two alternative patches of food, varyingthe predictability of food rewards and the "potential intakerate" at different spatial scales. Collared lemmings did notconsistently select patches that maximized their intake rateat either scale studied. Instead, they chose patches offeringthe least variation in food reward over the course of the experiment.Collared lemmings used prior knowledge gained from previousforaging bouts to assess food variability. We interpret theseresults as evidence for risk-averse foraging strategies, whichare predicted for continuous foragers aiming to minimize riskof starvation.     

Opportunity costs influence food selection and giving‐up density of dabbling ducks

The interaction of animals with their food can yield insights into habitat characteristics, such as perceived predation risk and relative quality. We deployed experimental foraging patches in wetlands used by migrating dabbling ducks Anas spp. in the central Illinois River Valley to estimate variation in seed removal and giving‐up density (GUD; i.e. density of food remaining in patches following abandonment) with respect to seed density, seed size, seed depth in the substrate, substrate firmness, perceived predation risk, and an energetic profitability threshold (i.e. critical food density). Seed depth and the density of naturally‐occurring seeds outside of experimental plots affected seed removal and GUD in experimental patches more than perceived predation risk, seed density, seed size or substrate firmness. The greatest seed removal and lowest GUDs in experimental patches occurred when food resources in alternative foraging locations outside of plots (i.e. opportunity costs) appeared to be near or below a critical food density (i.e. 119–181 kg ha^–1). Giving‐up densities varied substantially from a critical food density across a range of food densities in alternative foraging locations suggesting that fixed GUDs should not be used as surrogates for critical food densities in energetic carrying capacity models. Foraging and resting rates in and near experimental foraging patches did not reflect patterns of seed removal and were poor predictors of GUD and foraging habitat quality. Our results demonstrated the usefulness of GUDs as indicators of habitat quality for subsurface, benthic foragers relative to other available foraging patches and suggested that food may be limited for dabbling ducks during spring migration in some years in the midwestern USA.     

Spatial memory and foraging competition in captive western lowland gorillas (Gorilla gorilla gorilla)

Spatial memory and foraging competition were investigated in three mother/offspring pairs of western lowland gorillas,Gorilla gorilla gorilla, using a naturalistic foraging task at the Toronto Zoo. Sixteen permanent food sites were placed throughout the animals’ enclosures. All of the sites were baited and a pair of animals was free to visit the sites and collect the food. Five of the subjects collected the food with accuracy better than chance. Most of the subjects visited the sites using a pattern, and for half the subjects this was one of adjacency. The high accuracy of five of the subjects and the lack of a consistent adjacency pattern suggest that the animals did in fact use spatial memory. Furthermore, the gorillas tended to avoid visiting food sites that had been previously depleted by their partner. They also appeared to split their search of the enclosures, each visiting only a proportion of the food sites. This indicated that the animals were competing and altering their foraging behaviour based on the behaviour of their partner. Therefore, accuracy was recalculated to take this into account. When the sites depleted by either animal in a pair during a given trial were worked into the accuracy calculations for individual animals, three of the animals still maintained accuracy above chance. This suggests that the animals were not only able to remember which sites they had depleted, but those sites depleted by their foraging partner as well.