Does group foraging promote efficient exploitation of resources?

Increased avoidance of food patches previously exploited by other companions has been proposed as one adaptive benefit of group foraging. However, does group foraging really represent the most efficient way to exploit non- or slowly-renewing resources? Here, I used simulations to explore the costs and benefits of exploiting non-renewing resources by foragers searching for food patches independently or in groups in habitats with different types of resource distribution. Group foragers exploited resources in a patch more quickly and therefore spent proportionately more time locating new patches. Reduced avoidance of areas already exploited by others failed to overcome the increased time cost of searching for new food patches and group foragers thus obtained food at a lower rate than solitary foragers. Group foraging provided one advantage in terms of a reduction in the variance of food intake rate. On its own, reduced avoidance of exploitation competition through group foraging appears unlikely to increase mean food intake rate when exploiting non-renewing patches but may provide a way to reduce the risk of an energy shortfall.     

Long-Term Habitat Use by Mountain Gorillas (Gorilla gorilla beringei). 2. Reuse of Foraging Areas in Relation to Resource Abundance, Quality, and Depletion

Resource depression caused by current feeding and the rate of resource renewal should influence foragers' decisions about when to revisit foraging areas. Adjustment of foraging paths and revisit rates should be particularly important when resources renew slowly. Foragers can also benefit by returning more often to highly profitable than to less profitable foraging areas. Many highly frugivorous primates seem to time revisits to fruit sources so as to harvest fruit efficiently and, also, use efficient search paths. Fewer data on non-frugivores exist. Mountain gorillas are folivores that eat mostly perennially available, continuously growing herbs and vines. Vegetation regenerates slowly from the effects of gorilla trampling, though trampling can also facilitate food species productivity, at least in the short term. Adjustment of intervals between visits to foraging areas to the extent of previous use and to resource renewal rates should increase gorilla foraging efficiency. Long-term data on habitat use by 6 mountain gorilla social units show that revisit intervals vary in association with variation in the extent of previous use and in plant productivity. However, they also revisit areas more often, the higher the biomass and nutritional quality of food there. These data are generally consistent with the hypothesis that the gorillas crop resources on a sustained-yield basis, though more precise data on areal revisits and complementary long-term data on vegetation composition would be needed to test the hypothesis.     

The group-size paradox: effects of learning and patch departure rules

In many species, foraging in groups can enhance individual fitness.However, groups are often predicted to be larger than the sizethat maximizes individual fitness. This is because individualforagers are expected to continue joining a group until thefitness in the group falls to the level experienced by solitaryforagers. If such a process were pervasive, social foraging,paradoxically, would provide little evolutionary advantages.We propose a solution to the group-size paradox by allowingforagers to learn about habitat quality and leave food patcheswhen their current intake rate falls below that expected forthe whole habitat. By using a simulation model, we show thatunder a wide range of population sizes, foragers using suchrules abandon under- and overcrowded patches, ensuring thatgroup size remains close to the optimal value. The results holdin habitats with varying patch quality, but we note that thelack of food renewal in patches can disrupt the process of groupformation. We conclude that groups of optimal sizes can occurfrequently if fitness functions are peaked and resources patchilydistributed, without the need to invoke relatedness betweenjoiners and established group members, group defense againstjoiners, or other mechanisms that were proposed earlier to preventgroups from becoming too large.     

Efficient harvesting of renewing resources

Many foraging animals return to feeding sites to harvest replenishingresources, but little is known about efficient tactics for doingthis. Can animals with adequate cognitive abilities increasetheir efficiency by modifying their behavior according to memoriesof past experience at particular sites? We developed a simulationmodel of animals harvesting renewable resources from isolatedpatches in undefended, competitive situations. We compared fourforaging tactics: (1) moving stochastically without using anyinformation from past experiences (random searching); (2) movingstochastically, but going longer distances after encounteringlower reward (area-restricted searching); (3) repeatedly movingalong a fixed route (complete traplining); and (4) traplining,but sampling and shifting to neighboring rewarding patches afterencountering low reward (sample-and-shift traplining). FollowingPossingham, we tracked both the resources actually harvestedby a focal forager (i.e., rewards) and the standing crops ofresources that accumulated at patches. Complete traplining alwaysproduces less variation in elapsed time between visits thanrandom searching or area-restricted searching, which has threebenefits: increasing the reward crop harvested, if resourcerenews nonlinearly; reducing resource standing crop in patches;and reducing variation in reward crop per patch. Moreover, thesystematic revisitation schedule produced by complete trapliningmakes it more competitive, regardless of resource renewal scheduleor competitor frequency. By responding to their past experiences,using sample-and-shift traplining, foragers benefit only whenmany patches are left unvisited in the habitat. Otherwise, theexploratory component of sample-and-shift traplining, whichincreases the movement distance and the variation in elapsedtime between visits, makes it more costly than complete traplining.Thus, traplining will usually be beneficial, but foragers shouldswitch between "impatient" (sample-and-shift traplining) and"tenacious" (complete traplining) traplining, according to temporalchanges in surrounding situations.     

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.     

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.     

Effect of Group Size on Feeding Rate when Patches are Exhaustible

One benefit of group foraging is that individual foragers can join the food discoveries of companions and thus increase encounter rate with food patches. When food patches are exhaustible, however, individual shares of each patch will decrease with group size negating the effect of increased encounter rate. Mean feeding rate may actually decrease with group size as a result of aggression or time wasted joining already depleted patches, or when searching to join the food discoveries of others, which is referred to as scrounging, precludes finding food. I examined the relationship between mean feeding rate and group size in captive flocks of zebra finches (Taenopygia guttata) foraging for small clumps of seeds. Finches in groups of two or four fared better than solitary birds in terms of mean feeding rate despite the fact that birds in groups scrounged a large proportion of their food. Solitary birds initiated feeding activity after a longer delay, which led to their lower success. Early departures by food finders from food patches joined by others may have lessened the impact of scrounging on mean feeding rate. As a result of benefits from the presence of companions, group foraging in zebra finches appears a viable alternative to foraging alone despite the cost of sharing resources.     

Scale-dependent foraging and patch choice in fractal environments

Many spatially complex environments are fractal, and consumers in these environments face scale-dependent trade-offs between encountering high densities of small resource patches versus low densities of large resource patches. I address the effects of these trade-offs on foraging by incorporating scale-dependent encounter of resources in fractal landscapes into classical optimal foraging theory. This model is then used to predict optimal scales of perception (foraging scale) and patch choice in response to spatial features of landscapes. The model predicts that, for a given density of resources, landscapes with greater extent and fractal dimension and that contain patchy (low fractal dimension) resources favour large foraging scales and specialization on a small proportion of resource patches. Fragmented (low fractal dimension) landscapes of small extent with dispersed (high fractal dimension) resources favour smaller foraging scales and generalists that use a large proportion of available resource patches. These predictions synthesize the results of other spatially explicit consumer–resource models into a simple framework and agree reasonably well with results of several empirical studies. This study thus places optimal foraging theory in a spatial context and suggests evolutionary mechanisms of consumers' responses to important spatial phenomena (e.g. habitat fragmentation, resource aggregation). This revised version was published online in July 2006 with corrections to the Cover Date.     

A field assessment of optimal foraging in ants: trail patterns and seed retrieval by the European harvester ant Messor barbarus

Summary: The ant Messor barbarus is a major seed predator on annual grasslands of the Mediterranean area. This paper is an attempt to relate the foraging ecology of this species to resource availability and to address several predictions of optimal foraging theory under natural conditions of seed harvesting.¶Spatial patterns of foraging trails tended to maximise acquisition of food resources, as trails led the ants to areas where seeds were more abundant locally. Moreover, harvesting activity concentrated on highly frequented trails, on which seeds were brought into the nest in larger numbers and more efficiently, at a higher mean rate per worker.¶The predictions of optimal foraging theory that ants should be more selective in both more resource-rich and more distant patches were tested in the native seed background. We confirm that selectivity of ants is positively related to trail length and thus to distance from the nest of foraged seeds. Conversely, we fail to find a consistent relationship between selectivity and density or species diversity of seed patches. We discuss how selectivity assessed at the colony level may depend on factors other than hitherto reported behavioural changes in seed choice by individual foragers.     

The use of time and space by male and female gerbils exploiting a pulsed resource

Foraging theory postulates that interference is a foraging cost and affects patch exploitation and activity times. One such system contains two species of seed-eating gerbils inhabiting sandy habitats in the Negev Desert of Israel. Low population densities of the dominant species allowed us to examine the interaction between males and females of the subordinate species, Gerbillus andersoni allenbyi , as a function of interference and resource renewal. We used giving-up densities (GUDs; the amount of food left in a resource patch when a forager abandons the patch) in seed trays to quantify patch use by gerbils. By placing 6 trays at each foraging station and either presenting all 6 trays at the start of the night (pulse treatment) or presenting one tray at a station 6 times per night (renewal treatment), we were able to manipulate characteristics of resource renewal. We used radio telemetry to obtain an independent assessment of activity. Male and female G. a. allenbyi differed in their timing of activity, with males beginning earlier than females and remaining active later. This was most pronounced for the pulse treatment. For the renewal treatment, female activity in trays was more intense early in the night, but thereafter male activity was more intense. At the same time, telemetry showed that males and females did not differ in their total activity in or out of trays. This suggests that males begin their activity on the renewal treatment by exploiting the richest natural patches of seeds. Only later when these are depleted do they move to dominate the renewing seed trays. Finally, females exploited stabilized sand habitats more than did males, especially during the renewal treatment. Taken together, these findings suggest that male G. a. allenbyi interfere with foraging in females, causing temporal shifts in their use of space and resources.     

Exploration or exploitation: life expectancy changes the value of learning in foraging strategies

The acquisition of information is a fundamental part of individual foraging behaviour in heterogeneous and changing environments. We examine how foragers may benefit from utilizing a simple learning rule to update estimates of temporal changes in resource levels. In the model, initial expectation of resource conditions and rate of replacing past information by new experiences are genetically inherited traits. Patch-time allocation differs between learners and foragers that use a fixed patch-leaving threshold throughout the foraging season. It also deviates from foragers that obtain information about the environment at no cost. At the start of a foraging season, learners sample the environment by frequent movements between patches, sacrificing current resource intake for information acquisition. This is done to obtain more precise and accurate estimates of resource levels, resulting in increased intake rates later in season. Risk of mortality may alter the trade-off between exploration and exploitation and thus change patch sampling effort. As lifetime expectancy decreases, learners invest less in information acquisition and show lower foraging performance when resource level changes through time.     

Foraging rate versus sociality in the starling Sturnus vulgaris

It is well established that social conditions often modify foraging behaviour, but the theoretical interpretation of the changes produced is not straightforward. Changes may be due to alterations of the foraging currency (the mathematical expression that behaviour maximizes) and/or of the available resources. An example of the latter is when both solitary and social foragers maximize rates of gain over time, but competition alters the behaviour required to achieve this, as assumed by ideal free distribution models. Here we examine this problem using captive starlings Sturnus vulgaris. Subjects had access to two depleting patches that replenished whenever the alternative patch was visited. The theoretical rate-maximizing policy was the same across all treatments, and consisted of alternating between patches following a pattern that could be predicted using the marginal value theorem (MVT). There were three treatments that differed in the contents of an aviary adjacent to one of the two patches (called the 'social' patch). In the control treatment, the aviary was empty, in the social condition it contained a group of starlings, and in a non-specific stimulus control it contained a group of zebra finches. In the control condition both patches were used equally and behaviour was well predicted by the MVT. In the social condition, starlings foraged more slowly in the social than in the solitary patch. Further, foraging in the solitary patch was faster and in the social patch slower in the social condition than in the control condition. Although these changes are incompatible with overall rate maximization (gain rate decreased by about 24% by self-imposed changes), if the self-generated gain functions were used the MVT was a good predictor of patch exploitation under all conditions. We discuss the complexities of nesting optimal foraging models in more comprehensive theoretical accounts of behaviour integrating functional and mechanistic perspectives.     

Resource characteristics and competition affect colony and individual foraging strategies of the wood ant Formica integroides

Abstract.  1. Resource characteristics and competitive pressure can affect an ant colony's foraging strategy. This study examined the ability of the wood ant Formica integroides to respond, at both the colony and individual levels, to changes in competitive pressure for access to terrestrial and arboreal resources.
2. Because foraging behaviours depend on resource characteristics, foraging for different resource types (e.g. terrestrial and arboreal habitats) produces different spatial or territorial arrangements. In this study, terrestrial contests for resources followed an interference-exploitation tradeoff, while arboreal foragers defended entire trees as absolute territories.
3. Competitive pressure for access to arboreal resources was shown to increase with distance from F. integroides nests.
4. In this study, the ability of F. integroides to defend a resource varied with body size. Large foragers were better defenders than small foragers. For groups of foragers, the ability to defend a resource increased with the ratio of large to small foragers.
5. In response to competitive pressure, F. integroides colonies altered the size distribution of arboreal, but not terrestrial, foragers. An increase in competitive pressure was matched by an increase in the number of large foragers allocated to trees. This response to competition affected the relationship between body size and distance from the nest for arboreal foragers.
6. Foraging behaviours for individual arboreal foragers also varied with competitive pressure. As competition increased, large arboreal foragers spent more time in direct contact with the resource rather than standing between resource patches.     

Smart,smarter, smartest: foraging information states and coexistence

Animals possess different abilities to gain and use information about the foraging patches they exploit. When ignorant of the qualities of encountered patches, a smart forager should leave all patches after the same amount of fixed search time. A smarter forager can be Bayesian by using information on cumulative harvest and time spent searching a patch to better inform its patch‐departure decision. The smartest forager has immediate and continuous knowledge about patch quality, and can make a perfect decision about when to leave each patch. Here we let each of these three strategies harvest resources from a slowly regenerating environment. Eventually a steady‐state distribution of prey among patches arises where the environment‐wide resource renewal just balances the environment‐wide harvest of the foragers. The fixed time forager creates a distribution with the highest mean and highest variance of patch qualities, followed by the Bayesian and the prescient in that order. The less informed strategies promote distributions with both more resources and more exploitable information than the more informed strategies. While it is true that a better‐informed strategy will always out‐perform a less well‐informed, its increase in performance may not compensate it for any costs associated with being better informed. We imagine that the fixed time strategy may be least expensive and the prescient strategy most expensive in terms of sensory organs and associated assess and respond capabilities. To consider competition between such strategies with varying costs, we introduced a single individual of each of the strategies into the environments created by populations of the other strategies. There are threshold costs associated with the better‐informed strategy such that it can or cannot outcompete a less‐informed strategy. However, over a relatively narrow range of foraging costs, less‐informed and better‐informed strategies will coexist. Furthermore, for the prescient and the Bayesian strategies, some combinations of foraging costs produce alternate stable states – whichever strategy establishes first remains safe from invasion by the other.     

Competition in a group of equal foragers

Abstract Using techniques from renewal process theory, we build a stochastic model for gain accumulation in a group of equal competitors foraging in a patchy environment. The model for gain of the individuals is based on the waiting times between subsequent prey encounters by the group. These waiting times depend on the number of foragers in the group. A single parameter of this dependency encompasses a variety of foraging scenarios, from co-operation to scramble. With constant patch size, correlations between gains of any pair of foragers are negative. This dependency is most intense in small groups. Increased variation in patch size makes correlations in gains between group members positive irrespective of the group size. For a solitary forager, variance in gain approaches zero with increasing time in the patch. For an individual member in a group, variance grows monotonically. Thus, depending on the patch departure rule controlling the time to be spent in the patch, solitary foragers may have a smaller variance in gain than members in a group. As solitary foragers also potentially harvest all prey in the patch, it is hard to believe that grouping behavior would evolve solely on the basis of foraging.     

Urban gardens promote bee foraging over natural habitats and plantations

Increasing human land use for agriculture and housing leads to the loss of natural habitat and to widespread declines in wild bees. Bee foraging dynamics and fitness depend on the availability of resources in the surrounding landscape, but how precisely landscape related resource differences affect bee foraging patterns remains unclear. To investigate how landscape and its interaction with season and weather drive foraging and resource intake in social bees, we experimentally compared foraging activity, the allocation of foragers to different resources (pollen, nectar, and resin) and overall resource intake in the Australian stingless bee Tetragonula carbonaria (Apidae, Meliponini). Bee colonies were monitored in different seasons over two years. We compared foraging patterns and resource intake between the bees'' natural habitat (forests) and two landscapes differently altered by humans (suburban gardens and agricultural macadamia plantations). We found foraging activity as well as pollen and nectar forager numbers to be highest in suburban gardens, intermediate in forests and low in plantations. Foraging patterns further differed between seasons, but seasonal variations strongly differed between landscapes. Sugar and pollen intake was low in plantations, but contrary with our predictions, it was even higher in gardens than in forests. In contrast, resin intake was similar across landscapes. Consequently, differences in resource availability between natural and altered landscapes strongly affect foraging patterns and thus resource intake in social bees. While agricultural monocultures largely reduce foraging success, suburban gardens can increase resource intake well above rates found in natural habitats of bees, indicating that human activities can both decrease and increase the availability of resources in a landscape and thus reduce or enhance bee fitness.     

Tolerance of understory plants subject to herbivory by roe deer

Classical foraging theory states that animals feeding in a patchy environment can maximise their long term prey capture rates by quitting food patches when they have depleted prey to a certain threshold level. Theory suggests that social foragers may be better able to do this if all individuals in a group have access to the prey capture information of all other group members. This will allow all foragers to make a more accurate estimation of the patch quality over time and hence enable them to quit patches closer to the optimal prey threshold level. We develop a model to examine the foraging efficiency of three strategies that could be used by a cohesive foraging group to initiate quitting a patch, where foragers do not use such information, and compare these with a fourth strategy in which foragers use public information of all prey capture events made by the group. We carried out simulations in six different prey environments, in which we varied the mean number of prey per patch and the variance of prey number between patches. Groups sharing public information were able to consistently quit patches close to the optimal prey threshold level, and obtained constant prey capture rates, in groups of all sizes. In contrast all groups not sharing public information quit patches progressively earlier than the optimal prey threshold value, and experienced decreasing prey capture rates, as group size increased. This is more apparent as the variance in prey number between patches increases. Thus in a patchy environment, where uncertainty is high, although public information use does not increase the foraging efficiency of groups over that of a lone forager, it certainly offers benefits over groups which do not, and particularly where group size is large.     

Eavesdropping foragers use level of collective commotion as public information to target high quality patches

Public information offers a valuable means for social foragers to determine the relative quality of foraging patches. Despite much evidence that foragers use public information based on others’ feeding behavior, no experiments have examined whether foragers might use public information based on others’ competitive behavior, particularly the collective commotion that can be generated by aggregations. Such commotion could potentially provide a rich source of public information: as foragers compete in a patch with an especially high value resource, their heightened competition intensity could enable eavesdropping foragers to target this superior patch, based simply on its higher level of collective commotion. To test the hypothesis that the level of collective commotion is used as public information by eavesdropping foragers I conducted field experiments on terrestrial hermit crabs Coenobita compressus. These animals engage in collective competitive interactions in foraging patches for food and shells, generating variable levels of commotion across different quality patches. By experimentally manipulating the level of collective commotion in sham aggregations in the wild I show that a higher level of commotion is exploited by eavesdropping foragers to differentially target more valuable patches. Broadly, these results highlight an underappreciated significance of competitive by‐products and higher‐ order collective pheno mena as forms of public information for foragers.     

Quantifying the effect of colony size and food distribution on harvester ant foraging

Desert seed-harvester ants, genus Pogonomyrmex, are central place foragers that search for resources collectively. We quantify how seed harvesters exploit the spatial distribution of seeds to improve their rate of seed collection. We find that foraging rates are significantly influenced by the clumpiness of experimental seed baits. Colonies collected seeds from larger piles faster than randomly distributed seeds. We developed a method to compare foraging rates on clumped versus random seeds across three Pogonomyrmex species that differ substantially in forager population size. The increase in foraging rate when food was clumped in larger piles was indistinguishable across the three species, suggesting that species with larger colonies are no better than species with smaller colonies at collecting clumped seeds. These findings contradict the theoretical expectation that larger groups are more efficient at exploiting clumped resources, thus contributing to our understanding of the importance of the spatial distribution of food sources and colony size for communication and organization in social insects.     

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.