The effects of spatial food distribution and group size on foraging behaviour in a benthic fish

Animals foraging in heterogeneous environments benefit from information on local resource density because it allows allocation of foraging effort to rich patches. In foraging groups, this information may be obtained by individuals through sampling or by observing the foraging behaviour of group members. We studied the foraging behaviour of goldfish (Carassius auratus) groups feeding in pools on resources distributed in patches. First, we determined if goldfish use sampling information to distinguish between patches of different qualities, and if this allowed goldfish to benefit from a heterogeneous resource distribution. Then, we tested if group size affected the time dedicated to food searching and ultimately foraging success. The decision of goldfish to leave a patch was affected by whether or not they found food, indicating that goldfish use an assessment rule. Giving-up density was higher when resources were highly heterogeneous, but overall gain was not affected by resource distribution. We did not observe any foraging benefits of larger groups, which indicate that grouping behaviour was driven by risk dilution. In larger groups the proportion searching for food was lower, which suggests interactions among group members. We conclude that competition between group members affects individual investments in food searching by introducing the possibility for alternative strategies, such as scrounging or resource monopolisation.     

Skills,division of labour and economies of scale among Amazonian hunters and South Indian honey collectors

In foraging and other productive activities, individuals make choices regarding whether and with whom to cooperate, and in what capacities. The size and composition of cooperative groups can be understood as a self-organized outcome of these choices, which are made under local ecological and social constraints. This article describes a theoretical framework for explaining the size and composition of foraging groups based on three principles: (i) the sexual division of labour; (ii) the intergenerational division of labour; and (iii) economies of scale in production. We test predictions from the theory with data from two field contexts: Tsimane'' game hunters of lowland Bolivia, and Jenu Kuruba honey collectors of South India. In each case, we estimate the impacts of group size and individual group members'' effort on group success. We characterize differences in the skill requirements of different foraging activities and show that individuals participate more frequently in activities in which they are more efficient. We evaluate returns to scale across different resource types and observe higher returns at larger group sizes in foraging activities (such as hunting large game) that benefit from coordinated and complementary roles. These results inform us that the foraging group size and composition are guided by the motivated choice of individuals on the basis of relative efficiency, benefits of cooperation, opportunity costs and other social considerations.     

A marginal model of tolerated theft

Using marginal analysis to represent Blurton Jones's concept of tolerated theft, I show how equilibrium resource transfers among individuals might be affected by foraging behavior, resource qualities, and number of participants. The model applies to hominids and other species that exchange or share food or other resources. Among the results: Tolerated theft enhances the value to be derived from resources, packets intermediate in size are most likely to be subjected to tolerated theft, packet division is more likely to be unequal than equal, division is a function of group size, and tolerated theft is most likely in small groups. The model also suggests that among reciprocators the widest possible exchange or sharing is in the self-interest of the individual procuring the resource. In general, evolutionary cost-benefit accounting should track marginal changes in the value (fitness or utility) of resources. Marginal valuation is conceptually primary and may produce results that differ from direct measures of quantity.     

Competitive asymmetry, foraging area size and coexistence of annuals

Individuals allocate resources to the expansion of their foraging area and those resources are no longer available for the traits that determine how well those individuals are able to protect their foraging area against competitors. The resulting trade‐off between foraging area size and the traits associated with the ability to compete for the resources within the foraging area applies to ecological scenarios as different as territorial defence by individuals and colonies, and light competition in plants. Whether the trade‐off affects species performance in competition for resources at the area of overlap between foraging areas depends on the symmetry of resource division. In symmetric competition resources are divided equally between the competitors, while in asymmetric competition the individual with the smallest foraging area, and consequently the greatest competitive ability, gains all the resources. Competition may also be a combination of the symmetric and asymmetric processes. I studied the effects of competitive asymmetry on population dynamics and coexistence of two annual species with different sized foraging areas using an individual‐based spatially explicit simulation model. Symmetric competition favoured the species with the larger foraging area and did not allow coexistence. Competitive asymmetry favoured the species with smaller foraging area and allowed coexistence, which was due to the consequences of losing an asymmetric competition being more severe than losing a symmetric competition. The mechanism of coexistence is the larger foraging area's superiority in low population densities (little competition) and the smaller foraging area's ability to win a large foraging area when competition was intense. Competitive asymmetry and small size of both foraging areas led to population dynamics dominated by long‐term fluctuations of small intensity. Symmetric competition and large size of the foraging areas led to large short‐term fluctuations, which often resulted in the extinction of one or both of the species due to demographic stochasticity.     

Prey size affects the costs and benefits of group predation in nymphs of the predatory stink bug Andrallus spinidens (Heteroptera: Pentatomidae)

Group predation promotes foraging efficiency because it increases the size of prey that can be killed and improves hunting success compared to solitary predation. However, group predation may increase competition among group members during feeding. Earlier studies have focused on the advantages of group predation, but little is known about the costs and benefits of group predation for individual members of the group. Here, we show that the costs and benefits of group predation for individuals of the predatory stink bug Andrallus spinidens vary with prey size in laboratory experiments. We found that when A. spinidens fed on small prey, group predation did not significantly increase foraging efficiency but did increase competition for food among group members. In contrast, when prey was large, group predation promoted foraging efficiency, and competition over food was not detected. Our results suggest that group predation by A. spinidens nymphs is advantageous for individual members because it enables each member to hunt larger prey that could not be hunted alone. However, when group size was large or prey size was small, group predation increased competition among group members.     

Divergence in social foraging among morphs of the three‐spined stickleback,Gasterosteus aculeatus

Foraging in social groups has a number of benefits but can also increase the risk of exploitation. High tendency to shoal may be correlated with groups foraging, although facultatively social fish adjust both shoaling decisions and food resource defence based on intrinsic and extrinsic factors. The main aim of this study was to examine the relationships between shoaling, solitary foraging and aggression, forager tolerance of conspecifics joining at a discovered food patch and forager exploitation of resources discovered by others. We used two intra‐lacustrine three‐spined stickleback morph pairs, lava and mud, and monomorphic morphs from each of lava and mud habitats. The lava morph formed less cohesive shoals, was bolder during solitary foraging, approached and entered an occupied food patch less frequently than the mud morph, suggesting a link between shoaling and the propensity for social foraging. However, shoaling tendency and joiner tolerance were not correlated at a population level. Intralacustrine lava and mud morphs differed more markedly in joiner tolerance than morphs from single habitat lakes, whereas the opposite was true for shoaling tendency. We conclude that, in addition to differentiation in shoaling tendency, the lava and mud morphs differ in social foraging and these variations may act to promote population divergence. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 194–203.     

Social foragers adopt a riskier foraging mode in the centre of their groups

Foraging in groups provides many benefits that are not necessarily experienced the same way by all individuals. I explore the possibility that foraging mode, the way individuals exploit resources, varies as a function of spatial position in the group, reflecting commonly occurring spatial differences in predation risk. I show that semipalmated sandpipers (Calidris pusilla), a social foraging avian species, tended to adopt a riskier foraging mode in the central, more protected areas of their groups. Central birds effectively used the more peripheral group members as sentinels, allowing them to exploit a wider range of resources within the same group at the same time. This finding provides a novel benefit of living in groups, which may have a broad relevance given that social foraging species often exploit a large array of resources.     

Use of Social and Ecological Information in Tamarin Foraging Decisions

A major consequence of group living is that foragers may rely on social information in addition to ecological information to locate feeding sites. Although conspecifics can provide cues as to the spatial location of food patches, individual foraging decisions also must include some assessment of the likelihood of obtaining access to a resource other group members seek. This likelihood differs in the 2 models generally proposed to explain intragroup social foraging: the information-sharing model and the producer-scrounger model. We conducted an experimental field study on wild groups of emperor (Saguinus imperator) and saddleback (S. fuscicollis) tamarins to determine the foraging strategies adopted by individual group members and their relationship to social rank, food intake, and the ability to use ecological and social information in making intra-patch foraging decisions. Individual tamarins applied different behavioral strategies compatible with a finder-joiner paradigm to solve foraging problems. About half of the individuals in each study group initiated 74%–90% of all food searches and acted as finders. Most alpha individuals adopted a joiner strategy by monitoring the activities of others' to obtain a reward. The individual arriving first at a reward platform enjoyed a finder's advantage. Despite differences in search effort, both finders and joiners presented similar abilities in learning to associate ecological cues with the presence of food rewards at our experimental feeding stations. We conclude that within a group foraging context, tamarins integrate social and ecological information in decision-making.     

Decreasing functional responses as a result of adaptive consumer behavior

Summary Several different mechanisms that may produce decreasing functional responses are investigated using models that assume that an optimally foraging consumer is exploiting one or two resources. Decreasing functional responses are associated with situations in which there are costs to resource consumption. If the process of resource acquisition has costs, decreasing functional responses may occur when there is a single homogeneous resource. If the cost is solely a function of the amount of resource ingested, decreasing functional responses on a single resource do not occur. Both types of cost can produce decreasing functional responses when there are two resource types and a trade-off relationship between consumption of one and consumption of the other. Decreasing functional responses seem to be most likely to occur on a food that yields high benefits and costs per unit of foraging time or effort when there is an alternative resource which yields low benefits and costs. Given this type of foraging choice, the functional response is most likely to decrease when the benefits of ingestion increase at a decreasing rate, and the costs of ingestion increase at an increasing rate with amount ingested. An important and unique consequence of decreasing functional responses is the possibility of population cycles in differential equation models of consumer-resource systems with non-reproducing resources; this is illustrated with a simple comsumer-resource model.     

Florida Scrub-Jays (Aphelocoma coerulescens) are Sentinels More When Well-Fed (Even with no Kin Nearby)

Sentinels occupy high, exposed positions while other group members forage nearby. If sentinel behavior involves a foraging–predation risk trade‐off, animals should be sentinels more when fed supplemental food. When individual Florida scrub‐jays (Aphelocoma coerulescens) were fed fragments of peanuts, during the following 30 min they shifted 30% of their time from foraging to sentinel behavior. In a follow‐up experiment, we fed either one or two members in each group. As before, the jays reduced their foraging and spent much more time as sentinels when given supplemental food. In each treatment, pairs were sentinels simultaneously considerably less often than expected by chance. The dramatic shift from foraging to sentinel behavior suggests that for Florida scrub‐jays sentinel behavior brings substantial benefits for no greater cost than that of lost opportunities to forage. Because the results held for simple mated pairs of scrub‐jays, we argue that kin selection and social prestige are not necessary to explain sentinel behavior.     

Harvesting resources in groups or alone: the case of renewing patches

Group foraging has been proposed to be the most efficient mannerwith which to exploit habitats with renewing patches as individualsin groups are less likely to revisit patches that have alreadybeen exploited recently by others. However, to avoid a group-selectionargument, it is necessary to compare the success of solitaryand group foraging tactics when each competes with the other.We used a genetic algorithm approach to examine the costs andbenefits of exploiting renewing resources in a spatially andtemporally explicit habitat, thus controlling the time courseof resource renewal and including the time cost of travelingbetween patches, which may be a significant factor for groupforagers that deplete patches more quickly. Results indicatethat group foragers fare more poorly than an equivalent numberof solitary foragers in the same habitat unless the rate ofresource renewal is very low. The low revisitation rate by groupforagers allows resources to replenish more fully, thus maintainingthe resource level across the habitat at a higher level. Incontrast, solitary foragers, who revisit previously exploitedpatches more often, maintain the same resources at a lower level.Nevertheless, a pure population of group foragers can be readilyinvaded by solitary foragers even when the rate of renewal isat low levels. We conclude that while group foraging may bean efficient tactic to exploit renewing resources, it is nota stable strategy under the circumstances examined in this model.     

Cooperation and the common good

In this paper, we draw the attention of biologists to a result from the economic literature, which suggests that when individuals are engaged in a communal activity of benefit to all, selection may favour cooperative sharing of resources even among non-relatives. Provided that group members all invest some resources in the public good, they should refrain from conflict over the division of these resources. The reason is that, given diminishing returns on investment in public and private goods, claiming (or ceding) a greater share of total resources only leads to the actor (or its competitors) investing more in the public good, such that the marginal costs and benefits of investment remain in balance. This cancels out any individual benefits of resource competition. We illustrate how this idea may be applied in the context of biparental care, using a sequential game in which parents first compete with one another over resources, and then choose how to allocate the resources they each obtain to care of their joint young (public good) versus their own survival and future reproductive success (private good). We show that when the two parents both invest in care to some extent, they should refrain from any conflict over the division of resources. The same effect can also support asymmetric outcomes in which one parent competes for resources and invests in care, whereas the other does not invest but refrains from competition. The fact that the caring parent gains higher fitness pay-offs at these equilibria suggests that abandoning a partner is not always to the latter''s detriment, when the potential for resource competition is taken into account, but may instead be of benefit to the ‘abandoned’ mate.     

The ecology of cooperative breeding behaviour

Ecology is a fundamental driving force for the evolutionary transition from solitary living to breeding cooperatively in groups. However, the fact that both benign and harsh, as well as stable and fluctuating, environments can favour the evolution of cooperative breeding behaviour constitutes a paradox of environmental quality and sociality. Here, we propose a new model – the dual benefits framework – for resolving this paradox. Our framework distinguishes between two categories of grouping benefits – resource defence benefits that derive from group‐defended critical resources and collective action benefits that result from social cooperation among group members – and uses insider–outsider conflict theory to simultaneously consider the interests of current group members (insiders) and potential joiners (outsiders) in determining optimal group size. We argue that the different grouping benefits realised from resource defence and collective action profoundly affect insider–outsider conflict resolution, resulting in predictable differences in the per capita productivity, stable group size, kin structure and stability of the social group. We also suggest that different types of environmental variation (spatial vs. temporal) select for societies that form because of the different grouping benefits, thus helping to resolve the paradox of why cooperative breeding evolves in such different types of environments.     

Behavioral and energetic costs of group membership in a coral reef fish

Animals in social aggregations often spend more time foraging than solitary conspecifics. This may be a product of the relative safety afforded by aggregations: group members can devote more time to foraging and less time to antipredator behaviors than solitary animals (the “risk reduction” effect). All else being equal, risk reduction should result in higher food intake for grouped animals. However, intragroup competition may force group members to spend more time foraging in order to obtain the same food ration as solitary individuals (the “resource competition” effect). We compared these opposing explanations of foraging time allocation in a coral reef fish, bluehead wrasse (Thalassoma bifasciatum). Aggregations of juvenile bluehead wrasse experience safety-in-numbers, and preliminary observations suggested that juveniles in aggregations spent more time foraging for copepods in the water column than solitary juveniles. However, the risk reduction and resource competition hypotheses are indistinguishable on the basis of behavioral observations alone. Therefore, we collected behavioral, dietary, and growth data (using otolith growth rings) for bluehead wrasse at multiple reefs around a Caribbean island. Despite spending more time foraging in the water column, grouped fish did not capture more prey items and had slower growth rates than solitary fish. Thus, the increased foraging time of grouped fish appears to reflect resource competition, not risk reduction. This competition may limit the size and frequency of aggregations among juvenile bluehead wrasse, which have been shown to experience reduced mortality rates in larger groups. Bluehead wrasse recruits also spent less time foraging but grew faster at sites where planktonic copepod prey were more abundant. This suggests the possibility that large-scale spatiotemporal variability in the abundance of planktonic copepods over coral reefs may produce corresponding variability in the dynamics of reef fish populations. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The social cognition of social foraging: partner selection by underlying valuation

Humans and other animals have a variety of psychological abilities tailored to the demands of asocial foraging, that is, foraging without coordination or competition with other conspecifics. Human foraging, however, also includes a unique element: the creation of resource pooling systems. In this type of social foraging, people contribute when they have excess resources and receive provisioning when in need. Is this behavior produced by the same psychology as asocial foraging? If so, foraging partners should be judged by the same criteria used to judge asocial patches of resources: the net energetic benefits they provide. The logic of resource pooling speaks against this. Maintaining such a system requires the ability to judge others not on their short-term returns, but on the psychological variables that guide their behavior over the long term. We test this idea in a series of five studies using an implicit measure of categorization. Results showed that (a) others are judged by the costs they incur (a variable not relevant to asocial foraging), whereas (b) others are not judged by the benefits they provide when benefits provided are unrevealing of underlying psychological variables (despite this variable being relevant to asocial foraging). These results are suggestive of a complex psychology designed for both social and asocial foraging.     

Foraging responses ofFormica truncorum (Hymenoptera; Formicidae); exploiting stable vs spatially and temporally variable resources

Summary Social organization allows a division of labor between reproduction and foraging, as well as a task allocation among foraging individuals. Therefore, a colony simultaneously exploiting various resources can use different ways of getting them. This work analyzes wood ant foraging tactics both at the collective and the individual level. The main issues are: (1) How does a wood ant colony respond to stable vs temporally and spatially variable resources in terms of worker force allocation? (2) How is retrieval of ephemeral but rich resources effected at the individual level? The results show a correspondence between resource stability and behavior. Ants visited stable resources continually and recruited to ephemeral ones. They also visited empty food sites at a low frequency suggesting territoriality and constant scanning. The results suggest that recruitment may involve defence of the resource, since only a small fraction of all the workers available carry the food home. Additional ants arriving at the resource are recruited from inside the nest, not by reallocating outdoor workers. The foragers also form two separate groups, one site-persistent and another site-flexible. These two forager groups may reflect specialization on two different kinds of diet: honeydew or insect prey (spatially and temporally stable vs unstable resources, respectively).Formica truncorum is thus an example of how sociality allows the use of two different foraging strategies simultaneously: one that is based on recruitment to ephemeral resources and another that is based on search persistance and memory.     

Foraging sequence,energy intake and torpor: an individual‐based field study of energy balancing in desert golden spiny mice

We studied the relationship between sequence of foraging, energy acquired and use of torpor as an energy‐balancing strategy in diurnally active desert golden spiny mice. We hypothesised that individuals that arrive earlier to forage will get higher returns and consequently spend less time torpid. If that is the case, then early foragers can be viewed as more successful; if the same individuals arrive repeatedly early, they are likely to have higher fitness under conditions of resource limitation. For the first time, we show a relationship between foraging sequence and amount of resources removed, with individuals that arrive later to a foraging patch tending to receive lower energetic returns and to spend more time torpid. Torpor bears not only benefits but also significant costs, so these individuals pay a price both in lower energy intake and in extended periods of torpor, in what may well be a positive feedback loop.     

An asymmetric producer-scrounger game: body size and the social foraging behavior of coho salmon

A tension between cooperation and conflict characterizes the behavioral dynamics of many social species. The foraging benefits of group living include increased efficiency and reduced need for vigilance, but social foraging can also encourage theft of captured prey from conspecifics. The payoffs of stealing prey from others (scrounging) versus capturing prey (producing) may depend not only on the frequency of each foraging strategy in the group but also on an individual’s ability to steal. By observing the foraging behavior of juvenile coho salmon (Oncorhynchus kisutch), we found that, within a group, relatively smaller coho acted primarily as producers and took longer to handle prey, and were therefore more likely to be targeted by scroungers than relatively larger coho. Further, our observations suggest that the frequency of scrounging may be higher when groups contained individuals of different sizes. Based on these observations, we developed a model of phenotype-limited producer-scrounger dynamics, in which rates of stealing were structured by the relative size of producers and scroungers within the foraging group. Model simulations show that when the success of stealing is positively related to body size, relatively large predators should tend to be scroungers while smaller predators should be producers. Contrary to previous models, we also found that, under certain conditions, producer and scrounger strategies could coexist for both large and small phenotypes. Large scroungers tended to receive the highest payoff, suggesting that producer-scrounger dynamics may result in an uneven distribution of benefits among group members that—under the right conditions—could entrench social positions of dominance.     

Effects of group size and group density on trade-offs in resource selection by a group-territorial central-place foraging woodpecker

Trade-offs in resource selection by central-place foragers are driven by the need to balance the benefits of selecting resources against the costs of travel from the central place. For group-territorial central-place foraging birds, trade-offs in resource selection are likely to be complicated by a competitive advantage for larger groups at high group density that may limit accessibility of high-quality distant resources to small groups. We used the group-territorial, central-place foraging Red-cockaded Woodpecker Leuconotopicus borealis (RCW) as a case study to test predictions that increases in group density lead to differences in foraging distances and resource selection for groups of different sizes. We used GPS tracking and LiDAR-derived habitat data to model effects of group size on foraging distances and selection for high-quality pines (≥ 35.6 cm diameter at breast height (dbh)) and lower quality pines (25.4–35.6 cm dbh) by RCW groups across low (n = 14), moderate (n = 10) and high group density (n = 10) conditions. At low and moderate group density, all RCW groups selected distant high-quality pines in addition to those near the central place because competition for resources was low. In contrast, at high group density, larger groups travelled further to select high-quality pines, whereas smaller groups selected high-quality pines only when they were close to the central place and, conversely, were more likely to select lower quality pines at greater distances from the central place. Selection for high-quality pines only when close to the cavity tree cluster at high group density is important to long-term fitness of small RCW groups because it allows them to maximize benefits from both territorial defence and selecting high-quality resources while minimizing costs of competition. These relationships suggest that intraspecific competition at high group density entails substantive costs to smaller groups of territorial central-place foragers by limiting accessibility of distant high-quality foraging resources.     

Group foraging in wild brown hares: effects of resource distribution and social status

Wild brown hares (Lepus europaeus), though normally comparatively solitary, have the capacity to adjust their behaviour such that they can benefit when foraging in groups. They are able to allocate more time to feeding and have an increased corporate vigilance as group size increases. However, these benefits are conditional upon the food distribution. When food is spaced, all individuals benefit. When it is clumped into a small defendable patch, dominant hares attempt to monopolize the resource. They can successfully exclude subordinates when group size is small but, as group size increases, they must devote more time to defending the patch. Therefore, dominants spend less time feeding with increasing group size, while subordinates spend more, since they have more opportunity to feed while the dominant hare is off chasing other individuals. As a consequence, when more than two hares are present, all individuals do less well when food is clumped than when it is spaced.