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.     

Group size effects on foraging and vigilance in migratory Tibetan antelope

Large group sizes have been hypothesized to decrease predation risk and increase food competition. We investigated group size effects on vigilance and foraging behaviour during the migratory period in female Tibetan antelope Pantholops hodgsoni, in the Kekexili Nature Reserve of Qinghai Province, China. During June to August, adult female antelope and yearling females gather in large migratory groups and cross the Qinghai-Tibet highway to calving grounds within the Nature Reserve and return to Qumalai county after calving. Large groups of antelope aggregate in the migratory corridor where they compete for limited food resources and attract the attention of mammalian and avian predators and scavengers. We restricted our sampling to groups of less than 30 antelopes and thus limit our inference accordingly. Focal-animal sampling was used to record the behaviour of the free-ranging antelope except for those with lambs. Tibetan antelope spent more time foraging in larger groups but frequency of foraging bouts was not affected by group size. Conversely, the time spent vigilant and frequency of vigilance bouts decreased with increased group size. We suggest that these results are best explained by competition for food and risk of predation.     

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 Rules for Group Feeders: Area Copying Depends upon Food Density in Shoaling Goldfish

The decision rules governing forage area copying behaviour were investigated in shoaling fish. Shoaling goldfish were offered two equal food patches, one of which was adjacent to an equal-sized shoal feeding behind a transparent barrier. When food was low, goldfish foraged according to an area copying rule, but under high and zero food area copying disappeared. Only under high food density did equal numbers of fish feed at both sites as predicted by foraging theory. Under zero food the fish were less certain about where to forage. Precise visual cues from feeding fish were required: non-feeders did not attract area copiers. Furthermore, area copying was task-dependent since it reappeared strongly if fish were not able to forage on patches like their fellows. Control experiments eliminated an increase in group size for anti-predator advantage as an explanation. Two sequential decisions: to stay or move, and to join or leave may explain the results, which are not accommodated by simple optimality models. These decisions may be based on a comparison of current food intake with the anticipation of a higher reward by foraging socially.     

Context dependence in foraging behaviour of Achillea millefolium

Context-dependent foraging behaviour is acknowledged and well documented for a diversity of animals and conditions. The contextual determinants of plant foraging behaviour, however, are poorly understood. Plant roots encounter patchy distributions of nutrients and soil fungi. Both of these features affect root form and function, but how they interact to affect foraging behaviour is unknown. We extend the use of the marginal value theorem to make predictions about the foraging behaviour of roots, and test our predictions by manipulating soil resource distribution and inoculation by soil fungi. We measured plant movement as both distance roots travelled and time taken to grow through nutrient patches of varied quality. To do this, we grew Achillea millefolium in the centers of modified pots with a high-nutrient patch and a low-nutrient patch on either side of the plant (heterogeneous) or patch-free conditions (homogeneous). Fungal inoculation, but not resource distribution, altered the time it took roots to reach nutrient patches. When in nutrient patches, root growth decreased relative to homogeneous soils. However, this change in foraging behaviour was not contingent upon patch quality or fungal inoculation. Root system breadth was larger in homogeneous than in heterogeneous soils, until measures were influenced by pot edges. Overall, we find that root foraging behaviour is modified by resource heterogeneity but not fungal inoculation. We find support for predictions of the marginal value theorem that organisms travel faster through low-quality than through high-quality environments, with the caveat that roots respond to nutrient patches per se rather than the quality of those patches.     

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.     

Clash of kingdoms or why Drosophila larvae positively respond to fungal competitors

BACKGROUND: Competition with filamentous fungi has been demonstrated to be an important cause of mortality for the vast group of insects that depend on ephemeral resources (e.g. fruit, dung, carrion). Recent data suggest that the well-known aggregation of Drosophila larvae across decaying fruit yields a competitive advantage over mould, by which the larvae achieve a higher survival probability in larger groups compared with smaller ones. Feeding and locomotor behaviour of larger larval groups is assumed to cause disruption of fungal hyphae, leading to suppression of fungal growth, which in turn improves the chances of larval survival to the adult stage. Given the relationship between larval density, mould suppression and larval survival, the present study has tested whether fungal-infected food patches elicit communal foraging behaviour on mould-infected sites by which larvae might hamper mould growth more efficiently. RESULTS: Based on laboratory experiments in which Drosophila larvae were offered the choice between fungal-infected and uninfected food patches, larvae significantly aggregated on patches containing young fungal colonies. Grouping behaviour was also visible when larvae were offered only fungal-infected or only uninfected patches; however, larval aggregation was less strong under these conditions than in a heterogeneous environment (infected and uninfected patches). CONCLUSION: Because filamentous fungi can be deadly competitors for insect larvae on ephemeral resources, social attraction of Drosophila larvae to fungal-infected sites leading to suppression of mould growth may reflect an adaptive behavioural response that increases insect larval fitness and can thus be discussed as an anti-competitor behaviour. These observations support the hypothesis that adverse environmental conditions operate in favour of social behaviour. In a search for the underlying mechanisms of communal behaviour in Drosophila, this study highlights the necessity of investigating the role of inter-kingdom competition as a potential driving force in the evolution of spatial behaviour in insects.     

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.     

Behavioural Response of Juvenile Bluegill Sunfish to Variation in Predation Risk and Food Level

The behavioural response of juvenile bluegill sunfish (Lepomis macrochirus) to predation risk when selecting between patches of artificial vegetation differing in food and stem density was investigated. Bluegill foraging activity was significantly affected by all three factors. Regardless of patch stem density or risk of predation bluegills preferred patches with the highest prey number. During each trial bluegill foraging activity was clearly divided into a between- and within-patch component. In the presence of a predator bluegills reduced their between-patch foraging activity by an equivalent amount regardless of patch stem density or food level, apparently showing a risk-adjusting behavioural response to predation risk. Within patches, however, foraging activity was affected by both food level and patch stem density. When foraging in a patch offering a refuge from predation, the presence of a predator had no effect on bluegill foraging activity within this patch. However, if foraging in a patch with only limited refuge potential, bluegill foraging activity was reduced significantly in the presence of a predator. Further, this reduction was significantly greater if the patch contained a low versus a high food level, indicating a risk-balancing response to predation with respect to within-patch foraging activity. Both these responses differ from the risk-avoidance response to predation demonstrated by juvenile bluegills when selecting among habitats. Therefore, our results demonstrate the flexibility of juvenile bluegill foraging behaviour.     

东方田鼠家群个体体重、攻击性及胆量对其扩散的影响

扩散作为动物适应生存环境的重要特征之一,受到自身生物学特征及环境等方面的制约。以家群形式生活和子代雄鼠扩散为主的东方田鼠,其成员个体是否因领地食物和空间资源竞争导致体重小、攻击性弱及胆小个体先行扩散。以新鲜马唐叶片构建三块密集均质的食物斑块,在食物斑块周边以透明玻璃设置观测箱,采用透明塑胶管连接三块食物斑块作为动物扩散的通道,构建东方田鼠扩散行为观测装置。将东方田鼠家群子代成员投放至带有自身家群气味的食物斑块,测定成员个体在食物斑块上的觅食行为序列过程和参数,以及向其他食物斑块扩散的行为过程和参数以及家群中先行扩散个体体重、攻击性及胆量占所有家群数的比率,检验成员个体的体重、攻击性及胆量对扩散的影响。结果发现,体重小、攻击性弱和胆小个体的觅食启动时间极显著地大于体重大、攻击性强和胆大个体的,但其先行扩散的比率却显著地大于体重大、攻击性强和胆大个体的;然而体重小、攻击性弱及胆小个体的扩散开始时间显著或极显著地大于体重大、攻击性强及胆大个体的。结果揭示,东方田鼠家群成员随着年龄的增长和对食物及空间资源竞争的加剧,体重大、攻击性强及胆大个体会迫使体重小、攻击性弱及胆小个体先行扩散。     

Social networks predict patch discovery in a wild population of songbirds

Animals use social information in a wide variety of contexts. Its extensive use by individuals to locate food patches has been documented in a number of species, and various mechanisms of discovery have been identified. However, less is known about whether individuals differ in their access to, and use of, social information to find food. We measured the social network of a wild population of three sympatric tit species (family Paridae) and then recorded individual discovery of novel food patches. By using recently developed methods for network-based diffusion analysis, we show that order of arrival at new food patches was predicted by social associations. Models based only on group searching did not explain this relationship. Furthermore, network position was correlated with likelihood of patch discovery, with central individuals more likely to locate and use novel foraging patches than those with limited social connections. These results demonstrate the utility of social network analysis as a method to investigate social information use, and suggest that the greater probability of receiving social information about new foraging patches confers a benefit on more socially connected individuals.     

Patch use behaviour in benthic fish depends on their long-term growth prospects

Animals foraging in a heterogeneous environment may combine prior information on patch qualities and patch sample information to maximize intake rate. Prior information dictates the long-term expectations, whereas prior information in combination with patch sample information determines when to leave an individual food patch. We examined patch use behaviour of benthic feeding fish in their natural environment at different spatial scales to test if they could determine patch quality and if patch use behaviour was correlated with environmental quality. In seven lakes along a gradient of environmental quality (measured as maximum benthivore size), we made repeated measurements of giving-up density (GUD) in artificial food patches of different qualities. At the largest spatial scale, between lakes, we tested if giving-up densities revealed the long-term growth expectation of benthic fish. At the local scale of patches and micro patches we tested for the ability of benthic fish to assess patch quality, and how this ability depended on the patch exploitation levels between the different lakes. We found that GUD was positively related to maximum size of bream, suggesting that short-term behavioural decisions reflected long-term growth expectations. Benthic fish discriminated between nearby rich and poor patches, but not between rich and poor micropatches within a food patch. This suggests that the foraging scale of benthic fish lies between the patch and micro patch scale in our experiments. We conclude that patch use behaviour of benthic fish can provide a powerful measure of habitat quality that reveals how benthic fish perceive their environment.     

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.     

Patch time allocation and patch sampling by foraging great and blue tits

The rate at which parents deliver energy to their brood is an important factor in avian reproduction because poor condition caused by malnutrition may reduce the offspring's survival to breeding. Models of central place foraging predict that nesting parents should optimize their prey delivery rate by minimizing travelling distances and by selecting patches where the gain per unit cost is high. I investigated the allocation of searching time amongst food patches in the home ranges of breeding great tits, Parus major, and blue tits P. caeruleus, by radiotracking. The density of locations in individual trees was positively correlated with prey biomass within trees and negatively with the distance of the trees from the nest. These two factors explained 52% of the variance in the allocation of the birds' search time. In rich patches, food was reduced considerably within 20 m of the nests, and the birds' travelling distances increased significantly during the nestling period. In parallel to foraging selectively in rich resources near the nest, the birds continually sampled the trees in their territory. The average surplus search time due to resource exploration was 1.52 times (range 1.25-1.99) the expected search time if the birds had exclusively used the most profitable patch. Despite considerable effort in patch sampling, the overall search time per unit prey was 30% better than expected by an equal use of trees. The results suggest that foraging tit parents come close to the maximum rate of prey delivery possible in a given patch distribution. Copyright 2000 The Association for the Study of Animal Behaviour.     

Effects of food quality, diet preference and water on patch use by Nubian ibex

Measuring patch use of a forager can reveal not only its cost and benefits from foraging, but also the importance of environmental factors and the significance of energy, nutrients and predation risk to its fitness. In order to assess the effects of various variables that may affect the foraging behavior of free-ranging Nubian ibex in the Negev Desert, Israel, giving-up densities (GUD) in artificial food patches were measured following Kotler et al. In particular, we tested the effects of food quality and water availability on Nubian ibex foraging behavior. To do so, we (1) tested whether the tannic acid content of food affected diet preferences, (2) assayed their diet selection strategy, (3) tested if the foraging decisions of the Nubian ibex were affected by the availability of water and (4) determined the nutritional relationship between food resources and water. Nubian ibex had lower GUDs and used resources patches more intensively where water was available, the food quality was higher and the location was closer to the escape terrain. Nubian ibex showed an expanding specialist diet selection when exploiting resource patches with a mix of items that differ in quality. Overall, food and water were complementary resources for Nubian ibex, and tannins reduced food quality. These factors help to determine patch foraging behavior decisions in Nubian ibex and contribute to habitat quality.     

Does information sharing promote group foraging?

Individuals may join groups for several reasons, one of which is the possibility of sharing information about the quality of a foraging area. Sharing information in a patch-foraging scenario gives each group member an opportunity to make a more accurate estimate of the quality of the patch. In this paper we present a mathematical model in which we study the effect of group size on patch-leaving policy and per capita intake rate. In the model, group members share information equally in a random search for food. Food is distributed in patches according to a negative binomial distribution. A prediction from our model is that, the larger the group, the earlier each group member should leave the current patch. We also find that the benefit from enhanced exchange of information does not exceed the cost of sharing food with group members. The per capita intake rate decreases as the group size increases. Therefore, animals should only form groups when other factors outweigh the costs, which is easiest to achieve when the travelling time is short.     

Ecological constraints on group size in three species of neotropical primates

The foraging strategies and association patterns of 3 species of primates (Ateles geoffroyi, Alouatta palliata, Cebus capucinus) were studied over a 5-year period. The objective of the study was to provide a quantitative test of the hypothesis that the size, density and distribution of food resources influence the size of animal groups. In examining the assumptions of this hypothesis, it was shown that these primates used resources that occurred in patches, depleted the patches through their use, and that membership in large subgroups was associated with increased travel costs. The howler and spider monkey groups formed subgroups, the size of which could be predicted from the size, density and distribution of their plant food resources. When resources were clumped and at a low density, both the howler and spider monkeys were found in small subgroups, whereas when patches were uniformly distributed and at high density they formed larger subgroups. Capuchin monkeys, in contrast, did not respond to changes in these ecological variables by forming subgroups or changing the cohesion of their group.     

Detecting predators and locating competitors while foraging: an experimental study of a medium-sized herbivore in an African savanna

Vigilance allows individuals to escape from predators, but it also reduces time for other activities which determine fitness, in particular resource acquisition. The principles determining how prey trade time between the detection of predators and food acquisition are not fully understood, particularly in herbivores because of many potential confounding factors (such as group size), and the ability of these animals to be vigilant while handling food. We designed a fertilization experiment to manipulate the quality of resources, and compared awareness (distinguishing apprehensive foraging and vigilance) of wild impalas (Aepyceros melampus) foraging on patches of different grass height and quality in a wilderness area with a full community of predators. While handling food, these animals can allocate time to other functions. The impalas were aware of their environment less often when on good food patches and when the grass was short. The animals spent more time in apprehensive foraging when grass was tall, and no other variable affected apprehensive behavior. The probability of exhibiting a vigilance posture decreased with group size. The interaction between grass height and patch enrichment also affected the time spent in vigilance, suggesting that resource quality was the main driver when visibility is good, and the risk of predation the main driver when the risk is high. We discuss various possible mechanisms underlying the perception of predation risk: foraging strategy, opportunities for scrounging, and inter-individual interference. Overall, this experiment shows that improving patch quality modifies the trade-off between vigilance and foraging in favor of feeding, but vigilance remains ultimately driven by the visibility of predators by foragers within their feeding patches.     

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.     

The Causes of Resource Monopolization: Interaction Between Resource Dispersion and Mode of Competition

The degree of monopolization of resources is thought to be higher in groups that compete by interference than by exploitation. However, the monopolization of resources will presumably depend on (1) whether the dispersion of resources is economically defendable, and (2) whether some competitors have the ability to defend these resource distributions and hence capitalize on this potential. We tested for an interaction between the effects of temporal resource dispersion and aggressiveness on the degree of resource monopolization in a foraging system. Two species of fish differing markedly in aggressiveness (high: convict cichlids, Archocentrus nigrofasciatus; low: goldfish, Carrasius auratus) were allowed to compete intra‐specifically in groups of four for food that was either potentially defendable (arrived asynchronously) or not (arrived synchronously). As predicted, the monopolization of food, measured as the coefficient of variation of food eaten within groups, was significantly higher in the defendable than in the undefendable treatment for convict cichlids but not for goldfish. However, the monopolization of food was higher in the non‐aggressive goldfish than in the aggressive convict cichlids. Future studies should quantify and compare the monopolization in species that compete primarily via scramble competition to those that use primarily resource defence.