A test of the producer-scrounger foraging game in captive flocks of spice finches,Loncbura punctulata

Group foraging allows the co-existence of a strategy (producer) that involves searching for food, and its alternative (scrounger) exploiting the food of the producer. The use of producer and scrounger strategies has been modelled as an alternative-option scramble which assumes strong negative frequency-dependence of the scrounger's pay-offs. We tested this assumption in a flock feeding situation by manipulating the proportion of scroungers in flocks of spice finches, Lonchura punctulata. In a first experiment we found that: (1) the food intake of scroungers, and to a lesser extent producers, was negatively affected by an increase in the proportion of scroungers; (2) the food intake of producers and scroungers was equal when the proportion of scroungers was small, suggesting that producers, who exploited 35.4% of their patches by scrounging were opportunistically adjusting their use of the strategies until the pay-offs equalized. In a second experiment we tested whether finches could vary their use of the two strategies in response to changes in foraging conditions brought about by an increase in the cost of producing. As predicted by the game, finches reduced their use of the producer strategy and increased their use of the scrounger strategy when the cost of producing increased. These results suggest that spice finches can alter their allocation to each foraging alternative by experience and that the producer-scrounger game is a realistic model for predicting group foraging decisions. Correspondence to: L.-A. Giraldeau     

Family-related differences in social foraging tactic use in the zebra finch (Taeniopygia guttata)

When animals forage in groups, they can search for food themselves (producer tactic), or they can search for opportunities to exploit the food discoveries of others (scrounger tactic). Both theoretical and empirical work have shown that group-level use of these alternative tactics is influenced by environmental conditions including group size and food distribution, and individual tactic use can be influenced by several measures of individual state, including body condition. Because body condition has been shown to be heritable for various species, social foraging tactics may also be heritable. We looked for evidence of heritability in social foraging tactic use in the zebra finch (Taeniopygia guttata) by testing whether: (1) natural variation in body condition correlates with tactic use, (2) there are family-related differences in body condition, and (3) there are family-related differences in observed tactic use. Tactic use in the zebra finch was significantly related to body condition; individuals with lower body condition scores had a significantly higher use of the scrounger tactic as predicted from variance-sensitive producer–scrounger models. Body-condition scores differed significantly between families, suggesting that this aspect of individual state may have a heritable component. Finally, we recorded significant family-related differences in the use of producer and scrounger alternatives. These results are consistent with heritability in observed tactic use resulting from an inheritance of individual state, in this case body condition, which itself influences tactic use. Understanding how and why individuals differ in their use of alternative tactics is fundamental as it may provide important insights into inter-individual variation in fitness.     

Group foraging, patch exploitation time and the finder's advantage

 We use a combination of the marginal value theorem (MVT) of Charnov (1976), and a group foraging model featuring information sharing to address patch residence in an environment where food occurs in discrete patches. We shall show that among equal competitors the optimal patch time for the individual that finds the food patch is shorter than that for the non-finder among equal competitors, T _E < T _N. This is the case if the patch-finder commences food harvesting in the patch earlier and manages to monopolise a fraction of the prey items (finder's advantage) before the other individuals come to take their benefit. When individuals differ in their food-searching abilities so that some of them (producers) contribute proportionally more to food-searching than others (scroungers), and differ in ability to compete for the food found, a difference emerges between producer and scrounger individuals in the optimal patch time. Within a patch we always have the finder's advantage (T _E < T _N) regardless of phenotype. Between patches a suite of optimal patch times for encountering individuals emerges depending on the performance of producers and scroungers when changing from solitary feeding to feeding in a group. The optimal patch time for individuals that are affected more severely by competition is shorter than that for individuals of the phenotype with better competitive ability. When both phenotypes are affected similarly no difference in optimal patch times emerges. Received: 13 February 1996 / Accepted after revision: 28 September 1996     

Increasing foraging group size increases scrounger use and reduces searching efficiency in nutmeg mannikins (<Emphasis Type="Italic">Lonchura punctulata</Emphasis>)

Social foragers can benefit from others' success by joining and sharing their food discoveries. In a producer-scrounger (PS) system, foragers can either search for food themselves (play producer) or search for joining opportunities (play scrounger), but not both at the same time. Empirical evidence is accumulating to show that the joining decision of ground-feeding birds like nutmeg mannikins (Lonchura punctulata) can be modeled by a PS game. However some predictions remain to be tested. For instance, foragers are predicted to increase their use of the scrounger tactic as group size increases. Also, one consequence of the incompatibility between producer and scrounger tactics is that the per capita searching efficiency should decrease as the use of scrounger increases. I tested these predictions in an indoor aviary using four flocks of nutmeg mannikins. I manipulated the stable equilibrium frequency (SEF) of the scrounger tactic by varying group size and the finder's share. As predicted by PS games, birds increased their use of scrounger as group size increased. Also, the per capita interval between patch discoveries increased and the per capita finding rate decreased as conditions called for a higher SEF of scrounger. I discuss why the decreased searching efficiency observed likely follows from the incompatibility between producer and scrounger tactics rather than from artifacts of the conditions used or from any form of interference.     

A field investigation of scrounging in semipalmated sandpipers

Animals that forage in groups can produce their own food patches or scrounge the food discoveries of their companions. Mean tactic payoffs are expected to be the same at equilibrium for phenotypically equal foragers. Scrounging is also typically viewed as a risk-averse foraging strategy that provides a more even food intake rate over time. The occurrence of scrounging and the payoffs from different foraging modes have rarely been investigated in the field. Over two field seasons, I examined patch sharing in semipalmated sandpipers (Calidris pusilla) foraging on minute food items at the surface of the substrate. Birds could find patches on their own, a producing event, or join the food patches discovered by others, a scrounging event. I found that the average search time per patch did not differ between producing and scrounging but that the average time spent exploiting a patch was reduced nearly by half when scrounging. As a result, the proportion of time spent exploiting a patch, a measure of foraging payoffs, was significantly lower when scrounging. The variance in payoffs was similar for producing and scrounging. When producing their own patches, individuals that scrounged spent the same proportion of time exploiting a patch as those that only produced. However, within the same individuals, the search time for a scrounged patch was longer than the search time for a produced patch. The results show unequal payoffs for producing and scrounging in this system and suggest that low success in finding patches elicited scrounging.     

Division of labor between scouts and recruits: genetic influence and mechanisms

Every recruitment system in social insects requires some individuals that serve as scouts, foragers that search independently for food sources. It is not well understood which factors influence whether an individual becomes a scout or a recruit, nor how the division of labor between the two forager groups is regulated. It is shown here for honeybees (Apis mellifera), using two different molecular techniques, that there is a genetically based difference in the probability that individuals will scout independently for food. In contrast to earlier suggestions, experimental tests showed that the age of a bee does not seem to influence its probability of becoming a scout or a recruit. Furthermore, scout bees do not search opportunistically for either pollen or nectar but, rather, individuals have preferences that are genetically based. These findings are discussed in the framework of foraging regulation by specialization in honeybees and the adaptive significance of polyandry. Received: 23 October 1997 / Accepted after revision: 10 April 1998     

The effects of prior residence,competitive ability and food availability on the outcome of interactions between shrews (Sorex araneus L.)

Summary Interactions between pairs of shrews on a foraging grid were analysed to test for the effects of prior residence, food density and differences in competitive ability between individuals on the outcome.Prior residence was an important factor influencing the outcome of interactions but the degree of resident advantage varied both with food density on the grid and the difference in competitive ability between shrews.Payoff and resource holding power (R.H.P.) asymmetry effects can be implied from the influence of food density and competitive ability differences but prior residence may operate as an uncorrelated asymmetry only under certain conditions determined by combinations of the other two asymmetries.     

Social foraging and dominance relationships: the effects of socially mediated interference

In socially foraging animals, it is widely acknowledged that the position of an individual within the dominance hierarchy of the group has a large effect upon its foraging behaviour and energetic intake, where the intake of subordinates can be reduced through socially mediated interference. In this paper, we explore the effects of interference upon group dynamics and individual behaviour, using a spatially explicit individual-based model. Each individual follows a simple behavioural rule based upon its energetic reserves and the actions of its neighbours (where the rule is derived from game theory models). We show that dominant individuals should have larger energetic reserves than their subordinates, and the size of this difference increases when either food is scarce, the intensity of interference suffered by the subordinates increases, or the distance over which dominant individuals affect subordinates increases. Unlike previous models, the results presented in this paper about differences in reserves are not based upon prior assumptions of the effects of social hierarchy and energetic reserves upon predation risk, and emerge through nothing more than a reduction in energetic intake by the subordinates when dominants are present. Furthermore, we show that increasing interference intensity, food availability or the distance over which dominants have an effect also causes the difference in movement between ranks to increase (where subordinates move more than dominants), and the distance over which dominants have an effect changes the size of the groups that the different ranks are found in. These results are discussed in relation to previous studies of intra- and interspecific dominance hierarchies.     

Paying for information: partial loads in central place foragers

Information about food sources can be crucial to the success of a foraging animal. We predict that this will influence foraging decisions by group-living foragers, which may sacrifice short-term foraging efficiency to collect information more frequently. This result emerges from a model of a central-place forager that can potentially receive information on newly available superior food sources at the central place. Such foragers are expected to return early from food sources, even with just partial loads, if information about the presence of sufficiently valuable food sources is likely to become available. Returning with an incomplete load implies that the forager is at that point not achieving the maximum possible food delivery rate. However, such partial loading can be more than compensated for by an earlier exploitation of a superior food source. Our model does not assume cooperative foraging and could thus be used to investigate this effect for any social central-place forager. We illustrate the approach using numerical calculations for honeybees and leafcutter ants, which do forage cooperatively. For these examples, however, our results indicate that reducing load confers minimal benefits in terms of receiving information. Moreover, the hypothesis that foragers reduce load to give information more quickly (rather than to receive it) fits empirical data from social insects better. Thus, we can conclude that in these two cases of social-insect foraging, efficient distribution of information by successful foragers may be more important than efficient collection of information by unsuccessful ones.     

Individually different foraging methods in the desert ant Cataglyphis bicolor (Hymenoptera,Formicidae)

Summary Observations and field experiments on the foraging behaviour of individual workers of Cataglyphis bicolor in a Southern Tunisian shrub desert are reported. The workers search singly for their food (mostly animal carcasses) and are singleprey loaders. The individuals differ to a great extent in their persistence to re-search the place of a find on a previous foraging excursion. The differences range continuously from thoroughly researching a place to just walking by. If, in an experiment, the same reward is offered farther from the nest, each ant persists more in re-searching the place than if food is offered close to the nest. In a further experiment, some individuals persisted less in searching near the former finding site if they had collected a fly than after collecting a piece of cheese. There is, however, evidence that individuals do not differ in their food preference. Persistent individuals, which re-search the place of a former find, are faster than non-persistent ones in retrieving food that is experimentally arranged in an aggregated manner. The experiment failed to demonstrate the (reverse) superiority of non-persistent individuals foraging on homogeneously distributed food. The observations of unmanipulated foraging excursions in the field suggest such an advantage for non-persistent foragers under natural conditions where food in general occurs widely dispersed. The colony as a whole retrieves more food within the same time from an experimental lay-out that is homogeneous than from an aggregated one. The behavioural differences between individuals could be caused by a training bias of the short-lived foragers, leading to a different assessment of the profitability of a searching method which implies returning to a formerly rewarding place. Thus, each worker uses the most promising behaviour according to its individual experience. Alternatively, the individually different searching methods could mainly contribute to the welfare of the colony as a whole rather than leading to a maximal short-term efficiency of each individual. In particular, the colony, disposing of only a few highly persistent foragers, could quickly exploit occasional short-lived, but unpredictible, clumps of food within its foraging range.     

Collective foraging decision in a gregarious insect

Group foraging by eusocial insects implies sophisticated recruitment processes that often result in collective decisions to exploit the most profitable sources. These advanced levels of cooperation, however, remain limited to a small range of species, and we still know little about the mechanisms underlying group foraging behaviours in the great mass of animals exhibiting lower levels of social complexity. In this paper, we report, for the first time in a gregarious insect, the cockroach Blattella germanica (L.), a collective foraging decision whereby the selection of food sources is reached without requiring active recruitment. Groups of cockroaches given a binary choice between identical food sources exhibited exploitation asymmetries whose amplitude increases with group size. By coupling behavioural observations to computer simulations, we demonstrate that selection of food sources relies uniquely on a retention effect of feeding individuals on newcomers without comparison between available opportunities. This self-organised pattern presents similarities with the foraging dynamics of eusocial species, thus stressing the generic dimension of collective decision-making mechanisms based on social amplification rules despite fundamental differences in recruitment processes. We hypothesise that such parsimony could apply to a wide range of species and help understand the emergence of collective behaviours in simple social systems.     

Root plasticity buffers competition among plants: theory meets experimental data

Morphological plasticity is a striking characteristic of plants in natural communities. In the context of foraging behavior particularly, root plasticity has been documented for numerous species. Root plasticity is known to mitigate competitive interactions by reducing the overlap of the individuals' rhizospheres. But despite its obvious effect on resource acquisition, plasticity has been generally neglected in previous empirical and theoretical studies estimating interaction intensity among plants. In this study, we developed a semi-mechanistic model that addresses this shortcoming by introducing the idea of compensatory growth into the classical-zone-of influence (ZOI) and field-of-neighborhood (FON) approaches. The model parameters describing the belowground plastic sphere of influence (PSI) were parameterized using data from an accompanying field experiment. Measurements of the uptake of a stable nutrient analogue at distinct distances to the neighboring plants showed that the study species responded plastically to belowground competition by avoiding overlap of individuals' rhizospheres. An unexpected finding was that the sphere of influence of the study species Bromus hordeaceus could be best described by a unimodal function of distance to the plant's center and not with a continuously decreasing function as commonly assumed. We employed the parameterized model to investigate the interplay between plasticity and two other important factors determining the intensity of competitive interactions: overall plant density and the distribution of individuals in space. The simulation results confirm that the reduction of competition intensity due to morphological plasticity strongly depends on the spatial structure of the competitive environment. We advocate the use of semi-mechanistic simulations that explicitly consider morphological plasticity to improve our mechanistic understanding of plant interactions.     

Individual variation in winter foraging of black-capped chickadees

Summary Wintering black-capped chickadees (Paridae: Parus atricapillus) in northwestern Massachusetts showed a high degree of individual variation in foraging behavior. After accounting for the effects of different habitats and weather conditions, individual differences comprised 6–17% of the total observed variation in four measures of foraging location and rate of feeding. Differences between age and sex groups were not significant and explained comparatively little variation (0.0–1.4%). The chickadees did not fall into a few distinct behavioral categories but instead showed continuous variation on all measures of foraging behavior. It appeared that some variation among individuals was a consequence of behavioral convergence within social groups, since birds that were observed together were more similar in their foraging than expected by chance, after taking habitat differences into account. Our results therefore do not support the interpretation that individual variation in feeding behavior serves to reduce exploitation competition within social groups.     

Killer whale predation on marine mammals at Punta Norte,Argentina; food sharing,provisioning and foraging strategy

Summary The social dynamics of killer whales (Orcinus orca) that hunt marine mammals are apparently highly flexible, though strong individual associations do exist. The killer whales at Punta Norte offer an unusually detailed view of association patterns and foraging behaviour, and suggest a pattern of behaviour that optimizes hunting efficiency with exception only to strong associations between some individuals and the provisioning and training of offspring. The main points from this paper are as follows: First, hunting effort was concentrated where the capture rate was greatest. All pods selectively attacked the prey type for which they had the highest capture rate. The amount of southern sea lion prey captured was approximately equal to the estimated minimum energetic requirement for killer whales based on weight. Secondly, one whale in each pod did the majority of the hunting, and then provisioned the others in the pod. It was clear on numerous occasions that food was shared. A review of reported incidences of killer wales taking marine mammal prey suggests that it is common for a subset of the individuals in a pod to hunt. These results are discussed in the context of the evolution of foraging behaviour.Offprint requests to: A.R. Hoelzel at the first address     

Foraging in groups of water striders (Gerris remigis): effects of variability in prey arrivals and handling times

Summary In many species, socially subordinate individuals frequently remain in a group despite their lower priority of access to food and mates. This is expected to occur when the net benefits of staying in the group exceed those of a solitary existence. Analogously, territorial foragers surrounded by conspecific competitors are faced with tradeoffs as to patch tenacity. In either case, spatio-temporal variability in resource availability directly affects behavioral patterns and payoffs, particularly of individuals with low priority of access to the resource. However, such individuals may take advantage of natural environmental fluctuations in resource supply in cases where dominants are preoccupied. This arises regularly if two resource items cannot be handled simultaneously and if a second item arrives before handling of the first has been completed. I advance a heuristic model that predicts that foraging or mating success of individuals with low priority of access to resources may increase both with higher variance in inter-arrival times of the resource (given the same mean) and with an increase in the average handling time of the resource. I tested both predictions with two associations of individually marked, naturally foraging water striders (Gerris remigis) in the field. In natural streams individual water striders tend to occupy consistent positions that they may defend, resulting in priority of access to prey items floating downstream for individuals further up front. I manipulated the variance in prey inter-arrival times given the same mean, and the prey handling times by offering larger prey. The outcome was in qualitative agreement with the predictions of the model.     

Foraging tactics of two guilds of web-spinning spiders

Summary The foraging behavior of orbweaving and sheetweb weaving guilds of spiders was investigated. Orbweavers move among web-sites frequently, but sheetweb weavers do not. A simple model that examines the adaptive advantages of active foraging and sit-and-wait strategies leads to three predictions: 1) Orbweavers should have a simple decision rule for leaving web-sites, 2) Orbweavers' web-sites should have more variable payoffs than do the web-sites of sheetweb weavers, and 3) Orbweavers should have a lower cost of moving than do sheetweb weavers. Field and experimental data bear out each prediction. In addition, analysis of the residence times of spiders at web-sites that are used more than once reveals that members of the two guilds do not compete with each other for web-sites.The distinction between active foragers and sit-and-wait predators stems from the relative frequency of the decision to leave a foraging site. Thus active foragers are those predators that move often from patch to patch, and sit-and-wait predators are those that leave patches infrequently. In this sense, orbweaving spiders are active foragers and sheetweb weavers are sit-and-wait predators.     

Feeding and non-feeding aggression can be induced in invasive shore crabs by altering food distribution

Human activity can have a large impact on surrounding ecosystems. For example, humans alter resource distributions for other species, potentially modifying these species competitive dynamics. These changes in local competitive processes are frequently associated with species invasions. Here, we investigate how differences in resource distribution affect competitive behaviour using the highly invasive European shore crab (Carcinus maenas). Using a controlled laboratory experiment in combination with behaviour assays and social network analysis, we show that individuals feeding in habitats with clumped food distributions are more aggressive than individuals feeding in habitats where food is evenly dispersed, and this aggression is present even on days where crabs are not feeding. Additionally, this persistent aggression can be induced, suggesting that individuals of this invasive species possess the flexibility to modify their competitive behaviours in response to differences in food distributions. Furthermore, we show how these individual responses can lead to changes in overall organisation of aggressive interactions within a population. We discuss these results in relation to how human impacts can have long-term effects on competitive behavioural strategies, and how behavioural flexibility can allow invasive species to colonise and persist in highly impacted sites such as urban ecosystems.     

The honey bee shaking signal: function and design of a modulatory communication signal

This study explores the meaning and functional design of a modulatory communication signal, the honey bee shaking signal, by addressing five questions: (I) who shakes, (II) when do they shake, (III) where do they shake, (IV) how do receivers respond to shaking, and (V) what conditions trigger shaking. Several results confirm the work of Schneider (1987) and Schneider et al. (1986a): (I) most shakers were foragers (at least 83%); (II) shaking exhibited a consistent temporal pattern with bees producing the most signals in the morning (0810–1150 hours) just prior to a peak in waggle dancing activity; and (IV) bees moved faster (by 75%) after receiving a shaking signal. However, this study differs from previous work by providing a long-term, temporal, spatial, and vector analysis of individual shaker behavior. (III) Bees producing shaking signals walked and delivered signals in all areas of the hive, but produced the most shaking signals directly above the waggle dance floor. (IV) Bees responded to the signal by changing their direction of movement. Prior to receiving a signal, bees selected from the waggle dance floor moved, on average, towards the hive exit. After receiving a signal, some bees continued moving towards the exit but others moved directly away from the exit. During equivalent observation periods, non-shaken bees exhibited a strong tendency to move towards the hive exit. (V) Renewed foraging activity after food dearth triggered shaking signals, and, the level of shaking is positively correlated with the duration of food dearth. However, shaking signal levels also increased in the morning before foraging had begun and in the late afternoon after foraging had ceased. This spontaneous afternoon peak has not previously been reported. The shaking signal consequently appears to convey the general message “reallocate labor to different activities” with receiver context specifying a more precise meaning. In the context of foraging, the shaking signal appears to activate (and perhaps deactivate) colony foraging preparations. The generally weak response elicited by modulatory signals such as the shaking signal may result from a high receiver response threshold which allows the receiver to integrate multiple sources of information and which thereby increases the probability that receiver actions will be appropriate to colony needs. Received: 21 March 1997 / Accepted after revision: 30 August 1997     

The influence of kinship on foraging competition in Siberian jays

Foraging competition in Siberian jay groups was examined in relation to dominance and kinship to determine whether juvenile offspring, by associating with adults, gained in food acquisition relative to juvenile immigrants. Members of the adult pair were dominant over juvenile cohort members and males were dominant to females, although an inter-sexual hierarchy, with male juveniles occasionally overlapping adult females, was suggested. Few competitive asymmetries were found between adults and retained offspring or adults and immigrant juveniles when they were competing for food together, but in kin and non-kin foraging groups, respectively. Male offspring visited the bait site more frequently than adult males, and female immigrants spent less time at the bait site than adult females. Under these circumstances, hoarding activities may limit the ability of alpha members to control resources. In mixed groups containing both juvenile offspring and juvenile immigrants, no difference was found in the number of visits made to the bait site, although load sizes and foraging rates were lower for immigrant birds. Retained juveniles obtained greater load sizes and foraging rates when associating with adults. The social dominance of parents suggests that they control juvenile foraging. Although offspring benefit in the presence of adults, adults may incur a cost to their restraint by spending more time at the bait site when competing with immigrants. These results extend conclusions from previous work describing the role of selective tolerance by adults which relaxes competition with retained offspring in Siberian jay winter groups. The present findings suggest that offspring benefit in both immediate and future energy gains, which may have a direct influence on survival. Received: 18 September 1996 / Accepted after revision: 26 January 1997     

Effects of aggregation on feeding and survival in a communal wren

Summary Foraging by a social wren, Campylorhynchus nuchalis (Troglodytidae), in a tropical savanna habitat is not enhanced by aggregation. Data for marked individuals show that solitary foraging results in a higher capture rate than foraging near others. We find no evidence of imitative foraging, as individuals actively avoid successful foragers following a capture and successful foragers do not restrict their search to recently productive stations or techniques. Captures are seldom temporally clumped, and clumping is probably not pronounced enough to favor imitation. Juveniles show no greater tendency to respond to captures of others, or to succeed in foraging in a group, than do adults. Aggregation is probably disadvantageous for foraging because of dispersed, scarce, cryptic, and noneruptive prey and because of the searching technique of these foliage-gleaning insectivores. If predator avoidance is enhanced by aggregation, it does not result in either increased survival or increased foraging efficiency in large groups, even by juveniles.