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.     

Past experiences and future expectations generate context-dependent costs of foraging

Because environments can vary over space and time in non-predictable ways, foragers must rely on estimates of resource availability and distribution to make decisions. Optimal foraging theory assumes that foraging behavior has evolved to maximize fitness and provides a conceptual framework in which environmental quality is often assumed to be fixed. Another more mechanistic conceptual framework comes from the successive contrast effects (SCE) approach in which the conditions that an individual has experienced in the recent past alter its response to current conditions. By regarding foragers’ estimation of resource patches as subjective future value assessments, SCE may be integrated into an optimal foraging framework to generate novel predictions. We released Allenby’s gerbils (Gerbillus andersoni allenbyi) into an enclosure containing rich patches with equal amounts of food and manipulated the quality of the environment over time by reducing the amount of food in most (but not all) food patches and then increasing it again. We found that, as predicted by optimal foraging models, gerbils increased their foraging activity in the rich patch when the environment became poor. However, when the environment became rich again, the gerbils significantly altered their behavior compared to the first identical rich period. Specifically, in the second rich period, the gerbils spent more time foraging and harvested more food from the patches. Thus, seemingly identical environments can be treated as strikingly different by foragers as a function of their past experiences and future expectations.     

Vicarious sampling: the use of personal and public information by starlings foraging in a simple patchy environment

Group foragers may be able to assess patch quality more efficiently by paying attention to the sampling activities of conspecifics foraging in the same patch. In a previous field experiment, we showed that starlings foraging on patches of hidden food could use the successful foraging activities of others to help them assess patch quality. In order to determine whether a starling could also use another individual’s lack of foraging success to assess and depart from empty patches more quickly, we carried out two experimental studies which compared the behaviour of captive starlings sampling artificial patches both when alone and when in pairs. Solitary starlings were first trained to assess patch quality in our experimental two-patch system, and were then tested on an empty patch both alone and with two types of partner bird. One partner sampled very few holes and thus provided a low amount of public information; the other sampled numerous holes and thus provided a high amount of public information. In experiment 1, we found no evidence of vicarious sampling. Subjects sampled a similar number of empty holes when alone as when with the low and high information partners; thus they continued to rely on their own personal information to make their patch departure decisions. In experiment 2, we modified the experimental patches, increasing the ease with which a bird could watch another’s sampling activities, and increasing the difficulty of acquiring accurate personal sampling information. This time, subjects apparently did use public information, sampling fewer empty holes before departure when with the high-information partner than when with the low-information partner, and sampling fewer holes when with the low-information partner than when alone. We suggest that the degree to which personal and public information are used is likely to depend both on a forager’s ability to remember where it has already sampled and on the type of environment in which foraging takes place. Received: 31 January 1995/Accepted after revision: 11 September 1995     

Assessment of predation risk via illumination level: facultative central place foraging in the cricetid rodent Phyllotis darwini

It is well known that the risk of predation affects prey decision making. However, few studies have been concerned with the cues used by prey to assess this risk. Prey animals may use indirect environmental cues to assess predation hazard since direct evaluation may be dangerous. I studied the assessment of predation risk, manipulated via environmental illumination level, and the trade-off between foraging and predation hazard avoidance in the nocturnal rodentPhyllotis darwini (Rodentia: Cricetidae). In experimental arenas I simulated dark and full moon nights (which in nature correlate with low and high predation risk, respectively) and measured the immediate responses of animals to flyovers of a raptor model. Second, varying illumination only, I evaluated patch use, food consumption, central place foraging, and nocturnal variation of body weight. During flyover experiments, animals showed significantly more evasive reactions under full moon illumination than in moonless conditions. In the patch use experiments, rodents significantly increased their giving-up density and decreased their total food consumption under moonlight. On dark nights, rodents normally fed in the food patch, but when illumination was high they became central place foragers in large proportion. Moreover, the body weight of individuals decreased proportionately more during bright nights. These results strongly suggest thatP. darwini uses the level of environmental illumination as a cue to the risk of being preyed upon and may sacrifice part of its energy return to avoid risky situations.     

A patch use model to separate effects of foraging costs on giving-up densities: an experiment with white-tailed deer (<Emphasis Type="Italic">Odocoileus virginianus</Emphasis>)

The giving-up density of food (GUD), the amount of food remaining in a patch when a forager ceases foraging there, can be used to compare the costs of foraging in different food patches. But, to draw inferences from GUDs, specific effects of foraging costs (predation risk, metabolic and missed opportunities costs) on GUDs have to be identified. As high predation risk, high metabolic costs and abundant food all should produce high GUDs, this does not allow us to infer directly the quality of a habitat. In order to separate the effect of each foraging cost, we developed an optimal foraging model based on food supplementation. We illustrate the use of our model in a study where we assessed the impact of a power line right-of-way in a white-tailed deer (Odocoileus virginianus) winter yard by determining whether the negative effects of cover loss outweigh the positive effects of browse regeneration.     

Movement of foraging Tundra Swans explained by spatial pattern in cryptic food densities

We tested whether Tundra Swans use information on the spatial distribution of cryptic food items (below ground Sago pondweed tubers) to shape their movement paths. In a continuous environment, swans create their own food patches by digging craters, which they exploit in several feeding bouts. Series of short (<1 m) intra-patch movements alternate with longer inter-patch movements (>1 m). Tuber biomass densities showed a positive spatial auto-correlation at a short distance (<3 m), but not at a larger distance (3-8 m). Based on the spatial pattern of the food distribution (which is assumed to be pre-harvest information for the swan) and the energy costs and benefits for different food densities at various distances, we calculated the optimal length of an inter-patch movement. A swan that moves to the patch with the highest gain rate was predicted to move to the adjacent patch (at 1 m) if the food density in the current patch had been high (>25 g/m2) and to a more distant patch (at 7-8 m) if the food density in the current patch had been low (<25 g/m2). This prediction was tested by measuring the response of swans to manipulated tuber densities. In accordance with our predictions, swans moved a long distance (>3 m) from a low-density patch and a short distance (<3 m) from a high-density patch. The quantitative agreement between prediction and observation was greater for swans feeding in pairs than for solitary swans. The result of this movement strategy is that swans visit high-density patches at a higher frequency than on offer and, consequently, achieve a 38% higher long-term gain rate. Swans also take advantage of spatial variance in food abundance by regulating the time in patches, staying longer and consuming more food from rich than from poor patches. We can conclude that the shape of the foraging path is a reflection of the spatial pattern in the distribution of tuber densities and can be understood from an optimal foraging perspective.     

Searching behavior and use of sampling information by free-ranging bison (Bos bison)

I examined the searching behavior of free-ranging plains bison (Bos bison bison) in their natural habitat, and determined whether their assessment of food patch quality was influenced by the short-term sampling information acquired during search. Bison used area-concentrated search during their winter foraging activity. Their movements between areas of suitable food patches were influenced by local environmental conditions, being sometimes less sinuous, and at other times more sinuous, than expected from a correlated random walk model. Bison also systematically avoided digging in areas where plants of low profitability lay under the snow. Where they dug, there was evidence that a bison's perception of food quality varied during a foraging bout, and was therefore influenced by short-term sampling information. After controlling for forage quality, I found that small feeding craters were more likely to be preceded by samples of high quality food patches. My observations suggest that bison take advantage of the structural characteristics of their environment during searching activity, and base foraging decisions on local rather than global availability.     

Learning during competitive positioning in the nest: do nestlings use ideal free ‘foraging’ tactics?

Begging behaviour of nestling birds may involve more than a simple, honest source of information for parents to use in provisioning. Many aspects of begging behaviour relate instead to sibling competition for food items within the nest, and we might expect evidence of adaptive learning and behavioural adjustment in response to experience of the competitive environment. In this study, we consider begging in different locations within the nest as analogous to foraging in different patches, varying in food availability. Using hand-feeding trials, we created zones of differing profitability within an artificial nest by adjusting either the prey size or number of items delivered, and allowed only indirect competition between pairs of southern grey shrike (Lanius meridionalis) nestlings. Nestlings demonstrated the ability to detect differences in zone profitability and position themselves accordingly. By the end of both the prey size and delivery rate trials nestlings had increased the amount of time spent in the high quality zone. Such movement in response to differences in load quality, as well as frequency, demonstrates the ability of nestlings to learn about their environment and to facultatively adjust their begging in order to maximise energetic rewards.     

Distance to neighbours influences the trade-off between hiding after disturbance and defending food patches in convict cichlids (<Emphasis Type="Italic">Archocentrus nigrofasciatus</Emphasis>)

Territories are often aggregated. Because of this, distance to neighbours should influence how territory-holders balance safety from predators with the use and defence of resources. I examined the influence of distance to a neighbour on refuge use by pairs of convict cichlids (Archocentrus nigrofasciatus) faced with a conflict between hiding and defending food patches. Neighbours could reduce the rate of intrusions by strangers as a by-product of their own resource defence. This should allow fish with near neighbours to spend more time in the refuge. Neighbours could also steal from patches that are left undefended. This should lead to a reduction in use of the refuge. When one fish was confined to its refuge (so that its patch was undefended), theft by the other increased as inter-patch distance decreased. Distance between patches did not influence the rate of intrusion by non-territorial fish. When both fish defended patches, body mass influenced the effect of inter-patch distance on refuge use. Large fish rarely used the refuge, but small territory-holders spent more time in the refuge when patches were close together, as predicted. However, when one fish was dominant at both patches, distance between patches did not influence refuge use. These results suggest that, despite increased opportunity for theft, there is no realised foraging or defensive benefit to settling near neighbours that are of similar competitive ability.Communicated by J. Krause     

Behavioural responses to indirect and direct predator cues by a mammalian herbivore, the common brushtail possum

Cues for detecting and responding to perceived predation risk may be indirect, i.e., correlated with the probability of encountering a predator, or direct, i.e., produced by or related to the actual presence of a predator. Research shows, independently, both types of cues can influence anti-predator and foraging behaviours in prey species. However, since animals naturally encounter indirect and direct cues simultaneously, we were interested in quantifying their cumulative effect. Our aim was to evaluate food intake and behaviours (patch use, feeding (rate and time), vigilance) of a nocturnal mammalian herbivore to indirect (open vs. covered microhabitats; illumination) and direct (fox/owl odours) predator cues. We ran a preference trial with four paired treatments using a covered Safe food patch and an open Risk food patch, with one of four combinations of indirect and direct predator cues. Predation risk had a significant effect on both intake and behaviour (including feeding time, rate, and vigilance), but these effects differed depending on cues. No two combinations of cues produced exactly the same effects, illustrating the complexity of interactions that occur between cues. Covered patches were always perceived as less risky than open patches, but unexpectedly, open patches were perceived as riskier when dark rather than light. The strongest suite of (negative) responses to risk was associated with combined indirect and direct cues. These results highlight the importance of considering jointly, intake from a patch, intake rate, and behaviours, such as the proportion of time spent vigilant, when quantifying predation risk, rather than intake alone.     

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     

Predation risk and foraging behavior of the hoary marmot in Alaska

Summary I observed hoary marmots for three field seasons to determine how the distribution of food and the risk of predation influenced marmots' foraging behavior. I quantified the amount of time Marmota caligata foraged in different patches of alpine meadows and assessed the distribution and abundance of vegetation eaten by marmots in these meadows. Because marmots dig burrows and run to them when attacked by predators, marmot-toburrow distance provided an index of predation risk that could be specified for different meadow patches.Patch use correlated positively with food abundance and negatively with predation risk. However, these significant relationships disappeared when partial correlations were calculated because food abundance and risk were intercorrelated. Using multiple regression, 77.0% of the variance in patch use was explained by a combination of food abundance, refuge burrow density, and a patch's distance from the talus where sleeping burrows were located. Variations in vigilance behavior (look-ups to search for predators while feeding) according to marmots' ages, the presence of other conspecifics, and animals' proximity to their sleeping burrows all indicated that predation risk influenced foraging.In a forage-manipulation experiment, the use of forage-enhanced patches increased six-fold, verifying directly the role of food availability on patch used. Concomitant with increased feeding, however, was the intense construction of refuge burrows in experimental patches that presumably reduced the risk of feeding. Thus, I suggest that food and predation risk jointly influence patch use by hoary marmots and that both factors must be considered when modeling the foraging behavior of species that can be predator and prey simultaneously.     

Indirect effects of prey swamping: differential seed predation during a bamboo masting event

Resource pulses often involve extraordinary increases in prey availability that "swamp" consumers and reverberate through indirect interactions affecting other community members. We developed a model that predicts predator-mediated indirect effects induced by an epidemic prey on co-occurring prey types differing in relative profitability/preference and validated our model by examining current-season and delayed effects of a bamboo mass seeding event on seed survival of canopy tree species in mixed Patagonian forests. The model shows that predator foraging behavior, prey profitability, and the scale of prey swamping influence the character and strength of short-term indirect effects on various alternative prey. When in large prey-swamped patches, nonselective predators decrease predation on all prey types. Selective predators, instead, only benefit prey of similar quality to the swamping species, while very low or high preference prey remain unaffected. Negative indirect effects (apparent competition) may override such positive effects (apparent mutualism), especially for highly preferred prey, when prey-swamped patches are small enough to allow predator aggregation and/or predators show a reproductive numerical response to elevated food supply. Seed predation patterns during bamboo (Chusquea culeou) masting were consistent with predicted short-term indirect effects mediated by a selective predator foraging in large prey-swamped patches. Bamboo seeds and similarly-sized Austrocedrus chilensis (ciprés) and Nothofagus obliqua (roble) seeds suffered lower predation in bamboo flowered than nonflowered patches. Predation rates on the small-seeded Nothofagus dombeyi (coihue) and the large-seeded Nothofagus alpina (rauli) were independent of bamboo flowering. Indirect positive effects were transient; three months after bamboo seeding, granivores preyed heavily upon all seed types, irrespective of patch flowering condition. Moreover, one year after bamboo seeding, predation rates on the most preferred seed (rauli) was higher in flowered than in nonflowered patches. Despite rapid predator numerical responses, short-term positive effects can still influence community recruitment dynamics because surviving seeds may find refuge beneath the litter produced by bamboo dieback. Together, our theoretical analysis and experiments indicate that indirect effects experienced by alternative prey during and after prey-swamping episodes need not be universal but can change across a prey quality spectrum, and they critically depend on predator-foraging rules and the spatial scale of swamping.     

Look before you leap: is risk of injury a foraging cost?

Theory states that an optimal forager should exploit a patch so long as its harvest rate of resources from the patch exceeds its energetic, predation, and missed opportunity costs for foraging. However, for many foragers, predation is not the only source of danger they face while foraging. Foragers also face the risk of injuring themselves. To test whether risk of injury gives rise to a foraging cost, we offered red foxes pairs of depletable resource patches in which they experienced diminishing returns. The resource patches were identical in all respects, save for the risk of injury. In response, the foxes exploited the safe patches more intensively. They foraged for a longer time and also removed more food (i.e., had lower giving up densities) in the safe patches compared to the risky patches. Although they never sustained injury, video footage revealed that the foxes used greater care while foraging from the risky patches and removed food at a slower rate. Furthermore, an increase in their hunger state led foxes to allocate more time to foraging from the risky patches, thereby exposing themselves to higher risks. Our results suggest that foxes treat risk of injury as a foraging cost and use time allocation and daring—the willingness to risk injury—as tools for managing their risk of injury while foraging. This is the first study, to our knowledge, which explicitly tests and shows that risk of injury is indeed a foraging cost. While nearly all foragers may face an injury cost of foraging, we suggest that this cost will be largest and most important for predators.     

Current indirect fitness benefits associated with philopatry in juvenile prairie voles

Summary Prairie vole (Microtus ochrogaster) family groups were examined to determine possible indirect fitness benefits from the presence of juvenile helpers at the natal nest. The reproductive performance of family groups retaining two juveniles was compared with that of families in which all juveniles were removed when the subsequent litter was born. Litter sizes at birth of the second litter and pup survival rates were the same in the two treatments. Offspring quality was affected by the presence of juveniles however. Pups reared with juveniles weighed 13% more at weaning than pups reared without juveniles. Pups also opened their eyes sooner when juveniles were present. Differences in growth and development may have been affected by the amount of time pups were alone in the nest since pups in families with juveniles were left alone less frequently than were pups without juveniles. Maternal behavior patterns were not affected by the presence of juveniles. In contrast, fathers in families with juveniles spent more time in non-parental behaviors such as feeding, drinking, and foraging. In families with large litters, mothers delivered a subsequent litter sooner if juveniles were present. Subsequent litter sizes were the same in both treatments. Overall, both infants and parents benefited from the presence of juveniles, suggesting that helping may enhance the helper's indirect fitness in multiple ways.     

Food exploitation by a winter flock of greylag geese: behavioral dynamics,competition and social status

Summary To investigate the dynamics of the winter flock patch exploitation, feeding experiments were performed with 140 semi-tame, free-roaming greylag geese (Anser anser). Three different initial densities of barley were offered on a 50-m² patch: low (1600 grains/m²), intermediate (3300/m²) and high (I 1500/m²). Goose numbers on the patch, peck rates, and frequencies of agonistic encounters and of alert postures were observed by scanning the flock and in focal individuals. At low and intermediate initial food densities, peck rates decreased with food density, whereas at high food density, peck rates decreased only slightly over the feeding bout. Agonistic interactions increased as food decreased. A switch from exploitation to interference competition occurred at a threshold of approximately 900 remaining grains per square meter. With high initial food density, agonistic encounters began to increase gradually after 20–30 min. The same general patterns were observed for frequency of alertness. Family members occupied the patch for the longest time periods. The family female and the offspring fed most intensely. The family gander however, fed significantly less but was more aggressive and vigilant than all other social categories. Solitary geese spent their time on the patch doing little else but feeding and were the first to leave. It seems that high-ranking families and low-ranking singles adopt different competitive strategies, the former being superior interference competitors whereas the latter are forced scramble competitors.This study is dedicated to the memory of Konrad Lorenz Correspondence to: K. Kotrschal     

Trunk trails and the searching strategy of a leaf-cutter ant,Atta colombica

Summary Atta colombica uses chemical mass recruitment that allows the rapid exploitation of resources. Most foragers thus search only within patches. Accumulation of extra foragers at patches results in sampling of alternate food items and area-restricted search as patch resources are depleted.Individual workers have a higher probability of removing a leaf fragment the earlier they arrive at a bait. Workers that arrive when much of the resource is gone travel further on the bait (within the patch) but do not spend significantly more time at the patch. They give up after 50–80s.Foraging effort is centered on the extensive trail system, not on the nest a predicted by time and energy foraging models. Search effort is also trail centered. The probability that an item will be discovered decreases with distance from the trail and increasing litter depth. Trail traffic and trail quality together mave no significant effect although this may be because they act antagonistically.Economic considerations predict that trials should be built to high quality and very productive sites. If trails are built as a result of recruitment and recruitment reflects patch quality and productivity, characteristics of forage sites are physically embodied in the trail system.Leaf cutter foraging is better understood as a long term optimization that effectively exploits resources over the lifetime of the colony than as prudent predation that husbands resources.     

Males are faster foragers than females: intersexual differences of foraging behaviour in the Apennine chamois

Availability of food resources and individual characteristics can influence foraging behaviour, which can differ between males and females, leading to different patterns of food/habitat selection. In dimorphic species, females are usually more selective in food choice, show greater bite rates and spend more time foraging than males. We evaluated sexual differences in foraging behaviour in Apennine chamois Rupicapra pyrenaica ornata, during the warm season, before the rut. Both sexes selected nutritious vegetation patches and spent a comparable amount of time feeding. However, males had a significantly greater feeding intensity (bite rate) and a lower search effort for feeding (step rate), as well as they spent more time lying down than females. Females selected foraging sites closer to refuge areas than males. In chamois, sexual size dimorphism is seasonal, being negligible in winter–spring, but increasing to 30–40 % in autumn. Our results suggest that males enhance their energy and mass gain by increasing their food intake rate during the warm season, to face the costs of the mating season (November). Conversely, females seem to prioritize a fine-scale selection of vegetation and the protection of offspring. A great food intake rate of males in the warm season could have developed as a behavioural adaptation leading herbivores to the evolutionary transition from year-round monomorphism to permanent dimorphism, through seasonal dimorphism.     

Towards an ecological solution to the folivore paradox: patch depletion as an indicator of within-group scramble competition in red colobus monkeys (Piliocolobus tephrosceles)

A number of socioecological models assume that within-group food competition is either weak or absent among folivorous primates. This assumption is made because their food resources are presumed to be superabundant and evenly dispersed. However, recent evidence increasingly suggests that folivore group size is food-limited, that the primates prefer patchily distributed high-quality foods, and display some of the expected responses to within-group scramble competition. To investigate this apparent contradiction between theoretical models and recent empirical data, we examined the foraging behaviour of red colobus monkeys (Piliocolobus tephrosceles) in Kibale National Park, Uganda. We found that red colobus monkeys foraged in a manner that suggests they deplete patches of preferred foods: intake rate slowed significantly during patch occupancy while movement rate, an index of foraging effort, increased. Furthermore, patch occupancy was related to the size of the feeding group and the size of the patch. These results suggest that within-group scramble competition occurs, may limit folivore group size, and should be considered in models of folivore behavioural ecology.     

Female siskins choose mates by the size of the yellow wing stripe

Commonly, female birds use the brightly coloured patches on males to choose the best-quality mates. Coloured wing patches, however, have received little attention or have been previously related to social behaviour (as a signal to recruit conspecific individuals at feeding patches) or foraging (to flush prey) contexts, rather than to sexual selection. Here we provide evidence that in siskins (Carduelis spinus), wing patches function in mate choice. Mate-choice experiments showed that females were attracted by the size of the yellow wing stripe of the male, but not by the size of its black bib, body size, general plumage brightness or age. Experiments on birds with manipulated yellow wing stripes showed that females were sensitive to the size of this colour patch, irrespective of other male qualities. The preference of female siskins for males with larger wing patches when searching for a mate may be explained by the relationship of this trait to foraging ability, which would ensure females good parental investment from the chosen male.Communicated by W.A. Searcy