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.     

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.     

Giving-updensity and dietary shifts in the white-footed mouse, Peromyscus leucopus

Dietary shifts are commonly exhibited by omnivorous consumers when foraging from variable food resources. One advantage of dietary shifts for a consumer is the ability to gain complementary resources from different foods. In addition, dietary shifts often affect food-web dynamics. Despite the importance of dietary shifts to organismal, community, and ecosystem ecology, empirical studies of the ecological mechanisms that control dietary shifts are limited in scope. In this study, we tested the effects of complementary resources on dietary shifts of an omnivorous mammal, the white-footed mouse Peromyscus leucopus, in the context of depletable food patches in the natural environment. We used two complementary resources: seeds that provide a higher energy gain per unit handling time and mealworms that provide a higher protein gain per unit handling time. Stable isotopes of carbon and nitrogen (delta13C, delta15N) in mouse plasma were used to quantify dietary shifts, and we determined giving-up density (GUD), the food density at which a forager leaves a food patch (for an optimal forager, it should correspond to the quitting harvest rate that balances net fitness gain with various costs of foraging). The results showed that GUD increased most significantly when a mixture of seeds and mealworms was added, compared to when only seeds or mealworms were added. This suggests that, given a similar level of food availability, a patch with a mixture of complementary foods is of higher quality than a patch with only one type of food. Moreover, GUD measured with seeds (GUDs) correlated positively with seed availability, and GUD measured with mealworms (GUDmw) correlated positively with mealworm availability, indicating that the marginal value of seeds or mealworms decreases with their relatively availability in the environment. As GUDs increased, P. leucopus shifted their diets toward higher trophic levels, and as GUDmw increased, P. leucopus shifted their diets toward lower trophic levels, suggesting dietary shifts driven by food complementarity. This study demonstrated that the combination of giving-up density and stable-isotope analysis holds great potential for testing ecological mechanisms underlying dietary shifts.     

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.     

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.     

Spatial position and feeding success in ring-tailed coatis

The location of an animal within a social group has important effects on feeding success. When animals consume quickly eaten food items, individuals located at the front edge of a group typically have greater foraging success. When groups feed at large clumped resources, dominant individuals can often monopolize the resource, leading to higher feeding success in the center of the group. In order to test these predictions, behavioral data relating foraging success to within-group spatial position were recorded from two habituated groups of ring-tailed coatis (Nasua nasua) in Iguazu, Argentina. Foraging success did not fit expected patterns. When feeding on small ground litter invertebrates, coatis had the same foraging success at all spatial positions. This pattern likely resulted from an abundance of invertebrates in the ground litter. When feeding on fruit, individuals in the front of the group had greater feeding success, which was driven by the relatively quick depletion of fruit trees. Dominant juveniles were often located in the front of the group which led to increased access to food. This resulted in higher feeding success on fruits but simultaneously increased their risk of predation. Although groups typically became more elongated and traveled faster when feeding on fruit, it did not appear that the coatis were drastically changing their spacing strategies when switching between the two food types. Paradoxically, spatial position preferences during invertebrate foraging appeared to be driven by fruit trees. Because fruit trees were encountered so frequently, juveniles ranging at the front edge of the group during invertebrate foraging were the first to arrive at fruit trees and thus had higher foraging success. This study demonstrates the importance of how food patch size and depletion rate affect the spatial preferences of individuals.     

Autotomy shapes the trade-off between seeking cover and foraging in larval damselflies

Animals commonly choose between microhabitats that differ in foraging return and mortality hazard. I studied the influence of autotomy, the amputation of a body part, on the way larvae of the damselfly Lestes sponsa deal with the trade-off between foraging or seeking cover. Survival of Lestes larvae when confronted with the odonate predator Aeshna cyanea was higher in a complex than in a simple microhabitat, indicating that this more complex microhabitat was safer. Within the simple microhabitat, larvae without lamellae had a higher risk for mortality by predation than larvae with lamellae, showing a long-term cost of autotomy. When varying the foraging value (food present or absent) and predation risk (encaged predator or no predator) in the simple microhabitat, larvae with and without lamellae responded differentially to the imposed trade-off. All larvae spent more time in the simple microhabitat when food was present than when food was absent. Larvae without lamellae, however, only sporadically left the safe microhabitat, irrespective of the presence of the predator. In contrast, larvae with lamellae shifted more frequently towards the risky microhabitat than those without lamellae, and more often in the absence than in the presence of the predator. These decisions affected the foraging rates of the animals. I show for the first time that refuge use is higher after autotomy and that this is associated with the cost of reduced foraging success. The different microhabitat preferences for larvae with and without lamellae are consistent with their different vulnerabilities to predation and demonstrate the importance of intrinsic factors in establishing trade-offs. Received: 4 June 1999 / Received in revised form: 18 August 1999/ Accepted: 18 August 1999     

Is the antipredatory response in behaviour reflected in stress measured in faecal corticosteroids in a small rodent?

Predation risk has been shown to alter various behaviours in prey. Risk alters activity, habitat use and foraging, and weight decrease might be a consequence of that. In mammals, studies on physiological measures affected by risk of predation, other than weight, are rare. We studied in two separate laboratory experiments foraging, hoarding behaviour and expression of stress measured non-invasively from the faeces in the bank vole (Clethrionomys glareolus), a common boreal rodent. Voles were exposed to predation risk using odours of the least weasels (Mustela nivalis nivalis). Distilled water served as control. In the first experiment, we found that foraging effort, measured as sunflower seeds taken from seed trays filled with sand, was significantly lower in trays scented with weasel odour. Both immediate consumption of seeds and hoarding were affected negatively by the weasel odour. Females hoarded significantly more than males in autumn. In the second experiment, the negative effect of weasel odour on foraging was consistent over a 3-day experiment, but the strongest effect was observed in the first night. Foraging increased over the time of the experiment, which might reflect either energetic compensation during a longer period of risk, predicted in the predation risk allocation hypothesis, or habituation to the odour-simulated risk. Despite decreased foraging under predation risk, stress measured as corticosteroid metabolite concentration in vole faeces was not affected by the weasel odour treatment. In conclusion, we were able to verify predation-risk-mediated changes in the foraging effort of bank voles but no physiological stress response was measured non-invasively, probably due to great individual variation in secretion of stress hormones.     

Travel time affects optimal diets in depleting patches

Models of prey choice in depleting patches predict an expanding specialist strategy: Animals should start as specialists on the most profitable prey and then at some point during patch exploitation switch to a generalist foraging strategy. When patch residence time is long, the switch to a generalist diet is predicted to occur earlier than when patch residence time is short. We tested these predictions under laboratory conditions using female parasitoids (Aphidius colemani) exploiting patches of mixed instars aphid hosts (Myzus persicae, L1 and L4). The duration of patch exploitation was manipulated by changing travel time between patches. As predicted, patch residence times increase with travel time between patches. Our results provide empirical support for the expanding specialist prediction: Parasitoid females specialized initially on the more profitable hosts (L4), and as the patch depleted, they switched to a generalist diet by accepting more frequently the less profitable hosts (L1). The point at which they switched from specialist to generalist occurred later when travel times and hence patch residence times were short. By affecting the patch exploitation strategy, travel time also determines the composition of hosts left behind, the “giving up composition.” The change in the relative density of remaining host types alters aphid populations’ age structure.     

Costs and benefits of home range shifts among vervet monkeys (Cercopithecus aethiops) in Amboseli National Park,Kenya

Summary A decline in the density of trees and the deterioration of the habitat of vervet monkeys (Cercopithecus aethiops) in Amboseli National Park, Kenya, have been accompanied by movement of vervet groups into unfamiliar areas. These home range shifts were documented over a period of 26 months. The largest group moved into the home ranges of neighboring groups and acquired more trees, a resource that had become more limited. The groups whose home ranges were encroached upon did not share their home ranges with the intruders, but instead moved into areas that were unoccupied and had a lower density of trees. The largest group was thus more successful than smaller groups in competition for limited resources. For all but one group, movement into new areas was directly associated with the disappearances of female and immature monkeys. Most of the disappearances were attributable to predation. These results suggest that the cost of predation in unfamiliar areas may contribute to the evolution of philopatry in vervets. However, under unstable conditions, resource competition forces these animals to disperse (in groups) despite the high risk of predation. Offprint requests to: L.A. Isbell     

Subhabitat selection by foraging threespine stickleback (<Emphasis Type="Italic">Gasterosteus aculeatus</Emphasis>): previous experience and social conformity

Any mechanism that allows animals to increase their foraging efficiency is likely to be selected for, including the ability to learn to recognise and subsequently discriminate between habitat types based on their profitability. In a series of laboratory studies, we manipulated prey densities across two different experimental subhabitats and demonstrated that threespine stickleback (Gasterosteus aculeatus) can develop foraging preferences for subhabitats that have previously yielded prey. Fish were not recalling the spatial location of prey patches; rather, they were discriminating between subhabitats based on foraging experience there and allocating foraging effort accordingly. Foraging preferences took around 14 days to develop, and once established, they persisted independently of experimental prey density, suggesting that fish were using experience rather than real-time sampling to select foraging grounds. When we presented focal fish with social information cues, we found that they preferentially used local enhancement and current public information cues when they conflicted with previous experience, but that they did not use prior public information. This suggests that in the presence of conspecifics, individuals prioritise social conformity over the use of private information. We discuss our results in the context of optimal foraging and the trade-offs associated with balancing conflicting private and social information.     

Travel duration,energetic expenditure,and patch exploitation in the parasitic wasp <Emphasis Type="Italic">Venturia canescens</Emphasis>

Foraging animals usually keep track of how costly it is to reach new resource patches and adjust patch residence time and exploitation rate accordingly. There are at least two potential factors, which are not necessarily closely linked, that animals could measure to estimate costs of traveling: the time the forager needs to reach the next patch and the amount of energy it has to invest until arrival. In the parasitoid wasp Venturia canescens, females forage for hosts from which their offspring can develop. Two different types of this parasitoid exist. The thelytokous type lives in anthropogenic habitats where flight is not necessarily linked with foraging. The arrhenotokous type lives under field conditions and shows frequent flight activity. We tested whether the wasps would use time or energy needed to assess patch availability, by either confining them into vials or letting them travel actively in a flight mill between patch visits. Our results show that in thelytokous lines, time is a sufficient cue influencing patch exploitation and an additional effect of the energy needed was not visible. In the arrhenotokous wasps, however, only the number of rounds flown in the mill influenced subsequent behavior, while mere time spent traveling did not. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Predation risk makes bees reject rewarding flowers and reduce foraging activity

In the absence of predators, pollinators can often maximize their foraging success by visiting the most rewarding flowers. However, if predators use those highly rewarding flowers to locate their prey, pollinators may benefit from changing their foraging preferences to accept less rewarding flowers. Previous studies have shown that some predators, such as crab spiders, indeed hunt preferentially on the most pollinator-attractive flowers. In order to determine whether predation risk can alter pollinator preferences, we conducted laboratory experiments on the foraging behavior of bumble bees (Bombus impatiens) when predation risk was associated with a particular reward level (measured here as sugar concentration). Bees foraged in arenas containing a choice of a high-reward and a low-reward artificial flower. On a bee’s first foraging trip, it was either lightly squeezed with forceps, to simulate a crab spider attack, or was allowed to forage safely. The foragers’ subsequent visits were recorded for between 1 and 4 h without any further simulated attacks. Compared to bees that foraged safely, bees that experienced a simulated attack on a low-reward artificial flower had reduced foraging activity. However, bees attacked on a high-reward artificial flower were more likely to visit low-reward artificial flowers on subsequent foraging trips. Forager body size, which is thought to affect vulnerability to capture by predators, did not have an effect on response to an attack. Predation risk can thus alter pollinator foraging behavior in ways that influence the number and reward level of flowers that are visited.     

The role of internal and external information in foraging decisions of Melipona workers (Hymenoptera: Meliponinae)

Social insect foragers have to make foraging decisions based on information that may come from two different sources: information learned and memorised through their own experience (“internal” information) and information communicated by nest mates or directly obtained from their environment (“external” information). The role of these sources of information in decision-making by foragers was studied observationally and experimentally in stingless bees of the genus Melipona. Once a Melipona forager had started its food-collecting career, its decisions to initiate, continue or stop its daily collecting activity were mainly based upon previous experience (activity on previous days, the time at which foraging was initiated the day(s) before, and, during the day, the success of the last foraging flights) and mediated through direct interaction with the food source (load size harvested and time to collect a load). External information provided by returning foragers advanced the start of foraging of experienced bees. Most inexperienced bees initiated their foraging day after successful foragers had returned to the hive. The start of foraging by other inexperienced bees was stimulated by high waste-removal activity of nest mates. By experimentally controlling the entries of foragers (hence external information input) it was shown that very low levels of external information input had large effect on the departure of experienced foragers. After the return of a single successful forager, or five foragers together, the rate of forager exits increased dramatically for 15 min. Only the first and second entry events had large effect; later entries influenced forager exit patterns only slightly. The results show that Melipona foragers make decisions based upon their own experience and that communication stimulates these foragers if it concerns the previously visited source. We discuss the organisation of individual foraging in Melipona and Apis mellifera and are led to the conclusion that these species behave very similarly and that an information-integration model (derived from Fig. 1) could be a starting point for future research on social insect foraging. Received: 16 April 1997 / Accepted after revision: 30 August 1997     

Sea otter foraging on deep-burrowing bivalves in a California coastal lagoon

Sea otter, Enhydra lutris, predation had no detectable effect on abundance and size distribution of deep-burrowing bivalve prey in the Elkhorn Slough, California, USA. Up to 23 otters were present for 6 mo of the study period (March 1984 through April 1985). This is in contrast to previous studies of sea otter predation, especially on the shallow-burrowing Pismo clam Tivela stultorum, which can be found along the wave-exposed coast near the slough. The deep-burrowing clams Tresus nuttallii and Saxidomus nuttalli made up 61% of the prey taken in the slough, and are more difficult for otters to excavate than Pismo clams. The occurrence of foraging otters was highest in an area where the two bivalve prey were extremely abundant (18 individuals m^–2). However, the otters did not selectively prey on the largest clams available within the study sight, but foraged preferentially in a patch of smaller individuals where bivalve burrow depth was restricted by the presence of a dense clay layer. This foraging strategy maximized the amount of prey biomass obtained per unit volume of sediment excavated. Our findings suggest that in soft-sediment habitats deep-burrowing bivalves may be more resistant to otter predation than shallower burrowers.     

The establishment of foraging flocks in house sparrows: risk of predation and daily temperature

Summary The foraging decisions of animals often reflect a trade-off between the risk of predation and efficient foraging. One way an animal may reduce the risk of predation, and hence exploit a resource patch in relative safety, is by foraging in a group. Solitary pioneer sparrows often recruit others to a food source by making chirrup calls in order to establish foraging flocks. This study describes the decisions of house sparrows that arrive at food resources of different risks of predation. Four feeding sites at different distances from a perching site and from an observer were presented to sparrows. When the feeder was adjacent to the perching site and far from the observer, the pioneers chirruped less frequently and were more likely to forage alone than when the feeder was in the other three positions. There were differences in the scanning behaviour of sparrows at these sites, suggesting that they were responding to different risks of predation. Furthermore, the chirrup rates of pioneer sparrows in this study and a previous study were found to be negatively correlated with maximum daily temperature. This is consistent with the hypothesis that energy requirements may affect the flock establishment decisions of sparrows, and that the benefits of foraging in flocks may be greater at lower temperatures.     

Nest defense and central place foraging: A model and experiment

Summary A graphical model presented here indicates that a nest-defending forager should stay closer to its nest, forage for shorter times per patch, and deliver smaller loads than predicated for delivery rate maximization. The effect is more pronounced farther from the nest, so that if nest defense is especially important, the predator should leave far patches sooner than near ones, and deliver smaller loads from farther away. Moreover, if the attack rate at the nest is increased, the defending forager should move closer and deliver smaller prey.Experimental attacks with stuffed specimens at Gila woodpecker (Melanerpes uropygialis) nests produced the predicted changes in the foraging behavior of males, but not of females.Mated pairs may work as a team to pursue simultaneously two conflicting goals—food delivery and nest protection—both of which affect the survivorship of the young. Sexual dimorphism in monogamous species may result in part from specialization in these roles.     

Behavioral resource depression and decaying perceived risk of predation in two species of coexisting gerbils

Summary Behavioral resource depression occurs when the behavior of prey individuals changes in response to the presence of a predator, resulting in a reduction of the encounter rate of the predator with its prey. Here I present experimental evidence on the response of two species of gerbils (Gerbillus allenbyi and G. pyramidum) to the presence of barn owls. I conducted the experiments in a large aviary. Both gerbils responded to the presence of barn owl predators by foraging in fewer resource patches (seed trays) and by quitting foraged resource patches at a higher resource harvest rate (giving-up density of resource; GUD). This reduced the amount of time gerbils were exposed to owl predation, and hence the encounter rate of owls with gerbils, i.e., behavioral resource depression. Thus, the presence of owls imposes a foraging cost on gerbils due to risk of predation, and also on the owls themselves due to resource depression. I then examined how resource depression relaxed over time following exposure to owls. In the days following an encounter with the predator, the reduction in foraging activity for both gerbil species eased. Increasing numbers of trays were foraged each day, and GUDs in seed trays declined. The two gerbils differed in their rate of recovery, with G. pyramidum returning to prepredator levels of foraging after 1 or 2 nights and G. allenbyi taking 5 nights or longer. Interspecific differences in recovery rates may be based on differences between the species in vulnerability to predation and/or ability to detect the presence of predators. The differences in recovery rates may be due to optimal memory windows or decay rates, where differences between species are based on risk of predation or on how perceived risk changes with time since a predator was last encountered. Finally, differences between or among competitors in recovery from resource depression may provide foraging opportunities in time for the species which recover most quickly and may have implications for species coexistence.     

Within-group spatial position in ring-tailed coatis: balancing predation, feeding competition, and social competition

A variety of factors can influence an individual’s choice of within-group spatial position. For terrestrial social animals, predation, feeding success, and social competition are thought to be three of the most important variables. The relative importance of these three factors was investigated in groups of ring-tailed coatis (Nasua nasua) in Iguazú, Argentina. Different age/sex classes responded differently to these three variables. Coatis were found in close proximity to their own age/sex class more often than random, and three out of four age/sex classes were found to exhibit within-group spatial position preferences which differed from random. Juveniles were located more often at the front edge and were rarely found at the back of the group. Juveniles appeared to choose spatial locations based on feeding success and not predation avoidance. Since juveniles are the most susceptible to predation and presumably have less prior knowledge of food source location, these results have important implications in relation to predator-sensitive foraging and models of democratic group leadership. Subadults were subordinate to adult females, and their relationships were characterized by high levels of aggression. This aggression was especially common during the first half of the coati year (Nov–April), and subadults were more peripheralized during this time period. Subadults likely chose spatial positions to avoid aggression and were actively excluded from the center of the group by adult females. In the Iguazú coati groups, it appeared that food acquisition and social agonism were the major determinants driving spatial choice, while predation played little or no role. This paper demonstrates that within-group spatial structure can be a complex process shaped by differences in body size and nutritional requirements, food patch size and depletion rate, and social dominance status. How and why these factors interact is important to understanding the costs and benefits of sociality and emergent properties of animal group formation.     

A practical guide to avoid giving up on giving-up densities

The giving-up density (GUD) framework provides a powerful experimental approach with a strong theoretical underpinning to quantify foraging outcomes in heterogeneous landscapes. Since its inception, the GUD approach has been applied successfully to a vast range of foraging species and foraging scenarios. However, its application is not simple, as anyone who has tried to use it for the first time might attest. Limitations of the technique were noted at its conception, yet only the artificiality of the patches, the appropriateness of the food resource, and the possibility of multiple visiting foragers were identified. Here we show the current uses of GUD and outline the practical benefits as well as the often overlooked limitations of the technique. We define seven major points that need to be addressed when applying this methodology: (1) the curvilinearity between harvest rate and energy, (2) the energetic state of the forager, (3) the effect of group foraging, (4) food quality and substrate properties, (5) the predictability of the patch, (6) behavioral traits of the forager, and (7) nontarget species. We also suggest how GUD experiments can be enhanced by incorporating complementary methods (such as cameras) to better understand the foraging processes involved in the GUD itself. We conclude that the benefits of using GUD outweigh the costs, but that its limitations should not be ignored. Incorporating new methods when using GUD can potentially offer novel and important insights into the study of foraging behavior.