The fear diet: Risk,refuge, and biological control by omnivorous weed seed predators

Weed seed biocontrol by omnivorous mice and insects can limit weed seedbanks, but this ecosystem service can be difficult to predict given the broad diet breadth of seed predators and their potential for intraguild predation. Seed foraging behavior is further modified by fluctuating cues of predation risk from higher trophic levels and the availability of refuge habitat. Uncertainty about whether co-occurring insects and mice additively contribute to weed biocontrol or interfere with each other via intraguild predation limits our ability to recommend habitat management strategies that reliably promote seed destruction. Using seed removal assays, fluorescent powder tracking, and stable isotope analyses, we assessed effects of a predation risk cue (moonlight) on mouse foraging patterns in a patchwork of vegetated and exposed plots in a cultivated field. Mouse foraging activity decreased on exposed ground during the full moon, compared to dark nights, yet foraging movements were unaffected by moon cycle within refuge patches. Weed seed consumption was more than three times higher in cover than exposed soil, and 78% of that difference was attributable to invertebrate granivores. Mice and invertebrate granivores both exhibited higher foraging activity in cover, indicating co-occurrence of intraguild predators and prey. However, stable isotope analyses of fecal samples revealed that mice captured in refuge habitats fed at slightly lower trophic levels than those in exposed habitats (suggesting minimal intraguild predation in refuge habitat), and mouse diet was unaffected by moonlight. Despite increased availability of invertebrate prey in cover patches, mice do not appear to preferentially exploit prey when avoiding their own predators or interfere with weed seed predation. Therefore, functional redundancy of mice and invertebrate seed predators in cover crops and other refuge habitats may strengthen and stabilize weed seed biocontrol.     

Induced changes in island fox (Urocyon littoralis) activity do not mitigate the extinction threat posed by a novel predator

Prey response to novel predators influences the impacts on prey populations of introduced predators, bio-control efforts, and predator range expansion. Predicting the impacts of novel predators on native prey requires an understanding of both predator avoidance strategies and their potential to reduce predation risk. We examine the response of island foxes (Urocyon littoralis) to invasion by golden eagles (Aquila chrysaetos). Foxes reduced daytime activity and increased night time activity relative to eagle-na?ve foxes. Individual foxes reverted toward diurnal tendencies following eagle removal efforts. We quantified the potential population impact of reduced diurnality by modeling island fox population dynamics. Our model predicted an annual population decline similar to what was observed following golden eagle invasion and predicted that the observed 11% reduction in daytime activity would not reduce predation risk sufficiently to reduce extinction risk. The limited effect of this behaviorally plastic predator avoidance strategy highlights the importance of linking behavioral change to population dynamics for predicting the impact of novel predators on resident prey populations.     

Responses of a top and a meso predator and their prey to moon phases

We compared movement patterns and rhythms of activity of a top predator, the Iberian lynx Lynx pardinus, a mesopredator, the red fox Vulpes vulpes, and their shared principal prey, the rabbit Oryctolagus cuniculus, in relation to moon phases. Because the three species are mostly nocturnal and crepuscular, we hypothesized that the shared prey would reduce its activity at most risky moon phases (i.e. during the brightest nights), but that fox, an intraguild prey of lynx, would avoid lynx activity peaks at the same time. Rabbits generally moved further from their core areas on darkest nights (i.e. new moon), using direct movements which minimize predation risk. Though rabbits responded to the increased predation risk by reducing their activity during the full moon, this response may require several days, and the moon effect we observed on the rabbits had, therefore, a temporal gap. Lynx activity patterns may be at least partially mirroring rabbit activity: around new moons, when rabbits moved furthest and were more active, lynxes reduced their travelling distances and their movements were concentrated in the core areas of their home ranges, which generally correspond to areas of high density of rabbits. Red foxes were more active during the darkest nights, when both the conditions for rabbit hunting were the best and lynxes moved less. On the one hand, foxes increased their activity when rabbits were further from their core areas and moved with more discrete displacements; on the other hand, fox activity in relation to the moon seemed to reduce dangerous encounters with its intraguild predator.     

Predator-prey shell games: large-scale movement and its implications for decision-making by prey

Many classical models of food patch use under predation risk assume that predators impose patch-specific predation risks independent of prey behavior. These models predict that prey should leave a chosen patch only if and when the food depletes below some critical level. In nature, however, prey individuals may regularly move among food patches, even in the apparent absence of food depletion. We suggest that such prey movement is part of a predator-prey "shell game", in which predators attempt to learn prey location, and the prey attempt to be unpredictable in space. We investigate this shell game using an individual-based model that allows predators to update information about prey location, and permits prey to move with some random component among patches, but with reduced energy intake. Our results show the best prey strategy depends on what the predator does. A non-learning (randomly moving) predator favors non-moving prey – moving prey suffer higher starvation and predation. However, a learning predator favors prey movement. In general, the best prey strategy involves movement biased toward, but not completely committed to, the richer food patch. The strategy of prey movement remains beneficial even in combination with other anti-predator defenses, such as prey vigilance.     

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

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

The effect of predation pressure and predator adaptive foraging on the relative importance of consumptive and non‐consumptive predator net effects in a freshwater model system

An important challenge in community ecology is identifying the functional characteristics capable of predicting the nature and strength of predator effects on food webs. We developed an individual‐based model, based on a shallow lake model system, to evaluate the total, consumptive, and non‐consumptive indirect effect that predators have on basal resources when the predators differ in their foraging types (active adaptive foraging or sedentary foraging). Overall, both predator types caused similar total indirect effects on lower trophic levels. However, the nature net effects of predators diverged between predator foraging types. Active predators caused larger non‐consumptive effects, relative to the total indirect effect, irrespective of predation pressure levels. On the other hand, sedentary predators caused larger non‐consumptive effects for lower predation pressure levels, but consumptive effects became more important as predation pressure increased. Our simulations showed that the reliance on a particular mechanism driving consumer–resource interactions is altered by predator foraging behavior and highlight the importance of both prey and predator foraging behaviors to predict the causes and consequences of cascading effects observed in food webs.     

Implications of shared predation for space use in two sympatric leporids

Spatial variation in habitat riskiness has a major influence on the predator–prey space race. However, the outcome of this race can be modulated if prey shares enemies with fellow prey (i.e., another prey species). Sharing of natural enemies may result in apparent competition, and its implications for prey space use remain poorly studied. Our objective was to test how prey species spend time among habitats that differ in riskiness, and how shared predation modulates the space use by prey species. We studied a one‐predator, two‐prey system in a coastal dune landscape in the Netherlands with the European hare (Lepus europaeus) and European rabbit (Oryctolagus cuniculus) as sympatric prey species and red fox (Vulpes vulpes) as their main predator. The fine‐scale space use by each species was quantified using camera traps. We quantified residence time as an index of space use. Hares and rabbits spent time differently among habitats that differ in riskiness. Space use by predators and habitat riskiness affected space use by hares more strongly than space use by rabbits. Residence time of hare was shorter in habitats in which the predator was efficient in searching or capturing prey species. However, hares spent more time in edge habitat when foxes were present, even though foxes are considered ambush predators. Shared predation affected the predator–prey space race for hares positively, and more strongly than the predator–prey space race for rabbits, which were not affected. Shared predation reversed the predator–prey space race between foxes and hares, whereas shared predation possibly also released a negative association and promoted a positive association between our two sympatric prey species. Habitat riskiness, species presence, and prey species’ escape mode and foraging mode (i.e., central‐place vs. noncentral‐place forager) affected the prey space race under shared predation.     

Mesopredator suppression by an apex predator alleviates the risk of predation perceived by small prey

Predators can impact their prey via consumptive effects that occur through direct killing, and via non-consumptive effects that arise when the behaviour and phenotypes of prey shift in response to the risk of predation. Although predators'' consumptive effects can have cascading population-level effects on species at lower trophic levels there is less evidence that predators'' non-consumptive effects propagate through ecosystems. Here we provide evidence that suppression of abundance and activity of a mesopredator (the feral cat) by an apex predator (the dingo) has positive effects on both abundance and foraging efficiency of a desert rodent. Then by manipulating predators'' access to food patches we further the idea that apex predators provide small prey with refuge from predation by showing that rodents increased their habitat breadth and use of ‘risky′ food patches where an apex predator was common but mesopredators rare. Our study suggests that apex predators'' suppressive effects on mesopredators extend to alleviate both mesopredators'' consumptive and non-consumptive effects on prey.     

Predator behaviour and predation risk in the heterogeneous Arctic environment

1. Habitat heterogeneity and predator behaviour can strongly affect predator-prey interactions but these factors are rarely considered simultaneously, especially when systems encompass multiple predators and prey. 2. In the Arctic, greater snow geese Anser caerulescens atlanticus L. nest in two structurally different habitats: wetlands that form intricate networks of water channels, and mesic tundra where such obstacles are absent. In this heterogeneous environment, goose eggs are exposed to two types of predators: the arctic fox Vulpes lagopus L. and a diversity of avian predators. We hypothesized that, contrary to birds, the hunting ability of foxes would be impaired by the structurally complex wetland habitat, resulting in a lower predation risk for goose eggs. 3. In addition, lemmings, the main prey of foxes, show strong population cycles. We thus further examined how their fluctuations influenced the interaction between habitat heterogeneity and fox predation on goose eggs. 4. An experimental approach with artificial nests suggested that foxes were faster than avian predators to find unattended goose nests in mesic tundra whereas the reverse was true in wetlands. Foxes spent 3.5 times more time between consecutive attacks on real goose nests in wetlands than in mesic tundra. Their attacks on goose nests were also half as successful in wetlands than in mesic tundra whereas no difference was found for avian predators. 5. Nesting success in wetlands (65%) was higher than in mesic tundra (56%) but the difference between habitats increased during lemming crashes (15%) compared to other phases of the cycle (5%). Nests located at the edge of wetland patches were also less successful than central ones, suggesting a gradient in accessibility of goose nests in wetlands for foxes. 6. Our study shows that the structural complexity of wetlands decreases predation risk from foxes but not avian predators in arctic-nesting birds. Our results also demonstrate that cyclic lemming populations indirectly alter the spatial distribution of productive nests due to a complex interaction between habitat structure, prey-switching and foraging success of foxes.     

Predation cues rather than resource availability promote cryptic behaviour in a habitat-forming sea urchin

It is well known that predators often influence the foraging behaviour of prey through the so-called “fear effect”. However, it is also possible that predators could change prey behaviour indirectly by altering the prey’s food supply through a trophic cascade. The predator–sea urchin–kelp trophic cascade is widely assumed to be driven by the removal of sea urchins by predators, but changes in sea urchin behaviour in response to predators or increased food availability could also play an important role. We tested whether increased crevice occupancy by herbivorous sea urchins in the presence of abundant predatory fishes and lobsters is a response to the increased risk of predation, or an indirect response to higher kelp abundances. Inside two New Zealand marine reserves with abundant predators and kelp, individuals of the sea urchin Evechinus chloroticus were rarer and remained cryptic (i.e. found in crevices) to larger sizes than on adjacent fished coasts where predators and kelp are rare. In a mesocosm experiment, cryptic behaviour was induced by simulated predation (the addition of crushed conspecifics), but the addition of food in the form of drift kelp did not induce cryptic behaviour. These findings demonstrate that the ‘fear’ of predators is more important than food availability in promoting sea urchin cryptic behaviour and suggest that both density- and behaviourally mediated interactions are important in the predator–sea urchin–kelp trophic cascade.     

Complex tactics in a dynamic large herbivore–carnivore spatiotemporal game

The spatiotemporal game between predators and prey is a fundamental process governing their distribution dynamics. Players may adopt different tactics as the associated costs and benefits change through time. Yet few studies have investigated the potentially simultaneous and dynamic nature of movement tactics used by both players. It is particularly unclear to what extent perceived predation risk mediates the fine‐scale distribution of large and dangerous prey, which are mostly driven by bottom–up, resource‐related processes. We built habitat use and movement models based on 10 years of monitoring GPS‐collared grey wolves Canis lupus and plains bison Bison bison bison in Prince Albert National Park, Canada, to investigate the predator–large prey game in a multi‐prey system. Bison did not underuse patches of high‐quality vegetation at any time during the seasonal cycle even though wolves were selectively patrolling these areas. Rather, in at least one season, bison engaged in complex tactics comprised of proactive responses to the long‐term distribution (risky places) and reactive responses to the immediate proximity (risky times) of their opponent. In summer–autumn, bison reduced the time spent in food‐rich patches as both the long‐term use and the immediate proximity of wolves increased. By demonstrating that wolf distribution triggers patch abandonment by bison, we provide a key element in support of the shell game hypothesis – where prey move constantly to avoid predators attempting to anticipate their location. In winter, a season of relatively high energetic stress, bison no longer abandoned food‐rich patches as predation risk increased, while no bison responses to wolves were observed in spring–summer. Our work demonstrates the highly dynamic and complex nature of the predator–large prey spatiotemporal game, a key trait‐mediated mechanism by which trophic interactions structure ecological communities.     

The effect of habitat structure on prey mortality depends on predator and prey microhabitat use

Structurally complex habitats provide cover and may hinder the movement of animals. In predator–prey relationships, habitat structure can decrease predation risk when it provides refuges for prey or hinders foraging activity of predators. However, it may also provide shelter, supporting structures and perches for sit-and-wait predators and hence increase their predation rates. We tested the effect of habitat structure on prey mortality in aquatic invertebrates in short-term laboratory predation trials that differed in the presence or absence of artificial vegetation. The effect of habitat structure on prey mortality was context dependent as it changed with predator and prey microhabitat use. Specifically, we observed an ‘anti-refuge’ effect of added vegetation: phytophilous predators that perched on the plants imposed higher predation pressure on planktonic prey, while mortality of benthic prey decreased. Predation by benthic and planktonic predators on either type of prey remained unaffected by the presence of vegetation. Our results show that the effects of habitat structure on predator–prey interactions are more complex than simply providing prey refuges or cover for predators. Such context-specific effects of habitat complexity may alter the coupling of different parts of the ecosystem, such as pelagic and benthic habitats, and ultimately affect food web stability through cascading effects on individual life histories and trophic link strengths.     

The Effect of Moonlight on Scopoli's Shearwater Calonectris diomedea Colony Attendance Patterns and Nocturnal Foraging: A Test of the Foraging Efficiency Hypothesis

Moonlight is known to affect the nocturnal behaviour and activity rhythms of many organisms. For instance, predators active at night may take advantage from increased visibility afforded by the moon, while prey might regulate their activity patterns to become less detectable. Many species of pelagic seabirds attend their colony only at night, in complete darkness, avoiding approaching their nest sites under moonlight. This behaviour has been most often interpreted as an antipredator adaptation (‘predation avoidance’ hypothesis). However, it may also reflect a lower foraging efficiency during moonlit nights (‘foraging efficiency’ hypothesis). Indeed, moonlight may reduce prey availability because preferred seabird prey is known to occur at higher depths in moonlit nights. Using high‐accuracy behavioural information from data loggers, we investigated the effect of moonlight on colony attendance and at‐sea nocturnal foraging in breeding Scopoli's shearwaters Calonectris diomedea. We found that birds departing for self‐feeding trips around the full moon performed longer trips than those departing around the new moon. On nights when the moon was present only partly, nest burrow entrances took place largely in the moonless portion of the night. Moreover, contrary to predictions from the ‘foraging efficiency’ hypothesis, nocturnal foraging activity increased according to moonlight intensity, suggesting that birds increased their foraging activity when prey became more detectable. This study strengthens the idea that colony attendance behaviour is strictly controlled by moonlight in shearwaters, which is possibly related to the perception of a predation risk.     

Foraging behavior of an estuarine predator, the blue crab Callinectes sapidus in a patchy environment

To define general principles of predator‐prey dynamics in an estuarine subtidal environment, we manipulated predator density (the blue crab, Callinectes sapidus) and prey (the clam, Macoma balthica) patch distribution in large field enclosures in the Rhode River subestuary of the central Chesapeake Bay. The primary objectives were to determine whether predators forage in a way that maximizes prey consumption and to assess how their foraging success is affected by density of conspecifics. We developed a novel ultrasonic telemetry system to observe behavior of individual predators with unprecedented detail. Behavior of predators was more indicative of optimal than of opportunistic foraging. Predators appeared responsive to the overall quality of prey in their habitat. Rather than remaining on a prey patch until depletion, predators appeared to vary their patch use with quality of the surrounding environment. When multiple (two) prey patches were available, residence time of predators on a prey patch was shorter than when only a single prey patch was available. Predators seemed to move among the prey patches fairly regularly, dividing their foraging time between the patches and consuming prey from each of them at a similar rate. That predators more than doubled their consumption of prey when we doubled the number of prey (by adding the second patch) is consistent with optimizing behaviors ‐ rather than with an opportunistic increase in prey consumption brought about simply by the addition of more prey. Predators at high density, however, appeared to interfere with each other's foraging success, reflected by their lower rates of prey consumption. Blue crabs appear to forage more successfully (and their prey to experience higher mortality) in prey patches located within 15–20 meters of neighboring patch, than in isolated patches. Our results are likely to apply, at least qualitatively, to other crustacean‐bivalve interactions, including those of commercial interest; their quantitative applicability will depend on the mobility of other predators and the scale of patchiness they perceive.     

Behaviourally mediated indirect effects: interference competition increases predation mortality in foraging redshanks

1. The effect of competition for a limiting resource on the population dynamics of competitors is usually assumed to operate directly through starvation, yet may also affect survival indirectly through behaviourally mediated effects that affect risk of predation. Thus, competition can affect more than two trophic levels, and we aim here to provide an example of this. 2. We show that the foraging success of redshanks Tringa totanus (L.) foraging on active prey was highest in the front of flocks, whereas this was not the case for redshanks foraging on inactive prey. Also, when foraging on active prey, foraging success in a flock decreased as more birds passed through a patch, while overall foraging success was not lower on subsequent visits to the same patch. Thus, redshanks foraging on active prey suffered from interference competition, whereas this was not the case for redshanks foraging on inactive prey. 3. This interference competition led to differences in activity: redshanks attaining a lower foraging success had a higher walking rate. Greater activity was associated with wider flock spacing and shorter distances to cover, which has previously been shown to increase predation risk and mortality from sparrowhawks Accipiter nisus (L.). 4. We conclude that behavioural adaptations of prey species can lead to interference competition in foraging redshanks, and thus can affect their predation risk and mortality through increased activity. This study is one of the first to show how interference competition can be a mechanism for behaviourally mediated indirect effects, and provides further evidence for the suggestion that a single species occupying an intermediate trophic level may be simultaneously top-down controlled by a predator and bottom-up controlled by a behavioural response of its prey.     

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

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

Predator interference alters foraging behavior of a generalist predatory arthropod

Interactions between predators foraging in the same patch may strongly influence patch use and functional response. In particular, there is continued interest in how the magnitude of mutual interference shapes predator–prey interactions. Studies commonly focus on either patch use or the functional response without attempting to link these important components of the foraging puzzle. Predictions from both theoretical frameworks suggest that predators should modify foraging efforts in response to changes in feeding rate, but this prediction has received little empirical attention. We study the linkage between patch departure rates and food consumption by the hunting spider, Pardosa milvina, using field enclosures in which prey and predator densities were manipulated. Additionally, the most appropriate functional response model was identified by fitting alternative functional response models to laboratory foraging data. Our results show that although prey availability was the most important determinant of patch departure rates, a greater proportion of predators left enclosures containing elevated predator abundance. Functional response parameter estimation revealed significant levels of interference among predators leading to lower feeding rates even when the area allocated for each predator was kept constant. These results suggest that feeding rates determine patch movement dynamics, where interference induces predators to search for foraging sites that balance the frequency of agonistic interactions with prey encounter rates.     

Predation hazard and seed removal by small mammals: microhabitat versus patch scale effects

Predator avoidance may involve response strategies of prey species that are time and space specific. Many studies have shown that foraging individuals avoid predators by altering microhabitat usage; alternatively, sites may be selected according to larger-scale features of the habitat mosaic. We measured seed removal by two small mammal species (Peromyscus leucopus, and Microtus pennsylvanicus) at 474 stations over an experimentally created landscape of 12 patches, and under conditions of relatively high (full moon) and low (new moon) predatory hazard. Our objective was to determine whether predator avoidance involved the selection of small-, medium-, or large-scale features of the landscape (i.e., at the scale of microhabitats, habitats, or habitat patches). We found rates of seed removal to vary more with features of whole patches than according to variation in structural microhabitats within patches. Specific responses included: under-utilization of patch edge habitats during full moon periods, and microhabitat effects that were only significant when considered in conjunction with larger-scale features of the landscape. Individuals residing on large patches altered use of microhabitats/habitats to a greater extent than those on smaller patches. Studies just focusing on patterns of microhabitat use will miss responses at the larger scales, and may underestimate the importance of predation to animal foraging behavior.     

Habitat-dependent foraging in a classic predator–prey system: a fable from snowshoe hares

Current research contrasting prey habitat use has documented, with virtual unanimity, habitat differences in predation risk. Relatively few studies have considered, either in theory or in practice, simultaneous patterns in prey density. Linear predator–prey models predict that prey habitat preferences should switch toward the safer habitat with increasing prey and predator densities. The density‐dependent preference can be revealed by regression of prey density in safe habitat versus that in the riskier one (the isodar). But at this scale, the predation risk can be revealed only with simultaneous estimates of the number of predators, or with their experimental removal. Theories of optimal foraging demonstrate that we can measure predation risk by giving‐up densities of resource in foraging patches. The foraging theory cannot yet predict the expected pattern as predator and prey populations covary. Both problems are solved by measuring isodars and giving‐up densities in the same predator–prey system. I applied the two approaches to the classic predator–prey dynamics of snowshoe hares in northwestern Ontario, Canada. Hares occupied regenerating cutovers and adjacent mature‐forest habitat equally, and in a manner consistent with density‐dependent habitat selection. Independent measures of predation risk based on experimental, as well as natural, giving‐up densities agreed generally with the equal preference between habitats revealed by the isodar. There was no apparent difference in predation risk between habitats despite obvious differences in physical structure. Complementary studies contrasting a pair of habitats with more extreme differences confirmed that hares do alter their giving‐up densities when one habitat is clearly superior to another. The results are thereby consistent with theories of adaptive behaviour. But the results also demonstrate, when evaluating differences in habitat, that it is crucial to let the organisms we study define their own habitat preference.     

Risk management in optimal foragers: the effect of sightlines and predator type on patch use,time allocation,and vigilance in gerbils

In the foraging game between gerbils and their predators, gerbils manage risk of predation using the tools of time allocation (where, when and for how long to forage) and vigilance. The optimal level of a forager's vigilance should be affected by its encounter rate with predators and the effectiveness of its vigilance in reducing mortality risk. The physical structure of the environment can alter the effectiveness of its vigilance and therefore alter its foraging behaviour. We tested this for gerbils at risk of predation from barn owls or foxes in a large vivarium. In particular, we reduced the effectiveness of vigilance by placing obstructions around feeding trays that blocked sight lines along either the vertical (vigilance directed against owls) or horizontal axis (vigilance directed against foxes), thereby changing the physical structure of the environment. In addition, we manipulated the presence of foxes and owls. In general, gerbils harvested fewer seeds, allocated less time to foraging in dangerous patches, and used more vigilance while foraging where and when risks were higher (i.e. in the presence of predators and in bright moonlight). Vertical and horizontal sightline treatments interacted synergistically to further raise perceived risk. These results imply that blocking sight lines reduces the effectiveness of vigilance, causing gerbils to use it less. Moreover, in the presence of a predator, the gerbils’ response to the blocked sightlines was more severe – harvesting less food and spending less time and vigilance – in the patches with the increased risk. This was especially so in the presence of the predator that was expected to most benefit from blocking that particular type of sight line: cover that blocked vertical sight lines was riskiest in the presence of owls, and cover that blocked horizontal sight lines was riskiest in the presence of foxes. These results strongly indicate the importance of sightlines and landscape features such as bushes in the risk management and forging decisions of gerbils, demonstrating that bush cover provides mixed blessing to gerbils by providing cover, but making vigilance ineffective.