Performance level and efficiency of two differing predator-avoidance strategies depend on nutritional state of the prey fish

Animal prey has developed a variety of behavioural strategies to avoid predation. Many fish species form shoals in the open water or seek refuge in structurally complex habitats. Since anti-predator strategies bear costs and are energy-demanding, we hypothesised that the nutritional state of prey should modify the performance level and efficiency of such strategies. In aquaria either containing or lacking a structured refuge habitat, well-fed or food-deprived juvenile roach (Rutilus rutilus) were exposed to an open-water predator (pikeperch, Sander lucioperca). Controls were run without predators. In the presence of the predator, roach enhanced the performance of the anti-predator strategy and increased the use of the refuge habitat whereby food-deprived roach were encountered more often in the structure than well-fed roach. Nonetheless more starved than well-fed roach were fed upon by the predator. In the treatments offering only open-water areas, roach always formed dense shoals in the presence of the predator. The shoal density, however, was lower in starved roach. Starving fish in shoals experienced the highest predation mortality across all experimental treatments. The experiment confirmed the plasticity of the anti-predator behaviour in roach and demonstrated that food deprivation diminished the efficiency of shoaling more strongly than the efficiency of hiding. The findings may be relevant to spatial distribution of prey and predator–prey interactions under natural conditions because when prey are confronted with phases of reduced resource availability, flexible anti-predator strategies may lead to dynamic habitat use patterns.     

The anti-predator function of ‘eyespots’ on camouflaged and conspicuous prey

Animals utilise various strategies to reduce the risk of predation, including camouflage, warning colours and mimicry, and many of these protective signals promote avoidance behaviour in predators. For example, various species possess paired circular ‘eyespots’, which startle or intimidate predators, preventing or halting an attack. However, little is known of how the efficacy of such signals relates to the context in which they are found, and no studies have tested the relative effectiveness of anti-predator signals when on otherwise camouflaged and conspicuous prey. We find that the protective value of conspicuous wing spots, placed on artificial moth-like targets presented to wild birds in the field, is strongly affected by the attributes of the prey ‘animal’ on which they are found. Wing spots reduced predation when on conspicuous prey but were rendered ineffective when on otherwise camouflaged targets, indeed they increased the risk of predation compared to non-marked camouflaged controls. These results demonstrate how different anti-predator strategies may interact, and that protective signals can switch from being beneficial to costly under different contexts.     

Influences of divergent behavioral strategies upon risk allocation in juvenile flatfishes

Animals balance feeding and anti-predator behaviors at various temporal scales. When risk is infrequent or brief, prey can postpone feeding in the short term and temporally allocate feeding behavior to less risky periods. If risk is frequent or lengthy, however, prey must eventually resume feeding to avoid fitness consequences. Species may exhibit different behavioral strategies, depending on the fitness tradeoffs that exist in their environment or across their life histories. North Pacific flatfishes that share juvenile rearing habitat exhibit a variety of responses to predation risk, but their response to risk frequency has not been examined. We observed the feeding and anti-predator behaviors of young-of-the-year English sole (Parophrys vetulus), northern rock sole (Lepidopsetta polyxystra), and Pacific halibut (Hippoglossus stenolepis)—three species that exhibit divergent anti-predator strategies—following exposure to three levels of predation risk: no risk, infrequent (two exposures/day), and frequent (five exposures/day). The English sole responded to the frequent risk treatment with higher feeding rates than during infrequent risk, following a pattern of behavioral response that is predicted by the risk allocation hypothesis; rock sole and halibut did not follow the predicted pattern, but this may be due to the limited range of treatments. Our observations of unique anti-predator strategies, along with differences in foraging and species-specific ecologies, suggest divergent trajectories of risk allocation for the three species.     

How unpredictable is the individual scanning process in socially foraging mammals?

In group-forming prey species, theory assumes that individuals within groups should scan independently of one another, with vigilance sequences being relatively unpredictable, making interscan durations highly variable. We attempted to detect any divergence from randomness in the scanning process in three mammalian prey species phylogenetically and geographically separated and exposed to different levels of predation: waterbuck, Kobus ellipsiprymnus defassa, under a high observed predation risk, eastern grey kangaroo, Macropus giganteus, still experiencing occasional predation and European roe deer, Capreolus capreolus, under a very low natural predation risk. Our results revealed that the focal interscan duration increased when the duration of the preceding interscan increased, whatever the studied species and the predation risk that its individuals experienced, and decreased with the preceding scan duration in two species under, respectively, occasional and low predation risks. The exponential distribution was the tested model that fitted the observed distributions of interscan durations least well. We discuss what can trigger non-randomness in scanning, through a non-homogenous Poisson process, at both intra-individual and inter-individual levels, particularly with regard to previous studies that have demonstrated synchronisation of vigilance in such mammals. Our results suggest the need to reconsider any assumption of randomness in scanning in the basic model predicting form and frequency of scanning behaviour by prey species.     

The role of turbidity as a constraint on predator-prey interactions in aquatic environments

Many of the world's most productive aquatic ecosystems usually contain turbid water. Paradoxically, many fish species that live in these habitats are also those that often rely on vision to detect their predators and their prey. For these fish, turbidity will reduce the distance at which predator-prey interactions occur, and there should be a reduction in the opportunity for behavioural modification to control the risk of predation. Under these conditions, we predict that most antipredator behaviour will become ineffective and that predator-prey interactions in turbid water will be primarily characterised by the direct effects of predator consumption of prey, rather than behavioural modification reducing the growth rates of prey. This hypothesis was tested in two laboratory experiments. The first experiment investigated how water turbidity, risk of predation, and their interaction affect habitat choice decisions by fathead minnows (Pimephales promelas). These data demonstrate that fathead minnows reduce their use of dangerous habitats, but that this effect is diminished in turbid water. A second experiment examined mortality patterns when these fish were preyed upon by yellow perch (Perca flavescens) in clear and turbid water. Absolute mortality rates were unaffected by visibility, but patterns of mortality were random in turbid water and skewed towards smaller individuals in clear water. Combined, these results support our hypothesis and suggest that the impact of predation risk will be reduced in turbid aquatic ecosystems. Received: 23 May 1996 / Accepted after revision: 18 November 1996     

Time stress,predation risk and diurnal–nocturnal foraging trade-offs in larval prey

Insect larvae increase in size with several orders of magnitude throughout development making them more conspicuous to visually hunting predators. This change in predation pressure is likely to impose selection on larval anti-predator behaviour and since the risk of detection is likely to decrease in darkness, the night may offer safer foraging opportunities to large individuals. However, forsaking day foraging reduces development rate and could be extra costly if prey are subjected to seasonal time stress. Here we test if size-dependent risk and time constraints on feeding affect the foraging–predation risk trade-off expressed by the use of the diurnal–nocturnal period. We exposed larvae of one seasonal and one non-seasonal butterfly to different levels of seasonal time stress and time for diurnal–nocturnal feeding by rearing them in two photoperiods. In both species, diurnal foraging ceased at large sizes while nocturnal foraging remained constant or increased, thus larvae showed ontogenetic shifts in behaviour. Short night lengths forced small individuals to take higher risks and forage more during daytime, postponing the shift to strict night foraging to later on in development. In the non-seasonal species, seasonal time stress had a small effect on development and the diurnal–nocturnal foraging mode. In contrast, in the seasonal species, time for pupation and the timing of the foraging shift were strongly affected. We argue that a large part of the observed variation in larval diurnal–nocturnal activity and resulting growth rates is explained by changes in the cost/benefit ratio of foraging mediated by size-dependent predation and time stress.     

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.     

Do pronghorn mothers reveal the locations of their hidden fawns?

Summary Success of the hider strategy in ungulates depends, in part, on the mother's ability to minimize information she transmits about her young's hiding place while remaining close enough to distract or drive away a predator. We predicted that pronghorn (Antilocapra americana) mothers should: (1) maintain minimum distances from their hidden fawns sufficient to cause the expected energy gain for a ground predator, systematically searching around the mother, to fall below that expected when searching for some other prey; (2) orient the axes of either head or trunk towald the hidden fawn no more frequently than would be expected by chance; (3) schedule behavior so that no activity is more likely than another to occur when a visit to the hidden fawn is imminent. At the National Bison Range, where coyote (Canis latrans) predation on pronghorn fawns is frequent, pronghorn mothers conform to predictions (1) and (3), but not (2). Within the first 10% of their time away from fawns, mothers reached an average distance of 70.4 m from their fawns' biding places and remained at that distance until 95% of their time away was clapsed. At this mother-fawn distance a coyote, using the mother's position to begin a systematic search for the fawn, and searching at a rate of 4 m²/s, would gain energy at a lower rate than it would hunting ground squirrels (Spermophilus columbianus). Mothers pointed both head and trunk toward their hidden fawns more frequently than would be expected by chance and coyotes, able to use this information to establish a 90° quadrant to search, could expect rates of gain higher than those obtainable in 360° search. Mothers especially tended to orient both head and trunk toward their fawns when standing or moving. Coyotes that begin a 90° search based on the mother's head or trunk position only when mothers were standing or moving, could expect rates of energy gain almost double those expected in 360° search or ground squirrel hunting. Maternal activities (stand, feed, recline, or move) were distributed evenly across all mother-fawn distances and across percent total time away from the fawn. Thus, activity was not a good predictor of a mother's likelihood of soon returning to her fawn. Mothers also remained away from their fawns long enough to cause the expected rate of energy gain for a coyote hiding and watching for the mother's return to the fawn to fall well below the rate expected for searching or ground squirrel hunting.     

Mechanism of predator–prey detection and behavioral responses in some anuran tadpoles

Predator–prey relationship was studied in three sympatric species of anuran tadpoles. The study design consisted of allowing predaceous Hoplobatrachus tigerinus tadpoles to devour prey tadpoles (Sphaerotheca breviceps and Bufo melanostictus) placed in a plastic tub (five tadpoles of each species, stage ~27) in 30 min. In trials without refugia, more tadpoles of Bufo fell prey compared to Sphaerotheca. In contrast, provision of refugia using hydrilla plant reversed predation risk of the two species. The swimming speed (V _max = 64.55 ± 1.45 cm/s) of Hoplobatrachus tadpoles was much higher compared to the prey species (Bufo: 3.6 ± 0.4 cm/s; Sphaerotheca: 27.6 ± 1.6 cm/s). Poor swimming ability may account for the observed vulnerability of the Bufo tadpoles to predation especially in clear waters; refugia overcame predation to some extent. On the other hand, Sphaerotheca tadpoles that swim faster than the toad tadpoles were less vulnerable in open areas; refugia actually hindered swimming and increased predation. Experiments with association choice tests show that predaceous tadpoles detect prey based on both visual and chemical cues. On the other hand, the prey tadpoles detected predator based exclusively on chemical rather than visual cues. The antipredator defense strategy of the toad tadpoles is manifested in the form of reduced movements, remaining still for longer times and, increased burst speed. The present findings also suggest that in both prey species predator detection has a genetic basis since naive tadpoles with no prior exposure to predators exhibit fright response on first encounter with them.     

Habitat use and group size of pied cormorants (<Emphasis Type="Italic">Phalacrocorax varius</Emphasis>) in a seagrass ecosystem: possible effects of food abundance and predation risk

Both food abundance and predation risk may influence habitat use decisions. However, studies of habitat use by birds in marine environments have focused only on food abundance. I investigated the possible influences of food abundance and predation risk from tiger sharks (Galeocerdo cuvier) on habitat use by pied cormorants (Phalacrocorax varius) over two spatial scales and on cormorant group size. Cormorants were usually solitary, but group size was highest in shallow habitats during months when shark density was low. Regardless of season, cormorant density within shallow habitats was higher over seagrass than sand, and cormorants were distributed between these two microhabitats proportional to prey density. Therefore, cormorants appear to respond to prey abundance at a relatively narrow spatial scale (i.e., tens of meters). At the habitat-patch scale (~1 km), the density of cormorants and their prey (teleosts) was higher in shallow habitats than in deep ones, but the density of cormorants was influenced by an interaction between water temperature (i.e., season) and habitat. There was decreased use of shallow habitats as water temperature, and the density of tiger sharks, increased. When shark density was low, cormorants were distributed across habitats roughly in proportion to the abundance of fish, suggesting that cormorants respond to food abundance at the scale of habitat patches. However, as shark abundance increased, the relative density of cormorants dropped in the dangerous shallow habitats such that there was a greater density of cormorants relative to their food in deep habitats when sharks were abundant. This suggests that pied cormorants trade-off food and risk by accepting lower energetic returns to forage in safer habitats. This study provides the first evidence that marine habitat selection by birds may be influenced by such a trade-off, and provides further evidence that tiger sharks are important in determining habitat use of their prey and mediating indirect interactions within Shark Bay.Communicated by P. W. Sammarco, Chauvin     

Behavioural interference between ungulate species: roe are not on velvet with fallow deer

Interference is expected to occur at feeding areas between species with a similar diet, but few studies have tested this idea for wild ungulates. We analysed interactions between fallow deer, European roe deer and wild boar, in three sites, in a Mediterranean area. We expected that interference should be greater between deer than between them and wild boar. We documented the negative effects of behavioural interference by fallow on foraging behaviour of roe deer, under field conditions. Deer species built up 90% interference interactions, with fallow always dominant on roe, also through direct aggression. Although roe deer decreased feeding and increased vigilance levels in proximity (<50 m) of either fallow deer or wild boar, they were displaced significantly more often by the former than by the latter. Fallow deer were neither displaced nor alarmed by roe and rarely by wild boar. No deer species displaced wild boar. Interference was significantly greater on solitary roe deer, especially females, in spring and roe left the feeding ground most often in the smallest site (13 ha). Roe deer avoided areas where the local density of fallow deer was the highest. During our 4-year-study, roe deer density decreased whereas fallow deer numbers increased. Behavioural interference may explain how fallow deer outcompete roe deer through spatial exclusion from feeding sites and avoidance of areas with high densities of the former. Fallow deer evolved in semi-arid, relatively poor habitats of Asia Minor: interspecific defence of crucial resources could have developed as a beneficial tactic for its survival.     

Cercopagis pengoi and Mysis spp. alter their feeding rate and prey selection under predation risk of herring (Clupea harengus membras)

The anti-predator behaviour of Baltic crustacean planktivores was studied in feeding experiments under predation pressure of herring. The experiments were conducted with pelagic mysids: Mysis mixta and Mysis relicta, and with Cercopagis pengoi, a non-indigenous cladoceran, which invaded the Baltic Sea in 1992. Zooplankton was offered as prey. Two kinds of experiments were performed in the absence and presence of chemical predator cues: (1) two-prey experiments with prey, which have poor or good escape responses and all three planktivores and (2) natural prey experiments with mysids in natural zooplankton assemblages. The results showed that all three species reacted to the chemical cue of herring by decreasing their feeding rate and altering prey selection. C. pengoi selected easily captured prey (rotifers) in two-prey experiments under predation risk while selection for any prey was evident in mysids in natural prey experiments only in the absence of predator cues. This indicates that planktivores have different anti-predator strategies, which are modified by their own prey capture abilities. C. pengoi was a very efficient predator on small prey with size-specific prey consumption rate 5 to 18 times the rate of mysids. Results show that the studied planktivores are capable of adjusting their feeding behaviour to decrease their conspicuousness in order to increase survival under predation risk. Further, results support the view that C. pengoi has adapted well to the Baltic ecosystem, sharing food niche with pelagic mysids and most probably having a strong influence on the whole pelagic food web.     

Effects of small-scale habitat fragmentation on predator–prey interactions in a temperate sea grass system

During the last decades, fragmentation has become an important issue in ecological research. Habitat fragmentation operates on spatial scales ranging over several magnitudes from patches to landscapes. We focus on small-scale fragmentation effects relevant to animal foraging decision making that could ultimately generate distribution patterns. In a controlled experimental environment, we tested small-scale fragmentation effects in artificial sea grass on the feeding behaviour of juvenile cod (Gadus morhua). Moreover, we examined the influence of fragmentation on the distribution of one of the juvenile cod’s main prey resources, the grass shrimp (Palaemon elegans), in association with three levels of risk provided by cod (no cod, cod chemical cues and actively foraging cod). Time spent by cod within sea grass was lower in fragmented landscapes, but total shrimp consumption was not affected. Shrimp utilised vegetation to a greater extent in fragmented treatments in combination with active predation. We suggest that shrimp choose between sand and vegetation habitats to minimize risk of predation according to cod habitat-specific foraging capacities, while cod aim to maximize prey-dependent foraging rates, generating a habitat-choice game between predator and prey. Moreover, aggregating behaviour in grass shrimp was only found in treatments with active predation. Hence, we argue that both aggregation and vegetation use are anti-predator defence strategies applied by shrimp. We therefore stress the importance of considering small-scale behavioural mechanisms when evaluating consequences from habitat fragmentation on trophic processes in coastal environments.     

Female guppies shorten brood retention in response to predator cues

Predation risk influences the duration of offspring development in many species where embryos develop from externally shed eggs. Surprisingly, such predator-mediated effects on offspring development have rarely been explored in live-bearers. In this paper, we use the guppy (Poecilia reticulata), a live-bearing freshwater fish, to test whether the duration of brood retention (the time from mating to parturition) is influenced by experimental changes in the perceived level of predation. Because the swimming performance of female guppies is impaired during late pregnancy, we predicted that females would withhold broods for shorter periods when they are exposed to cues that signal a heightened risk of predation on adults rather than on juveniles. We therefore simulated increased risk of predation on adults by using a combination of pike-shaped models (resembling natural predators that prey on adult guppies) and ‘alarm substances’ derived from the skin extracts of adult conspecific females. Our results revealed that, under simulated predation risk, female guppies produced broods significantly more quickly than their counterparts assigned to a control group where predator cues were absent. A subsequent evaluation of offspring swimming performance revealed a significant positive association between neonate swimming speeds and the duration of brood retention, suggesting that by accelerating parturition, females may produce offspring with impaired locomotor skills. These findings, in conjunction with similar results from other live-bearing species, suggest that the conditions experienced by gestating females can generate significant variation in the timing of offspring development with potentially important implications for offspring fitness.     

Background level of risk determines the intensity of predator neophobia in juvenile convict cichlids

Behavioural ecology is rife with examples of prey animals that are able to adjust the intensity of their anti-predator response to match background risk levels. Often, preys need experience with predators before they will invest in costly anti-predator responses. This means that prey animals often fail to respond to predators during their first encounter. Recently, we have shown that prey raised under high-risk conditions may exhibit avoidance of potential predation cues independent of experience (neophobia). Such phenotypically plastic neophobic predator responses may reduce the initial costs of learning ecologically relevant threats while maintaining sufficient behavioural plasticity to respond to variation in local conditions. Here, we test if induced neophobia results in threat-sensitive behavioural trade-offs in response to a novel chemosensory cue. Our first experiment shows that while juvenile convict cichlids (Amatitlania nigrofasciata) pre-exposed to high (but not low) risk conditions exhibited predator avoidance to a novel odour (rainbow trout, Oncorhynchus mykiss), the response intensity was not influenced by the concentration of trout odour detected. Our second experiment demonstrated that the intensity of anti-predator response towards a novel predator cue was dependent upon the level of background risk. Convict cichlids pre-exposed to high-risk conditions showed stronger responses than those pre-exposed to low-risk conditions, while cichlids pre-exposed to intermediate-risk conditions exhibited intermediate response intensities. Together, these data demonstrate that background levels of risk and not the concentration of novel cues detected shape the induced neophobic response pattern of juvenile convict cichlids.     

Food webs and intraguild predation: community interactions of a native mesocarnivore

Trophic level interactions between predators create complex relationships such as intraguild predation. Theoretical research has predicted two possible paths to stability in intraguild systems: intermediate predators either outcompete higher-order predators for shared resources or select habitat based on security. The effects of intraguild predation on intermediate mammalian predators such as swift foxes (Vulpes velox) are not well understood. We examined the relationships between swift foxes and both their predators and prey, as well the effect of vegetation structure on swift fox-coyote (Canis latrans) interactions, between August 2001 and August 2004. In a natural experiment created by the Pinon Canyon Maneuver Site in southeastern Colorado, USA, we documented swift fox survival and density in a variety of landscapes and compared these parameters in relation to prey availability, coyote abundance, and vegetation structure. Swift fox density varied significantly between study sites, while survival did not. Coyote abundance was positively related to the basal prey species and vegetation structure, while swift fox density was negatively related to coyote abundance, basal prey species, and vegetation structure. Our results support the prediction that, under intraguild predation in terrestrial systems, top predator distribution matches resource availability (resource match), while intermediate predator distribution inversely matches predation risk (safety match). While predation by coyotes may be the specific cause of swift fox mortality in this system, the more general mechanism appears to be exposure to predation moderated by shrub density.     

Behavioral response races, predator-prey shell games, ecology of fear, and patch use of pumas and their ungulate prey

The predator-prey shell game predicts random movement of prey across the landscape, whereas the behavioral response race and landscape of fear models predict that there should be a negative relationship between the spatial distribution of a predator and its behaviorally active prey. Additionally, prey have imperfect information on the whereabouts of their predator, which the predator should incorporate in its patch use strategy. I used a one-predator-one-prey system, puma (Puma concolor)-mule deer (Odocoileus hemionus) to test the following predictions regarding predator-prey distribution and patch use by the predator. (1) Pumas will spend more time in high prey risk/low prey use habitat types, while deer will spend their time in low-risk habitats. Pumas should (2) select large forage patches more often, (3) remain in large patches longer, and (4) revisit individual large patches more often than individual smaller ones. I tested these predictions with an extensive telemetry data set collected over 16 years in a study area of patchy forested habitat. When active, pumas spent significantly less time in open areas of low intrinsic predation risk than did deer. Pumas used large patches more than expected, revisited individual large patches significantly more often than smaller ones, and stayed significantly longer in larger patches than in smaller ones. The results supported the prediction of a negative relationship in the spatial distribution of a predator and its prey and indicated that the predator is incorporating the prey's imperfect information about its presence. These results indicate a behavioral complexity on the landscape scale that can have far-reaching impacts on predator-prey interactions.     

Making sense of predator scents: investigating the sophistication of predator assessment abilities of fathead minnows

According to the threat-sensitive predator avoidance hypothesis, selection favors prey that accurately assess the degree of threat posed by a predator and adjust their anti-predator response to match the level of risk. Many species of animals rely on chemical cues to estimate predation risk; however, the information content conveyed in these chemical signatures is not well understood. We tested the threat-sensitive predator avoidance hypothesis by determining the specificity of the information conveyed to prey in the chemical signature of their predator. We found that fathead minnows (Pimephales promelas) could determine the degree of threat posed by northern pike (Esox lucius) based on the concentration of chemical cues used. The proportion of minnows that exhibited an anti-predator response when exposed to a predator cue increased as the concentration of the pike cue used increased. More surprisingly, the prey could also distinguish large pike from small pike based on their odor alone. The minnows responded more intensely to cues of small pike than to cues of large pike. In this predator–prey system small pike likely represent a greater threat than large pike.Communicated by A. Mathis     

The effect of hunger on the strength and duration of the antipredator behavioral response of green frog (<Emphasis Type="Italic">Rana clamitans</Emphasis>) tadpoles

The activity level of prey reflects a trade-off between predation risk and foraging gain. A number of theoretical and empirical studies have shown that a prey's energetic state or the level of its resource should influence this trade-off (i.e., what the optimal activity level at a level of predation risk is). Here, I show that the energetic state of prey may also influence the duration of their antipredator behavioral response. Green frog tadpoles (Rana clamitans) reduced their activity level for a shorter time during exposure to the chemical cue of predatory larval dragonflies (Anax spp.) as their time since last feeding increased (i.e., as their energetic state decreased). Interestingly, the tadpoles strongly reduced their activity level upon cue exposure in all treatments. Thus, the relative activity level of tadpoles at different energetic states varied over time.     

Patch use and vigilance by sympatric lemmings in predator and competitor-driven landscapes of fear

Prey living in risky environments can adopt a variety of behavioral tactics to reduce predation risk. In systems where predators regulate prey abundance, it is reasonable to assume that differential patterns of habitat use by prey species represent adaptive responses to spatial variation in predation. However, patterns of habitat use also reflect interspecific competition over habitat. Collared (Dicrostonyx groenlandicus) and brown (Lemmus trimucronatus) lemmings represent such a system and possess distinct upland tundra versus mesic meadow habitat preferences consistent with interspecific competition. Yet, we do not know whether this habitat preference might also reflect differences in predation risk or whether the two species differ in their behavioral tactics used to avoid predation. We performed experiments where we manipulated putative predation risk perceived by lemmings by increasing protective cover in upland and meadow habitats while we recorded lemming activity and behavior. Both lemming species preferentially used cover more than open patches, but Dicrostonyx was more vigilant than Lemmus. Both species also constrained their activity to protective patches in upland and meadow habitats, but during different periods of the day. Use of cover and vigilance were independent of habitat, suggesting that both species live in a fearsome but flattened landscape of fear at Walker Bay (Nunavut, Canada), and that their habitat preference is a consequence of competition rather than predation risk. Future studies aiming to map the contours of fear in multi-prey–predator systems should consider how predation and competition interact to modify prey species’ habitat preference, patch use, and vigilance.