Antipredator strategies of house finches: are urban habitats safe spots from predators even when humans are around?

Urbanization decreases species diversity, but it increases the abundance of certain species with high tolerance to human activities. The safe-habitat hypothesis explains this pattern through a decrease in the abundance of native predators, which reduces predation risk in urban habitats. However, this hypothesis does not consider the potential negative effects of human-associated disturbance (e.g., pedestrians, dogs, cats). Our goal was to assess the degree of perceived predation risk in house finches (Carpodacus mexicanus) through field studies and semi-natural experiments in areas with different levels of urbanization using multiple indicators of risk (flock size, flight initiation distance, vigilance, and foraging behavior). Field studies showed that house finches in more urbanized habitats had a greater tendency to flock with an increase in population density and flushed at larger distances than in less urbanized habitats. In the semi-natural experiment, we found that individuals spent a greater proportion of time in the refuge patch and increased the instantaneous pecking rate in the more urbanized habitat with pedestrians probably to compensate for the lower amount of foraging time. Vigilance parameters were influenced in different ways depending on habitat type and distance to flock mates. Our results suggest that house finches may perceive highly urbanized habitats as more dangerous, despite the lower number of native predators. This could be due to the presence of human activities, which could increase risk or modify the ability to detect predators. House finches seem to adapt to the urban environment through different behavioral strategies that minimize risk.     

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.     

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.     

Habitat quality and heterogeneity influence distribution and behavior in African buffalo (Syncerus caffer)

Top-down effects of predators on prey behavior and population dynamics have been extensively studied. However, some populations of very large herbivores appear to be regulated primarily from the bottom up. Given the importance of food resources to these large herbivores, it is reasonable to expect that forage heterogeneity (variation in quality and quantity) affects individual and group behaviors as well as distribution on the landscape. Forage heterogeneity is often strongly driven by underlying soils, so substrate characteristics may indirectly drive herbivore behavior and distribution. Forage heterogeneity may further interact with predation risk to influence prey behavior and distribution. Here we examine differences in spatial distribution, home range size, and grouping behaviors of African buffalo as they relate to geologic substrate (granite and basalt) and variation in food quality and quantity. In this study, we use satellite imagery, forage quantity data, and three years of radio-tracking data to assess how forage quality, quantity, and heterogeneity affect the distribution and individual and herd behavior of African buffalo. We found that buffalo in an overall poorer foraging environment keyed-in on exceptionally high-quality areas, whereas those foraging in a more uniform, higher-quality area used areas of below-average quality. Buffalo foraging in the poorer-quality environment had smaller home range sizes, were in smaller groups, and tended to be farther from water sources than those foraging in the higher-quality environment. These differences may be due to buffalo creating or maintaining nutrient hotspots (small, high-quality foraging areas) in otherwise low-quality foraging areas, and the location of these hotspots may in part be determined by patterns of predation risk.     

Field evidence for pervasive indirect effects of fishing on prey foraging behavior

The indirect, ecosystem-level consequences of ocean fishing, and particularly the mechanisms driving them, are poorly understood. Most studies focus on density-mediated trophic cascades, where removal of predators alternately causes increases and decreases in abundances of lower trophic levels. However, cascades could also be driven by where and when prey forage rather than solely by prey abundance. Over a large gradient of fishing intensity in the central Pacific's remote northern Line Islands, including a nearly pristine, baseline coral reef system, we found that changes in predation risk elicit strong behavioral responses in foraging patterns across multiple prey fish species. These responses were observed as a function of both short-term ("acute") risk and longer-term ("chronic") risk, as well as when prey were exposed to model predators to isolate the effect of perceived predation risk from other potentially confounding factors. Compared to numerical prey responses, antipredator behavioral responses such as these can potentially have far greater net impacts (by occurring over entire assemblages) and operate over shorter temporal scales (with potentially instantaneous response times) in transmitting top-down effects. A rich body of literature exists on both the direct effects of human removal of predators from ecosystems and predators' effects on prey behavior. Our results draw together these lines of research and provide the first empirical evidence that large-scale human removal of predators from a natural ecosystem indirectly alters prey behavior. These behavioral changes may, in turn, drive previously unsuspected alterations in reef food webs.     

Rats on the run: removal of alien terrestrial predators affects bush rat behaviour

Predators can strongly influence the microhabitat use and foraging behaviour of prey. In a large-scale replicated field experiment in East Gippsland, Australia, we tested the effects of reduced alien red fox (Vulpes vulpes) and alien wild dog (Canis lupus familiaris) abundance (treatment) on native bush rat (Rattus fuscipes) behaviour. Bush rats are exposed to two main guilds of predators, namely mammalian carnivores and birds of prey. Tracking rat movements using the spool-and-line technique revealed that, in treatment sites, rats used ground cover, which provides shelter from predators, less often than at unmanipulated fox and wild dog abundance (non-treatment sites). In treatment sites, rats more frequently moved on logs where they would have been exposed to hunting foxes and dogs than in non-treatment sites. Furthermore, in treatments, rats showed a preference for understorey but not in non-treatments. Hence, bush rats adapted their behaviour to removal of alien terrestrial predators. Giving-up densities (GUDs) indicated no treatment effects on the marginal feeding rate of bush rats. Interestingly, GUDs were higher in open patches than in sheltered patches, suggesting higher perceived predation risk of bush rats during foraging at low versus high cover. The lack of treatment effects on GUDs but the clear response of bush rats to cover may be explained by the impact of predators other than foxes and wild dogs.     

Habitat use and foraging behavior of tiger sharks (<Emphasis Type="Italic">Galeocerdo cuvier</Emphasis>) in a seagrass ecosystem

Understanding the foraging behavior and spatial distribution of top predators is crucial to gaining a complete understanding of communities. However, studies of top predators are often logistically difficult and it is important to develop appropriate methods for identifying factors influencing their spatial distribution. Sharks are top predators in many marine communities, yet no studies have quantified the habitat use of large predatory sharks or determined the factors that might influence shark spatial distributions. We used acoustic telemetry and animal-borne video cameras ("Crittercam") to test the hypothesis that tiger shark (Galeocerdo cuvier) habitat use is determined by the availability of their prey. We also used Crittercam to conduct the first investigation of foraging behavior of tiger sharks. To test for habitat preferences of sharks, the observed proportion of time in each habitat for each individual was compared to the predicted values for that individual based on correlated random walk and track randomization methods. Although there was individual variation in habitat use, tiger sharks preferred shallow seagrass habitats, where their prey is most abundant. Despite multiple encounters with potential prey, sharks rarely engaged in prolonged high-speed chases, and did not attack prey that were vigilant. We propose that the tiger sharks' foraging tactic is one of stealth, and sharks rely upon close approaches to prey in order to be successful. This study shows that using appropriate analysis techniques and a variety of field methods it is possible to elucidate the factors influencing habitat use and gain insights into the foraging behavior of elusive top predators.     

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.     

Tests of landscape influence: nest predation and brood parasitism in fragmented ecosystems

The effects of landscape fragmentation on nest predation and brood parasitism, the two primary causes of avian reproductive failure, have been difficult to generalize across landscapes, yet few studies have clearly considered the context and spatial scale of fragmentation. Working in two river systems fragmented by agricultural and rural-housing development, we tracked nesting success and brood parasitism in > 2500 bird nests in 38 patches of deciduous riparian woodland. Patches on both river systems were embedded in one of two local contexts (buffered from agriculture by coniferous forest, or adjacent to agriculture), but the abundance of agriculture and human habitation within 1 km of each patch was highly variable. We examined evidence for three models of landscape effects on nest predation based on (1) the relative importance of generalist agricultural nest predators, (2) predators associated with the natural habitats typically removed by agricultural development, or (3) an additive combination of these two predator communities. We found strong support for an additive predation model in which landscape features affect nest predation differently at different spatial scales. Riparian habitat with forest buffers had higher nest predation rates than sites adjacent to agriculture, but nest predation also increased with increasing agriculture in the larger landscape surrounding each site. These results suggest that predators living in remnant woodland buffers, as well as generalist nest predators associated with agriculture, affect nest predation rates, but they appear to respond at different spatial scales. Brood parasitism, in contrast, was unrelated to agricultural abundance on the landscape, but showed a strong nonlinear relationship with farm and house density, indicating a critical point at which increased human habitat causes increased brood parasitism. Accurate predictions regarding landscape effects on nest predation and brood parasitism will require an increased appreciation of the multiple scales at which landscape components influence predator and parasite behavior.     

Functional responses: a question of alternative prey and predator density

Throughout the study of ecology, there has been a growing realization that indirect effects among species cause complexity in food webs. Understanding and predicting the behavior of ecosystems consequently depends on our ability to identify indirect effects and their mechanisms. The present study experimentally investigates indirect interactions arising between two prey species that share a common predator. In a natural field experiment, we introduced different densities of mealworms (Tenebrio molitor), an alternative prey, to a previously studied predator-prey system in which paper wasps (Polistes dominulus) preyed on shield beetle larvae (Cassida rubiginosa). We tested if alternative prey affects predation on the first prey (i.e., the predator-dependent functional response of paper wasps) by modifying either interference among predators or the effective number of predators foraging on shield beetles. Presence of mealworms significantly reduced the effective number of predators, whereas predator interference was not affected. In this way, the experimentally introduced alternative prey altered the wasps' functional response and thereby indirectly influenced C. rubiginosa density. In all prey-density combinations offered, paper wasps constantly preferred T. molitor. This led to an asymmetrical, indirect interaction between both prey species: an increase in mealworm density significantly relaxed predation on C. rubiginosa, whereas an increase in C. rubiginosa density intensified predation on mealworms. Such asymmetrical outcomes of a fixed food preference can significantly affect the population dynamics of the species involved. In spite of the repeated finding of a Type III functional response in this system, our experiment did not reveal switching behavior in paper wasps. The variety of mechanisms underlying direct and indirect interactions within our study system exemplifies the importance of incorporating alternative prey when investigating the impact of a generalist predator on a focal prey population under realistic field conditions.     

Patterns of shell repair in articulate brachiopods indicate size constitutes a refuge from predation

The cost of overcoming prey defenses relative to the value of internal tissues is a key criterion in predator/prey interactions. Optimal foraging theory predicts: (1) specific sizes of prey will result in the best returns to predators, and (2) there will often be a size at which the cost/benefit balance is low enough to effectively exclude predation. Data presented here on styles of repaired shell damage and size at which injury had been sustained was collected from samples of terebratulide brachiopods from the Antarctic Peninisula (Liothyrella uva), Falkland Islands (Magellania venosa and Terebratella dorsata) and Chile (M. venosa). The predominant form of damage on shells was indicative of predators attacking the valve margins. The modal size for repaired damage was more than 10 mm smaller than the modal size for the overall size distribution in each species and there were no repaired attacks in the largest size classes of any species. These data suggest that size forms a refuge from predation, as would be predicted by optimal foraging theory. The optimal sizes that predators appeared to attack vary between species, as do the sizes that provided a refuge from predation. High levels of multiple repairs (19% of the M. venosa population from the Falkland Islands sampled had 2 or more repairs) suggest that the mortality following attack is low, suggesting that many predators abandon their attacks.     

Outbreak suppression by predators depends on spatial distribution of prey

The capacity of a predator population to suppress a prey population that varies in abundance and spatial distribution is explored in a lattice simulation model. The model is based on empirically derived parameters for particular species. Within season predation by Pterostichus cupreus (Coleoptera: Carabidae) of varying densities and distributions of the prey Rhopalosiphum padi (Homoptera: Aphididae) in spring cereals was simulated. From these spatially explicit simulations prey population suppression was found to be largely dependent on the spatial distribution of the prey. A possible mechanism was that high degrees of prey aggregation provided refuge for the prey that, when aggregated, escaped detection by P. cupreus. In contrast, P. cupreus was found to efficiently suppress incipient outbreaks for evenly distributed prey populations, even at high prey densities. A higher predator density compensated for the lowered control ability of the predators for highly aggregated prey populations and hastened the decline of the prey population.     

Adaptive prey behavior and the dynamics of intraguild predation systems

Intraguild predation constitutes a widespread interaction occurring across different taxa, trophic positions and ecosystems, and its endogenous dynamical properties have been shown to affect the abundance and persistence of the involved populations as well as those connected with them within food webs. Although optimal foraging decisions displayed by predators are known to exert a stabilizing influence on the dynamics of intraguild predation systems, few is known about the corresponding influence of adaptive prey decisions in spite of its commonness in nature. In this study, we analyze the effect that adaptive antipredator behavior exerts on the stability and persistence of the populations involved in intraguild predation systems. Our results indicate that adaptive prey behavior in the form of inducible defenses act as a stabilizing mechanism and show that, in the same direction that adaptive foraging, enhances the parameter space in which species can coexist through promoting persistence of the IG-prey. At high levels of enrichment, the intraguild predation system exhibits unstable dynamics and zones of multiples attractors. In addition, we show that the equilibrium density of the IG-predator could be increased at intermediate values of defense effectiveness. Finally we conclude that adaptive prey behavior is an important mechanism leading to species coexistence in intraguild predation systems and consequently enhancing stability of food webs.     

Defining ecologically relevant scales for spatial protection with long‐term data on an endangered seabird and local prey availability

Human activities are important drivers of marine ecosystem functioning. However, separating the synergistic effects of fishing and environmental variability on the prey base of nontarget predators is difficult, often because prey availability estimates on appropriate scales are lacking. Understanding how prey abundance at different spatial scales links to population change can help integrate the needs of nontarget predators into fisheries management by defining ecologically relevant areas for spatial protection. We investigated the local population response (number of breeders) of the Bank Cormorant (Phalacrocorax neglectus), a range‐restricted endangered seabird, to the availability of its prey, the heavily fished west coast rock lobster (Jasus lalandii). Using Bayesian state‐space modeled cormorant counts at 3 colonies, 22 years of fisheries‐independent data on local lobster abundance, and generalized additive modeling, we determined the spatial scale pertinent to these relationships in areas with different lobster availability. Cormorant numbers responded positively to lobster availability in the regions with intermediate and high abundance but not where regime shifts and fishing pressure had depleted lobster stocks. The relationships were strongest when lobsters 20–30 km offshore of the colony were considered, a distance greater than the Bank Cormorant's foraging range when breeding, and may have been influenced by prey availability for nonbreeding birds, prey switching, or prey ecology. Our results highlight the importance of considering the scale of ecological relationships in marine spatial planning and suggest that designing spatial protection around focal species can benefit marine predators across their full life cycle. We propose the precautionary implementation of small‐scale marine protected areas, followed by robust assessment and adaptive‐management, to confirm population‐level benefits for the cormorants, their prey, and the wider ecosystem, without negative impacts on local fisheries.     

Species-specific antipredator capacities and prey refuges: interactions between piscivorous perch (Perca fluviatilis) and juvenile perch and roach (Rutilus rutilus)

The outcome of predator-prey interactions depends on the characteristics of predators and prey as well as the structure of the environment. In a replicated field enclosure experiment, we tested the effects of quantity and quality of different prey refuges (no structure, structure forming a partial refuge, and structure forming a complete refuge) on the interaction between piscivorous perch (Perca fluviatilis) and juvenile perch and roach (Rutilus rutilus). We quantified the behaviour of the predators and the prey and predator-induced prey mortality. The piscivores stayed in or close to the prey refuge and were more dispersed in the presence than in the absence of prey refuges. Survival of juvenile perch and roach decreased in the presence of predators and was higher for juvenile roach than for juvenile perch. In addition, juvenile perch survival increased with refuge efficiency Roach formed schools which were denser in the presence of predators, had a higher swimming speed (both in the open water and in the refuge) and used a larger area than juvenile perch. Both prey species decreased their distance to the prey refuge and increased the proportion of their time spent in the refuge in the presence of predators. The number of switches between the open-water habitat and the prey refuge was higher for juvenile roach than for juvenile perch. Juvenile perch used different parts of the prey refuge in a flexible way depending both on presence of predators and refuge type whereas juvenile roach used the different parts of the prey refuge in fixed proportions over all refuge treatments. Our results suggest that juvenile roach had a overall higher capacity to avoid predation than juvenile perch. However, in the presence of qualitatively different prey refuges juvenile perch responded to predators with more flexible refuge use than juvenile roach. The differences in antipredator capacities of juvenile perch and roach when subjected to piscivorous perch predation may depend on differences in life history patterns of the two species.     

Linking marine predator diving behavior to local prey fields in contrasting habitats in a subarctic glacial fjord

Foraging theory predicts that animals will adjust their foraging behavior in order to maximize net energy intake and that trade-offs may exist that can influence their behavior. Although substantial advances have been made with respect to the foraging ecology of large marine predators, there is still a limited understanding of how predators respond to temporal and spatial variability in prey resources, primarily due to a lack of empirical studies that quantify foraging and diving behavior concurrently with characteristics of prey fields. Such information is important because changes in prey availability can influence the foraging success and ultimately fitness of marine predators. We assessed the diving behavior of juvenile female harbor seals (Phoca vitulina richardii) and prey fields near glacial ice and terrestrial haulout sites in Glacier Bay (58°40′N, ?136°05′W), Alaska. Harbor seals captured at glacial ice sites dived deeper, had longer dive durations, lower percent bottom time, and generally traveled further to forage. The increased diving effort for seals from the glacial ice site corresponded to lower prey densities and prey at deeper depths at the glacial ice site. In contrast, seals captured at terrestrial sites dived shallower, had shorter dive durations, higher percent bottom time, and traveled shorter distances to access foraging areas with much higher prey densities at shallower depths. The increased diving effort for seals from glacial ice sites suggests that the lower relative availability of prey may be offset by other factors, such as the stability of the glacial ice as a resting platform and as a refuge from predation. We provide evidence of differences in prey accessibility for seals associated with glacial ice and terrestrial habitats and suggest that seals may balance trade-offs between the costs and benefits of using these habitats.     

Local interactions between predators and prey call into question commonly used functional responses

Commonly used functional response models (Holling’s type I and type II models) assume that the encounter rate of a predator increases linearly with prey density, provided that the predator is searching for prey. In other other words, aN (a is the baseline encounter rate and N is prey density) describes the encounter rate. This study examined whether the models are adequate when predators and prey interact locally by using a spatially explicit individual based model because local interactions affect the spatial distribution of predators and prey, which also affects the encounter rate. Predators were assumed to possess a spatial perception range that influenced their foraging behavior (e.g., if a prey is in the perception range, the predator moves towards the prey). The effect of antipredator behavior by prey was also examined. The results suggest that prey and predator densities as well as handling time affect the baseline rate (i.e., parameter a) as opposed to the common assumption that the parameter is constant. The nature of model deviations depended on both the antipredator behavior and the predators’ perception range. Understanding these deviations is important as they qualitatively affect community dynamics.     

Patterns of Predation Risk and Survival of Bird Nests in a Chilean Agricultural Landscape

Abstract: We used experimental nests baited with California Quail (  Callipepla californica ) eggs or clay eggs to examine relative risks of nest predation in an agricultural landscape and in two large forest preserves in a south-temperate rainforest in Chile. The most common predators, as identified by marks on clay eggs, were a caracara (   Milvago chimango ), a blackbird ( Curaeus curaeus ), and rodents. Nest losses from predation were similar in large and small forest patches and lower in patches than in extensive forest. In general, predation risk was higher (and nest survival therefore lower) on forest edges than in forest interior, in short-grass pasture than in tall-grass pasture, in narrow corridors than in wide corridors, and on visible nests than on concealed nests. High predation risks in pasture habitat tended to increase the risk of nest predation in adjacent forest edges. For open-cup nesters, the risk of nest predation was relatively high in the present agricultural landscape, indicating that much of the available wooded habitat (  forest edges, narrow corridors) offers poor nesting habitat, although it may be suitable for foraging and traveling. The numerous bird-plant mutualisms in this landscape may be at risk if nesting success of the principal mutualists is consistently low.     

Trade-offs between maternal foraging and fawn predation risk in an income breeder

The choice of neonatal hiding place is critical for ungulates adopting hiding anti-predator strategies, but the consequences of different decisions have rarely been evaluated with respect to offspring survival. First, we investigated how landscape-scale choices made by roe deer fawns and their mothers affected predation risk by red foxes in a forest–farmland mosaic in southeastern Norway. After, we examined the effect of site-specific characteristics and behaviour (i.e. visibility, mother–fawn distance and abundance of the predator’s main prey item—small rodents) on predation risk. The study of habitat use, selection and habitat-specific mortality revealed that roe deer utilised the landscape matrix in a functional way, with different habitats used for feeding, providing maternal care and as refugia from predation. Mothers faced a trade-off between foraging and offspring survival. At the landscape-scale decisions were primarily determined by maternal energetic constraints and only secondarily by risk avoidance. Indeed, forage-rich habitats were strongly selected notwithstanding the exceptionally high densities of rodents which increased fawn predation. At fine spatial scales, a high visibility of the mother was the major factor determining predation risk; however, mothers adjusted their behaviour to the level of risk at the bed site to minimise predation. Fawns selected both landscape-scale refugia and concealed bed sites, but failure to segregate from the main prey of red foxes led to higher predation. This study provides evidence for the occurrence of spatial heterogeneity in predation risk and shows that energetically stressed individuals can tackle the foraging-safety trade-off by adopting scale-dependent anti-predator responses.     

The growth-predation risk trade-off under a growing gape-limited predation threat

Growth is a critical ecological trait because it can determine population demography, evolution, and community interactions. Predation risk frequently induces decreased foraging and slow growth in prey. However, such strategies may not always be favored when prey can outgrow a predator's hunting ability. At the same time, a growing gape-limited predator broadens its hunting ability through time by expanding its gape and thereby creates a moving size refuge for susceptible prey. Here, I explore the ramifications of growing gape-limited predators for adaptive prey growth. A discrete demographic model for optimal foraging/growth strategies was derived under the realistic scenario of gape-limited and gape-unconstrained predation threats. Analytic and numerical results demonstrate a novel fitness minimum just above the growth rate of the gape-limited predator. This local fitness minimum separates a slow growth strategy that forages infrequently and accumulates low but constant predation risk from a fast growth strategy that forages frequently and experiences a high early predation risk in return for lower future predation risk and enhanced fecundity. Slow strategies generally were advantageous in communities dominated by gape-unconstrained predators whereas fast strategies were advantageous in gape-limited predator communities. Results were sensitive to the assumed relationships between prey size and fecundity and between prey growth and predation risk. Predator growth increased the parameter space favoring fast prey strategies. The model makes the testable predictions that prey should not grow at the same rate as their gape-limited predator and generally should grow faster than the fastest growing gape-limited predator. By focusing on predator constraints on prey capture, these results integrate the ecological and evolutionary implications of prey growth in diverse predator communities and offer an explanation for empirical growth patterns previously viewed to be anomalies.