Under pressure: Short- and long-term response to predation varies in two populations of a live-bearing fish

The non-lethal effects of predation can significantly influence animal behavior and population composition. Research has often centered around prey response to predator exposure in the short term, but fewer studies have highlighted the effects of long-term predator exposures. In addition, studies of responses to predation risk are not always calibrated against the ecological history of predation risk in specific populations. We address these gaps by examining the effects of both long- and short-term predator exposure on the behavior of individuals from populations that have different ecological histories of predation risk. We exposed individuals from high-predation and low-predation populations of the live-bearing freshwater poeciliid, Heterandria formosa, to predators to assess changes in male reproductive behavior toward females. We also assessed longer-term reproductive responses by exposing male and female H. formosa to predators at a random time of day, every day, for 30 days. In the presence of a predator, in the short term, males changed the frequency of their behaviors and females varied in their concentration of cortisol, demonstrating immediate responses to the perceived risk. The magnitude of these changes was larger in the population without a long history of predator exposure. However, we found that males and females did not change their reproductive output when exposed to predators over longer periods of time, suggesting that individuals acclimatize to the level of predation risk they experience. These results also suggest that short-term variation in behavior or stress hormone responses should not be used as proxies for long-term responses or fitness effects. Future work should assess both short-term behavior and long-term responses while simultaneously considering the ecological history of populations.     

Sexual Differences in the Behavioral Response of Túngara Frogs, Physalaemus pustulosus, to Cues Associated with Increased Predation Risk

Engaging in mating behaviors usually increases exposure to predators for both males and females. Anti‐predator strategies during reproduction may have important fitness consequences for prey. Previous studies have shown that individuals of several species adjust their reproductive behavior according to their assessment of predation risk, but few studies have explored potential sexual differences in these strategies. In this study, we investigate whether the acoustic cues associated with predatory attacks or those associated with predators themselves affect the mating behavior of female and male túngara frogs, Physalaemus pustulosus. We compared the responses of females approaching a mate and those of calling males when exposed to mating calls associated with sounds representing increased hazard. When presented with mating calls that differed only in whether or not they were followed by a predation‐related sound, females preferentially approached the call without predation‐related sounds. In contrast to females, calling males showed greater vocal response to calls associated with increased risk than to a call by itself. We found significant differences in the responses of females and males to several sounds associated with increased hazard. Females behaved more cautiously than males, suggesting that the sexes balance the risk of predation and the cost of cautious mating strategies differently.     

Manipulating Individual Decisions and Environmental Conditions Reveal Individual Quality in Decision-Making and Non-Lethal Costs of Predation Risk

Habitat selection is a crucial decision for any organism. Selecting a high quality site will positively impact survival and reproductive output. Predation risk is an important component of habitat quality that is known to impact reproductive success and individual condition. However, separating the breeding consequences of decision-making of wild animals from individual quality is difficult. Individuals face reproductive decisions that often vary with quality such that low quality individuals invest less. This reduced reproductive performance could appear a cost of increased risk but may simply reflect lower quality. Thus, teasing apart the effects of individual quality and the effect of predation risk is vital to understand the physiological and reproductive costs of predation risk alone on breeding animals. In this study we alter the actual territory location decisions of pied flycatchers by moving active nests relative to breeding sparrowhawks, the main predators of adult flycatchers. We experimentally measure the non-lethal effects of predation on adults and offspring while controlling for effects of parental quality, individual territory choice and initiation of breeding. We found that chicks from high predation risk nests (<50 m of hawk) were significantly smaller than chicks from low risk nests (>200 m from hawk). However, in contrast to correlative results, females in manipulated high risk nests did not suffer decreased body condition or increased stress response (HSP60 and HSP70). Our results suggest that territory location decisions relative to breeding avian predators cause spatial gradients in individual quality. Small adjustments in territory location decisions have crucial consequences and our results confirm non-lethal costs of predation risk that were expressed in terms of smaller offspring produced. However, females did not show costs in physiological condition which suggests that part of the costs incurred by adults exposed to predation risk are quality determined.     

Precipitation and predation risk alter the diversity and behavior of pollinators and reduce plant fitness

Biotic and abiotic factors may individually or interactively disrupt plant–pollinator interactions, influencing plant fitness. Although variations in temperature and precipitation are expected to modify the overall impact of predators on plant–pollinator interactions, few empirical studies have assessed if these weather conditions influence anti-predator behaviors and how this context-dependent response may cascade down to plant fitness. To answer this question, we manipulated predation risk (using artificial spiders) in different years to investigate how natural variation in temperature and precipitation may affect diversity (richness and composition) and behavioral (visitation) responses of flower-visiting insects to predation risk, and how these effects influence plant fitness. Our findings indicate that predation risk and an increase in precipitation independently reduced plant fitness (i.e., seed set) by decreasing flower visitation. Predation risk reduced pollinator visitation and richness, and altered species composition of pollinators. Additionally, an increase in precipitation was associated with lower flower visitation and pollinator richness but did not alter pollinator species composition. However, maximum daily temperature did not affect any component of the pollinator assemblage or plant fitness. Our results indicate that biotic and abiotic drivers have different impacts on pollinator behavior and diversity with consequences for plant fitness components. Even small variation in precipitation conditions promotes complex and substantial cascading effects on plants by affecting both pollinator communities and the outcome of plant–pollinator interactions. Tropical communities are expected to be highly susceptible to climatic changes, and these changes may have drastic consequences for biotic interactions in the tropics.     

Behavioral responses of a sex-role reversed pipefish to a gradient of perceived predation risk

Conspicuous behaviors such as courtship and mating often makeanimals susceptible to predation. When perceiving themselvesat an elevated level of risk, animals frequently reduce conspicuousbehaviors in tradeoff for a decrease in probability of beingpreyed upon. In the present study, we used two experiments toexamine the effect of perceived predation risk from cod (Gadusmorhud) on nonreproductive and reproductive behaviors in thesex-role reversed pipefish (Syngnathus typhle). In the firstexperiment, no differences due to predation risk were detectedin the nonreproductive behaviors of either males or females.In the second experiment, predation risk had significant effectson reproductive behaviors. Pipefish were allowed to court andcopulate at four different predation levels. We created predationlevels differing in perceived predation risk by controllingthe number of sensory modes through which pipefish could detectthe presence of a cod. As predation risk increased, pipefishcopulated and courted less frequently, swam alone (displayedand searched for conspecifics) less often, and waited longerbefore commencing courtship. These changes in behavior minimizedthe amount of time spent above the eelgrass and presumably reducedconspicuousness to visual predators. Pipefish also copulatedafter a smaller amount of courtship as predation risk increased,indicating that they may trade information concerning mate qualityfor a reduction in predation risk. No differences were foundin any response variable between males and females. The roleof operational sex ratios and intersexual competition in determiningwhich sex assumes greater costs in mate acquisition is questioned.     

Vigilance and fitness in grey partridges Perdix perdix: the effects of group size and foraging-vigilance trade-offs on predation mortality

1. Vigilance increases fitness by improving predator detection but at the expense of increasing starvation risk. We related variation in vigilance among 122 radio-tagged overwintering grey partridges Perdix perdix (L.) across 20 independent farmland sites in England to predation risk (sparrowhawk Accipiter nisus L., kill rate), use of alternative antipredation behaviours (grouping and use of cover) and survival. 2. Vigilance was significantly higher when individuals fed in smaller groups and in taller vegetation. In the covey period (in early winter when partridges are in flocks), vigilance and use of taller vegetation was significantly higher at sites with higher sparrowhawk predation risk, but tall vegetation was used less by larger groups. Individuals were constrained in reducing individual vigilance by group size and habitat choice because maximum group size was determined by overall density in the area during the covey period and by the formation of pairs at the end of the winter (pair period), when there was also a significant twofold increase in the use of tall cover. 3. Over the whole winter individual survival was higher in larger groups and was lower in the pair period. However, when controlling for group size, mean survival decreased as vigilance increased in the covey period. This result, along with vigilance being higher at sites with increasing with raptor risk, suggests individual vigilance increases arose to reduce short-term predation risk from raptors but led to long-term fitness decreases probably because high individual vigilance increased starvation risk or indicated longer exposure to predation. The effect of raptors on survival was less when there were large groups in open habitats, where individual partridges can probably both detect predators and feed efficiently. 4. Our study suggests that increasing partridge density and modifying habitat to remove the need for high individual vigilance may decrease partridge mortality. It demonstrates the general principle that antipredation behaviours may reduce fitness long-term via their effects on the starvation-predation risk trade-off, even though they decrease predation risk short-term, and that it may be ecological constraints, such as poor habitat (that lead to an antipredation behaviour compromising foraging), that cause mortality, rather than the proximate effect of an antipredation behaviour such as vigilance.     

Reproduction under predation risk and the trade-off between current and future reproduction in the threespine stickleback

An increasing number of studies show that animals adjust their reproductive effort to the risk of predation. However, to maximize lifetime reproductive success this adjustment should depend on the animals'' current and future reproductive potential. Here I tested this hypothesis by allowing threespine stickleback males (Gasterosteus aculeatus), differing in current and future mating probabilities, to reproduce in pools in both the presence and absence of predators. As expected, males adjusted their reproductive effort to the risk of predation. Fewer males bred, and all males developed less nuptial coloration in the presence of predators. However, males with a low current mating probability took less risk than males with a higher mating probability, whereas all males increased risk taking when future reproductive opportunities decreased. The results thus support the hypothesis that males are able to assess both the risk of predation and their current versus future mating probability, and adjust their reproductive decisions accordingly. The study further suggests that predation risk may have less effect on sexual selection than previously assumed, as the males which refrained from reproducing in the presence of predators were mainly males with a low mating probability.     

Optimization of reproductive effort and foraging time in mammals: The influence of resource level and predation risk

Summary The trade-off between fitness benefits from foraging and associated costs in terms of predation risk is analysed by a simple model which takes into account the differential predation risk for reproducing and non-reproducing individuals. The currency that animals are assumed to maximize is their expected absolute fitness (probability of survival plus half of the expected litter size) after a potential reproductive period. Depending on resource levels and predation risk, this maximization can be achieved by (1) opting for individual survival and behaving as a strict time minimizer, (2) by reproducing at the maximal rate and behaving as a strict energy maximizer or (3) by submaximal reproductive effort and a behaviour intermediate between time minimization and energy maximization. Small changes in the availability of food or cover or in the density of predators can shift the optimum from one strategy to another. The shift is particularly abrupt, if predation pressure increases and the availability of resources remains high. This could explain the spatial and temporal variation in the reproductive effort and body weight observed in boreal small mammals with sustained, multiannual population fluctuations.     

Predation-Related Costs and Benefits of Conspecific Attraction in Songbirds—An Agent-Based Approach

Songbirds that follow a conspecific attraction strategy in the habitat selection process prefer to settle in habitat patches already occupied by other individuals. This largely affects the patterns of their spatio-temporal distribution and leads to clustered breeding. Although making informed settlement decisions is expected to be beneficial for individuals, such territory clusters may potentially provide additional fitness benefits (e.g., through the dilution effect) or costs (e.g., possibly facilitating nest localization if predators respond functionally to prey distribution). Thus, we hypothesized that the fitness consequences of following a conspecific attraction strategy may largely depend on the composition of the predator community. We developed an agent-based model in which we simulated the settling behavior of birds that use a conspecific attraction strategy and breed in a multi-predator landscape with predators that exhibited different foraging strategies. Moreover, we investigated whether Bayesian updating of prior settlement decisions according to the perceived predation risk may improve the fitness of birds that rely on conspecific cues. Our results provide evidence that the fitness consequences of conspecific attraction are predation-related. We found that in landscapes dominated by predators able to respond functionally to prey distribution, clustered breeding led to fitness costs. However, this cost could be reduced if birds performed Bayesian updating of prior settlement decisions and perceived nesting with too many neighbors as a threat. Our results did not support the hypothesis that in landscapes dominated by incidental predators, clustered breeding as a byproduct of conspecific attraction provides fitness benefits through the dilution effect. We suggest that this may be due to the spatial scale of songbirds’ aggregative behavior. In general, we provide evidence that when considering the fitness consequences of conspecific attraction for songbirds, one should expect a trade-off between the benefits of making informed decisions and the costs of clustering.     

Does an opportunistic predator preferentially attack nonvigilant prey?

The dilution effect as an antipredation behaviour is the main theoretical reason for grouping in animals and states that all individuals in a group have an equal risk of being predated if equally spaced from each other and the predator. Stalking predators, however, increase their chance of attack success by preferentially targeting nonvigilant individuals, potentially making relative vigilance rates in a group relatively important in determining predation compared with the dilution effect. Many predators, however, attack opportunistically without stalking, when targeting of nonvigilant individuals may be less likely, so that the dilution effect will then be a relatively more important antipredation reason for grouping. We tested whether an opportunistically hunting predator, the sparrowhawk, Accipiter nisus, preferentially attacked vigilant or feeding prey models presented in pairs. We found that sparrowhawks attacked vigilant and feeding mounts at similar frequencies. Our results suggest that individuals should prioritize maximizing group size or individual vigilance dependent on the type of predator from which they are at risk. When the most likely predator is a stalker, individuals should aim to have the highest vigilance levels in a group, and there may be relatively little selective advantage to being in the largest group. In contrast, if the most likely predator is an opportunist, then individuals should simply aim to be in the largest group and can also spend more time foraging without compromising predation risk. For most natural systems this will mean a trade-off between the two strategies dependent on the frequency of attack of each predator type. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.     

Fear factor: prey habitat selection and its consequences in a predation risk landscape

Predation risk influences prey use of space. However, little is known about how predation risk influences breeding habitat selection and the fitness consequences of these decisions. The nest sites of central-place foraging predators may spatially anchor predation risk in the landscape. We explored how the spatial dispersion of avian predator nests influenced prey territory location and fitness related measures. We placed 249 nest boxes for migrant pied flycatchers Ficedula hypoleuca , at distances between 10 and 630 m, around seven different sparrowhawk nests Accipiter nisus . After closely monitoring flycatcher nests we found that flycatcher arrival dates, nest box occupation rates and clutch size showed a unimodal relationship with distance from sparrowhawk nests. This relationship suggested an optimal territory location at intermediate distances between 330 and 430 m from sparrowhawk nests. Furthermore, pied flycatcher nestling quantity and quality increased linearly with distance from sparrowhawk nests. These fitness related measures were between 4 and 26% larger in flycatcher nestlings raised far from, relative to those raised nearby, sparrowhawk nests. Our results suggest that breeding sparrowhawk affected both flycatcher habitat selection and reproductive success. We propose that nesting predators create predictable spatial variation in predation risk for both adult prey and possibly their nests, to which prey individuals are able to adaptively respond. Recognising predictable spatial variation in perceived predation risk may be fundamental for a proper understanding of predator-prey interactions and indeed prey species interactions.     

The interaction between predation risk and food ration on behavior and morphology of Eurasian perch

The risk of both predation and food level has been shown to affect phenotypic development of organisms. However, these two factors also influence animal behavior that in turn may influence phenotypic development. Hence, it might be difficult to disentangle the behavioral effect from the predator or resource‐level effects. This is because the presence of predators and high resource levels usually results in a lower activity, which in turn affects energy expenditure that is used for development and growth. It is therefore necessary to study how behavior interacts with changes in body shape with regard to resource density and predators. Here, we use the classic predator‐induced morphological defense in fish to study the interaction between predator cues, resource availability, and behavioral activity with the aim to determine their relative contribution to changes in body shape. We show that all three variables, the presence of a predator, food level, and activity, both additively and interactively, affected the body shape of perch. In general, the presence of predators, lower swimming activity, and higher food levels induced a deep body shape, with predation and behavior having similar effect and food treatment the smallest effect. The shape changes seemed to be mediated by changes in growth rate as body condition showed a similar effect as shape with regard to food‐level and predator treatments. Our results suggests that shape changes in animals to one environmental factor, for example, predation risk, can be context dependent, and depend on food levels or behavioral responses. Theoretical and empirical studies should further explore how this context dependence affects fitness components such as resource gain and mortality and their implications for population dynamics.     

Predation risk effects on fitness related measures in a resident bird

Predation risk is thought to be highly variable in space and time. However, breeding avian predators may create locally fixed and spatially fairly predictable predation risk determined by the distance to their nest. From the prey perspective, this creates predation risk gradients that potentially have an effect on fitness and behavioural decisions of prey. We studied how breeding avian predators affect habitat selection (nest location) and the resulting fitness consequences in a northern population of resident willow tit ( Parus montanus ). Data included 429 willow tit nests over a four year period in a landscape containing a total of 33 avian predator nests. Willow tit nests were located randomly in the landscape and no predator avoidance in habitat selection or emptying of territories in proximity to predators was observed. Nestling size, however, was positively associated with distance from predator nests (n=252). Nestling mass and wing length were about 4.5% smaller close to predator nests compared to nestlings raised far from predator nests. Tarsus length also exhibited a positive relationship with increasing distance from predator nest but this was limited to habitats of young forests and pine bogs or dense mixed forests (4% increase). It is likely that habitat structural complexity influenced the perception of predation risk in different habitats. Our results indicate that willow tits do not provide reliable cues of predator free habitats for settling migrants. Nonetheless, breeding avian predators may create predictable predation risk in the landscape which is an important factor affecting reproductive success and potentially the demography of prey populations.     

Squeezing out the last egg—annual fish increase reproductive efforts in response to a predation threat

Both constitutive and inducible antipredator strategies are ubiquitous in nature and serve to maximize fitness under a predation threat. Inducible strategies may be favored over constitutive defenses depending on their relative cost and benefit and temporal variability in predator presence. In African temporary ponds, annual killifish of the genus Nothobranchius are variably exposed to predators, depending on whether larger fish invade their habitat from nearby rivers during floods. Nonetheless, potential plastic responses to predation risk are poorly known. Here, we studied whether Nothobranchius furzeri individuals adjust their life history in response to a predation threat. For this, we monitored key life history traits in response to cues that signal the presence of predatory pumpkinseed sunfish (Lepomis gibbosus). While growth rate, adult body size, age at maturation, and initial fecundity were not affected, peak and total fecundity were higher in the predation risk treatment. This contrasts with known life history strategies of killifish from permanent waters, which tend to reduce reproduction in the presence of predators. Although our results show that N. furzeri individuals are able to detect predators and respond to their presence by modulating their reproductive output, these responses only become evident after a few clutches have been deposited. Overall our findings suggest that, in the presence of a predation risk, it can be beneficial to increase the production of life stages that can persist until the predation risk has faded.     

Spatial decomposition of predation risk using resource selection functions: an example in a wolf–elk predator–prey system

Predation is a fundamental ecological and evolutionary process that varies in space, and the avoidance of predation risk is of central importance in foraging theory. While there has been a recent growth of approaches to spatially model predation risk, these approaches lack an adequate mechanistic framework that can be applied to real landscapes. In this paper we show how predation risk can be decomposed into encounter and attack stages, and modeled spatially using resource selection functions (RSF) and resource selection probability functions (RSPF). We use this approach to compare the effects of landscape attributes on the relative probability of encounter, the conditional probability of death given encounter, and overall wolf and elk resource selection to test whether predation risk is simply equivalent to location of the predator. We then combine the probability of encounter and conditional probability of death into a spatially explicit function of predation risk following Lima and Dill's reformulation of Holling's functional response. We illustrate this approach in a wolf–elk system in and adjacent to Banff National Park, Alberta, Canada. In this system we found that the odds of elk being encountered by wolves was 1.3 times higher in pine forest and 4.1 times less in grasslands than other habitats. The relative odds of being killed in pine forests, given an encounter, increased by 1.2. Other habitats, such as grasslands, afforded elk reduced odds (4.1 times less) of being encountered and subsequently killed (1.4 times less) by wolves. Our approach illustrates that predation risk is not necessarily equivalent to just where predators are found. We show that landscape attributes can render prey more or less susceptible to predation and effects of landscape features can differ between the encounter and attack stages of predation. We conclude by suggesting applications of our approach to model predator–prey dynamics using spatial predation risk functions in theoretical and applied settings.     

Defending the Weak: Parental Defense Peaks When Chick Vulnerability is Greatest in the Herring Gull (Larus argentatus)

Successful reproduction often depends upon parents providing offspring with resources and protection. In birds, reproductive success can often be enhanced by parents engaging in antipredator behaviors, but these behaviors can be costly. Theoretically, individuals should temporally modify the intensity of nest defense behavior to balance the costs and benefits of current and future reproductive success. More specifically, nest defense should vary throughout a nesting attempt to maximize fitness of the adults. Here, we consider the relationship between nest defense behavior and chick vulnerability in the herring gull (Larus argentatus), where chicks are under high predation risk. We estimated chick vulnerability by quantifying survival probabilities at different periods of the nestling stage. Simultaneously, we quantified changes in parental aggression throughout the nesting cycle by simulating predation attempts using a human predator model. We found that chick survival probabilities were lowest (i.e., vulnerability was highest) and parental aggression in nest defense was greatest during the first 10 days after hatching. Thus, we show that parents are most defensive when chicks are most vulnerable and that adults optimize nest defense behaviors in a way that maximizes their fitness.     

Behavioural Response of Bats to Perceived Predation Risk While Foraging

The ability to detect and respond to predation risk while foraging may have important fitness consequences for prey organisms. Anti‐predator behaviours may reduce the probability of mortality because of predation, but they may also be associated with reduced foraging efficiency. Several behaviours of bats have been suggested to serve as anti‐predator responses, and there is evidence that predation, particularly by avian predators such as owls, may be an important cause of bat mortality. Previous studies have attempted to determine whether predator presence affects bat behaviour when emerging from roost sites, but few have examined effects of predator presence on bat behaviour while foraging. In this study, we investigated whether foraging bats respond to predator cues by presenting bats with an acoustic cue simulating the presence of an owl. Within matched trials, which were conducted at different locations each of 18 nights, significantly fewer bat detections were recorded at owl playback stations than at control stations (no auditory cue), suggesting an avoidance response by bats. An acoustic control (i.e. station playing woodpecker calls), however, did not have significantly more detections than the stations playing the owl calls, suggesting that bats may simply be avoiding noise and more detailed investigation is warranted. Although evidence for owl predation on bats is minimal in North America, results of this study may indicate that the perceived presence of owls may represent a factor influencing the behaviour of bats while foraging.     

Breeding Phenology of Birds: Mechanisms Underlying Seasonal Declines in the Risk of Nest Predation

Seasonal declines in avian clutch size are well documented, but seasonal variation in other reproductive parameters has received less attention. For example, the probability of complete brood mortality typically explains much of the variation in reproductive success and often varies seasonally, but we know little about the underlying cause of that variation. This oversight is surprising given that nest predation influences many other life-history traits and varies throughout the breeding season in many songbirds. To determine the underlying causes of observed seasonal decreases in risk of nest predation, we modeled nest predation of Dusky Flycatchers (Empidonax oberholseri) in northern California as a function of foliage phenology, energetic demand, developmental stage, conspecific nest density, food availability for nest predators, and nest predator abundance. Seasonal variation in the risk of nest predation was not associated with seasonal changes in energetic demand, conspecific nest density, or predator abundance. Instead, seasonal variation in the risk of nest predation was associated with foliage density (early, but not late, in the breeding season) and seasonal changes in food available to nest predators. Supplemental food provided to nest predators resulted in a numerical response by nest predators, increasing the risk of nest predation at nests that were near supplemental feeders. Our results suggest that seasonal changes in foliage density and factors associated with changes in food availability for nest predators are important drivers of temporal patterns in risk of avian nest predation.     

Threat-sensitive anti-intraguild predation behaviour: maternal strategies to reduce offspring predation risk in mites

Predation is a major selective force for the evolution of behavioural characteristics of prey. Predation among consumers competing for food is termed intraguild predation (IGP). From the perspective of individual prey, IGP differs from classical predation in the likelihood of occurrence because IG prey is usually more rarely encountered and less profitable because it is more difficult to handle than classical prey. It is not known whether IGP is a sufficiently strong force to evolve interspecific threat sensitivity in antipredation behaviours, as is known from classical predation, and if so whether such behaviours are innate or learned. We examined interspecific threat sensitivity in antipredation in a guild of predatory mite species differing in adaptation to the shared spider mite prey (i.e. Phytoseiulus persimilis, Neoseiulus californicus and Amblyseius andersoni). We first ranked the players in this guild according to the IGP risk posed to each other: A. andersoni was the strongest IG predator; P. persimilis was the weakest. Then, we assessed the influence of relative IGP risk and experience on maternal strategies to reduce offspring IGP risk: A. andersoni was insensitive to IGP risk. Threat sensitivity in oviposition site selection was induced by experience in P. persimilis but occurred independently of experience in N. californicus. Irrespective of experience, P. persimilis laid fewer eggs in choice situations with the high- rather than low-risk IG predator. Our study suggests that, similar to classical predation, IGP may select for sophisticated innate and learned interspecific threat-sensitive antipredation responses. We argue that such responses may promote the coexistence of IG predators and prey.     

Optimal diving under the risk of predation

Many air-breathing aquatic foragers may be killed by aerial or subsurface predators while recovering oxygen at the surface; yet the influence of predation risk on time allocation during dive cycles is little known in spite of numerous studies on optimal diving. We modeled diving behavior under the risk of predation at the surface. The relationship between time spent at the surface and the risk of death is predicted to influence the optimal surface interval, regardless of whether foragers accumulate energy at a constant rate while at the food patch, deplete food resources over the course of the dive, or must search for food during the dive. When instantaneous predation risk during a single surface interval decreases with time spent at the surface, a diver should increase its surface interval relative to that which maximizes energy intake, thereby increasing dive durations and reducing the number of surfacings per foraging bout. When instantaneous risk over a single surface interval does not change or increases with increasing time at the surface, divers should decrease their surface interval (and consequently their dive duration) relative to that which maximizes energy intake resulting in more dives per foraging bout. The fitness consequences of selecting a suboptimal surface interval vary with the risk function and the way divers harvest energy when at depth. Finally, predation risk during surface intervals should have important consequences for habitat selection and other aspects of the behavioral ecology of air-breathing aquatic organisms.