From process to pattern: how fluctuating predation risk impacts the stress axis of snowshoe hares during the 10-year cycle

Predation is a central organizing process affecting populations and communities. Traditionally, ecologists have focused on the direct effects of predation—the killing of prey. However, predators also have significant sublethal effects on prey populations. We investigated how fluctuating predation risk affected the stress physiology of a cyclic population of snowshoe hares (Lepus americanus) in the Yukon, finding that they are extremely sensitive to the fluctuating risk of predation. In years of high predator numbers, hares had greater plasma cortisol levels at capture, greater fecal cortisol metabolite levels, a greater plasma cortisol response to a hormone challenge, a greater ability to mobilize energy and poorer body condition. These indices of stress had the same pattern within years, during the winter and over the breeding season when the hare:lynx ratio was lowest and the food availability the worst. Previously we have shown that predator-induced maternal stress lowers reproduction and compromises offspring’s stress axis. We propose that predator-induced changes in hare stress physiology affect their demography through negative impacts on reproduction and that the low phase of cyclic populations may be the result of predator-induced maternal stress reducing the fitness of progeny. The hare population cycle has far reaching ramifications on predators, alternate prey, and vegetation. Thus, predation is the predominant organizing process for much of the North American boreal forest community, with its indirect signature—stress in hares—producing a pattern of hormonal changes that provides a sensitive reflection of fluctuating predator pressure that may have long-term demographic consequences.     

The consequences of predator control for brown hares (<Emphasis Type="Italic">Lepus europaeus</Emphasis>) on UK farmland

The brown hare Lepus europaeus is a valued game species but also a species of conservation concern owing to its severe decline in abundance on farmland throughout Europe during the twentieth century. Changes in the farmland habitat and predation have both been cited as causative factors. Their relative roles have been unclear, but most conservation action has focused on improving habitat. We analyse data from a sequence of three unique studies (one experiment and two demonstrations) covering the period 1985–2006 in which control of several common predator species was undertaken to increase densities of wild game on farmland in England. Across the three studies, regression modelling of the proportional change in hare numbers between successive years showed that—after site, year differences and harvesting were accounted for—predator control was a significant determinant of hare population change. Where habitat improvement also took place, hares reached autumn densities that were exceptional for the UK and which could sustain substantial harvests. When predation control was stopped, hare densities fell, even where habitat improvements remained in place. This analysis demonstrates that even where farmland habitat is greatly improved, uncontrolled predation prevents hares making full use of its carrying capacity. This helps explain the mixed—and at best modest—success of agri-environment schemes in the UK and elsewhere in Europe to increase hare densities. Game-shooting estates, on which effective predator control takes place, probably have a special significance within the landscape as source areas for brown hares.     

Predator-Mediated Indirect Effects of Snowshoe Hares on Dall's Sheep in Alaska

ABSTRACT Indirect interactions among species can strongly influence population dynamics and community structure but are often overlooked in management of large mammals. We estimated survival of Dall's sheep (Ovis dalli) in the central Alaska Range, USA, during years of differing snowshoe hare (Lepus americanus) abundance to test whether indirect interactions with a cyclic hare population affect Dall's sheep either negatively, by subsidizing predators (apparent competition), or positively, by diverting predation (apparent commensalism). Annual survival of adult female sheep was consistently high (0.85 for all yr and age classes combined). In contrast, annual estimates of lamb survival ranged from 0.15 to 0.63. The main predators of lambs were coyotes (Canis latrans) and golden eagles (Aquila chrysaetos), which rely on hares as their primary food and prey on lambs secondarily. Coyotes and eagles killed 78% of 65 radiocollared lambs for which cause of death was known. Lamb survival was negatively related to hare abundance during the previous year, and lamb survival rates more than doubled when hare abundance declined, supporting the hypothesis of predator-mediated apparent competition between hares and sheep. However, stage-specific predation and delays in predator responses to changes in hare numbers led to a positive relationship between abundance of adult Dall's sheep and hares. Lacking reliable estimates of survival, a manager might erroneously conclude that hares benefit sheep. Thus, support for different indirect effects can be obtained from different types of data, which demonstrates the need to determine the mechanisms that create indirect interactions. Long-term survey data suggest that predation by coyotes is limiting this sheep population below levels typical when coyotes were rare or absent. Understanding the nature of indirect interactions is necessary to effectively manage complex predator–prey communities.     

Implications of shared predation for space use in two sympatric leporids

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

Coyote prey selection and community stability during a decline in food supply

The foraging behavior of predators can have a large influence on community dynamics and has been shown to increase stability in some cases and decrease stability in others. I studied the foraging behavior of coyotes ( Canis latrans ) in the Alaska Range during the peak and decline of a snowshoe hare ( Lepus americanus ) population cycle (1999–2002). Coyote diet was compared with prey availability to test for changes in prey selection and to examine the effect of coyote predation on the vertebrate prey community. Coyotes responded to the hare decline by increasing selection for hares and porcupines, whereas selection for voles, ground squirrels and Dall sheep did not change. Although the study area was characterized by considerable habitat heterogeneity, coyotes utilized the area as a fine-grained environment. Coyote foraging behavior was driven primarily by changes in snowshoe hare abundance, and their sensitivity to change in alternative prey density was low. Predation by coyotes may therefore decrease the stability of alternative prey populations rather than dampening fluctuations. In order for predation to enhance the stability of prey populations, I hypothesize that prey profitability must be determined primarily by abundance.     

Modelling the Canada lynx and snowshoe hare population cycle: the role of specialist predators

Mathematical models of the snowshoe hare (Lepus americanus) and Canada lynx (Lynx canadensis) population cycles in the boreal forest have largely focused on the interaction between a single specialist predator and its prey. Here, we consider the role that other hare predators play in shaping the cycles, using a predator–prey model for up to three separate specialist predators. We consider the Canada lynx, coyote (Canis latrans) and great horned owl (Bubo virginianus). Our model improves on past modelling efforts in two ways: (1) our model solutions more closely represent the boreal hare and predator cycles with respect to the cycle period, maximum and minimum hare densities and maximum and minimum predator densities for each predator, and (2) our model sheds light on the role each specialist plays in regulation of the hare cycle, in particular, the dynamics of the raptor appear to be crucial for characterising the low hare densities correctly.     

Viable populations in a prey–predator system

 Lotka–Volterra equations are considered a dynamical game, where the phenotypes of the predator and of the prey can vary. This differs from the usual procedure of specifying as a priori laws according to which strategies are supposed to change. The question at stake is the survival of each of the species, instead of the maximization of a given pay-off by each player, as it is commonly discussed in games. The predator needs the prey, while the prey can survive without the predator. These obvious and simplistic constraints are enough to shape the regulation of the system: notably, the largest closed set of initial conditions can be delineated, from which there exists at least one evolutionary path where the population can avoid extinction forever. To these so-called viable trajectories, viable strategies are associated, respectively for the prey or for the predator. A coexistence set can then be defined. Within this set and outside the boundary, strategies can vary arbitrarily within given bounds while remaining viable, whereas on the boundary, only specific strategies can guarantee the viability of the system. Thus, the largest set can be determined, outside of which strategies will never be flexible enough to avoid extinction. Received 2 May 1995; received in revised form 15 August 1995     

Optimal foraging: the difficulty of exploiting different feeding strategies simultaneously

Summary The foraging efficiency of a visually feeding fish, perch (Perca fluviatilis) was studied on two prey species (Daphnia magna and Chaoborus obscuripus) presented either separately or combined. It is shown that when both prey species are present, the foraging efficiency of the predator is reduced. This is due to the predator's inability to simultaneously cope with prey species with different anti-predatory behaviour. In the mixed-meal experiment the predator captured both prey species in equal proportions in disagreement with optimal foraging models assuming that handling time and encounter rate for a prey species are independent of other prey species. The results are, however, in agreement with optimal foraging models assuming that handling time and encounter rate are influenced by short time learning.     

Escape and alerting responses by Balearic lizards (Podarcis lilfordi) to movement and turning direction by nearby predators

Escape theory predicts that prey monitoring an approaching predator delay escape until predation risk outweighs costs of fleeing. However, if a predator is not detected until it is closer than the optimal flight initiation distance (FID = distance between predator and prey when escape begins), escape should begin immediately. Similarly, if a change in a nearby predator’s behavior indicates increased risk, the optimal FID increases, sometimes inducing immediate escape. If a predator that has been standing immobile near a prey suddenly turns toward the prey, greater risk is implied than if the predator turns away. If the immobile predator suddenly moves its foot without turning, it might be launching an attack. Therefore, we predicted that frequency of fleeing and preparation to flee are greater when a predator turns toward than away from prey and that frequency of fleeing when a predator suddenly moves decreases as distance between predator and prey increases. We verified these predictions in the Balearic lizard Podarcis lilfordi in field experiments in which an investigator simulated the predator. Lizards fled and performed alerting responses indicating readiness to flee more frequently when the predator turned toward than away from them, and fled more frequently the nearer the predator.     

Snowshoe hare demography during a cyclic population low

1. Snowshoe hare ( Lepus americanus Erxleben) populations were studied in south-west Yukon during the low phase of the 10-year population cycle. Food availability and predator abundance were manipulated in a factorial design to determine the importance of each factor in hare dynamics during this phase.
2. Food was abundant during the low phase, and snowshoe hares were not food limited.
3. Survival of hares was higher than at any other phase of the cycle, and predators were scarce, but >75% of hare deaths resulted from predation.
4. Food addition resulted in higher hare densities and better body condition than on control sites. There were no observable effects of food addition on population rate of increase, recruitment, survival or age structure.
5. Mammalian predator reduction resulted in higher hare densities, higher survival, better body condition and an older age structure. Relative to control populations, recruitment was lower and population rates of increase similar.
6. The joint manipulation of food addition + predator reduction had greater positive effects on hare density and body condition than either single factor manipulation. Survival was better than on control sites, and the age structure was older than on control sites. Population rates of increase were similar, but recruitment was higher on the control areas.
7. We conclude that snowshoe hare dynamics at the low of the cycle are dominated by the interaction of food and predation. Risk of predation also had indirect effects on snowshoe hare age structure and body condition.     

Better safe than sorry: The response to a simulated predator and unfamiliar scent by the European hare

Predator presence can create a “landscape of fear,” which is defined as the spatially explicit distribution of perceived predation risk as seen by prey. Prey species can alter their behavior and space use as a response to increased predation risk, which might be traded off with energetic requirements. Thus, whether or not an anti-predator behavior is performed might depend on the perceived risk. In this study, we investigated the behavioral and spatial response of the European hare (Lepus europaeus) toward the presence of a predator scent, using red fox (Vulpes vulpes) scat, an unfamiliar control scent (butyric acid), and a true control (no scent). We collected data on hare activity times and behavior using 50 camera trap locations and spatial data using GPS telemetry (30,481 GPS positions of 12 hares). Hares showed spatial anti-predator behaviors within their home range, for example, the local avoidance of areas treated with scent and remaining further from field edges, in response to sympatric predator scent and partly also in response to unfamiliar butyric acid. Conversely, more costly anti-predator behaviors, that is, increased vigilance at the expense of foraging, were only shown in response to the predator scent. Our results suggest that prey species respond flexibly toward scent cues, utilizing less costly anti-predator behaviors independent of the perceived threat, whereas costly anti-predator responses are only used in the presence of “real” threat. Further, our findings emphasize that a combination of camera trap and GPS data can provide detailed information on animal behavior and space use, and caution that interpretation of one data source alone might lead to incomplete conclusions.     

Intensity of livestock grazing in relation to habitat use by brown hares (Lepus europaeus)

There is very limited information concerning livestock (sheep and goats) and brown hare Lepus europaeus interaction when both coexist. The effect of the intensity of livestock grazing on seasonal habitat use by hares, in a typical Mediterranean rangeland, was evaluated using the pellet-count method. Lightly grazed pastures were less preferred by hares compared with moderately grazed ones, whereas ungrazed pastures were used less intensively than grazed ones. Because livestock grazing reduces the quantity of standing biomass proportionally to its grazing intensity, forage resource was not the driving force for pasture selection. The increased use of moderately grazed pastures by hares in relation to lightly and ungrazed ones, where vegetation was more abundant, could be attributed to their reduced herbage height and density. This behaviour is probably a tactic that hares follow for predator avoidance, because they are more likely to detect visually approaching predators when feeding in a biotope with a limited herbaceous layer. The conclusion of this research is that livestock and brown hare coexistence may be compatible and beneficial rather than competitive when stocking rates do not exceed grazing capacity, leading to the conclusion that proper livestock grazing and hare population management can be feasible in practice.     

Multiscale habitat relationships of snowshoe hares (Lepus americanus) in the mixed conifer landscape of the Northern Rockies,USA: Cross‐scale effects of horizontal cover with implications for forest management

Snowshoe hares (Lepus americanus) are an ecologically important herbivore because they modify vegetation through browsing and serve as a prey resource for multiple predators. We implemented a multiscale approach to characterize habitat relationships for snowshoe hares across the mixed conifer landscape of the northern Rocky Mountains, USA. Our objectives were to (1) assess the relationship between horizontal cover and snowshoe hares, (2) estimate how forest metrics vary across the gradient of snowshoe hare use and horizontal cover, and (3) model and map snowshoe hare occupancy and intensity of use. Results indicated that both occupancy and intensity of use by snowshoe hares increased with horizontal cover and that the effect became stronger as intensity of use increased. This underscores the importance of dense horizontal cover to achieve high use, and likely density, of snowshoe hares. Forest structure in areas with high snowshoe hare use and horizontal cover was characterized as multistoried with dense canopy cover and medium‐sized trees (e.g., 12.7–24.4 cm). The abundance of lodgepole pine (Pinus contorta) was associated with snowshoe hare use within a mixed conifer context, and the only species to increase in abundance with horizontal cover was Engelmann spruce (Picea engelmannii) and subalpine fir (Abies lasiocarpa). Our landscape‐level modeling produced similar patterns in that we observed a positive effect of lodgepole pine and horizontal cover on both occupancy and use by snowshoe hares, but we also observed a positive yet parabolic effect of snow depth on snowshoe hare occupancy. This work is among the first to characterize the multiscale habitat relationships of snowshoe hares across a mixed conifer landscape as well as to map their occupancy and intensity of use. Moreover, our results provide stand‐ and landscape‐level insights that directly relate to management agencies, which aids in conservation efforts of snowshoe hares and their associated predators.     

Dynamics of herbivores and resources on a landscape with interspersed resources and refuges

A tradeoff between energy gain from foraging and safety from predation in refuges is a common situation for many herbivores that are vulnerable to predation while foraging. This tradeoff affects the population dynamics of the plant–herbivore–predator interaction. A new functional response is derived based on the Holling type 2 functional response and the assumption that the herbivore can forage at a rate that maximizes its fitness. The predation rate on the herbivore is assumed to be proportional to the product of the time that the herbivore spends foraging and a risk factor that reflects the habitat complexity; where greater complexity means greater interspersion of high food quality habitat and refuge habitat, which increases the amount of the edge zone between refuge and foraging areas, making foraging safer. The snowshoe hare is chosen as an example to demonstrate the resulting dynamics of an herbivore that has been intensely studied and that undergoes well-known cycling. Two models are studied in which the optimal foraging by hares is assumed, a vegetation–hare–generalist predator model and a vegetation–hare–specialist predator model. In both cases, the results suggest that the cycling of the snowshoe hare population will be greatly moderated by optimal foraging in a habitat consisting of interspersed high quality foraging habitat and refuge habitat. However, there are also large differences in the dynamics produced by the two models as a function of predation pressure.     

Fear and lethality in snowshoe hares: the deadly effects of non‐consumptive predation risk

Predators play a critical, top–down role in shaping ecosystems, driving prey population and community dynamics. Traditionally, studies of predator‐prey interactions have focused on direct effects of predators, namely the killing of prey. More recently, the non‐consumptive effects of predation risk are being appreciated; e.g. the ‘ecology of fear’. Prey responses to predation risk can be morphological, behavioural, and physiological, and are assumed to come at a cost to prey fitness. However, few studies have examined the relationship between predation risk and survival in wild animals. We tested the hypothesis that predation risk itself could reduce survival in wild‐caught snowshoe hares. We exposed female snowshoe hares to a simulated predator (a trained dog) during gestation only, and measured adult survival and, in surviving females, their ability to successfully wean offspring. We show for the first time in a wild mammal that the risk of predation can itself be lethal. Predation risk reduced adult female survival by 30%, and had trans‐generational effects, reducing offspring survival to weaning by over 85% – even though the period of risk ended at birth. As a consequence of these effects the predator‐exposed group experienced a decrease in number, while the control group substantially increased. Challenges remain in determining the importance of risk‐induced mortality in natural field settings; however, our findings show that non‐lethal predator encounters can influence survival of both adults and offspring. Future work is needed to test these effects in free‐living animals.     

A quantitative analysis of the abundance and demography of European hares Lepus europaeus in relation to habitat type, intensity of agriculture and climate

1. European hares Lepus europaeus have declined throughout Europe since the 1960s. Possible reasons for this include agricultural intensification and changes in climate and predator numbers, but no clear consensus has been reached as to the relative importance of each of these. We aimed to identify factors associated with high and low hare numbers throughout Europe, to determine which could have caused population declines. 2. Results of 77 research papers from 12 European countries were summarized. Relationships between hare density and demographics and habitat, climate, hunting and predator variables were examined and quantified where possible. Temporal changes in factors identified as being associated with high or low numbers of hares were then examined to see if they could explain population declines. 3. Data from pastural habitats were limited, but densities of hares were low. Arable habitats had higher densities than mixed areas in spring, unless farming was intensive in which case densities were similar. In autumn the two habitats had similar densities. Field size, temperature, precipitation and hunting had no effect on density throughout Europe. Fecundity was affected by climate. 4. Arable land, various crops, fallow habitat and temperature were positively associated, and monoculture, precipitation and predators negatively associated with hare abundance. The relationship of field size, pasture and woodland with abundance depended on spatial scale. 5. Habitat changes caused by agricultural intensification are the ultimate cause of hare population declines. Effects of changes in climate or predator numbers are magnified by the loss of high-quality year-round forage and cover. Further research is required on how habitat changes affect fecundity and survival, and to identify which parameters have the greatest impact on population numbers. Farmland management policies that target the re-establishment of some of the habitat diversity lost within fields, farms and landscapes will help to reverse the decline of the European hare.     

An Inverse Problem: Trappers Drove Hares to Eat Lynx

The Canadian lynx and snowshoe hare pelt data by the Hudson Bay Company did not fit the classical predator–prey theory. Rather than following the peak density of the hare, that of the lynx leads it, creating the hares-eat-lynx (HEL) paradox. Although trappers were suspected to play a role, no mathematical model has ever demonstrated the HEL effect. Here we show that the long-held assumption that the pelt number is a proxy of the wild populations is false and that when the data are modeled by the harvest rates by the trappers, the problem is finally resolved: both the HEL paradox and the classical theory are unified in our mechanistic hare-lynx-competitor-trapper (HLCT) model where competitor stands for all predators of the hares other than the lynx. The result is obtained by systematically fitting the data to various models using Newton’s inverse problem method. Main findings of this study include: the prey-eats-predator paradox in kills by an intraguild top-predator can occur if the top-predator prefers the predator to the prey; the benchmark HLCT model is more sensitive to all lynx-trapper interactions than to the respective hare-trapper interactions; the Hudson Bay Company’s hare pelt number maybe under-reported; and, the most intriguing of all, the trappers did not interfere in each other’s trapping activities.     

Habitat selection and spatiotemporal interactions of a reintroduced mesocarnivore

Habitat quality and quantity are key factors in evaluating the potential for success of a wildlife translocation. However, because of the difficulty or cost of evaluating these factors, habitat assessments may not include valuable information on important habitat attributes including the abundance and distribution of prey, predators, and competitors. Fishers (Pekania pennanti) are one of the most commonly reintroduced carnivores in North America, and they are a species of conservation concern in their western range. We examined the relative importance of landscape features and species interactions in determining habitat use of a reintroduced population of fishers in the southern Cascade Mountains, Washington, USA. We used detections of prey and predators at 134 remote camera stations, remotely sensed forest structure data, and telemetry locations of fishers in a resource selection function to assess the relative importance of prey, predators, and forest structure in fisher habitat selection. Fishers selected habitats based on forest conditions and activity levels of snowshoe hares (Lepus americanus), whereas bobcat (Lynx rufus) and coyote (Canis latrans) activity levels did not directly affect habitat selection. The probability of fisher use increased in older stands, close to recently disturbed stands, and in areas with intermediate levels of hare activity. Bobcat and hare activity levels were positively correlated, and fishers avoided areas with the greatest hare activity, suggesting that fishers may experience a food-safety tradeoff in the study area. Temporal activity patterns in photo detections indicate that fishers may mediate this danger by avoiding bobcats temporally. Our findings suggest that fishers in Washington prefer habitat mosaics of old and recently disturbed stands where they have greater access to resting structures and hares. Management that maintains mosaics of young and old forest across large landscapes is likely to support fisher recovery. Future reintroduction efforts would benefit from an assessment of prey and predator abundance in proposed reintroduction areas before project initiation. © 2019 The Wildlife Society.     

Influence of forest structure on the abundance of snowshoe hares in western Wyoming

Snowshoe hares (Lepus americanus) are a primary prey species for Canada lynx (Lynx canadensis) in western North America. Lynx management plans require knowledge of potential prey distribution and abundance in the western United States. Whether even-aged regenerating forests or multi-storied forests contain more snowshoe hares is currently unknown. During 2006–2008, we estimated snowshoe hare density in 3 classes of 30–70-year-old lodgepole pine (Pinus contorta) and 4 classes of late seral multi-storied forest with a spruce (Picea engelmannii)-fir (Abies lasiocarpa) component in the Bridger-Teton National Forest, Wyoming. We recorded physiographic variables and forest structure characteristics to understand how these factors influence abundance of snowshoe hares. In many instances, snowshoe hares were more abundant in late seral multi-storied forests than regenerating even-aged forests. Forest attributes predicting hare abundance were often more prevalent in multi-storied forests. Late seral multi-storied forests with a spruce–fir component and dense horizontal cover, as well as 30–70-year-old lodgepole pine with high stem density, were disproportionately influential in explaining snowshoe hare densities in western Wyoming. In order to promote improved habitat conditions for snowshoe hares in this region, management agencies should consider shifting their focus towards maintaining, enhancing, and promoting multi-storied forests with dense horizontal cover, as well as developing 30–70-year-old lodgepole pine stands with high stem density that structurally mimic multi-storied forests. © 2012 The Wildlife Society.     

Spatial dynamics of communities with intraguild predation: the role of dispersal strategies

I investigate the influence of dispersal strategies on intraguild prey and predators (competing species that prey on each other). I find an asymmetry between the intraguild prey and predator in their responses to each other's dispersal. The intraguild predator's dispersal strategy and dispersal behavior have strong effects on the intraguild prey's abundance pattern, but the intraguild prey's dispersal strategy and behavior have little or no effect on the intraguild predator's abundance pattern. This asymmetry arises from the different constraints faced by the two species: the intraguild prey has to acquire resources while avoiding predation, but the intraguild predator only has to acquire resources. It leads to puzzling distribution patterns: when the intraguild prey and predator both move away from areas of high density, they become aggregated to high-density habitats, but when they both move toward areas of high resource productivity, they become segregated to resource-poor and resource-rich habitats. Aggregation is more likely when dispersal is random or less optimal, and segregation is more likely as dispersal becomes more optimal. The crucial implication is that trophic constraints dictate the fitness benefits of using dispersal strategies to sample environmental heterogeneity. A strategy that affords greater benefits to an intraguild predator can lead to a more optimal outcome for both the intraguild predator and prey than a strategy that affords greater benefits to an intraguild prey.