Titrating the cost of plant toxins against predators: determining the tipping point for foraging herbivores

1. Foraging herbivores must deal with plant characteristics that inhibit feeding and they must avoid being eaten. Principally, toxins limit food intake, while predation risk alters how long animals are prepared to harvest resources. Each of these factors strongly affects how herbivores use food patches, and both constraints can pose immediate proximate costs and long-term consequences to fitness. 2. Using a generalist mammalian herbivore, the common brushtail possum (Trichosurus vulpecula), our aim was to quantitatively compare the influence of plant toxin and predation risk on foraging decisions. 3. We performed a titration experiment by offering animals a choice between non-toxic food at a risky patch paired with food with one of five toxin concentrations at a safe patch. This allowed us to identify the tipping point, where the cost of toxin in the safe food patch was equivalent to the perceived predation risk in the alternative patch. 4. At low toxin concentration, animals ate more from the safe than the risky patch. As toxin concentration increased at the safe patch, intake shifted until animals ate mainly from the risky patch. This shift was associated with behavioural changes: animals spent more time and fed longer at the risky patch, while vigilance increased at both risky and safe patches. 5. Our results demonstrate that the variation in toxin concentration, which occurs intraspecifically among plants, can critically influence the relative cost of predation risk on foraging. We show that herbivores quantify, compare and balance these two different but proximate costs, altering their foraging patterns in the process. This has potential ecological and evolutionary implications for the production of plant defence compounds in relation to spatial variation in predation risk to herbivores.     

The Importance of Predation Risk and Missed Opportunity Costs for Context-Dependent Foraging Patterns

Correct assessment of risks and costs of foraging is vital for the fitness of foragers. Foragers should avoid predation risk and balance missed opportunities. In risk-heterogeneous landscapes animals prefer safer locations over riskier, constituting a landscape of fear. Risk-uniform landscapes do not offer this choice, all locations are equally risky. Here we investigate the effects of predation risk in patches, travelling risk between patches, and missed social opportunities on foraging decisions in risk-uniform and risk-heterogeous landscapes. We investigated patch leaving decisions of 20 common voles (M. arvalis) in three experimental landscapes: safe risk-uniform, risky risk-uniform and risk-heterogeneous. We varied both the predation risk level and the predation risk distribution between two patches experimentally and in steps, assuming that our manipulation consequently yield different distributions and levels of risk while foraging, risk while travelling, and costs of missed, social opportunities (MSOCs). We measured mean GUDs (giving-up density of food left in the patch) for both patches as a measure of foraging gain, and delta GUD, the differences among patches, as a measure of the spatial distribution of foraging effort over a period of six hours. Distribution of foraging effort was most even in the safe risk-uniform landscapes and least even in the risk-heterogeneous landscape, with risky risk-uniform landscapes in between. Foraging gain was higher in the safe than in the two riskier landscapes (both uniform and heterogeneous). Results supported predictions for the effects of risk in foraging patches and while travelling between patches, however predictions for the effects of missed social opportunities were not met in this short term experiment. Thus, both travelling and foraging risk contribute to distinct patterns observable high risk, risk-uniform landscapes.     

Food quality and quantity are more important in explaining foraging of an intermediate‐sized mammalian herbivore than predation risk or competition

During times of high activity by predators and competitors, herbivores may be forced to forage in patches of low‐quality food. However, the relative importance in determining where and what herbivores forage still remains unclear, especially for small‐ and intermediate‐sized herbivores. Our objective was to test the relative importance of predator and competitor activity, and forage quality and quantity on the proportion of time spent in a vegetation type and the proportion of time spent foraging by the intermediate‐sized herbivore European hare (Lepus europaeus). We studied red fox (Vulpes vulpes) as a predator species and European rabbit (Oryctolagus cuniculus) as a competitor. We investigated the time spent at a location and foraging time of hare using GPS with accelerometers. Forage quality and quantity were analyzed based on hand‐plucked samples of a selection of the locally most important plant species in the diet of hare. Predator activity and competitor activity were investigated using a network of camera traps. Hares spent a higher proportion of time in vegetation types that contained a higher percentage of fibers (i.e., NDF). Besides, hares spent a higher proportion of time in vegetation types that contained relatively low food quantity and quality of forage (i.e., high percentage of fibers) during days that foxes (Vulpes vulpes) were more active. Also during days that rabbits (Oryctolagus cuniculus) were more active, hares spent a higher proportion of time foraging in vegetation types that contained a relatively low quality of forage. Although predation risk affected space use and foraging behavior, and competition affected foraging behavior, our study shows that food quality and quantity more strongly affected space use and foraging behavior than predation risk or competition. It seems that we need to reconsider the relative importance of the landscape of food in a world of fear and competition.     

Hazardous duty pay and the foraging cost of predation

We review the concepts and research associated with measuring fear and its consequences for foraging. When foraging, animals should and do demand hazardous duty pay. They assess a foraging cost of predation to compensate for the risk of predation or the risk of catastrophic injury. Similarly, in weighing foraging options, animals tradeoff food and safety. The foraging cost of predation can be modelled, and it can be quantitatively and qualitatively measured using risk titrations. Giving‐up densities (GUDs) in depletable food patches and the distribution of foragers across safe and risky feeding opportunities are two frequent experimental tools for titrating food and safety. A growing body of literature shows that: (i) the cost of predation can be big and comprise the forager's largest foraging cost, (ii) seemingly small changes in habitat or microhabitat characteristics can lead to large changes in the cost of predation, and (iii) a forager's cost of predation rises with risk of mortality, the forager's energy state and a decrease in its marginal value of energy. In titrating for the cost of predation, researchers have investigated spatial and temporal variation in risk, scale‐dependent variation in risk, and the role of predation risk in a forager's ecology. A risk titration from a feeding animal often provides a more accurate behavioural indicator of predation risk than direct observations of predator‐inflicted mortality. Titrating for fear responses in foragers has some well‐established applications and holds promise for novel methodologies, concepts and applications. Future directions for expanding conceptual and empirical tools include: what are the consequences of foraging costs arising from interference behaviours and other sources of catastrophic loss? Are there alternative routes by which organisms can respond to tradeoffs of food and safety? What does an animal's landscape of fear look like as a spatially explicit map, and how do various environmental factors affect it? Behavioural titrations will help to illuminate these issues and more.     

Integrating the costs of plant toxins and predation risk in foraging decisions of a mammalian herbivore

Foraging herbivores must satisfy their nutrient requirements in a world of toxic plants while also avoiding predators. Plant toxins and perceived predation risk at food patches should both reduce patch residency time, but the relative strengths of these factors on feeding decisions has rarely been quantified. Using an arboreal generalist herbivore, the common brushtail possum Trichosurus vulpecula, we tested the effects on food intake of the plant toxin, cineole, and regurgitated pellets from one of its predators, the powerful owl Ninox strenua at the small spatial scale of the food patch. We used the giving-up density (GUD) framework, with animals harvesting food items (sultanas) in an inedible matrix (small pebbles). We ran two consecutive field experiments in a eucalypt woodland in eastern Australia, 1 month apart in the same location. In experiment 1, there was a significant interaction between cineole [at 17% of dry matter (DM)] and owl pellets. The GUD was lowest in the absence of both cineole and owl pellet, intermediate in the presence of owl pellet; and highest with cineole ± owl pellet. The effect of owl pellet diminished over time. In experiment 2, only cineole (at 10% DM) increased the GUD significantly. The difference in effect of owl pellet was probably due to both habituation and freshness of the cue. Our study demonstrates the importance of synthesising predator–prey and plant–herbivore ecology to better understand the complex set of constraints influencing foraging herbivores. The greater effect of toxin than fear on possums is likely to be due to its high, but ecologically relevant concentration. This highlights the need to explore the relative and net impacts of a range of concentrations of plant toxins and predation risks.     

Response of pollinators to the tradeoff between resource acquisition and predator avoidance

Although the behaviour of animals facing the conflicting demands of increasing foraging success and decreasing predation risk has been studied in many taxa, the response of pollinators to variations in both factors has only been studied in isolation. We compared visit rates of two pollinator species, hoverflies and honeybees, to 40 Chrysanthemum segetum patches in which we manipulated predation risk (patches with and without crab spiders) and nectar availability (rich and poor patches) using a full factorial design. Pollinators responded differently to the tradeoff between maximising intake rate and minimising predation risk: honeybees preferred rich safe patches and avoided poor risky patches while the number of hoverflies was highest at poor risky patches. Because honeybees were more susceptible to predation than hoverflies, our results suggest that, in the presence of competition for resources, less susceptible pollinators concentrate their foraging effort on riskier resources, where competition is less severe. Crab spiders had a negative effect on the rate at which inflorescences were visited by honeybees. This effect was mediated through changes in the foraging strategy of honeybees, and could, in principle, be reversed by increasing nectar productivity of inflorescences. Our study shows that both pollinator species responded simultaneously and differently to variations in food reward and predation risk, and highlights the importance of studying the foraging strategies of pollinators in order to fully understand how plant–pollinator interactions are established.     

Long- and short-term state-dependent foraging under predation risk: an indication of habitat quality

Animals living in environments of different quality will have different expectations of their future reproductive success and survival. This may affect the individual's risk-taking behaviour as manifest in the cost of predation. We investigated the foraging behaviour of starlings, Sturnus vulgaris, when perceived predation risk varied between patches. Short-term food availability varied between treatments and long-term differences in perceptions of environmental quality varied between groups of individuals. This corresponds to variation in the three components of the cost of predation (P): the predation risk (μ); the change in reproductive value with energy gain (∂F /∂e); and the reproductive value or fitness factor (F). The birds showed that they experienced a higher cost of predation while using the risky food patches (μ component) and in the high food treatment (∂F /∂e component). Furthermore, birds from a high-reward habitat revealed a higher P than birds from a poor habitat (F component). The results show that the costs of predation are possible to tease apart by using behavioural indicators. The method presented allows measurement of fitness prospects of individuals, which may have consequences for conservation, for example, to identify low-quality habitat. Copyright 2002 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour     

Scaling from plot experiments to landscapes: studying grasshoppers to inform forest ecosystem management

Ecologists studying food web interactions routinely conduct their experiments at scales of 1–10 m² whereas real-world landscape-level management problems exist on scales of 10⁶ m² or larger. It is often asserted that the experimental tradition in ecology has little to offer to environmental management because small scale empirical insights are not easily, if at all, translatable to the large scale problems. Small scale experiments are very local in nature and they are conducted in ways that tend to homogenize background environmental variation. Real world management is conducted across vast landscapes. Managers routinely must wrestle with complexity that is introduced by the heterogeneous structure of those landscapes and they often have limited recourse to do careful experimentation. How then is empirical ecological science ever to inform landscape-level management? The solution to this dilemma lies in arriving at good working conceptualizations of ecosystem structure and function that embody principles that are relatively scale independent. In this paper, the evolutionary ecological principle of foraging versus predation risk avoidance trade-offs is proffered as one central organizing conceptualization for plant-herbivore interactions across all systems. The utility of this conceptualization is first illustrated by presenting results of detailed experiments involving spider predators, grasshopper herbivores, and two classes of plant resources that afford grasshoppers differential protection from predators: nutritionally superior but risky grasses and less nutritious but safer herbs. The paper then shows how the foraging versus predation risk avoidance conceptualization in the context of a “landscape of fear” can be applied to manage large herbivore impacts of forest regeneration following forest harvesting. I present results of landscape-scale experiments that mediate predation risk of the herbivores through manipulation of safe habitat in order to enlist herbivores to facilitate boreal forest mixed species regeneration through preferential foraging of certain woody species.     

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.     

Patch Use in Free‐Ranging Goats: Does a Large Mammalian Herbivore Forage like Other Central Place Foragers?

Classic central place foraging theory does not focus on the foraging of central place herbivores. This is especially true with regard to large mammalian herbivores. To understand the foraging dynamics of these neglected foragers, we measured giving‐up densities (GUDs) in artificial food patches. We did this at different distances away from the central point (i.e. corral) for a herd of free‐ranging domestic goats. To determine temporal changes, we conducted the study over a 3‐mo period during an extended dry season. Throughout our study, goats foraged across a gradient of food availability where forage was more available farther away from the central point. In contrast to the prediction that predation risk and/or increased travel costs were the main drivers of foraging decisions, we found that the goats increased their feeding effort (i.e. achieved lower GUDs) the farther away they moved from the central point. This suggests that either metabolic or missed opportunity costs were the main factors that influenced foraging decisions. In addition, we suggest that social foraging may have also played a role. With increases in foraging opportunities away from the central point, a herd will likely move slowly while foraging. As a result, individuals can feed intensively from patches but remain part of the group. Ironically, owing to the sustained close proximity of other group members, individuals may perceive patches farther from the central point as being safer. Temporally, the goats increased their feeding effort throughout the dry season. This suggests there was a decline in food quality and/or availability across the environment as the study progressed. Despite this increase in feeding effort, the negative relationship with distance did not change. Ultimately, our results provide key insight into how metabolic, missed opportunity and perceived predation costs influence the feeding decisions of large central place herbivores.     

The benefits of being in a bad neighbourhood: plant community composition influences red deer foraging decisions

Diet selection by mammalian herbivores is often influenced by plant community composition, and numerous studies have focused on the relationships between herbivore foraging decisions and food/plant species abundance. However, few have examined the role of neighbour palatability in affecting foraging of a target plant by large mammalian herbivores. We used a large-scale field dataset on diet selection by red deer Cervus elaphus in Fiordland National Park, New Zealand to: (1) estimate the palatability of native forest plant species to introduced deer from observed patterns of browse damage; and (2) examine whether intraspecific variation in browsing of plants can be related to variation in the local abundance of alternative forage species. Overall, 21 of the 53 forest species in our dataset were never browsed by deer. At a community level, plants were more likely to be browsed if they were in a patch of vegetation of high forage quality, containing high abundances of highly palatable species and/or low abundances of less-palatable species. Our findings suggest that deer make foraging decisions at both a coarse-grain level, selecting vegetation patches within a landscape based on the overall patch quality, and at a fine-grain level by choosing among individual plants of different species.     

Herbivore host plant selection: whitefly learns to avoid host plants that harbour predators of her offspring

Evidence is accumulating that herbivorous arthropods do not simply select host plants based on their quality, but also on the predation risk associated with different host plants. It has been suggested that herbivores exclude plant species with high predation risk from their host range. This assumes a constant, predictable predation risk as well as a rather static behaviour on the part of the herbivore; plants are ignored irrespective of the actual predation risk. We show that adult females of a small herbivore, the whitefly Bemisia tabaci, can learn to avoid plants with predatory mites that attack only juvenile whiteflies, while they accept host plants of the same species without predators. Predatory mites disperse more slowly than whiteflies; they cannot fly and walk from plant to plant. Hence, by avoiding plants with predators, the whiteflies create a temporary refuge for their offspring. We suggest that the experience of arthropod herbivores with risks associated with host plants plays an important role in their host plant selection.     

Seasonality and edge effect determine herbivory risk according to different plant association models

We report evidence of hierarchical resource selection by large herbivores and plant neighbouring effects in a Mediterranean ecosystem. Plant palatability was assessed according to herbivore foraging decisions. We hypothesize that under natural conditions large herbivores follow a hierarchical foraging pattern, starting at the landscape scale, and then selecting patches and individual plants. A between- and within-patch selection study was carried out in an area formed by scrubland and pasture patches, connected by habitat edges. With regard to between-patch selection, quality-dependent resource selection is reported: herbivores mainly consume pasture in spring and woody plants in winter. Within-patch selection was also observed in scrub habitats, influenced by season, relative patch palatability and edge effect. We defined a Proximity Index (PI) between palatable and unpalatable plants, which allowed verification of neighbouring effects. In spring, when the preferred food resource (i.e. herbs) is abundant, we observed that in habitat edges large herbivores basically select the relatively scarce palatable shrubs, whereas inside scrubland, unpalatable shrub consumption was related to increasing PI. In winter, a very different picture was observed; there was low consumption of palatable species surrounded by unpalatable species in habitat edges, where the latter were more abundant. These outcomes could be explained though different plant associations described in the literature. We conclude that optimal foraging theory provides a conceptual framework behind the observed interactions between plants and large herbivores in Mediterranean ecosystems.     

Foraging efficiency of <Emphasis Type="Italic">Akodon azarae</Emphasis> under different plant cover and resource levels

The goal of this work was to determine how the foraging behaviour of Akodon azarae changes with predation risk and food availability in cropfield borders of Buenos Aires, Argentina. Our hypotheses were that A. azarae has a greater foraging efficiency in safe areas than in risky ones and that the foraging behaviour of A. azarae also depends on the level of resources. We measured giving-up densities (GUDs) and food consumption twice a year in artificial foraging patches (bottles with known amounts of millet seed) in covered and open areas and with two different levels of seed abundance. In both periods, GUDs were lower in the covered areas than in the open ones independently of food level. Consumption increased with food level in covered areas but not in open areas. Based on these results, we conclude that A. azarae appears to maximize its consumption depending on predation risk.     

Context dependence in foraging behaviour of Achillea millefolium

Context-dependent foraging behaviour is acknowledged and well documented for a diversity of animals and conditions. The contextual determinants of plant foraging behaviour, however, are poorly understood. Plant roots encounter patchy distributions of nutrients and soil fungi. Both of these features affect root form and function, but how they interact to affect foraging behaviour is unknown. We extend the use of the marginal value theorem to make predictions about the foraging behaviour of roots, and test our predictions by manipulating soil resource distribution and inoculation by soil fungi. We measured plant movement as both distance roots travelled and time taken to grow through nutrient patches of varied quality. To do this, we grew Achillea millefolium in the centers of modified pots with a high-nutrient patch and a low-nutrient patch on either side of the plant (heterogeneous) or patch-free conditions (homogeneous). Fungal inoculation, but not resource distribution, altered the time it took roots to reach nutrient patches. When in nutrient patches, root growth decreased relative to homogeneous soils. However, this change in foraging behaviour was not contingent upon patch quality or fungal inoculation. Root system breadth was larger in homogeneous than in heterogeneous soils, until measures were influenced by pot edges. Overall, we find that root foraging behaviour is modified by resource heterogeneity but not fungal inoculation. We find support for predictions of the marginal value theorem that organisms travel faster through low-quality than through high-quality environments, with the caveat that roots respond to nutrient patches per se rather than the quality of those patches.     

Quantifying the response of free-ranging mammalian herbivores to the interplay between plant defense and nutrient concentrations

While trying to achieve their nutritional requirements, foraging herbivores face the costs of plant defenses, such as toxins. Teasing apart the costs and benefits of various chemical constituents in plants is difficult because their chemical defenses and nutrient concentrations often co-vary. We used an approach derived from predator–prey studies to quantitatively compare the foraging response of a free-ranging mammalian herbivore, the swamp wallaby (Wallabia bicolor), through three feeding trials with artificial diets that differed in their concentrations of (1) the terpene 1,8-cineole, (2) primary constituents (including nitrogen and fiber), and (3) both the terpene and the primary constituents. Applying the giving-up density (GUD) framework, we demonstrated that the foraging cost of food patches increases with higher dietary cineole concentration and decreases with higher dietary nutrient concentration. The effect of combined differences in nutrients and cineole concentrations on GUD was interactive, and high nutrient food required more cineole to achieve the same patch value as low nutrient food. Our results indicate that swamp wallabies equate low nutrient, poorly defended food with high nutrient, highly defended food, providing two contrasting diets with similar cost–benefit outcomes. This behavior suggests that equal concentrations of chemical defenses provide nutrient-poor plants with relatively greater protection as nutrient-rich plants. Nutrient-rich plants may therefore face the exacerbated problem of being preferred by herbivores and therefore need to produce more defense compounds to achieve the same level of defense as nutrient-poor plants. Our findings help explain the difference in anti-herbivore strategy of nutrient-poor and rich plants, i.e., tolerance versus defense.     

Foraging patterns of voles at heterogeneous avian and uniform mustelid predation risk

Temporal variation of antipredatory behavior and a uniform distribution of predation risk over refuges and foraging sites may create foraging patterns different from those anticipated from risk in heterogenous habitats. We studied the temporal variation in foraging behavior of voles exposed to uniform mustelid predation risk and heterogeneous avian predation risk of different levels induced by vegetation types in eight outdoor enclosures (0.25 ha). We manipulated mustelid predation risk with weasel presence or absence and avian predation risk by reducing or providing local cover at experimental food patches. Foraging at food patches was monitored by collecting giving-up densities at artificial food patches, overall activity was automatically monitored, and mortality of voles was monitored by live-trapping and radiotracking. Voles depleted the food to lower levels in the sheltered patches than in the exposed ones. In enclosures with higher avian predation risk caused by lower vegetation height, trays were depleted to lower levels. Unexpectedly, voles foraged in more trays and depleted trays to lower levels in the presence of weasels than in the absence. Weasels match their prey's body size and locomotive abilities and therefore increase predation risk uniformly over both foraging sites and refuge sites that can both be entered by the predator. This reduces the costs of missing opportunities other than foraging. Voles changed their foraging strategy accordingly by specializing on the experimental food patches with predictable returns and probably reduced their foraging in the matrix of natural food source with unpredictable returns and high risk to encounter the weasel. Moreover, after 1 day of weasel presence, voles shifted their main foraging activities to avoid the diurnal weasel. This behavior facilitated bird predation, probably by nocturnal owls, and more voles were killed by birds than by weasels. Food patch use of voles in weasel enclosures increased with time. Voles had to balance the previously missed feeding opportunities by progressively concentrating on artificial food patches.     

Plant Defenses and Predation Risk Differentially Shape Patterns of Consumption,Growth, and Digestive Efficiency in a Guild of Leaf-Chewing Insects

Herbivores are squeezed between the two omnipresent threats of variable food quality and natural enemy attack, but these two factors are not independent of one another. The mechanisms by which organisms navigate the dual challenges of foraging while avoiding predation are poorly understood. We tested the effects of plant defense and predation risk on herbivory in an assemblage of leaf-chewing insects on Solanum lycopersicum (tomato) that included two Solanaceae specialists (Manduca sexta and Leptinotarsa decemlineata) and one generalist (Trichoplusia ni). Defenses were altered using genetic manipulations of the jasmonate phytohormonal cascade, whereas predation risk was assessed by exposing herbivores to cues from the predaceous stink bug, Podisus maculiventris. Predation risk reduced herbivore food intake by an average of 29% relative to predator-free controls. Interestingly, this predator-mediated impact on foraging behavior largely attenuated when quantified in terms of individual growth rate. Only one of the three species experienced lower body weight under predation risk and the magnitude of this effect was small (17% reduction) compared with effects on foraging behavior. Manduca sexta larvae, compensated for their predator-induced reduction in food intake by more effectively converting leaf tissue to body mass. They also had higher whole-body lipid content when exposed to predators, suggesting that individuals convert energy to storage forms to draw upon when risk subsides. In accordance with expectations based on insect diet breadth, plant defenses tended to have a stronger impact on consumption and growth in the generalist than the two specialists. These data both confirm the ecological significance of predators in the foraging behavior of herbivorous prey and demonstrate how sophisticated compensatory mechanisms allow foragers to partially offset the detrimental effects of reduced food intake. The fact that these mechanisms operated across a wide range of plant resistance phenotypes suggests that compensation is not always constrained by reduced food quality.     

Non‐consumptive effects of a top‐predator decrease the strength of the trophic cascade in a four‐level terrestrial food web

The fear of predators can strongly impact food web dynamics and ecosystem functioning through effects on herbivores morphology, physiology or behaviour. While non‐consumptive predator effects have been mostly studied in three‐level food chains, we lack evidence for the propagation of non‐consumptive indirect effects of apex predators in four level food‐webs, notably in terrestrial ecosystems. In experimental mesocosms, we manipulated a four‐level food chain including top‐predator cues (snakes), mesopredators (lizards), herbivores (crickets), and primary producers (plants). The strength of the trophic cascade induced by mesopredators through the consumption of herbivores decreased in the presence of top‐predator cues. Specifically, primary production was higher in mesocosms where mesopredators were present relative to mesocosms with herbivores only, and this difference was reduced in presence of top‐predator cues, probably through a trait‐mediated effect on lizard foraging. Our study demonstrates that non‐consumptive effects of predation risk can cascade down to affect both herbivores and plants in a four‐level terrestrial food chain and emphasises the need to quantify the importance of such indirect effects in natural communities.     

Opportunity costs influence food selection and giving‐up density of dabbling ducks

The interaction of animals with their food can yield insights into habitat characteristics, such as perceived predation risk and relative quality. We deployed experimental foraging patches in wetlands used by migrating dabbling ducks Anas spp. in the central Illinois River Valley to estimate variation in seed removal and giving‐up density (GUD; i.e. density of food remaining in patches following abandonment) with respect to seed density, seed size, seed depth in the substrate, substrate firmness, perceived predation risk, and an energetic profitability threshold (i.e. critical food density). Seed depth and the density of naturally‐occurring seeds outside of experimental plots affected seed removal and GUD in experimental patches more than perceived predation risk, seed density, seed size or substrate firmness. The greatest seed removal and lowest GUDs in experimental patches occurred when food resources in alternative foraging locations outside of plots (i.e. opportunity costs) appeared to be near or below a critical food density (i.e. 119–181 kg ha^–1). Giving‐up densities varied substantially from a critical food density across a range of food densities in alternative foraging locations suggesting that fixed GUDs should not be used as surrogates for critical food densities in energetic carrying capacity models. Foraging and resting rates in and near experimental foraging patches did not reflect patterns of seed removal and were poor predictors of GUD and foraging habitat quality. Our results demonstrated the usefulness of GUDs as indicators of habitat quality for subsurface, benthic foragers relative to other available foraging patches and suggested that food may be limited for dabbling ducks during spring migration in some years in the midwestern USA.