Seed predation and selection exerted by a seed predator influence subalpine tree densities

Strongly interacting species often have pronounced direct and indirect effects on other species. Here we focus of the effects of pine squirrels (Tamiasciurus spp.), which are a dominant pre-dispersal seed predator of many conifers including limber pines (Pinus flexilis) and whitebark pines (P. albicaulis). Pine squirrels depress seed abundance by harvesting most limber and whitebark pine cones on their territories. Pine squirrels further reduce seed availability for Clark's Nutcrackers (Nucifraga columbiana), the primary seed disperser of these pines, because selection exerted by pine squirrels has reduced the number of seeds per cone and causes seeds to be less accessible. We predicted that, if fewer seeds were available for dispersal by nutcrackers, pine recruitment should be suppressed in areas with pine squirrels. In support of this prediction, stand densities were about two times greater in areas where pine squirrels are absent than in areas where they are present. Alternative explanations that we considered do not account for these differences; however, precipitation may limit stand densities in the absence of seed limitation by pine squirrels. In sum, pine squirrels apparently depress limber and whitebark pine stand densities, with the potential for ecosystem impacts because these pines are foundation species within Western subalpine ecosystems.     

On the relative contributions of wind vs. animals to seed dispersal of four Sierra Nevada pines

Selective pressures that influence the form of seed dispersal syndromes are poorly understood. Morphology of plant propagules is often used to infer the means of dispersal, but morphology can be misleading. Several species of pines, for example, have winged seeds adapted for wind dispersal but owe much of their establishment to scatter-hoarding animals. Here the relative importance of wind vs. animal dispersal is assessed for four species of pines of the eastern Sierra Nevada that have winged seeds but differed in seed size: lodgepole pine (Pinus contorta murrayana, 8 mg); ponderosa pine (Pinus ponderosa ponderosa, 56 mg); Jeffrey pine (Pinus jeffreyi, 160 mg); and sugar pine (Pinus lambertiana, 231 mg). Pre-dispersal seed mortality eliminated much of the ponderosa pine seed crop (66%), but had much less effect on Jeffrey pine (32% of seeds destroyed), lodgepole pine (29%), and sugar pine (7%). When cones opened most filled seeds were dispersed by wind. Animals removed > 99% of wind-dispersed Jeffrey and sugar pine seeds from the ground within 60 days, but animals gathered only 93% of lodgepole pine seeds and 38% of ponderosa pine seeds during the same period. Animals gathered and scatter hoarded radioactively labeled ponderosa, Jeffrey, and sugar pine seeds, making a total of 2103 caches over three years of study. Only three lodgepole pine caches were found. Caches typically contained 1-4 seeds buried 5-20 mm deep, depths suitable for seedling emergence. Although Jeffrey and sugar pine seeds are initially wind dispersed, nearly all seedlings arise from animal caches. Lodgepole pine is almost exclusively wind dispersed, with animals acting as seed predators. Animals treated ponderosa pine in an intermediate fashion. Two-phased dispersal of large, winged pine seeds appears adaptive; initial wind dispersal helps to minimize pre-dispersal seed mortality whereas scatter hoarding by animals places seeds in sites with a higher probability of seedling establishment.     

The combined effects of seed perishability and seed size on hoarding decisions by Pére David’s rock squirrels

The food perishability hypothesis reasons that the perishability of nondormant acorns through rapid germination is the primary determinant of hoarding decisions (e.g., embryo removal in nondormant acorns in particular) by scatter-hoarding squirrels. However, we do not know whether seed size and its interactions with seed germination schedule affect squirrel’s hoarding decisions. By presenting pairs of acorns with contrasting germination/dormancy conditions and seed size, we investigated the relative importance of each target trait in determining the hoarding decisions of free-ranging Pére David’s rock squirrel (Sciurotamias davidianus) in Central China. Consistent with the food perishability hypothesis, the squirrels were highly sensitive to subtle differences of acorn germination status either within nondormant acorns or between nondormant and dormant acorns. Though there were no significant differences in seed hoarding and dispersal distance, the embryo-removal probability of nondormant acorns (especially those germinated) was much higher than that of dormant acorns prior to hoarding. Our results also support the seed size hypothesis. Large acorns were often hoarded more and moved farther than small acorns, and large nondormant acorns also had a higher probability of having their embryos removed. Moreover, the interactions between seed size and seed germination schedule had a large effect on whether a given acorn was hoarded or hoarded with its embryo removed. Our study indicates that the combined effects from seed germination schedule and seed size have the potential to determine hoarding decisions of scatter-hoarding squirrels.     

Variation in the food hoarding behaviour of red squirrels

Summary To investigate variation in food storing behaviour within a population of red squirrels (Tamiasciurus hudsonicus) we intensively sampled the size and spatial dispersion of hoards. Single-item and large hoards were far more prevalent than expected by chance (Figs. 1 and 2) and appeared to be distinct components of the hoarding tactic employed in this population. Scattered single-item hoards represented more than half of all hoards constructed and approximately one-fifth of all cones stored. Large hoards (> 10 cones) comprised less than 15% of all hoards, and about 50% of all cones stored. In response to experimental manipulations, squirrels altered their hoarding behaviour in order to concentrate cones in the litter and brush heaps we provided, but still scattered a large proportion of their stores. We conclude that hoarding patterns in red squirrels are more variable and plastic than has previously been recognized, and discuss this variability in terms of adaptive explanations. Offprint requests to: R.J. Robertson     

Anticipation and tracking of pulsed resources drive population dynamics in eastern chipmunks

Pulsed systems are characterized by boom and bust cycles of resource production that are expected to cascade through multiple trophic levels. Many of the consumers within pulsed resource systems have specific adaptations to cope with these cycles that may serve to either amplify or dampen their community-wide consequences. We monitored a seed predator, the eastern chipmunk (Tamias striatus), in an American beech (Fagus grandifolia) dominated forest, and used capture-mark-recapture analyses to estimate chipmunk vital rates and relate them to interannual variation in beech seed production. The summer activity and reproduction of adults anticipated autumn beech production, with high activity and intense reproduction occurring in summers prior to beech masts. Chipmunks also reproduced every spring following a beech mast. However, adult survival was independent of beech production. In contrast, juvenile survival was lower in years of mast failure than in years of mast production, but their activity was consistently high and independent of beech production. Population growth was strongly affected by the number of juveniles and therefore by beech seed production, which explains nearly 70% of variation in population growth. Our results suggest that a combination of resource-dependent reproduction and variable activity levels associated with anticipation and response to resource pulses allows consumers to buffer potential deleterious effects of low food abundance on their survival.     

Interspecific competition in tree squirrels: do introduced grey squirrels (<Emphasis Type="Italic">Sciurus carolinensis</Emphasis>) deplete tree seeds hoarded by red squirrels (<Emphasis Type="Italic">S. vulgaris</Emphasis>)?

Red squirrels (Sciurus vulgaris) and introduced eastern grey squirrels (S. carolinensis) scatterhoard seeds of broadleaf trees. Scatterhoarded seeds are an essential resource in spring and their consumption increases red-squirrel fitness. We examined whether grey squirrels partly deplete the high-energy food resources cached by red squirrels, reducing their consumption, in two ways: (1) at the population level, comparing energy intake of feeding on cached seeds between a study site with red and grey squirrels and one with only red squirrels present; and (2) at the individual level, in the study site where species co-exist, relating hoard recovery of red squirrels to the amount of core-area overlap with grey squirrels. There were no significant site differences in the mean daily energy intake of red squirrels feeding on seeds recovered from caches. However, in the red-grey site, during spring, red squirrels that had a high percentage of their home-range core area overlapped by grey squirrels had a lower daily energy intake than low-overlap red squirrels. Body mass of red squirrels in spring was negatively correlated with the percentage of interspecific core-area overlap, but not with core-area overlap with other red squirrels. Our data suggest that interspecific competition for scatterhoarded seeds, with grey squirrels pilfering red squirrels' food caches, caused a reduced energy intake in red squirrels with a high degree of interspecific core-area overlap, and reduced body mass in spring. Therefore, cache pilfering is likely to reduce reproductive output in red squirrels, and thus play a role in the replacement of red by grey squirrels. Electronic Publication     

Interspecific associations of Cape ground squirrels with two mongoose species: benefit or cost?

Mixed-species associations have been described in many vertebrate species, but few behavioral studies have investigated associations between species from different mammalian orders. Cape ground squirrels (Xerus inauris) are highly social rodents that inhabit burrows with two species of mongoose, but the benefits of these interspecific associations to ground squirrels remain unresolved. We compared the behavior of squirrels while solitary, with conspecifics, and in the presence of suricates (Suricata suricatta) and yellow mongooses (Cynictis pencillatus). Squirrels spent less time alert and more time feeding when suricates were present, but increased vigilance in the presence of yellow mongooses. In a series of mobbing trials with a puff adder (Bitis arietans), a common predator of all three species, Cape ground squirrels were the most active in mobbing the snake. Our results suggest that Cape ground squirrels benefit from associating with suricates, but not necessarily with yellow mongooses. Both mongoose species benefit from the burrowing activities of the squirrels for thermoregulation and escape from predators, and a suite of other organisms may similarly benefit from the habitat modifications by Cape ground squirrels, suggesting they could be considered ecosystem engineers of the arid and semi-arid regions of southern Africa. Thus, the association between Cape ground squirrels and suricates appears mutually beneficial, whereas yellow mongooses may merely be commensals of the squirrels.     

Antipredator responses of koomal (Trichosurus vulpecula hypoleucus) against introduced and native predators

Antipredator behavior studies generally assess prey responses to single predator species although most real systems contain multiple species. In multi-predator environments prey ideally use antipredator responses that are effective against all predator species, although responses may only be effective against one predator and counterproductive for another. Multi-predator systems may also include introduced predators that the prey did not co-evolve with, so the prey may either fail to recognize their threat (level 1 naiveté), use ineffective responses (level 2 naiveté) or succumb to their superior hunting ability (level 3 naiveté). We analyzed microhabitat selection of an Australian marsupial (koomal, Trichosurus vulpecula hypoleucus) when faced with spatiotemporal differences in the activity/density levels of one native (chuditch, Dasyurus geoffroii) and two introduced predators (red fox, Vulpes vulpes; feral cat, Felis catus). From this, we inferred whether koomal recognized introduced predators as a threat, and whether they minimized predation risk by either staying close to trees and/or using open or dense microhabitats. Koomal remained close to escape trees regardless of the predator species present, or activity/density levels, suggesting koomal employ this behavior as a first line of defense. Koomal shifted to dense cover only under high risk scenarios (i.e., with multiple predator species present at high densities). When predation risk was low, koomal used open microhabitats, which likely provided benefits not associated with predator avoidance. Koomal did not exhibit level 1 naiveté, although further studies are required to determine if they exhibit higher levels of naiveté (2–3) against foxes and cats.     

Communal nesting in an ‘asocial’ mammal: social thermoregulation among spatially dispersed kin

Communal nesting can help defray the high cost of endothermic heat production in cold environments, but such social behavior is generally thought to be incompatible with the persistent defense of exclusive territories in typically ‘asocial’ animals. We examined the propensity for communal nesting in female red squirrels (Tamiasciurus hudsonicus), which maintain individual year-round territories, through intensive monitoring of litters over 22 years and by radio-tracking females during 3 years in late winter/early spring. Communal nesting was exceptionally rare during lactation: of 1,381 litters tracked to emergence, we observed a single instance in which two closely related (r?=?0.5) females pooled their litters into a single nest. In contrast, nest sharing between 2–3 females was relatively common in the late winter/early spring, prior to mating; at least 12 of 63 females (19 %) engaged in communal nesting during a year of systematic tracking of radio-collared females from late February to April. Communal nesting occurred more frequently when temperatures were colder, suggesting that such aggregations might function to reduce thermoregulatory costs. These social associations were typically, though not exclusively, between closely related individuals (r?≥?0.25 for seven of eight cases; mother–daughter dyads: four of eight), suggesting this cooperative behavior might evolve through kin selection and/or may reflect extended parental care. Our results demonstrate that female red squirrels engage in communal nesting, typically with closely related kin, despite a dispersed population structure that stems from the persistent defense of individual territories.     

Counterintuitive effects of large-scale predator removal on a midlatitude rodent community

Historically, small mammals have been focal organisms for studying predator-prey dynamics, principally because of interest in explaining the drivers of the cyclical dynamics exhibited by northern vole, lemming, and hare populations. However, many small-mammal species occur at relatively low and fairly stable densities at temperate latitudes, and our understanding of how complex predator assemblages influence the abundance and dynamics of these species is surprisingly limited. In an intact grassland ecosystem in western Montana, USA, we examined the abundance and dynamics of Columbian ground squirrels (Spermophilus columbianus), deer mice (Peromyscus maniculatus), and montane voles (Microtus montanus) on 1-ha plots where we excluded mammalian and avian predators and ungulates, excluded ungulates alone, or allowed predators and ungulates full access. Our goal was to determine whether the relatively low population abundance and moderate population fluctuations of these rodents were due to population suppression by predators. Our predator-exclusion treatment was divided into two phases: a phase where we excluded all predators except weasels (Mustela spp.; 2002-2005), and a phase where all predators including weasels were excluded (2006-2009). Across the entire duration of the experiment, predator and/or ungulate exclusion had no effect on the abundance or overall dynamics of ground squirrels and deer mice. Ground squirrel survival (the only species abundant enough to accurately estimate survival) was also unaffected by our experimental treatments. Prior to weasel exclusion, predators also had no impacts on montane vole abundance or dynamics. However, after weasel exclusion, vole populations reached greater population peaks, and there was greater recruitment of young animals on predator-exclusion plots compared to plots open to predators during peak years. These results suggest that the impacts of predators cannot be generalized across all rodents in an assemblage. Furthermore, they suggest that specialist predators can play an important role in suppressing vole abundance even in lower-latitude vole populations that occur at relatively low densities.     

Threat-sensitive foraging by larval threespine sticklebacks (Gasterosteus aculeatus)

Summary The threat-sensitive predator avoidance hypothesis predicts that prey can assess the relative threat posed by a predator and adjust their behaviour to reflect the magnitude of the threat. We tested the ability of larval threespine sticklebacks to adjust their foraging in the presence of predators by exposing them to conspecific predators of various sizes and recording their foraging and predator avoidance behaviours. Larvae (<30 days post-hatch) displayed predator escape behaviours only towards attacking predators. At 3 weeks post-hatch larvae approached the predator after fleeing, a behaviour which may be the precursor to predator inspection. Larvae reduced foraging and spent less time in the proximity of large and medium-sized predators compared to small predators. The reduction in foraging was negatively correlated to the predator/larva size ratio, indicating that larvae increased their foraging as they increased in size relative to the predator. We conclude that larval sticklebacks can assess the threat of predation early in their ontogeny and adjust their behaviour accordingly.Correspondence to: J.A. Brown     

Risk assessment in western mosquitofish (Gambusia affinis): do multiple cues have additive effects?

The threat-sensitivity hypothesis predicts that prey individuals will increase antipredator behaviors as apparent predator risk increases. An implicit assumption of the threat-sensitivity hypothesis is that predator risk is additive. In other words, all characteristics of a predator that indicate risk should contribute in an additive way to determine the degree of risk-sensitive behavior. We tested this assumption in the laboratory by presenting live predators (green sunfish, Lepomis cyanellus) to groups of western mosquitofish (Gambusia affinis). We examined effects of predator diet, hunger level, and size on predator avoidance and inspection behavior of mosquitofish. Both predator diet and predator hunger level were significant and additive determinants of distance maintained from a predator, resulting in a graded response to combinations of these predator cues. In contrast, whereas predator diet was the most important determinant of general avoidance distances, predator hunger level was more important in determining mosquitofish vertical distribution and inspection behavior. Thus, the relationship between predator cue and the antipredator behavior that it elicits is dependent on which cues and behaviors are examined. Our data suggest that during risky behaviors, such as predator inspection, mosquitofish rely mainly on visual cues (behavior differences between hungry and satiated predators), whereas general avoidance behavior is determined by additive responses from visual and chemical cues.     

Improving escape responses of hatchery-reared scallops <Emphasis Type="Italic">Argopecten purpuratus</Emphasis>

Hatchery rearing of the scallop Argopecten purpuratus has resulted in successive generations of scallops not exposed to predators that are less sensitive to and escape more slowly from predators than wild scallops. The present study examined whether conditioning hatchery-reared A. purpuratus to its natural predator, the sea star Meyenaster gelatinosus, improved its escape responses. Both juvenile and adult A. purpuratus from Tongoy Bay, Chile, were exposed for 7 days to different conditions: (1) continuous predator odor, (2) predator contact for 30 min three times a day, (3) a combination of the two previous conditions, and (4) no exposure to the predator (control). After conditioning, we evaluated scallop’s escape responses: reaction time, total clap number, duration of the clapping response, clapping rate, and the time scallops spent closed when exhausted. Conditioning with contact and odor plus contact (i.e., high predation risk) resulted in 25 and 50% shorter reaction times of juveniles and adults, respectively. Further, these stimuli caused juveniles to increase the number of claps and clapping rate. For adults, the time spent closed after exhaustion decreased by 50 and 63% after conditioning with contact and odor plus contact, respectively. Therefore, it is shown for the first time that exposure of scallops to increasing predator stimuli enhances escape responses, evidence of threat-sensitive predator avoidance.     

Causes and consequences of unequal seedling production in forest trees: a case study in red oaks

Inequality in reproductive success has important implications for ecological and evolutionary dynamics, but lifetime reproductive success is challenging to measure in long-lived species such as forest trees. While seed production is often used as a proxy for overall reproductive success, high mortality of seeds and the potential for trade-offs between seed number and quality draw this assumption into question. Parentage analyses of established seedlings can bring us one step closer to understanding the causes and consequences of variation in reproductive success. In this paper we demonstrate a new method for estimating individual seedling production and average percentage germination, using data from two mixed-species populations of red oaks (Quercus rubra, Q. velutina, Q. falcata, and Q. coccinea). We use these estimates to examine the distribution of female reproductive success and to test the relationship between seedling number and individual seed production, age, and growth rate. We show that both seed and seedling production are highly skewed, roughly conforming to zero-inflated lognormal distributions, rather than to the Poisson or negative-binomial distributions often assumed by population genetics analyses. While the number of established offspring is positively associated with mean annual seed production, a lower proportion of seeds from highly fecund individuals become seedlings. Our red oak populations also show evidence of trade-offs between growth rate and reproductive success. The high degree of inequality in seedling production shown here for red oaks, and by previous studies in other species, suggests that many trees may be more vulnerable to genetic drift than previously thought, if immigration in limited by fragmentation or other environmental changes.     

Different escape tactics of two vole species affect the success of the hunting predator, the least weasel

In the ongoing evolutionary arms race between predators and their prey, successful escape from the predator leads to the evolution of improved escape tactics in prey, but also predators become more effective in following and attacking the prey. Antipredatory behavior of prey is considered to be the strongest towards their most dangerous predators. However, prey species can differ both in vulnerability and efficiency of escape to a shared predator. We studied escape reactions of two vole species, the bank vole (Myodes glareolus) and the field vole (Microtus agrestis), under a simulated predation risk of the least weasel (Mustela nivalis nivalis). We conducted a laboratory experiment where a vole was given a possibility to escape from a weasel by fleeing to a horizontal tunnel or climbing the tree. Subsequently to the vole escape decision, we released a weasel to the same tunnel system to test how the weasel succeeded in following the vole. Weasel presence changed the behavior of voles as especially bank voles escaped by climbing. Instead, the majority of field voles fled into the ground-layer tunnel. The different escape tactics of the voles affected the success of the weasel, because climbing voles were less often successfully followed. We suggest that the difference in escape tactics has evolved as an adaptation to different habitats; meadow-exploiting field voles using ground-level escape while bank voles living in three-dimensional forest habitat frequently use arboreal escape tactics. This is likely to lead to different habitat-dependent vulnerabilities to predation in Microtus and Myodes vole species.     

Logging, Conifers, and the Conservation of Crossbills

A survey of the forestry literature shows that as the age and area of coniferous forests decline, decreased conifer seed production and increased frequencies of cone failures can be expected. This would, in turn, cause declines in crossbills ( Loxia ), which specialize on the seeds held in conifer cones. In western North America, at least five different species of Red Crossbills ( L. curvirostra ) have recently been distinguished (Groth 1990); each specializes on a different species or even a single variety of conifer (Benkman 1993). Measures for conserving this diversity of crossbills include protecting mature and old-growth stands, and increasing rotation ages throughout the range of each of the required conifers. These recommendations are not unique to crossbills, but rather the loss of crossbill diversity is another reason to employ such measures.     

The Paradox of the Long‐Term Positive Effects of a North American Crayfish on a European Community of Predators

Abstract: Invasions of non‐native species are one of the major causes of losses of native species. In some cases, however, non‐natives may also have positive effects on native species. We investigated the potential facilitative effects of the North American red swamp crayfish (Procambarus clarkii) on the community of predators in southwestern Spain. To do so, we examined the diets of predators in the area and their population trends since introduction of the crayfish. Most predator species consumed red swamp crayfish, which sometimes occurred in over 50% of their diet samples. Moreover, the abundance of species preying on crayfish increased significantly in the area as opposed to the abundance of herbivores and to predator populations in other areas of Europe, where those predators are even considered threatened. Thus, we report the first case in which one non‐native species is both beneficial because it provides prey for threatened species and detrimental because it can drive species at lower trophic levels to extinction. Increases in predator numbers that are associated with non‐native species of prey, especially when some of these predators are also invasive non‐natives, may increase levels of predation on other species and produce cascading effects that threaten native biota at longer temporal and larger spatial scales. Future management plans should include the complexity of interactions between invasive non‐natives and the entire native community, the feasibility of successful removal of non‐native species, and the potential social and economic interests in the area.     

Seed predation by Neotropical rain forest mammals increases diversity in seedling recruitment

Seed dispersal and seedling recruitment (the transition of seeds to seedlings) set the spatiotemporal distribution of new individuals in plant communities. Many terrestrial rain forest mammals consume post-dispersal seeds and seedlings, often inflicting density-dependent mortality. In part because of density-dependent mortality, diversity often increases during seedling recruitment, making it a critical stage for species coexistence. We determined how mammalian predators, adult tree abundance, and seed mass interact to affect seedling recruitment in a western Amazonian rain forest. We used exclosures that were selectively permeable to three size classes of mammals: mice and spiny rats (weighing <1 kg), medium-sized rodents (1-12 kg), and large mammals (20-200 kg). Into each exclosure, we placed seeds of 13 tree species and one canopy liana, which varied by an order of magnitude in adult abundance and seed mass. We followed the fates of the seeds and resulting seedlings for at least 17 months. We assessed the effect of each mammalian size class on seed survival, seedling survival and growth, and the density and diversity of the seedlings that survived to the end of the experiment. Surprisingly, large mammals had no detectable effect at any stage of seedling recruitment. In contrast, small- and medium-sized mammals significantly reduced seed survival, seedling survival, and seedling density. Furthermore, predation by small mammals increased species richness on a per-stem basis. This increase in diversity resulted from their disproportionately intense predation on common species and large-seeded species. Small mammals thereby generated a rare-species advantage in seedling recruitment, the critical ingredient for frequency dependence. Predation by small (and to a lesser extent, medium-sized) mammals on seeds and seedlings significantly increases tree species diversity in tropical forests. This is the first long-term study to dissect the effects of various mammalian predators on the recruitment of a diverse set of tree species.     

The influence of intraguild predation on prey suppression and prey release: a meta-analysis

Intraguild predation (IGP) occurs when one predator species consumes another predator species with whom it also competes for shared prey. One question of interest to ecologists is whether multiple predator species suppress prey populations more than a single predator species, and whether this result varies with the presence of IGP. We conducted a meta-analysis to examine this question, and others, regarding the effects of IGP on prey suppression. When predators can potentially consume one another (mutual IGP), prey suppression is greater in the presence of one predator species than in the presence of multiple predator species; however, this result was not found for assemblages with unidirectional or no IGP. With unidirectional IGP, intermediate predators were generally more effective than the top predator at suppressing the shared prey, in agreement with IGP theory. Adding a top predator to an assemblage generally caused prey to be released from predation, while adding an intermediate predator caused prey populations to be suppressed. However, the effects of adding a top or intermediate predator depended on the effectiveness of these predators when they were alone. Effects of IGP varied across different ecosystems (e.g., lentic, lotic, marine, terrestrial invertebrate, and terrestrial vertebrate), with the strongest patterns being driven by terrestrial invertebrates. Finally, although IGP theory is based on equilibrium conditions, data from short-term experiments can inform us about systems that are dominated by transient dynamics. Moreover, short-term experiments may be connected in some way to equilibrium models if the predator and prey densities used in experiments approximate the equilibrium densities in nature.     

Tidal regime dictates the cascading consumptive and nonconsumptive effects of multiple predators on a marsh plant

Prey perception of predators can dictate how prey behaviorally balance the need to avoid being eaten with the need to consume resources, and this perception and consequent behavior can be strongly influenced by physical processes. Physical factors, however, can also alter the density and diversity of predators that pursue prey. Thus, it remains uncertain to what extent variable risk perception and antipredator behavior vs. variation in predator consumption of prey underlie prey-resource dynamics and give rise to large-scale patterns in natural systems. In an experimental food web where tidal inundation of marsh controls which predators access prey, crab and conch (predators) influenced the survivorship and antipredator behavior of snails (prey) irrespective of whether tidal inundation occurred on a diurnal or mixed semidiurnal schedule. Specifically, cues of either predator caused snails to ascend marsh leaves; snail survivorship was reduced more by unrestrained crabs than by unrestrained conchs; and snail survivorship was lowest with multiple predators than with any single predator despite interference. In contrast to these tidally consistent direct consumptive and nonconsumptive effects, indirect predator effects differed with tidal regime: snail grazing of marsh leaves in the presence of predators increased in the diurnal tide but decreased in the mixed semidiurnal tidal schedule, overwhelming the differences in snail density that resulted from direct predation. In addition, results suggest that snails may increase their foraging to compensate for stress-induced metabolic demand in the presence of predator cues. Patterns from natural marshes spanning a tidal inundation gradient (from diurnal to mixed semidiurnal tides) across 400 km of coastline were consistent with experimental results: despite minimal spatial variation in densities of predators, snails, abiotic stressors, and marsh productivity, snail grazing on marsh plants increased and plant biomass decreased on shorelines exposed to a diurnal tide. Because both the field and experimental results can be explained by tidal-induced variation in risk perception and snail behavior rather than by changes in snail density, this study reinforces the importance of nonconsumptive predator effects in complex natural systems and at large spatial scales.