Seeing is believing: information content and behavioural response to visual and chemical cues

Predator avoidance and foraging often pose conflicting demands. Animals can decrease mortality risk searching for predators, but searching decreases foraging time and hence intake. We used this principle to investigate how prey should use information to detect, assess and respond to predation risk from an optimal foraging perspective. A mathematical model showed that solitary bees should increase flower examination time in response to predator cues and that the rate of false alarms should be negatively correlated with the relative value of the flower explored. The predatory ant, Oecophylla smaragdina, and the harmless ant, Polyrhachis dives, differ in the profile of volatiles they emit and in their visual appearance. As predicted, the solitary bee Nomia strigata spent more time examining virgin flowers in presence of predator cues than in their absence. Furthermore, the proportion of flowers rejected decreased from morning to noon, as the relative value of virgin flowers increased. In addition, bees responded differently to visual and chemical cues. While chemical cues induced bees to search around flowers, bees detecting visual cues hovered in front of them. These strategies may allow prey to identify the nature of visual cues and to locate the source of chemical cues.     

Incidental nest predation in freshwater turtles: inter- and intraspecific differences in vulnerability are explained by relative crypsis

There has long been interest in the influence of predators on prey populations, although most predator–prey studies have focused on prey species that are targets of directed predator searching. Conversely, few have addressed depredation that occurs after incidental encounters with predators. We tested two predictions stemming from the hypothesis that nest predation on two sympatric freshwater turtle species whose nests are differentially prone to opportunistic detection—painted turtles (Chrysemys picta) and snapping turtles (Chelydra serpentina)—is incidental: (1) predation rates should be density independent, and (2) individual predators should not alter their foraging behavior after encountering nests. After monitoring nest survival and predator behavior following nest depredation over 2 years, we confirmed that predation by raccoons (Procyon lotor), the primary nest predators in our study area, matched both predictions. Furthermore, cryptic C. picta nests were victimized with lower frequency than more detectable C. serpentina nests, and nests of both species were more vulnerable in human-modified areas where opportunistic nest discovery is facilitated. Despite apparently being incidental, predation on nests of both species was intensive (57% for painted turtles, 84% for snapping turtles), and most depredations occurred within 1 day of nest establishment. By implication, predation need not be directed to affect prey demography, and factors influencing prey crypsis are drivers of the impact of incidental predation on prey. Our results also imply that efforts to conserve imperiled turtle populations in human-modified landscapes should include restoration of undisturbed conditions that are less likely to expose nests to incidental predators.     

Evolution mediates the effects of apex predation on aquatic food webs

Ecological and evolutionary mechanisms are increasingly thought to shape local community dynamics. Here, I evaluate if the local adaptation of a meso-predator to an apex predator alters local food webs. The marbled salamander (Ambystoma opacum) is an apex predator that consumes both the spotted salamander (Ambystoma maculatum) and shared zooplankton prey. Common garden experiments reveal that spotted salamander populations which co-occur with marbled salamanders forage more intensely than those that face other predator species. These foraging differences, in turn, alter the diversity, abundance and composition of zooplankton communities in common garden experiments and natural ponds. Locally adapted spotted salamanders exacerbate prey biomass declines associated with apex predation, but dampen the top-down effects of apex predation on prey diversity. Countergradient selection on foraging explains why locally adapted spotted salamanders exacerbate prey biomass declines. The two salamander species prefer different prey species, which explains why adapted spotted salamanders buffer changes in prey composition owing to apex predation. Results suggest that local adaptation can strongly mediate effects from apex predation on local food webs. Community ecologists might often need to consider the evolutionary history of populations to understand local diversity patterns, food web dynamics, resource gradients and their responses to disturbance.     

Relationship between moonlight and nightly activity patterns of the ocelot (Leopardus pardalis) and some of its prey species in Formosa,Northern Argentina

The moon can profoundly influence the activity patterns of animals. If predators are more successful under bright moonlight, prey species are likely to respond by shifting their own activity patterns (predator-avoidance hypothesis). However, the assumption that prey will necessarily avoid full-moon nights does not take into account that moonlight also allows prey to more easily detect predators, and to forage more efficiently. Thus, nightly activity patterns could depend on night vision capabilities (visual-acuity hypothesis). To consider the possible influences of moonlight and to distinguish between these hypotheses, we used camera-trapping records of a predator, the ocelot (Leopardus pardalis), and several of its night-active prey to compare activity patterns under different moonlight conditions. The ocelots' activity patterns were not strongly related to moonlight, but showed a slight tendency for higher activity during brighter nights. Tapeti rabbits (Sylvilagus brasiliensis) and brocket deer (Mazama americana) showed a clear preference for brighter nights. White-eared opossums (Didelphis albiventris) also showed a trend to be less active in new moon light. In contrast, smaller grey four-eyed opossums (Philander opossum) and the poor eye-sight nine-banded armadillo (Dasypus novemcinctus) showed similar activity patterns across all moon phases. Since activity patterns of most prey species were not shifted away from the activity of the ocelot, the differences between species are probably linked to their night vision capabilities, and emphasise the need for more information on the visual system of these taxa. Their activity patterns seem to be less strongly linked to avoidance of predation than previously thought, suggesting that foraging and predator detection benefits may play a more important role than usually acknowledged.     

Predator identity and time of day interact to shape the risk–reward trade-off for herbivorous coral reef fishes

Non-consumptive effects (NCEs) of predators occur as prey alters their habitat use and foraging decisions to avoid predation. Although NCEs are recognized as being important across disparate ecosystems, the factors influencing their strength and importance remain poorly understood. Ecological context, such as time of day, predator identity, and prey condition, may modify how prey species perceive and respond to risk, thereby altering NCEs. To investigate how predator identity affects foraging of herbivorous coral reef fishes, we simulated predation risk using fiberglass models of two predator species (grouper Mycteroperca bonaci and barracuda Sphyraena barracuda) with different hunting modes. We quantified how predation risk alters herbivory rates across space (distance from predator) and time (dawn, mid-day, and dusk) to examine how prey reconciles the conflicting demands of avoiding predation vs. foraging. When we averaged the effect of both predators across space and time, they suppressed herbivory similarly. Yet, they altered feeding differently depending on time of day and distance from the model. Although feeding increased strongly with increasing distance from the predators particularly during dawn, we found that the barracuda model suppressed herbivory more strongly than the grouper model during mid-day. We suggest that prey hunger level and differences in predator hunting modes could influence these patterns. Understanding how context mediates NCEs provides insight into the emergent effects of predator–prey interactions on food webs. These insights have broad implications for understanding how anthropogenic alterations to predator abundances can affect the spatial and temporal dynamics of important ecosystem processes.     

Hydrodynamic mediation of predator-prey interactions: differential patterns of prey susceptibility and predator success explained by variation in water flow

In most shallow water marine systems, fluid movements vary on scales that may influence local community dynamics both directly, through changes in the abundance of species, and indirectly, by modifying important behaviors of organisms. We examined how differences in current speed affect the outcome of predator-prey interactions for two species of marine benthic predators (knobbed whelks, Busycon carica, and blue crabs, Callinectes sapidus) foraging on two common prey species (bay scallops, Argopecten irradians, and hard clams, Mercenaria mercenaria). The predators differ in their foraging strategies and prey in their potential escape responses. Predation by blue crabs, highly mobile predators/scavengers that rely upon chemical odors transported in the water column to locate prey, could be strongly affected by changes in current speed and turbulent mixing because their foraging strategy relies on a high degree of spatial integration of prey odor plumes. Whelks, slow moving, predatory gastropods that often forage with their bodies buried in the sediment, may be less susceptible to flow-induced distortion of prey odor plumes because their sluggish movements result in a high degree of temporal integration of prey odors. Bay scallops, relatively mobile bivalves capable of rapid short-distance swimming burst, and hard clams, sedentary bivalves, have been shown to respond to varying degrees to predator odors that are dispersed in the water column. Flow regime for the predator-prey experiments was manipulated in situ using large channels. Predation by blue crabs on both juvenile hard clams and bay scallops decreased with increases in water flow (0-12 vs. 0-30 cm s⁻¹). Whelk predation on bay scallops increased with increases in water flow, whereas predation by whelks on hard clams did not differ between flow regimes. For blue crabs movement decreased at periods of high water flow. Because blue crabs locate prey through chemolocation of water-borne cues, which are diluted rapidly at higher flows, decreases in foraging may result from the inability to successfully detect prey at enhanced flows. Differences in predation by whelks could not be explained by a similar mechanism. Visual observations of foraging whelks revealed no differences in whelk behavior between the two flow regimes. The pattern of higher whelk predation on scallops at enhanced flow is likely to be related to a flow-inhibiting ability of scallops to detect predator approach. Thus, flow enhancement interferes with three of the predator-prey systems but the effect on predator success depends on whether the predator or prey is most affected.     

An island-wide predator manipulation reveals immediate and long-lasting matching of risk by prey

Anti-predator behaviour affects prey population dynamics, mediates cascading effects in food webs and influences the likelihood of rapid extinctions. Predator manipulations in natural settings provide a rare opportunity to understand how prey anti-predator behaviour is affected by large-scale changes in predators. Here, we couple a long-term, island-wide manipulation of an important rodent predator, the island fox (Urocyon littoralis), with nearly 6 years of measurements on foraging by deer mice (Peromyscus maniculatus) to provide unequivocal evidence that prey closely match their foraging behaviour to the number of fox predators present on the island. Peromyscus maniculatus foraging among exposed and sheltered microhabitats (a measure of aversion to predation risk) closely tracked fox density, but the nature of this effect depended upon nightly environmental conditions known to affect rodent susceptibility to predators. These effects could not be explained by changes in density of deer mice over time. Our work reveals that prey in natural settings are cognizant of the dynamic nature of their predators over timescales that span many years, and that predator removals spanning many generations of prey do not result in a loss of anti-predator behaviour.     

Effects of Chemical Cues on Foraging in Damselfly Larvae, <Emphasis Type="Italic">Enallagma antennatum</Emphasis>

Animals experiencing a trade-off between predation risk and resource acquisition must accurately predict ambient levels of predation risk to maximize fitness. We measure this trade-off explicitly in larvae of the damselfly Enallagma antennatum, comparing consumption rates in the presence of chemical cues from predators and injured prey. Damselflies distinguished among types of chemical cues based on species of prey injured or eaten. Injured coexisting heterospecific and unknown heterospecific chemical cues did not reduce foraging relative to starved predator cues, while cues arising from predators eating a coexisting heterospecific did decrease foraging. This study shows a cost in terms of reduced foraging in response to chemical cues and further defines the ability of prey to respond discerningly to chemical cues.     

Turbidity amplifies the non‐lethal effects of predation and affects the foraging success of characid fish shoals

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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.     

Active foraging for toxic prey during gestation in a snake with maternal provisioning of sequestered chemical defences

Many animals sequester dietary defensive compounds and incorporate them into the offspring, which protects the young against predation. One possible but poorly investigated question is whether females of such species actively prey upon toxic diets. The snake Rhabdophis tigrinus sequesters defensive steroids from toads consumed as prey; it also feeds on other amphibians. Females produce chemically armed offspring in direct proportion to their own level of toad-derived toxins by provisioning the toxins to their eggs. Our field observations of movements and stomach contents of radio-tracked R. tigrinus showed that gravid snakes preyed upon toads by actively foraging in the habitat of toads, even though toads were a scarce resource and toad-searching may incur potential costs. Our Y-maze experiments demonstrated that gravid females were more likely to trail the chemical cues of toads than were males or non-gravid females. These results showed behavioural switching in females and active foraging for scarce, toxic prey during gestation. Because exploitation of toads by gravid females results in their offspring being more richly endowed with prey-derived toxins, active foraging for toxic prey is expected to be an adaptive antipredator trait, which may enhance chemical defence in offspring.     

Linking Morphological and Behavioural Defences: Prey Fish Detect the Morphology of Conspecifics in the Odour Signature of their Predators

Predation is a strong selective force acting on both morphology and behaviour of prey animals. While morphological defences (e.g. crypsis, presence of armours or spines or specific body morphologies) and antipredator behaviours (e.g. change in foraging or reproductive effort, or hiding and fleeing behaviours) have been widely studied separately, few studies have considered the interplay between the two. The question raised in our study is whether antipredator behaviours of a prey fish to predator odours could be influenced by the morphology of prey conspecifics in the diet of the predator. We used goldfish (Carassius auratus) as our test species; goldfish exposed to predation risk significantly increase their body depth to length ratio, which gives them a survival advantage against gape‐limited predators. We exposed shallow‐bodied and deep‐bodied goldfish to the odour of pike (Esox lucius) fed either form of goldfish. Deep‐bodied goldfish displayed lower intensity antipredator responses than shallow‐bodied ones, consistent with the hypothesis that individuals with morphological defences should exhibit less behavioural modification than those lacking such defences. Moreover, both shallow‐ and deep‐bodied goldfish displayed their strongest antipredator responses when exposed to the odour of pike fed conspecifics of their own morphology, indicating that goldfish are able to differentiate the morphology of conspecifics through predator diet cues. For a given individual, predator threat increases as the prey become more like the individual eaten, revealing a surprising level of sophistication of chemosensory assessment by prey fish.     

Prey bacteria shape the community structure of their predators

Although predator–prey interactions among higher organisms have been studied extensively, only few examples are known for microbes other than protists and viruses. Among the bacteria, the most studied obligate predators are the Bdellovibrio and like organisms (BALOs) that prey on many other bacteria. In the macroscopical world, both predator and prey influence the population size of the other''s community, and may have a role in selection. However, selective pressures among prey and predatory bacteria have been rarely investigated. In this study, Bacteriovorax, a predator within the group of BALOs, in environmental waters were fed two prey bacteria, Vibrio vulnificus and Vibrio parahaemolyticus. The two prey species yielded distinct Bacteriovorax populations, evidence that selective pressures shaped the predator community and diversity. The results of laboratory experiments confirmed the differential predation of Bacteriovorax phylotypes on the two bacteria species. Not only did Bacteriovorax Cluster IX exhibit the versatility to be the exclusive efficient predator on Vibrio vulnificus, thereby, behaving as a specialist, but was also able to prey with similar efficiency on Vibrio parahaemolyticus, indicative of a generalist. Therefore, we proposed a designation of versatilist for this predator. This initiative should provide a basis for further efforts to characterize the predatory patterns of bacterial predators. The results of this study have revealed impacts of the prey on Bacteriovorax predation and in structuring the predator community, and advanced understanding of predation behavior in the microbial world.     

Activity pattern as a mechanism of predator avoidance in two species of acmaeid limpet

Activity patterns in the foraging activity of two species of acmaeid limpets, Collisella limatula (Carpenter, 1864) and C. scabra) (Gould, 1846) were documented over the entire tidal cycle. C. limatula was found to be active only while awash by the tide and during night-time emergence; periods of inactivity were spent in crevices and on the undersides of boulders. C. scabra, which homes to a scar on the upper surfaces of rocks, foraged only while awash in daylight. The activity pattern of C. limatula was demonstrated to be an effective mechanism for avoiding predation by Octopus, a major visual predator of limpets, in both laboratory and field experiments. The homing habit of Collisella scabra was also shown to reduce the rate of predation in the laboratory. It is suggested that the observed activity patterns have evolved in response to predation pressure from swift-moving visual predators.     

Behavioral “bycatch” from camera trap surveys yields insights on prey responses to human‐mediated predation risk

Human disturbance directly affects animal populations and communities, but indirect effects of disturbance on species behaviors are less well understood. For instance, disturbance may alter predator activity and cause knock‐on effects to predator‐sensitive foraging in prey. Camera traps provide an emerging opportunity to investigate such disturbance‐mediated impacts to animal behaviors across multiple scales. We used camera trap data to test predictions about predator‐sensitive behavior in three ungulate species (caribou Rangifer tarandus; white‐tailed deer, Odocoileus virginianus; moose, Alces alces) across two western boreal forest landscapes varying in disturbance. We quantified behavior as the number of camera trap photos per detection event and tested its relationship to inferred human‐mediated predation risk between a landscape with greater industrial disturbance and predator activity and a “control” landscape with lower human and predator activity. We also assessed the finer‐scale influence on behavior of variation in predation risk (relative to habitat variation) across camera sites within the more disturbed landscape. We predicted that animals in areas with greater predation risk (e.g., more wolf activity, less cover) would travel faster past cameras and generate fewer photos per detection event, while animals in areas with less predation risk would linger (rest, forage, investigate), generating more photos per event. Our predictions were supported at the landscape‐level, as caribou and moose had more photos per event in the control landscape where disturbance‐mediated predation risk was lower. At a finer‐scale within the disturbed landscape, no prey species showed a significant behavioral response to wolf activity, but the number of photos per event decreased for white‐tailed deer with increasing line of sight (m) along seismic lines (i.e., decreasing visual cover), consistent with a predator‐sensitive response. The presence of juveniles was associated with shorter behavioral events for caribou and moose, suggesting greater predator sensitivity for females with calves. Only moose demonstrated a positive behavioral association (i.e., longer events) with vegetation productivity (16‐day NDVI), suggesting that for other species bottom‐up influences of forage availability were generally weaker than top‐down influences from predation risk. Behavioral insights can be gleaned from camera trap surveys and provide complementary information about animal responses to predation risk, and thus about the indirect impacts of human disturbances on predator–prey interactions.     

Modeling changes in predator functional response to prey across spatial scales

Extrapolation of predator functional responses from laboratory observations to the field is often necessary to predict predation rates and predator-prey dynamics at spatial and temporal scales that are difficult to observe directly. We use a spatially explicit individual-based model to explore mechanisms behind changes in functional responses when the scale of observation is increased. Model parameters were estimated from a predator-prey system consisting of the predator Delphastus catalinae (Coleoptera: Coccinellidae) and Bemisia tabaci biotype B (Hemiptera: Aleyrodidae) on tomato plants. The model explicitly incorporates prey and predator distributions within single plants, the search behavior of predators within plants, and the functional response to prey at the smallest scale of interaction (within leaflets) observed in the laboratory. Validation revealed that the model is useful in scaling up from laboratory observations to predation in whole tomato plants of varying sizes. Comparing predicted predation at the leaflet scale, as observed in laboratory experiments, with predicted predation on whole plants revealed that the predator functional response switches from type II within leaflets to type III within whole plants. We found that the magnitude of predation rates and the type of functional response at the whole plant scale are modulated by (1) the degree of alignment between predator and prey distributions and (2) predator foraging behavior, particularly the effect of area-concentrated search within plants when prey population density is relatively low. The experimental and modeling techniques we present could be applied to other systems in which active predators prey upon sessile or slow-moving species.     

Linking predator risk and uncertainty to adaptive forgetting: a theoretical framework and empirical test using tadpoles

Hundreds of studies have examined how prey animals assess their risk of predation. These studies work from the basic tennet that prey need to continually balance the conflicting demands of predator avoidance with activities such as foraging and reproduction. The information that animals gain regarding local predation risk is most often learned. Yet, the concept of ‘memory’ in the context of predation remains virtually unexplored. Here, our goal was (i) to determine if the memory window associated with predator recognition is fixed or flexible and, if it is flexible, (ii) to identify which factors affect the length of this window and in which ways. We performed an experiment on larval wood frogs, Rana sylvatica, to test whether the risk posed by, and the uncertainty associated with, the predator would affect the length of the tadpoles'' memory window. We found that as the risk associated with the predator increases, tadpoles retained predator-related information for longer. Moreover, if the uncertainty about predator-related information increases, then prey use this information for a shorter period. We also present a theoretical framework aiming at highlighting both intrinsic and extrinsic factors that could affect the memory window of information use by prey individuals.     

Habitat overlap between predatory benthic fish and their invertebrate prey in streams: the relative influence of spatial and temporal factors on predation risk

1. The spatial heterogeneity of ecosystems as well as temporal activity patterns of organisms can have far‐reaching effects on predator–prey relationships. We hypothesised that spatiotemporal constraints in mesohabitat use by benthic fish predators would reduce habitat overlap with benthic invertebrates and lead to mesohabitat‐specific predation risks. 2. We analysed the spatiotemporal activity patterns of two small‐bodied benthivorous fishes, gudgeon (Gobio gobio) and stone loach (Barbatula barbatula), and of benthic invertebrates in a small temperate stream during three 24‐h field experiments. By applying a novel method of field video observation, we monitored the spatiotemporal foraging behaviour of the fish in their natural environment. A parallel analysis of invertebrate mesohabitat use by means of small area Hess sampling allowed a direct estimation of habitat overlap at a pool–riffle scale. 3. Gudgeon showed a dominant spatial activity pattern preferring pools at all times of day, whereas stone loach used both mesohabitats but with a distinct temporal (nocturnal) activity pattern. The patterns of residence were not identical with those of active foraging. Invertebrate community composition differed significantly between mesohabitats but not between times of day. More than half of the total dissimilarity between pools and riffles was accounted for by six invertebrate taxa. Five of these were subject to higher fish predation in pools than in riffles. The total prey consumption of the two fish species together in pools was about three times as high as in riffles. Trophic niche breadth of stone loach and thus its predation range was broader than that of gudgeon. 4. These results indicate that the potential predation risk for stream invertebrates depends on the combination of spatial and temporal patterns of both predator and prey. Given the distinct differences in predation risk found between pools and riffles, we conclude that spatial heterogeneity at the mesohabitat scale can influence mechanisms and consequences of selective predation. We also suggest that the analysis of spatiotemporal predator–prey relationships should not be based on the premise that the main residence habitat and active foraging habitat of a predator are identical.     

Vulnerability of age-0 pallid sturgeon Scaphirhynchus albus to predation; effects of predator type,turbidity, body size,and prey density

Predation can play an important role in the recruitment dynamics of fishes with intensity regulated by behavioral (i.e., prey selectivity) and/or environmental conditions that may be especially important for rare or endangered fishes. We conducted laboratory experiments to quantify prey selection and capture efficiency by three predators employing distinct foraging strategies: pelagic piscivore (walleye Sander vitreus); benthic piscivore (flathead catfish Pylodictis olivaris) and generalist predator (smallmouth bass Micropterus dolomieu) foraging on two size classes of age-0 pallid sturgeon: large (75–100 mm fork length [FL]) and small (40–50 mm FL). Experiments at high (> 70 nephalometric turbidity units [NTU]) and low (< 5 NTU) turbidity for each predator were conducted with high and low densities of pallid sturgeon and contrasting densities of an alternative prey, fathead minnow Pimephales promelas. Predator behaviors (strikes, captures, and consumed prey) were also quantified for each prey type. Walleye and smallmouth bass negatively selected pallid sturgeon (Chesson’s α?=?0.04–0.1) across all treatments, indicating low relative vulnerability to predation. Relative vulnerability to predation by flathead catfish was moderate for small pallid sturgeon (α?=?0.44, neutral selection), but low for large pallid sturgeon (α?=?0.11, negative selection). Turbidity (up to 100 NTU) did not affect pallid sturgeon vulnerability, even at low density of alternative prey. Age-0 pallid sturgeon were easily captured by all predators, but were rarely consumed, suggesting mechanisms other than predator capture efficiency govern sturgeon predation vulnerability.     

Foraging time and spatial patterns of predation in experimental populations

Summary Responses of the predaceous mites Phytoseiulus persimilis, Typhlodromus (=Metaseiulus) occidentalis, and Amblyseius andersoni to spatial variation in egg density of the phytophagous mite, Tetranychus urticae, were studied in the laboratory.The oligophagous predator P. persimilis showed initially a direct density dependent foraging time allocation and variation in foraging time increased with prey density. With changes in prey density due to predation, predator foraging rates (per hour) decreased with time and density dependent foraging gradually became density independence, because P. persimilis continued to respond to initial prey density, instead of the changing prey density and distribution. The consequent spatial pattern of predation by P. persimilis was density independent, although slopes of predation rate-prey density regressions increased with time.Compared with P. persimilis, the narrowly polyphagous predator T. occidentalis responded relatively slowly to the the presence or absence of prey eggs but not to prey density: the mean and variation of foraging time spent in patches with prey did not differ with prey density, but was significantly greater in patches with prey eggs than in patches without eggs. Prey density and distribution changed only slightly due to predation and overall foraging rates remained more or less constant. The consequent spatial pattern of predation by T. occidentalis was inversely density dependent. As with P. persimilis, slopes of predation rate-prey density regressions increased with time (i.e. the inverse density dependence in T. occidentalis became weaker through time).The broadly polyphagous predator A. andersoni showed density independent foraging time allocation with variation independent of prey density. With changes in prey density over time due to prey depletion, overall foraging rates decreased. The consequent spatial pattern of predation by A. andersoni also changed through time; it initially was inversely density dependent, but soon became density independent.Overall, P. persimilis and T. occidentalis spent more time in prey patches than A. andersoni, suggesting that A. andersoni tended to spend more time moving outside patches. The overall predation rates and searching efficiency were higher in P. persimilis than in A. andersoni and T. occidentalis. Predator reproduction was highest in P. persimilis, lower in T. occidentalis and the lowest A. andersoni.The differences in response to prey distribution among the three predaceous species probably reflect the evolution of these species in environments with different patterns of prey distribution. The degree of polyphagy is a major determinant of the aggregative response, but other attributes such as handling time are also important in other aspects of phytoseiid foraging behavior (e.g. searching efficiency or predation rate).