Does risk of intraspecific interactions induce shifts in prey-size preference in aquatic predators?

Interactions between foragers may seriously affect individual foraging efficiency. In a laboratory study of handling time, prey value and prey-size preference in northern pike and signal crayfish, we show that risk of intraspecific interactions between predators does not affect handling time or value of prey. However, the presence of agonistic intraspecific interactors shifts prey-size preference in these predators. Neither northern pike nor signal crayfish foraging alone show a prey-size preference, while pike foraging among conspecifics prefer small prey, and crayfish foraging in groups prefer large prey. We ascribe the different outcomes in prey preference to differences in susceptibility to interactions: northern pike under risk avoid large prey to avoid long handling times and the associated risk of interactions, while signal crayfish foraging among conspecifics may defend themselves and their prey during handling, and thus select prey to maximise investment. In addition, the value of pike prey (roach) is low for very small prey, maximises for small prey, and then decreases monotonically for larger prey, while crayfish prey (pond snail) value is low for very small prey, has a maximum at small prey, but does not decrease as much for larger prey. Therefore, a large and easily detected snail prey provides a crayfish with as much value as a small prey. We conclude that interaction risk and predator density affect prey-size preference differently in these aquatic predators, and therefore has different potential effects on prey-size structure and population and community dynamics. Received: 4 October 1999 / Revised: 20 March 2000 / Accepted: 27 May 2000     

Feeding behavior of the sea star Astropecten articulatus (Echinodermata: Asteroidea): an evaluation of energy-efficient foraging in a soft-bottom predator

The energy efficiency of the foraging behavior of Astropecten articulatus (Say) was evaluated in the laboratory. Individuals utilized in the study were collected in the northern Gulf of Mexico from 1990 to 1992. Sea stars presented with equal numbers and weights of low-quality and high-quality prey consistently selected prey of the higher quality. Choice of prey appeared to be mediated by contact chemoreception. Individuals presented with equal weights or equal numbers of different-sized prey models demonstrated a significant preference for smaller prey. Size-selective feeding may be attributable to the ease of manipulation and ingestion of smaller prey, which mazimizes food intake per unit time. In the absence of prey, A. articulatus exhibited a directional pattern of movement. However, as prey were encountered, both the frequency and magnitude of angular deviations in the foraging path increased, resulting in increased foraging in areas of higher prey density. This response to prey availability may increase foraging efficiency by maximizing the rate of prey encounter. Like four other species of the genus Astropecten, A. articulatus exhibited two peaks of activity corresponding with dawn and dusk. Diurnal activities with periods of increased prey availability or periods during which predators are diminished or absent. The sea star A. articulatus exhibits foraging behaviors consistent with the maximization of net energy intake per unit foraging time.     

Individual specialization in the hunting wasp <Emphasis Type="Italic">Trypoxylon</Emphasis> (<Emphasis Type="Italic">Trypargilum</Emphasis>) <Emphasis Type="Italic">albonigrum</Emphasis> (Hymenoptera,Crabronidae)

Individual-level variation in resource use occurs in a broad array of vertebrate and invertebrate taxa and may have important ecological and evolutionary implications. In this study, we measured the degree of individual-level variation in prey preference of the hunting wasp Trypoxylon albonigrum, which inhabits the Atlantic Forest in southeastern Brazil. This wasp captures several orb-weaving spider genera to provision nests. Individuals consistently specialized on a narrow subset of the prey taxa consumed by the population, indicating the existence of significant individual-level variation in prey preferences. The population niche was broader in the wet season in terms of both prey size and taxa. In the case of prey size, the population niche expansion was achieved via increased individual niche breadths, whereas in the case of prey taxa, individual niches remained relatively constrained, and the population niche expanded via increased interindividual variation. The observed pattern suggests the possibility of functional trade-offs associated with the taxon of the consumed prey. The nature of the trade-offs remains unknown, but they are likely related to learning in searching and/or handling prey. We hypothesize that by specializing on specific prey taxa, individuals increase foraging efficiency, reducing foraging time and ultimately increasing reproductive success.     

What limits predator detection in blue tits (<Emphasis Type="Italic">Parus caeruleus</Emphasis>): posture,task or orientation?

To detect threats and reduce predation risk prey animals need to be alert. Early predator detection and rapid anti-predatory action increase the likelihood of survival. We investigated how foraging affects predator detection and time to take-off in blue tits (Parus caeruleus) by subjecting them to a simulated raptor attack. To investigate the impact of body posture we compared birds feeding head-down with birds feeding head-up, but could not find any effect of posture on either time to detection or time to take-off. To investigate the impact of orientation we compared birds having their side towards the attacking predator with birds having their back towards it. Predator detection, but not time to take-off, was delayed when the back was oriented towards the predator. We also investigated the impact of foraging task by comparing birds that were either not foraging, foraging on chopped mealworms, or foraging on whole ones. Foraging on chopped mealworms did not delay detection compared to nonforaging showing that foraging does not always restrict vigilance. However, detection was delayed more than 150% when the birds were foraging on whole, live mealworms, which apparently demanded much attention and handling skill. Time to take-off was affected by foraging task in the same way as detection was. We show that when studying foraging and vigilance one must include the difficulty of the foraging task and prey orientation.Communicated by P.A. Bednekoff     

Pursuit-deterrent communication between prey animals and timber rattlesnakes (Crotalus horridus): the response of snakes to harassment displays

A thorough understanding of communication requires an evaluation of both the signaler and receiver. Most analyses of prey–predator communication are incomplete because they examine only the behavior of the prey. Predators in these systems may be understudied because they are perceived as less tractable research subjects, due to their more cryptic hunting behaviors and secretive lifestyles. For example, research on interactions between rodents and rattlesnakes has focused on the behavior of rodent signalers, while responses of snakes have been virtually unexamined. Rattlesnakes are ambush predators, and capture rodents by waiting at foraging sites for long periods of time. In this study, I take advantage of the sedentary nature of this foraging strategy and use fixed videography to record natural encounters between timber rattlesnakes (Crotalus horridus) and their prey. Three different prey species were found to exhibit conspicuous visual displays to snakes, both when snakes were actively foraging, and when they were basking. After receiving displays, foraging snakes left their ambush sites and moved long distances before locating subsequent ambush sites, indicating that they responded to displays by abandoning attempts to ambush prey in the vicinity of signalers. This study represents the first quantitative analysis of the response of free-ranging snakes to signals from their prey, and elucidates a technique by which such quantitative data can be more easily obtained.     

Going deep: common murres dive into frigid water for aggregated,persistent and slow-moving capelin

Owing to the necessity of delivering food to offspring at colonies, breeding seabirds are highly constrained in their foraging options. To minimize constraints imposed by central-place foraging and to optimize foraging behavior, many species exhibit flexible foraging tactics. Here we document the behavioral flexibility of pursuit-diving common murres Uria aalge when foraging on female capelin Mallotus villosus in the northwest Atlantic. Quite unexpectedly, being visual foragers, we found that common murres dived throughout the day and night. Twenty-one percent of recorded dives (n = 272 of 1,308 dives) were deep (≥50 m; maximum depth = 152 m, maximum duration = 212 s), bringing murres into sub-0°C water in the Cold Intermediate Layer (CIL; 40–180 m) of the Labrador Current. Deep dives occurred almost exclusively during the day when murres would have encountered spatially predictable aggregations of capelin between 100 and 150 m in the water column. Temperatures within the CIL shaped trophic interactions and involved trade-offs for both predators and prey. Sub-0°C temperatures limit a fish’s ability to escape from endothermic predators by reducing burst/escape speeds and also lengthening the time needed to recover from burst-type activity. Thus, while deep diving may be energetically costly, it likely increases certainty of prey capture. Decreased murre foraging efficiency at night (indicated by an increase in the number of dives per bout) reflects both lower light conditions and changing prey behavior, as capelin migrate to warmer surface waters at night where their potential to escape from avian predators could increase.     

Prey caloric value and predator energy needs: foraging predictions for wild spinner dolphins

Spinner dolphins (Stenella longirostris) feed on individual small (2–10 cm long) prey that undergo diel vertical migrations, presumably making them inaccessible to dolphins during the day. To examine how time, prey behavior, prey distribution, and energy needs constrain dolphin foraging, a calorimeter was used to measure the caloric content of prey items. These data were combined with information on prey distribution in the field and the energetic needs of dolphins to construct basic bioenergetic models predicting the total prey consumption and mean feeding rates of wild dolphins as well as potential prey preferences. The mean caloric density of mesopelagic animals from Hawaii was high (2,837 cal/g wet weight for shrimps, squids, and myctophid fishes). Their total caloric content, however, was low because of their small size. Energy value of prey and energetic needs of spinner dolphins were used to examine the effect of time and energy constraints on dolphin foraging. The results predict that spinner dolphins need to consume an estimated minimum of 1.25 large prey items per minute to meet their maintenance energy needs. If the additional energy costs of foraging are considered, the estimated necessary foraging rate is predicted to increase only slightly when large prey are consumed. If smaller prey are consumed, the total energy demand may be twice the basic maintenance value. Prey density and size are predicted to be important in determining if dolphins can forage successfully, meeting their energetic needs. The prey size predictions compare well with results from previous gut content studies and from stomach contents of a recently stranded spinner dolphin that had enough prey in its stomach to meet its estimated basic maintenance energy needs for a day. Finally, the results suggest that spinner dolphins are time and therefore efficiency limited rather than being limited by the total amount of available prey. This may explain the diel migration exhibited by spinner dolphins that allows them to follow the movements of their prey and presumably maximizes their foraging time.Communicated by P.W. Sammarco, Chauvin     

Maximization of foraging efficiency and resource defense by group retrieval in the ant Formica schaufussi

Summary Workers of the ant Formica schaufussi forage as individuals and cooperate in groups to retrieve arthropod prey. In 2 sample years, group-transported prey were on average 6.8 and 4.7 times heavier than individually retrieved items, and the average loading ratios of groups were greater than the loading ratios of single foragers. Retrieval group size was adjusted to prey size, and prey transport velocity for individuals and groups tended to decrease with increasing prey weight. The efficiency of individual and group retrieval, estimated from calculations of the prey delivery rate to the nest (PDR) achieved by each foraging mode, varied as a function of prey size. Individual retrieval maximized PDR at a prey weight of 19.5 mg, and group transport maximized PDR at 190 mg. Although the PDR maxima of an individual in a group and a solitary forager were approximately equal, depending on prey size, group transport may maximize foraging efficiency. Group transport also decreased interference competition from sympatric ant species. Group-transported prey having a greater likelihood of successful retrieval were within the size range of prey that maximized foraging efficiency. Transport group size appeared to be more important in prey defense than in increasing prey transport velocity, suggesting an important role of group size in competitive ability.Offprint requests to: J.F.A. Traniello     

Parent-offspring aggression in moorhens

Summary Colonial orb-weaving spiders from Mexico were studied to test predictions of risksensitive foraging theory: 1. group foraging increases prey capture/individual, and reduces prey variance; 2. spiders should be expected to exhibit risk-averse behavior (forage in groups) when the average level of prey exceeds individual needs, and exhibit risk-prone behavior (forage solitarily) when prey are searce. Laboratory and field studies show that group foraging increases capture efficiency and reduces variability in prey captured per spider. In desert/mesquite grassland habitat, where prey availability is low, M. atascadero forage solitarily in most cases. In tropical rainforest/agriculture sites, M. increassata forage in large colonies of thousands of webs. In intermediate habitats, M. spinipes forages solitarily or in groups, depending on prey availability. Over a range of sites with varying levels of prey, M. spinipes shifts from a risk-prone to a risk-averse group foraging strategy as prey increases.Group foraging behavior observed in colonial Metepeira fits the predictions of risk-sensitive foraging models. These findings explain why spiders tend to group webs together only in areas of superabundant prey. The role of risk-sensitivity in the evolution of coloniality in spiders is discussed.     

Prey selection in <Emphasis Type="Italic">Octopus rubescens</Emphasis>: possible roles of energy budgeting and prey nutritional composition

This study explores the relationship between energy budgeting and prey choice of Octopus rubescens. Seventeen male Octopus rubescens were collected between June 2006 and August 2007 from Admiralty Bay, Washington. Prey choices made by individuals in the laboratory deviated widely from those expected from a simple optimal foraging model. O. rubescens chose the crab Hemigrapsus nudus over the clam Nuttallia obscurata as prey by a ratio of 3:1, even though prey energy content and handling times suggested that this octopus could obtain 10 times more energy intake per unit time when choosing the latter compared to the former prey species. Octopus energy budgets were similar when consuming either of the prey species except for lipid extraction efficiency that was significantly higher in octopuses consuming H. nudus. This suggests that lipid digestibility may play an important role in the prey choice of O. rubescens.     

Foraging in groups of water striders (Gerris remigis): effects of variability in prey arrivals and handling times

Summary In many species, socially subordinate individuals frequently remain in a group despite their lower priority of access to food and mates. This is expected to occur when the net benefits of staying in the group exceed those of a solitary existence. Analogously, territorial foragers surrounded by conspecific competitors are faced with tradeoffs as to patch tenacity. In either case, spatio-temporal variability in resource availability directly affects behavioral patterns and payoffs, particularly of individuals with low priority of access to the resource. However, such individuals may take advantage of natural environmental fluctuations in resource supply in cases where dominants are preoccupied. This arises regularly if two resource items cannot be handled simultaneously and if a second item arrives before handling of the first has been completed. I advance a heuristic model that predicts that foraging or mating success of individuals with low priority of access to resources may increase both with higher variance in inter-arrival times of the resource (given the same mean) and with an increase in the average handling time of the resource. I tested both predictions with two associations of individually marked, naturally foraging water striders (Gerris remigis) in the field. In natural streams individual water striders tend to occupy consistent positions that they may defend, resulting in priority of access to prey items floating downstream for individuals further up front. I manipulated the variance in prey inter-arrival times given the same mean, and the prey handling times by offering larger prey. The outcome was in qualitative agreement with the predictions of the model.     

Effects of aggregation on feeding and survival in a communal wren

Summary Foraging by a social wren, Campylorhynchus nuchalis (Troglodytidae), in a tropical savanna habitat is not enhanced by aggregation. Data for marked individuals show that solitary foraging results in a higher capture rate than foraging near others. We find no evidence of imitative foraging, as individuals actively avoid successful foragers following a capture and successful foragers do not restrict their search to recently productive stations or techniques. Captures are seldom temporally clumped, and clumping is probably not pronounced enough to favor imitation. Juveniles show no greater tendency to respond to captures of others, or to succeed in foraging in a group, than do adults. Aggregation is probably disadvantageous for foraging because of dispersed, scarce, cryptic, and noneruptive prey and because of the searching technique of these foliage-gleaning insectivores. If predator avoidance is enhanced by aggregation, it does not result in either increased survival or increased foraging efficiency in large groups, even by juveniles.     

Diet of young Atlantic bluefin tuna (<Emphasis Type="Italic">Thunnus thynnus</Emphasis>) in eastern and western Atlantic foraging grounds

Atlantic bluefin tuna (Thunnus thynnus) are highly migratory predators whose abundance, distribution, and somatic condition have changed over the past decades. Prey community composition and abundance have also varied in several foraging grounds. To better understand underlying food webs and regional energy sources, we performed stomach content and stable isotope analyses on mainly juvenile (60–150 cm curved fork length) bluefin tuna captured in foraging grounds in the western (Mid-Atlantic Bight) and eastern (Bay of Biscay) Atlantic Ocean. In the Mid-Atlantic Bight, bluefin tuna diet was mainly sand lance (Ammodytes spp., 29% prey weight), consistent with historic findings. In the Bay of Biscay, krill (Meganyctiphanes norvegica) and anchovy (Engraulis encrasicolus) made up 39% prey weight, with relative consumption of each reflecting annual changes in prey abundance. Consumption of anchovies apparently declined after the local collapse of this prey resource. In both regions, stable isotope analysis results showed that juvenile bluefin tuna fed at a lower trophic position than indicated by stomach content analysis. In the Mid-Atlantic Bight, stable isotope analyses suggested that >30% of the diet was prey from lower trophic levels that composed <10% of the prey weights based upon traditional stomach content analyses. Trophic position was similar to juvenile fish sampled in the NW Atlantic but lower than juveniles sampled in the Mediterranean Sea in previous studies. Our findings indicate that juvenile bluefin tuna targeted a relatively small range of prey species and regional foraging patterns remained consistent over time in the Mid-Atlantic Bight but changed in relation to local prey availability in the Bay of Biscay.     

How the mountain chickadee procures more food in less time for its nestlings

Summary Parent mountain chickadees (Parus gambeli) visit the same foraging location on consecutive flights more frequently than expected by chance. This pattern of repetitive return to a foraging location does not increase the frequency of capturing the same prey on consecutive flights. The quantity of prey captured on a visit also does not change significantly with repeated visits to a foraging location. Foraging time, however, is significantly shorter during repeated visits to a foraging location. Foraging time is therefore a major determinant of whether a run of consecutive flights to a location occurs. Decreased foraging time is also a benefit of selecting the same prey type on consecutive flights. Over the course of a day, the prey types returned on the greatest percentage of flights are returned multiply per flight. These prey are usually small and are probably present at high densities in the environment. Large prey size is the second best predictor of frequency of a prey type in the diet. Short foraging time to capture a prey is the third best predictor. Thus prey which can be readily obtained in large numbers, even if they are small prey, are preferentially returned to older nestlings. The mountain chickadee uses information on foraging time to decide when to move to a new foraging location or when to pursue a different type of prey. These decisions should increase the amount of time available for prey gathering.     

Foraging behavior in early Atlantic cod larvae (Gadus morhua) feeding on a protozoan (Balanionsp.) and a copepod nauplius (Pseudodiaptomussp.)

Food limitation is likely to be a source of mortality for fish larvae in the first few weeks after hatching. In the laboratory, we analyzed all aspects of foraging in cod larvae (Gadus morhua Linnaeus) from 5 to 20 d post-hatching using protozoa (Balanion sp.) and copepod nauplii (Pseudodiaptomus sp.) as prey. A camera acquisition system with two orthogonal cameras and a digital image analysis program was used to observe patterns of foraging. Digitization provided three-dimensional speeds, distances, and angles for each foraging event, and determined prey and fish larval head and tail positions. Larval cod swimming speeds, perception distances, angles, and volumes increased with larval fish size. Larval cod swam in a series of short intense bursts interspersed with slower gliding sequences. In 94% of all foraging events prey items were perceived during glides. Larval cod foraging has three possible outcomes: unsuccessful attacks, aborted attacks, and successful attacks. The percentage of successful attacks increased with fish size. In all larval fish size classes, successful attacks had smaller attack distances and faster attack speeds than unsuccessful attacks. Among prey items slowly swimming protozoans were the preferred food of first-feeding cod larvae; larger larvae had higher swimming speeds and captured larger, faster copepod nauplii. Protozoans may be an important prey item for first-feeding larvae providing essential resources for growth to a size at which copepod nauplii are captured. Received: 20 April 1999 / Accepted: 12 January 2000     

The influence of subsurface thermal structure on the diving behavior of northern fur seals (<Emphasis Type="Italic">Callorhinus ursinus</Emphasis>) during the breeding season

In the heterogeneous marine environment, predators can increase foraging success by targeting physical oceanographic features, which often aggregate prey. For northern fur seals (Callorhinus ursinus), two prevalent oceanographic features characterize foraging areas during summer in the Bering Sea: a stable thermocline and a subsurface “cold pool”. The objective of this study was to examine the influence of these features on foraging behavior by equipping fur seals from St. Paul Island (Alaska, USA) with time-depth recorders that also measured water temperature. Foraging bout variables (e.g., mean dive depth and percent time diving in a bout) were compared with respect to subsurface thermal characteristics (thermocline presence and strength and cold pool presence). Over 74% of bouts occurred in association with strong thermoclines (temperature change > 5°C). Few differences were found for dive behavior in relation to the presence of a thermocline and the cold pool, but for epipelagic bouts, a strong thermocline resulted in increased bottom times, number of dive wiggles, and percent time diving when compared to moderate thermoclines. There was also a positive relationship between mean dive depth and thermocline depth. The combination of increasing foraging effort in areas with strong thermoclines and diving to depths closely related to the thermocline indicates this feature is important foraging habitat for northern fur seals and may act to concentrate prey and increase foraging success. By recognizing the environmental features northern fur seals use to find prey, managers will be better equipped to identify and protect foraging habitat that is important to northern fur seals, and possibly other marine predators in the Bering Sea.     

An individual-oriented model for ecological risk assessment of wading birds

The contribution of certain contaminants to reproductive failure in many avian species has been an ongoing concern. Appropriate quantitative techniques have focused either on the individual organisms by providing explicit bioaccumulation dynamics or on whole ecosystems by looking at the fate of the contaminant but fail to make the necessary link via population dynamics of interacting individuals. We used the individual-oriented approach in an effort to quantify effects of chronic contaminant exposure on individual birds. This was made possible by the use of an object-oriented model, where individual birds are interacting objects, and their actions are implemented by passing to them appropriate messages. Using this modeling approach a breeding colony of Great Blue Herons (Ardea herodias) is simulated as an assemblage of interacting individuals whose daily actions (foraging, growth, feeding of the young) are simultaneously followed over short time intervals for a nesting season. Spatial distribution of the contaminants in prey resources is used on a cell by cell basis and their effects on certain behavior characteristics of adult birds (e.g. foraging efficiency, effects on flying efficiency, parental care) are taken into account. Results showed that sublethal effects could have a considerable effect on colony success. Appropriate selection of endpoints for risk assessment yields a variety of scenarios for colony success.     

Search behavior of the checker-throated antwren foraging in aerial leaf litter

Summary Checker-throated antwrens (Formicariidae: Myrmotherula fulviventris) live in lowland neotropical forests and forage from dead curled leaves in the understory. Because they search each leaf individually they provide an opportunity to study the use of potential visual cues by an insectivorous bird. Long and highly curled leaves contain the most arthropods and checker-throated antwrens were more successful when foraging at those leaves. Yet, they used leaves at random with respect to these potential cues. Antwrens spent longer searching for arthropods in each highly curled leaf than in less curled leaves. Because of this additional search time, prey capture success per unit foraging time was only slightly greater for highly curled leaves than at the average dead leaf in the aerial leaf litter. Thus, the cues that antwrens could use to locate richer leaves are those features that obscure the prey from avian predators. Unlike other foraging systems, the antwrens appear to have no reliable cues indicating more profitable foraging sites.Address for correspondence     

Diurnal and nocturnal feeding rate in Kentish plovers <Emphasis Type="Italic">Charadrius alexandrinus</Emphasis> on an intertidal flat as recorded by telescopic video systems

To elucidate the effects of light, site, temperature, time after emersion, and wind speed on foraging attempt rate (FAR) (attempts/unit time) and feeding success (FS) (captures/attempts) in the obligate visual foraging shorebird, Kentish plovers Charadrius alexandrinus, field observations were performed at a sandflat in Tokyo Bay, using a visible-light telescope camcorder during the day and a thermal infrared telescope camcorder at night. The re-analysis capability and frame-step replay of highly magnified video-images can contribute to accurate measurement of feeding behaviour over conventional focal observation techniques. Kentish plovers increased both FAR and FS in areas of high prey (polychaetes and crabs) density, resulting in a synergistic increase in feeding rate (captures/unit time). In areas of high prey abundance, FAR was higher at lower wind speed. FS increased with increasing time after emersion. Increasing temperatures induced a positive effect on FAR but a negative effect on FS. The effect of light on FAR was not observed; however, time-to-defecation occurrence was 2.2-fold shorter at night than during the day, indicating that feeding rate and FS are higher at night. These results are attributable to an increase in availability of cues due to higher nocturnal activity in polychaetes. Since available foraging time (emersion time) at night was 1.7-fold longer than during the day in the present study period, Kentish plovers could capture 3.7-fold (2.2 × 1.7) more prey at night than during the day. Results from this study indicate that nocturnal feeding in overwintering plovers is not a compensation but a major foraging activity to meet their energy requirements. An erratum to this article can be found at     

Cryptic prey colouration increases search time in brown trout (<Emphasis Type="Italic">Salmo trutta</Emphasis>): effects of learning and body size

Little is known about how cryptic colouration influences prey search in near-surface aquatic habitats, although such knowledge is critical for understanding the adaptive value of colour crypsis as well as the perceptive constraints influencing foraging behaviour in these environments. This study had two main aims: (1) to investigate how background colour matching by prey affects foraging efficiency by brown trout parr and (2) to investigate how foraging ability on cryptic and conspicuous prey is affected by fish size at age (reflecting dominance). We addressed these questions by training wild brown trout parr to forage individually on live brown-coloured maggots on a cryptic (brown) or conspicuous (green) background. A separate experiment confirmed the absence of trout preference for brown or green substrate. The results show that prey background colour matching increases search time in brown trout. Search time generally decreased by learning, but conspicuous prey remained an easier prey to find throughout the six training trials. Thus, perceptive constraints appear to limit search efficiency for cryptic prey, suggesting that cryptic colouration can confer survival benefits to prey in natural environments. Smaller fish generally found conspicuous prey faster than larger individuals, whereas search time for cryptic prey was not influenced by body size. This suggests that smaller individuals compensate for inferior competitive ability by increasing foraging activity rather than improving cognitive ability. The technique of varying cognitive demands in behavioural tasks could be used more in future studies aimed at distinguishing motivational effects from cognitive explanations for variation in behavioural performance.