Predator perception of detritus and eggsac decorations spun by orb-web spiders Cyclosa octotuberculata: Do they function to camouflage the sniders?

Camouflage is one of the most widespread and powerful strategies that animals use to make detection/recognition more difficult. Many orb-web spiders of the genus Cyclosa add prey remains, plant debris, moults, and/or eggsacs to their webs called web decorations. Web decorations resembling spider body colour pattern have been considered to camouflage the spider from predators. While this camouflage is obvious from a human's perspective, it has rarely been investigated from a predator's perspective. In this study, we tested the visibility of web decorations by calculating chromatic and achromatic contrasts of detritus and eggsac decorations built by Cyclosa octotuberculata, against four different backgrounds viewed by both bird (e.g., blue tits) and hymenopteran (e.g. Wasps) predators. We showed that both juvenile and adult spiders on webs with detritus or egg-sac deco-rations were undetectable by both hymenopteran and bird predators over short and long distances. Our results thus suggest that decorating webs with detritus or eggsacs by C. Octotuberculata may camouflage the spiders from both hymenopteran and bird predators in their common habitats.     

The multiple disguises of spiders: web colour and decorations, body colour and movement

Diverse functions have been assigned to the visual appearance of webs, spiders and web decorations, including prey attraction, predator deterrence and camouflage. Here, we review the pertinent literature, focusing on potential camouflage and mimicry. Webs are often difficult to detect in a heterogeneous visual environment. Static and dynamic web distortions are used to escape visual detection by prey, although particular silk may also attract prey. Recent work using physiological models of vision taking into account visual environments rarely supports the hypothesis of spider camouflage by decorations, but most often the prey attraction and predator confusion hypotheses. Similarly, visual modelling shows that spider coloration is effective in attracting prey but not in conveying camouflage. Camouflage through colour change might be used by particular crab spiders to hide from predator or prey on flowers of different coloration. However, results obtained on a non-cryptic crab spider suggest that an alternative function of pigmentation may be to avoid UV photodamage through the transparent cuticle. Numerous species are clearly efficient locomotory mimics of ants, particularly in the eyes of their predators. We close our paper by highlighting gaps in our knowledge.     

Spider movement, UV reflectance and size, but not spider crypsis, affect the response of honeybees to Australian crab spiders

According to the crypsis hypothesis, the ability of female crab spiders to change body colour and match the colour of flowers has been selected because flower visitors are less likely to detect spiders that match the colour of the flowers used as hunting platform. However, recent findings suggest that spider crypsis plays a minor role in predator detection and some studies even showed that pollinators can become attracted to flowers harbouring Australian crab spider when the UV contrast between spider and flower increases. Here we studied the response of Apis mellifera honeybees to the presence of white or yellow Thomisus spectabilis Australian crab spiders sitting on Bidens alba inflorescences and also the response of honeybees to crab spiders that we made easily detectable painting blue their forelimbs or abdomen. To account for the visual systems of crab spider's prey, we measured the reflectance properties of the spiders and inflorescences used for the experiments. We found that honeybees did not respond to the degree of matching between spiders and inflorescences (either chromatic or achromatic contrast): they responded similarly to white and yellow spiders, to control and painted spiders. However spider UV reflection, spider size and spider movement determined honeybee behaviour: the probability that honeybees landed on spider-harbouring inflorescences was greatest when the spiders were large and had high UV reflectance or when spiders were small and reflected little UV, and honeybees were more likely to reject inflorescences if spiders moved as the bee approached the inflorescence. Our study suggests that only the large, but not the small Australian crab spiders deceive their preys by reflecting UV light, and highlights the importance of other cues that elicited an anti-predator response in honeybees.     

Predator hunting mode influences patterns of prey use from grazing and epigeic food webs

Multichannel omnivory by generalist predators, especially the use of both grazing and epigeic prey, has the potential to increase predator abundance and decrease herbivore populations. However, predator use of the epigeic web (soil surface detritus/microbe/algae consumers) varies considerably for reasons that are poorly understood. We therefore used a stable isotope approach to determine whether prey availability and predator hunting style (active hunting vs. passive web-building) impacted the degree of multichannel omnivory by the two most abundant predators on an intertidal salt marsh, both spiders. We found that carbon isotopic values of herbivores remained constant during the growing season, while values for epigeic feeders became dramatically more enriched such that values for the two webs converged in August. Carbon isotopic values for both spider species remained midway between the two webs as values for epigeic feeders shifted, indicating substantial use of prey from both food webs by both spider species. As the season progressed, prey abundance in the grazing food web increased while prey abundance in the epigeic web remained constant or declined. In response, prey consumption by the web-building spider shifted toward the grazing web to a much greater extent than did consumption by the hunting spider, possibly because passive web-capture is more responsive to changes in prey availability. Although both generalist predator species engaged in multichannel omnivory, hunting mode influenced the extent to which these predators used prey from the grazing and epigeic food webs, and could thereby influence the strength of trophic cascades in both food webs.     

Background evolution in camouflage systems: A predator-prey/pollinator-flower game

A common predator or anti-predator strategy involves camouflage based on background matching. In some systems, the background is an organism whose fitness is affected by the predator-prey interaction. In these cases, the phenotype of the background species may evolve to affect the degree of background matching in the predator-prey interaction. For example, some flower species (the background) are inhabited by camouflaged ambush predators that attack visiting pollinators. These flowers have a fitness interest in the outcome of the predator-prey interaction because flowers depend on pollinator visitations for reproduction. Therefore, floral colour might evolve relative to predator colour so as to influence the detectability of resident predators. I have created a three-player game, based on Signal Detection Theory, to model the co-evolution of predator and prey/pollinator behavioural strategies with floral colour. This model makes two general predictions: (1) Constraints on predator distributions favour the evolution of flowers that match the predators’ colour because they prevent predators from overexploiting these flowers; (2) factors that produce less discriminating pollinators also favour the evolution of flowers that match the predators’ colour because these pollinators are willing to land on these flowers even if the safety of the flower is in doubt.     

Camouflage and colour change: antipredator responses to bird and snake predators across multiple populations in a dwarf chameleon

Potential prey are often exposed to multiple predators that vary in their foraging tactics and ability to detect prey. For animals that rely on crypsis to avoid predators, one solution is to alter their behaviour or appearance to maximize crypsis in ways that are specific to different types of predator. We tested whether dwarf chameleons ( Bradypodion transvaalense ) showed different behavioural responses, including colour change, towards multiple predators (bird and snake models) that detect and capture prey in different ways, and whether these antipredator responses varied geographically. Chameleons consistently used the same body postures (lateral compression and flipping to the opposite side of the branch) and displayed similar chromatic (colour) contrast against the natural background in response to both predator types. However, they became significantly more achromatically contrasting (brighter) in the presence of the snake compared to the bird. This relative difference in achromatic contrast towards the two types of predator was consistent among populations. There were also significant differences in both absolute achromatic and chromatic contrast among populations despite very similar light environment, background coloration and habitat structure. Our results highlight facultative crypsis as one type of flexible antipredator tactic and emphasize the importance of visual ecology in understanding prey–predator interactions.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 437–446.     

The effectiveness of post-contact defenses in a prey with no pre-contact detection

Most empirical and theoretical papers on prey–predator interactions are for animals with long-range detection, animals that can detect and react to predators long before these touch the prey. Heavy-bodied and chemically defended harvestmen (Arachnida, Opiliones) are an exception to this general pattern and rely on contact to detect arthropod predators. We examined the interactions between the Brazilian wandering spider Ctenus ornatus with harvestmen (Mischonyx cuspidatus) or control prey (Gryllus sp. and M. cuspidatus immature, both with soft integuments). Considering a prey–predator system in which fleeing from or reacting to a predator at a distance is not possible, we predicted both a high survival value of near-range defense mechanisms and that mortality would be higher in the absence of such defense mechanisms. We also expected the predator to behave differently when interacting with harvestmen or with a control prey without such defense mechanisms. Our results from laboratory experiments partially matched our predictions: First of all, histological sections showed that the integument of adult harvestmen is thicker than that of immature harvestmen and that of crickets. Adult harvestmen were less preyed upon than the control prey; the heavy armature increases the survival rate but the secretions from the scent glands do not. The predator did behave differently when attacking harvestmen compared to crickets. Despite the large size difference between predator and harvestmen, the protection provided by the armature allowed some of the harvestmen to survive encounters without pre-contact detection, thus greatly reducing the reliance on long-range detection to survive encounters with predators. Harvestmen call for theoretical and empirical work on prey–predator interactions that take into account the possibility that prey may not detect the predator before contact is established.     

Attraction of a predator to chemical information related to nonprey: when can it be adaptive?

Information specificity can be important to animals in makingoptimal decisions. However, it is not always necessary to useevery level of specificity. We analyzed the response of thepredatory mite Phytoseiulus persimilis to plant-produced informationrelated to a nonprey herbivore. This predator is a specialistfeeding on spider mites in the genus Tetranychus. Caterpillarsof Spodoptera exigua cannot serve as prey. Plants respond toan infestation by herbivores with the emission of volatilesthat attract carnivorous enemies of the herbivores. Conspecific plants infested with different herbivore species can emit blendsthat are qualitatively identical, while differing in the ratiosof blend components. However, different plant species emitvolatile blends that differ qualitatively. We demonstratedthat the predator P. persimilis is attracted to volatiles frombean plants infested with S. exigua caterpillars, but thatthis attraction is affected by predator starvation and host-plantexperience. One-hour and 24-h starved predators were made to represent predators that just lost a prey patch versus predatorsthat have totally lost a prey patch. Predators reared on spidermites on bean were attracted to bean plants infested with caterpillarswhen starved for 1 h but not when starved for 24 h. Both predatorgroups were attracted to bean plants infested with prey (i.e.,spider mites). One-hour starved predators can use the odorto relocate the rewarding prey patch they just lost contactwith, and using a general olfactory representation of the blendis sufficient for relocation. In contrast, for 24-h starvedpredators, the perception of a plant's odor blend is unlikelyto represent the prey patch lost, and discriminating betweenan odor blend representing prey or nonprey will avoid investingtime in finding a nonprey herbivore. In contrast, predatorsthat had been reared on spider mites on cucumber and thus hadexperienced a qualitatively different odor blend were not attractedto volatiles from caterpillar-infested bean plants. They wereattracted to spider mite-infested bean plants, irrespectiveof starvation level. To cucumber-experienced predators, theperception of bean plant odor cannot represent the prey patch lost, but only a new prey patch. Being discriminative and onlyresponding to prey-infested plants is adaptive in this situation.Our results are discussed in the context of optimal informationprocessing.     

Benefit by contrast: an experiment with live aposematic prey

Aposematic prey often have a coloration that contrasts withthe background. One beneficial effect of such conspicuous colorationis that it produces faster and more durable avoidance by predators.Another suggested benefit is that prey that contrast with thebackground are more quickly discerned and recognized as unpalatableby experienced predators. To further investigate the effectsof prey contrast on predator behavior, I conducted an experimentwith young chicks (Gallus gallus domesticus) as predators onlive aposematic and nonaposematic prey. Birds with prior experienceof both prey types were allowed into an arena with both palatableprey and aposematic prey on backgrounds that either closelymatched or contrasted with the coloration of the aposematicprey. Also, the time a bird had available to decide to attacka prey was manipulated by including a competing chick or not.The experienced birds showed greater attack latencies for aposematicprey on more contrasting backgrounds, and aposematic prey werealso attacked to a greater extent when on a matching background.The presence of a competitor generated similar effects, wherebirds in high competition attacked more and faster comparedto birds subjected to lower degree of competition, but therewas no interaction between competition and contrast. Thus,the experiment provides evidence that prey contrast againstthe background may produce better recognition and avoidance,independently of predator viewing time.     

Ineffective crypsis in a crab spider: a prey community perspective

Cryptic coloration is assumed to be beneficial to predators because of an increased encounter rate with unwary prey. This hypothesis is, however, very rarely, if ever, studied in the field. The aim of this study was to quantify the encounter rate and capture success of an ambush predator, in the field, as a function of its level of colour-matching with the background. We used the crab spider Misumena vatia, which varies its body colour and can thereby match the colour of the flower it hunts upon. We carried out a manipulative field experiment using a complete factorial design resulting in six different colour combinations of crab spiders and flowers differing in their degree of colour-matching. A rich and diverse set of naturally occurring insects visited the flowers while we continuously video-recorded the spider''s foraging activity. This enabled us to test the crypsis, the spider avoidance and the flower visitor attraction hypotheses, all three supported by previous studies. Flower visitors of different groups either avoided crab spiders independent of colour-matching, such as solitary bees and syrphid flies, or ignored them, such as bumble-bees and honeybees. Moreover, colour-matched spiders did not have a higher encounter rate and capture success compared to the visually apparent ones. Thus, our results support the spider avoidance hypothesis, reject the two other hypotheses and uncovered a fourth behaviour: indifference to predators. Because flower visitors reacted differently, a community approach is mandatory in order to understand the function of background colour-matching in generalist predators. We discuss our results in relation to the size and sociality of the prey and in relation to the functional significance of colour change in this predator.     

The effects of predation risk from crab spiders on bee foraging behavior

Recent studies have suggested that top–down effects ofpredation on plant–pollinator interactions may not be,as previously thought, rare and/or weak. In this paper, we explorethe effects of crab spiders (Araneae: Thomisidae) on the behaviorof 2 species of bee (Hymenoptera: Apidae) foraging for nectarand pollen on 3 different plant species in central Portugal.In 2 experiments, we found that the eusocial bee Apis melliferawas significantly less likely to inspect and accept a floweror inflorescence if it harbored a spider. In contrast, we foundno such effects of spiders on the behavior of the solitary beeEucera notata. Further experiments showed that the effects ofenvironmental cues associated with predators on flower visitationby A. mellifera were detectable even when no spider was presentat the moment a flower was encountered. Such indirect effectswere only identified, however, in bees foraging on 1 of 2 plantspecies studied. In a final experiment, A. mellifera was shownto respond negatively to the presence of the corpses of conspecificsglued to flowers. This suggests that prey corpses left exposedon petals or bracts by spiders provide an obvious cue that beescan use to avoid predators. These results add to a growing bodyof evidence that plant–pollinator interactions are notimmune to the effects of predation and suggest that the strengthof such effects vary both between and within species.     

Hunting‐mediated predator facilitation and superadditive mortality in a European ungulate

Predator‐prey theory predicts that in the presence of multiple types of predators using a common prey, predator facilitation may result as a consequence of contrasting prey defense mechanisms, where reducing the risk from one predator increases the risk from the other. While predator facilitation is well established in natural predator‐prey systems, little attention has been paid to situations where human hunters compete with natural predators for the same prey. Here, we investigate hunting‐mediated predator facilitation in a hunter‐predator‐prey system. We found that hunter avoidance by roe deer (Capreolus capreolus) exposed them to increase predation risk by Eurasian lynx (Lynx lynx). Lynx responded by increasing their activity and predation on deer, providing evidence that superadditive hunting mortality may be occurring through predator facilitation. Our results reveal a new pathway through which human hunters, in their role as top predators, may affect species interactions at lower trophic levels and thus drive ecosystem processes.     

Camouflage in predators

Camouflage – adaptations that prevent detection and/or recognition – is a key example of evolution by natural selection, making it a primary focus in evolutionary ecology and animal behaviour. Most work has focused on camouflage as an anti‐predator adaptation. However, predators also display specific colours, patterns and behaviours that reduce visual detection or recognition to facilitate predation. To date, very little attention has been given to predatory camouflage strategies. Although many of the same principles of camouflage studied in prey translate to predators, differences between the two groups (in motility, relative size, and control over the time and place of predation attempts) may alter selection pressures for certain visual and behavioural traits. This makes many predatory camouflage techniques unique and rarely documented. Recently, new technologies have emerged that provide a greater opportunity to carry out research on natural predator–prey interactions. Here we review work on the camouflage strategies used by pursuit and ambush predators to evade detection and recognition by prey, as well as looking at how work on prey camouflage can be applied to predators in order to understand how and why specific predatory camouflage strategies may have evolved. We highlight that a shift is needed in camouflage research focus, as this field has comparatively neglected camouflage in predators, and offer suggestions for future work that would help to improve our understanding of camouflage.     

Fine-scale substrate use by a small sit-and-wait predator

Substrate choice is one of the most important decisions thatsit-and-wait predators must make. Not only may it dictate theprey available but also the cover for the predator which mayconceal it from prey or its own predators. However, while ona particular substrate the behavior and use of that substratemay vary widely. When naïve, newly emerged crab spiderlingsMisumena vatia (Thomisidae) occupied flowering goldenrod Solidagocanadensis, their behavior differed markedly on inflorescenceswith relatively sparse and densely packed flower heads as wellas on experimentally thinned and unthinned inflorescences. Initially,the spiderlings most often hunted at the thinned sites and hidamong the dense flower heads at the unthinned sites, a differencethat disappeared in all broods tested after 2–3 h, possiblybecause of the growing hunger of the initially concealed individuals.Prey capture (dance flies) in the thinned sites initially significantlyexceeded that in unthinned sites but subsequently did not differ.However, spiderlings encountered their principal predator, thejumping spider Pelegrina insignis, significantly more oftenon unthinned than thinned inflorescences. Even though usagepatterns initially differed strikingly, spiderlings did notdiffer in their rates of quitting the two types of sites. Theseresults suggest a trade-off between foraging and predator avoidancethat changes in response to increasing hunger over time.     

Indirect cues in selecting a hunting site in a sit‐and‐wait predator

Sit‐and‐wait predators use relatively simple rules for their decisions to choose and leave a patch, such as using the direct presence of prey to select a hunting site. However, the direct presence of prey can only be used when there is a highly visited patch in the proximity of the predator. Therefore, it is plausible that sit‐and‐wait predators also exploit indirect cues of prey presence and, consequently, use associative learning to select a hunting site. The present study tests for the role of associative learning in a sit‐and‐wait predator species for which the ecology is well understood: Misumena vatia Clerck crab spiders. An ecologically relevant scenario is used by selecting flower colour as the conditioned stimulus and prey presence as the unconditioned stimulus. The results provide no evidence that M. vatia crab spiders use the association between flower colour and food presence for selecting a hunting site. After a training phase of being exposed to a colourful artificial flower highly visited by bees, spiders select a hunting site independently of its colour during the testing phase. Investigations of similar scope and ecological relevance are required with other sit‐and‐wait predators to identify the conditions promoting the use of associative learning for foraging site selection when animals face an unpredictable food supply.     

Predator Detection is Limited in Microhabitats with High Light Intensity: An Experiment with Brown‐Headed Cowbirds

Variations in ambient light conditions across different microhabitats can modify the detectability of predators and prey. Prey have been shown to be more visible in sunlit than in shaded patches, leading to higher predation risk and more investment in vigilance (predation risk hypothesis). Additionally, prey have been hypothesized to take longer to detect predators in sunlit compared to shaded patches because of the excess of sunlight causing glare effects (disability glare hypothesis). We tested the predictions of these two non‐mutually exclusive hypotheses in a seminatural experiment with brown‐headed cowbirds by measuring vigilance behavior and detection of a ground predator in patches under the shade of vegetation and in the open. Light intensity and achromatic contrast were higher in the sunlit patches, which could enhance glare effects, but chromatic contrast was higher in the shaded patches. Brown‐headed cowbirds took longer to show alert reactions to and flee from a ground predator in sunlit compared to shaded patches. However, the two parameters associated with perceived predation risk (vigilance prior to the predator exposure and time to resume foraging after the attack) did not differ between sunlit and shaded patches. Our findings support to a greater extent the disability glare hypothesis than the predation risk hypothesis. Overall, ambient light conditions can affect two critical components of behavioral predator–prey interactions in terrestrial habitats: detection of and escape from predators. The effects of disability glare are expected to be more pronounced in bird species with wider visual fields or without sun‐shading structures; however, species may compensate through various behaviors (e.g. avoidance of sunlit patches and changes in head orientation).     

Multiple intraguild predators reduce mortality risk of a mutual agricultural pest prey in simple,but not in complex,experimental settings

Multiple predator species that coexist with each other and their mutual prey can have combined effects on prey mortality that are similar to the sum of each predator's individual impact (linear effects), greater than the sum of each predator's individual impact (risk enhancement), or less than the sum of each predator's individual impact (risk reduction). Understanding multiple predator effects is important to determine the impact of predators on pest prey in agroecosystems. If two predators share the same broad spatial domain and hunting mode and engage in intraguild predation, then their combination is expected to result in risk reduction for a mutual prey. We tested this hypothesis using both additive and replacement experimental designs on two species of generalist wolf spider predators (Tasmanicosa leuckartii and Hogna crispipes) that hunt in the same domain, and a mutual insect prey (cotton bollworm Helicoverpa armigera). We used two types of enclosures: a small simple laboratory enclosure, and a larger more complex cotton plant enclosure. We found that in the small simple laboratory enclosures, the presence of two spiders led to risk reduction of Helicoverpa larva mortality as expected, but in larger more complex cotton plant enclosures the presence of both species resulted in linear effects rather than risk reduction on Helicoverpa mortality. Furthermore, intraguild predation did not change multiple predator effects in laboratory or plant enclosures. This study has implications for managing arthropod predators in agroecosystems; contrary to predictions of ecological frameworks, coexistence of predators that share the same hunting mode and hunting domain may not lead to risk reduction on a mutual prey in more complex environments, where encounters among predators can be lower. Conservation of multiple predators of a single guild can play an essential role on biological control of insect pests.     

Predator interference alters foraging behavior of a generalist predatory arthropod

Interactions between predators foraging in the same patch may strongly influence patch use and functional response. In particular, there is continued interest in how the magnitude of mutual interference shapes predator–prey interactions. Studies commonly focus on either patch use or the functional response without attempting to link these important components of the foraging puzzle. Predictions from both theoretical frameworks suggest that predators should modify foraging efforts in response to changes in feeding rate, but this prediction has received little empirical attention. We study the linkage between patch departure rates and food consumption by the hunting spider, Pardosa milvina, using field enclosures in which prey and predator densities were manipulated. Additionally, the most appropriate functional response model was identified by fitting alternative functional response models to laboratory foraging data. Our results show that although prey availability was the most important determinant of patch departure rates, a greater proportion of predators left enclosures containing elevated predator abundance. Functional response parameter estimation revealed significant levels of interference among predators leading to lower feeding rates even when the area allocated for each predator was kept constant. These results suggest that feeding rates determine patch movement dynamics, where interference induces predators to search for foraging sites that balance the frequency of agonistic interactions with prey encounter rates.     

The re-emergence of felid camouflage with the decay of predator recognition in deer under relaxed selection

When a previously common predator disappears owing to local extinction, the strong source of natural selection on prey to visually recognize that predator becomes relaxed. At present, we do not know the extent to which recognition of a specific predator is generalized to similar looking predators or how a specific predator-recognition cue, such as coat pattern, degrades under prolonged relaxed selection. Using predator models, we show that deer exhibit a more rapid and stronger antipredator response to their current predator, the puma, than to a leopard displaying primitive rosettes similar to a locally extinct predator, an early jaguar. Presentation of a novel tiger with a striped coat engendered an intermediate speed of predator recognition and strength of antipredator behaviour. Responses to the leopard model slightly exceeded responses to a non-threatening deer model, suggesting that thousands of years of relaxed selection have led to the loss of recognition of the spotted coat as a jaguar-recognition cue, and that the spotted coat has regained its ability to camouflage the felid form. Our results shed light on the evolutionary arms race between adoption of camouflage to facilitate hunting and the ability of prey to quickly recognize predators by their formerly camouflaging patterns.     

The struggle for safety: effectiveness of caterpillar defenses against bird predation

The effectiveness of anti‐predator traits, such as warning signals and camouflage, has rarely been quantified from a phylogenetic community ecology perspective. Here we use a phylogenetic comparative analysis to test the association between several putative anti‐predator traits and bird predation risk in an assemblage of caterpillar species. We synthesize eight years of field and laboratory study of a temperate forest community, including a four‐year bird exclusion experiment that provided comparative measures of bird predation risk for 38 caterpillar species from a phylogenetic community. We then conducted a phylogenetic generalized least‐squares and information‐theoretic model selection analysis of warning signals (aposematism or mimicry), camouflage (crypsis or masquerade), and behavioral responses to physical attack as predictors of bird predation, while also accounting for putatively important effects of the abundance, mean body size, and phenology of caterpillar species. The most behaviorally specialized caterpillar species possessing warning signals experienced the lowest bird predation risk, supporting aposematism theory and highlighting the role of prey behavior in the visual signaling of predators. Among the camouflaged caterpillar species, those with the greatest latency to detection by human proxy predators experienced the lowest bird predation risk, supporting camouflage theory. Caterpillar behavioral responses to physical attack, however, predicted increased bird predation risk among camouflaged caterpillars. Although caterpillar abundance, body size, and phenology were expected to be important based on inference from optimal foraging theory and previous field observations, these factors had limited predictive power. This study provides methodologically unique evidence for the importance of morphological and behavioral components of primary, visual defenses of caterpillars against their avian predators in a natural community.