Wolf spider predator avoidance tactics and survival in the presence of diet-associated predator cues (Araneae: Lycosidae)

Some prey can distinguish between chemical cues from predators fed different diets. Here we document the first evidence of diet-based chemical discrimination of predators in a terrestrial arthropod and measure the survival value of behavioural responses to predator chemical cues. We tested activity level and avoidance behaviour of the wolf spider, Pardosa milvina, to faeces and silk associated with the predatory wolf spider, Hogna helluo, fed either P. milvina or crickets (Acheta domesticus). We then measured survival of Pardosa in the presence of Hogna when placed on blank paper or paper previously occupied by Hogna fed either crickets or Pardosa. Filter paper previously occupied by Hogna from each diet treatment or a blank control were simultaneously presented to adult female Pardosa among four treatment pairs (N=15/treatment): (1) blank paper/blank paper, (2) Hogna fed crickets/blank, (3) Hogna fed Pardosa /blank and (4) Hogna fed Pardosa / Hogna fed crickets. Cues from Hogna fed either crickets or Pardosa elicited significantly less activity relative to blank controls. Cues from Hogna fed Pardosa elicited a significantly greater reduction in activity than Hogna fed crickets. When given a choice, Pardosa initially chose the blank substrate significantly more often than either substrate with Hogna cues. Spiders survived longer in the presence of cues from either Hogna diet treatment relative to blank paper, but there was no significant effect of predator diet on survival. Results suggest diet-based predator cues elicit different levels of activity in Pardosa that reduce predation in the presence of Hogna. Copyright 2001 The Association for the Study of Animal Behaviour.     

Knowing the Risk: Crickets Distinguish between Spider Predators of Different Size and Commonness

Predators unintentionally release chemical and other cues into their environment that can be used by prey to assess predator presence. Prey organisms can therefore perform specific antipredator behavior to reduce predation risk, which can strongly shape the outcome of trophic interactions. In contrast to aquatic systems, studies on cue‐driven antipredator behavior in terrestrial arthropods cover only few species to date. Here, we investigated occurrence and strength of antipredator behavior of the wood cricket Nemobius sylvestris toward cues of 14 syntopic spider species that are potential predators of wood crickets. We used two different behavioral arena experiments to investigate the influence of predator cues on wood cricket mobility. We further tested whether changes in wood cricket mobility can be explained by five predator‐specific traits: hunting mode, commonness, diurnal activity, predator–prey body–size ratio, and predator–prey life stage differences. Crickets were singly recorded (1) in separate arenas, either in presence or absence of spider cues, to analyze changes in mobility on filter paper covered with cues compared with normal mobility on filter paper without cues; and (2) in subdivided arenas partly covered with spider cues, where the crickets could choose between cue‐bearing and cue‐less areas to analyze differences in residence time and mobility when crickets are able to avoid cues. Crickets either increased or reduced their mobility in the presence of spider cues. In the experiments with cues and controls in separate arenas, the magnitude of behavioral change increased significantly with increasing predator–prey body size ratio. When crickets could choose between spider cues and control, their mobility was significantly higher in the presence of cues from common spider species than from rare spiders. We therefore conclude that wood crickets distinguish between cues from different predator species and that spiders unintentionally release a species‐specific composition and size‐dependent quantity of cues, which lead to distinct antipredator behavior in wood crickets.     

Chemical‐Mediated Predator Avoidance in the European House Cricket (Acheta domesticus) is Modulated by Predator Diet

Chemical information often mediates interactions between predators and prey, and threat‐sensitivity theory includes predictions that prey species should respond to chemical signatures of predators in a manner that is commensurate with the level of the assessed threat. Using the European house cricket (Acheta domesticus), we explored the influence of diet‐derived cues from the centipede Scolopocryptops sexspinosus on anti‐predator behavior in three laboratory experiments. In experiment 1, we compared the amount of time that adult female crickets spent on untreated filter paper and filter paper exposed to centipedes fed either the larvae of Hermetia illucens (black soldier fly), crickets, or a mixture of fly larvae and crickets. We discovered that crickets spent significantly less time on filter paper exposed to centipedes fed crickets only or a mixture of crickets and fly larvae compared with blank filter paper or filter paper exposed to centipedes fed fly larvae only. In our second experiment, we compared the amount of time that crickets spent on blank filter paper and filter paper exposed to adult female conspecifics to rule out the possibility that crickets simply avoid all filter paper exposed to metabolic by‐products, and crickets exhibited no discrimination. In our third experiment, we tested the potential effects of diet order on anti‐predatory behaviors. Specifically, we compared the amount of time that adult female crickets spent on filter paper exposed to centipedes fed fly larvae followed by crickets and filter paper exposed to centipedes fed crickets followed by fly larvae. We discovered no diet sequence effect. Our study demonstrates that European house crickets are sensitive to the chemical cues of their centipede predators, but only when centipedes have fed upon crickets.     

Tradeoffs involved in site selection and foraging in a wolf spider: effects of substrate structure and predation risk

Understanding how animals weigh habitat features, exposure to predators and access to resources is important to determining their life history and distribution across the landscape. For example, when predators accumulate in structurally complex habitats, they face an environment with different competitive interactions, foraging opportunities and predatory risks. The wolf spider Pardosa milvina inhabits the soil surface of highly disturbed habitats such as agricultural fields throughout eastern North America. Pardosa displays effective antipredator behavior in the presence of chemical cues produced by a larger coexisting wolf spider, Hogna helluo . We used those cues to simulate predation risk in laboratory and field experiments designed to test the effects of habitat substrate and predation risk on site selection and prey consumption of Pardosa . In general, Pardosa preferred more complex substrates over bare dirt but those preferences were eliminated or reversed when cues from Hogna were present. Feeding trials revealed that substrate alone had few effects on Pardosa prey consumption, which we measured by documenting the change in the abdomen width. Although the presence of Hogna cues reduced prey consumption overall in field feeding trials, the negative effect of predation risk on prey consumption was only observed in grass and bare dirt substrates in the laboratory. We also found that prey capture was negatively affected by habitat complexity for both spider species but that same complexity offered Pardosa protection from predation by Hogna. This study provides insight into how two predator species interact to balance site selection and feeding in order to avoid predation. Shifts in foraging and distributional patterns of predators can have profound implications for their role in the food web.     

Presampling sensory information and prey density assessment by wolf spiders (Araneae, Lycosidae)

Decisions regarding foraging patch residence time and the assessmentof patch quality may be mediated by various sources of information.This study examined the use of sensory cues by hunting spidersto assess prey density in the absence of prey capture. Adultfemale wolf spiders [Schizocosa ocreata (Hentz); Lycosidae]had food withheld for 4 days and then were exposed to artificialforaging patches containing four densities of crickets (0, 3,10, 20) with different sensory stimuli (visual and vibratoryinformation, visual only, and vibratory only). The spiders werenot allowed to feed during trials, and patch residence timewas recorded. The spiders varied patch residence time basedon sensory cues alone and spent more time in patches with higherprey density. With visual information only, spiders could apparentlydistinguish among prey densities almost as well as with visualand vibratory cues combined, but residence time did not differamong prey densities when only vibratory information was presented.Measurements of vibration levels produced by cricket activityunder experimental sensory treatments conform to test results,suggesting that visual detection of crickets is important inpatch assessment used in determining patch residence time.     

Hunger effects on foraging responses to perceptual cues in immature and adult wolf spiders (Lycosidae)

The wolf spider, Schizocosa ocreata (Hentz), varies foraging patch residence time in the presence of different sensory cues from prey, even without food rewards. This study examines the influence and interaction of hunger state, age and sex on the use of different types of sensory information to determine foraging patch sampling duration. In a series of two-chambered artificial foraging patches, I tested 26 S. ocreata once as immatures, and again as adults, under two hunger states (satiated and 7 days without food). Patches varied in the type of sensory information provided by live prey (crickets) as follows: visual cues alone; vibratory cues alone; combined visual/vibratory cues; and control (no prey). Without feeding in patches, the type of sensory stimuli available from prey strongly affected patch residence time, with spiders using primarily visual rather than vibratory cues. Hunger level as a main effect had no influence on residence time, but hunger state did mediate the importance of visual or vibratory information. Significant age- and sex-related differences in patch residence time in the presence of different sensory cues were found.These data suggest that ontogenetic and sex-specific foraging strategies are influenced by use of prefeeding perceptual cues rather than hunger state in wolf spiders. Copyright 1999 The Association for the Study of Animal Behaviour.     

Spider cues stimulate feeding,weight gain and survival of crickets

1. To avoid predation, prey often change their behaviour upon encountering cues of predator presence. Such behavioural changes should enhance individual survival, but are likely to be energy‐demanding. This should deplete energy reserves of the prey, unless it increases food intake. 2. These hypotheses were studied by conducting two microcosm experiments. In the first, crickets were kept on plants previously occupied by a spider or on control plants. After 3 days leaf consumption and weight gain of the crickets were quantified. In the second experiment, crickets were kept in the presence or absence of spider cues for 3 days. Spiders were then added and predation of the crickets was recorded during 24 h. 3. Crickets that had previously experienced spider cues were more successful in avoiding predation. Moreover, crickets under predation risk tended to increase foraging in the first microcosm experiment and gained more weight in the second microcosm experiment. 4. The results demonstrate that previous experience of predator cues decreases predation rate. Furthermore, they suggest that crickets are able to compensate for increased energy demands caused by antipredator behaviour. In more natural situations, moving to cue‐free plants may play an addition role.     

Effects of Spider Chemotactile Cues on Arthropod Behavior

Predation risk can strongly affect the behavior of prey species. However, empirical evidence for changes in behavior driven by spider cues is restricted to relatively few taxa. Here, we conducted a series of behavioral experiments to test for changes in activity among a wide range of terrestrial arthropods. We confronted 13 insect and eight spider species with chemotactile cues of three spider species. We applied two different experimental setups: In the ‘no-choice experiment’ prey individuals were either put on control filter papers or on filter papers previously occupied by a spider. In the ‘choice experiment’, the prey individuals were able to choose between filter paper halves with and without spider cues. In both setups, the response to spider cues depended significantly on prey species, with some species increasing and others decreasing their activity. Surprisingly few prey species responded to the spider cues at all. Our results indicate that predator recognition upon contact with cue bearing filter papers is strongly prey-specific and that behavioral effects driven by spider chemotactile cues are an exception rather than the rule among terrestrial arthropods.     

Two ecologically-divergent generalist predators have different responses to landscape fragmentation

We conducted three replicated field experiments to test the population response of two ecologically-divergent wolf spider species ( Hogna helluo and Pardosa milvina ) to three correlates of landscape fragmentation: area reduction, spatial subdivision, and increased edge to core ratio. We selected these two species because they differ in vagility and habitat selectivity. Hogna helluo is relatively large, averse to disturbed substrata, and has poor colonization abilities. Conversely, Pardosa is small, vagile, and will use barren, disturbed areas. In a test for the effect of area reduction on populations of the two wolf spiders, we destroyed 0%, 20% or 80% of randomly selected habitat islands in replicated experimental landscapes. We found that population densities of Hogna declined significantly, even at the lowest level of area reduction (20%), and that there was an increase in numbers of Pardosa . In a test for the response to an increase in landscape subdivision, we created four levels of habitat fragmentation in replicate plots. We found a significant decline in Hogna populations with increasing fragmentation. Pardosa populations did not respond to the fragmentation. In the third experiment we kept landscape area and subdivision constant, but manipulated the edge-to-core ratio. We found that populations of Hogna declined sharply with increasing edge, and that populations of Pardosa did not respond. These two syntopic wolf spiders have distinctly different responses to landscape fragmentation.     

Food imprinting and subsequent prey preference in the lynx spider,Oxyopes salticus (Araneae: Oxyopidae)

This object of this study was to determine if early feeding experience can affect subsequent prey choice in lynx spiderlings (Oxyopes salticus), and if so, is it evidence for food/olfactory imprinting. After emergence from the egg sac, three groups of 10 spiderlings were each fed exclusively for a 1-week period on one of three naturally-occurring prey species (crickets): group 1 fed on nymphs of Gryllotalpa hexadactyla; group 2 (Dialeurodes citrifoli); group 3 (Microcentrum rhombifolium). Following this, they were tested for subsequent prey preference in choice tests conducted in a plastic arena. Each spiderling was presented simultaneously with one individual of each prey species in a randomized design. Spiderlings exhibited a significant first preference for the original diet. Thus, experience with certain foods encountered by newly hatched spiderlings can affect subsequent prey preference in this species. Given the primacy of the early experience, the results are best interpreted in terms of imprinting and represent the first demonstration of olfactory imprinting in a spider.     

Habitat complexity dampens selection on prey activity level

Conspecific prey individuals often exhibit persistent differences in behavior (i.e., animal personality) and consequently vary in their susceptibility to predation. How this form of selection varies across environmental contexts is essential to predicting ecological and evolutionary dynamics, yet remains currently unresolved. Here, we use three separate predator–prey systems (sea star–snail, wolf spider–cricket, and jumping spider–cricket) to independently examine how habitat structural complexity influences the selection that predators impose on prey behavioral types. Prior to conducting staged predator–prey interaction encounters, we ran prey individuals through multiple behavioral assays to determine their average activity level. We then allowed individual predators to interact with groups of prey in either open or structurally complex habitats and recorded the number and individual identity of prey that were eaten. Habitat complexity had no effect on overall predation rates in any of the three predator–prey systems. Despite this, we detected a pervasive interaction between habitat structure and individual prey activity level in determining individual prey survival. In open habitats, all predators imposed strong selection on prey behavioral types: sea stars preferentially consumed sedentary snails, while spiders preferentially consumed active crickets. Habitat complexity dampened selection within all three systems, equalizing the predation risk that active and sedentary prey faced. These findings suggest a general effect of habitat complexity that reduces the importance of prey activity level in determining individual predation risk. We reason this occurs because activity level (i.e., movement) is paramount in determining risk within open environments, whereas in complex habitats, other behavioral traits (e.g., escape ability to a refuge) may take precedence.     

Predator-induced flow disturbances alert prey,from the onset of an attack

Many prey species, from soil arthropods to fish, perceive the approach of predators, allowing them to escape just in time. Thus, prey capture is as important to predators as prey finding. We extend an existing framework for understanding the conjoint trajectories of predator and prey after encounters, by estimating the ratio of predator attack and prey danger perception distances, and apply it to wolf spiders attacking wood crickets. Disturbances to air flow upstream from running spiders, which are sensed by crickets, were assessed by computational fluid dynamics with the finite-elements method for a much simplified spider model: body size, speed and ground effect were all required to obtain a faithful representation of the aerodynamic signature of the spider, with the legs making only a minor contribution. The relationship between attack speed and the maximal distance at which the cricket can perceive the danger is parabolic; it splits the space defined by these two variables into regions differing in their values for this ratio. For this biological interaction, the ratio is no greater than one, implying immediate perception of the danger, from the onset of attack. Particular attention should be paid to the ecomechanical aspects of interactions with such small ratio, because of the high degree of bidirectional coupling of the behaviour of the two protagonists. This conclusion applies to several other predator–prey systems with sensory ecologies based on flow sensing, in air and water.     

Age and Sex-Based Differences in the Use of Prey Sensory Cues in Wolf Spiders (Araneae: Lycosidae)

Differences in foraging patterns mediated by sensory cues were examined between adult and juvenile male and female wolf spiders (Schizocosa rovneri; Lycosidae). Patch residence time for thirty-one spiders were tested among juveniles and adults in artificial foraging patches. Patches varied in sensory information provided by live prey (crickets) as follows: visual stimuli alone; vibratory stimuli alone; visual and vibratory stimuli together; and control (no stimuli). Spiders moved between patches for one hour, but could not feed. Adult Schizocosa rovneri use primarily visual information to determine patch residence time, but juveniles use vibratory cues as well. Significant age and sex-based differences in the use of sensory cues suggest that observed divergent foraging strategies are partly due to the use of different perceptual cues in prey detection.     

The Nutritional Content of Prey Affects the Foraging of a Generalist Arthropod Predator

While foraging theory predicts that predatory responses should be determined by the energy content and size of prey, it is becoming increasingly clear that carnivores regulate their intake of specific nutrients. We tested the hypothesis that prey nutrient composition and predator nutritional history affects foraging intensity, consumption, and prey selection by the wolf spider, Pardosa milvina. By altering the rearing environment for fruit flies, Drosophila melanogaster, we produced high quality flies containing more nitrogen and protein and less lipid than low quality fruit flies. In one experiment, we quantified the proportion of flies taken and consumption across a range of densities of either high or low quality flies and, in a second experiment, we determined the prey capture and consumption of spiders that had been maintained on contrasting diets prior to testing. In both cases, the proportion of prey captured declined with increasing prey density, which characterizes the Type II functional response that is typical of wolf spiders. Spiders with similar nutritional histories killed similar numbers of each prey type but consumed more of the low quality prey. Spiders provided high quality prey in the weeks prior to testing killed more prey than those on the low quality diet but there was no effect of prior diet on consumption. In the third experiment, spiders were maintained on contrasting diets for three weeks and then allowed to select from a mixture of high and low quality prey. Interestingly, feeding history affected prey preferences: spiders that had been on a low quality diet showed no preference but those on the high quality diet selected high quality flies from the mixture. Our results suggest that, even when prey size and species identity are controlled, the nutritional experience of the predator as well as the specific content of the prey shapes predator-prey interactions.     

The Effect of Predator Hunger on Chemically Mediated Antipredator Responses and Survival in the Wolf Spider Pardosa milvina (Araneae: Lycosidae)

The wolf spider Pardosa milvina exhibits effective antipredator behavior (reduced movement) in the presence of silk and excreta from a larger co‐occurring predatory wolf spider Hogna helluo. As the quantity and quality of the silk and excreta may vary with the hunger state of the predator, we tested if chemical cues from a hungry vs. satiated H. helluo affected the activity level and survival of P. milvina. Pardosa milvina response was measured on substrates containing chemical cues from (1) a satiated H. helluo, (2) a H. helluo that had been withheld food for 2 wk, (3) cues from adult domestic crickets (Acheta domesticus), or (4) a blank test substrate (20 replicates/treatment). Pardosa milvina activity level was recorded on each substrate over a 30‐min period using video‐tracking equipment (Videomex‐I^®; Columbus Instruments, Columbus, OH, USA). We then measured P. milvina survival in the presence of hungry or satiated H. helluo on cues from a hungry H. helluo, satiated H. helluo, or a blank control (2 × 3 design). Pardosa milvina significantly reduced activity in the presence of H. helluo cues and showed significantly less activity in the presence of cues from a hungry H. helluo compared with a satiated one. Cue type and predator hunger state significantly affected P. milvina survival in the presence of live predators. However, cues from hungry vs. satiated H. helluo resulted in no difference in P. milvina survival. Pardosa milvina can discriminate between hungry vs. satiated predators based on silk and excreta cues alone, but differences in behavior as a result of this discrimination did not translate into increased survival in the presence of a live predator.     

Males make poor meals: a comparison of nutrient extraction during sexual cannibalism and predation

Cannibalism is hypothesized to have evolved as a way to obtain a high-quality meal. We examined the extraction of lipid and protein by female wolf spiders, Hogna helluo, during sexual cannibalism of males and predation of crickets. Most food-limited females did not cannibalize males but immediately consumed a size-matched cricket. When consuming male H. helluo and crickets, female H. helluo only consumed 51% of the male body while they consumed 72% of the cricket body. While males had higher protein content in their bodies than crickets and other insects, female H. helluo ingested similar amounts of protein from male H. helluo and crickets. Female H. helluo extracted 47% of the protein present in male H. helluo and 67% of the protein present in crickets. Females were able to extract nearly all of the lipid present in male H. helluo and crickets. However, crickets and other insects had almost 4 times higher lipid content than male H. helluo. The ratio of lipid to protein consumed from crickets appeared more similar to the nutritional requirements of egg production than that of males. Taken together, female hesitancy to engage in cannibalism, low extraction of nutrients from males and a low ratio of lipid to protein in the food extracted from males suggest that males may be poor-quality prey items compared to common insects such as crickets.     

Are you what you eat? A highly transient and prey‐influenced gut microbiome in the grey house spider Badumna longinqua

Stable core microbial communities have been described in numerous animal species and are commonly associated with fitness benefits for their hosts. Recent research, however, highlights examples of species whose microbiota are transient and environmentally derived. Here, we test the effect of diet on gut microbial community assembly in the spider Badumna longinqua. Using 16S rRNA gene amplicon sequencing combined with quantitative PCR, we analyzed diversity and abundance of the spider's gut microbes, and simultaneously characterized its prey communities using nuclear rRNA markers. We found a clear correlation between community similarity of the spider's insect prey and gut microbial DNA, suggesting that microbiome assembly is primarily diet‐driven. This assumption is supported by a feeding experiment, in which two types of prey—crickets and fruit flies—both substantially altered microbial diversity and community similarity between spiders, but did so in different ways. After cricket consumption, numerous cricket‐derived microbes appeared in the spider's gut, resulting in a rapid homogenization of microbial communities among spiders. In contrast, few prey‐associated bacteria were detected after consumption of fruit flies; instead, the microbial community was remodelled by environmentally sourced microbes, or abundance shifts of rare taxa in the spider's gut. The reshaping of the microbiota by both prey taxa mimicked a stable core microbiome in the spiders for several weeks post feeding. Our results suggest that the spider's gut microbiome undergoes pronounced temporal fluctuations, that its assembly is dictated by the consumed prey, and that different prey taxa may remodel the microbiota in drastically different ways.     

Predators buffer the effects of variation in prey nutrient content for nutrient deposition

Predator feeding behavior and digestion regulate the flow of nutrients through ecosystems by determining the fate of prey nutrients. Most predators feed on a diversity of prey items, which differ widely in traits including their nutrient content. Yet, relatively little is known of the mechanisms through which variation in prey nutrient content affects the form by which nutrients are deposited into the environment. The overall goal of this study was to test how variation in the nutrient content of prey affected the fate of nutrients following predation by an arthropod carnivore, the Carolina wolf spider Hogna carolinensis. We manipulated the macronutrient content of prey by varying the diet on which crickets were fed to produce prey treatments that differed in lipid and protein content. Nutrients were measured as both macronutrients and elements in prey and elements in excreta. We found that there was no effect of diet treatment on the amounts of elements or macronutrients in prey carcasses and excreta despite significant variation in the nutrient content of those prey. This is in contrast to studies of some aquatic systems where mass balance by consumers results in variation in excreta content depending on the nutrient content of food. Wolf spiders assimilated the majority of prey nutrients and deposited relatively small and similar amounts of nutrients following feeding. Hence, while prey can vary widely in nutrient content, our findings suggest that this variation has little effect on the amounts of nutrients deposited by predators.     

Predicting Predatory Outcomes in the Context of Carryover Effects: Interactions between Juvenile Frogs and Spider Predators

As representatives of organisms with complex life histories, frogs provide an ideal system to study predator‐induced carryover effects: how the risk of predation in one life stage can impact predator–prey interactions in a later stage. Invertebrate predation on frogs has been widely reported, although studies of the behavioral mechanisms underlying their interactions in the terrestrial stage have been lacking. We made detailed observations of interactions between a wolf spider (Tigrosa helluo) and Blanchard's cricket frog (Acris blanchardi) to determine factors that predict capture success and to evaluate potential carryover effects from aquatic predation risk. Juvenile frogs, reared with or without dragonfly predator cues, were placed in an arena with or without spider cues and allowed to interact with a spider. Spiders captured frogs, and an interaction between frog size and activity predicted frog survival. We found no evidence that either aquatic or terrestrial cues altered frog behavior or survival. By preying upon a demographically important life stage, spiders may contribute to population dynamics in frogs.     

Chemical Mediation of Prey Recognition by Spider-Hunting Wasps

Predator–prey interactions are important in maintaining the structure and dynamics of ecological communities. Both predators and prey use cues from a range of sensory modalities to detect and assess one another; identification of these cues is necessary to understand how selection operates to shape predator–prey interactions. Mud-dauber wasps (Sphecidae) provision their larval nests with paralyzed spiders, and different genera of wasps specialize on particular spider taxa. Sceliphron caementarium (Drury 1773) wasps preferentially capture spiders that build two-dimensional (2D) webs, rather than those that construct three-dimensional (3D) webs, but the basis of this preference is not clear. Wasps may choose spiders based on an assessment of their web architecture, as 3D webs may provide better defenses against wasp predation than do 2D webs. However, because many hymenopterans use chemical cues to locate and recognize prey, it is also possible that mud-dauber wasps rely on chemical cues associated with the spider and/or the web to assess prey suitability. When we offered foraging S. caementarium wasps 2D and 3D spiders both on and off their webs, we found that in both cases the wasps took 2D spiders and avoided 3D spiders, demonstrating that the web itself is not the impediment. Results of a series of behavioral choice assays involving filter paper discs containing spider cues and chemically manipulated spiders or spider dummies corroborated the importance of spider chemical cues in mediation of prey recognition by mud-dauber wasps. We also discuss the relative importance of visual and chemical cues for prey recognition by wasps, examine the anti-predator behaviors of 2D and 3D spiders, and consider the role of wasp predation in spider diversification.