Effects of ant competition and bird predation on the spider assemblage of a citrus grove

Characterizing intraguild interactions is key to improving understanding of food webs because they are major forces in the structuring of communities. Spiders are generalist predators with intermediate positions in the food web that establish intraguild interactions with ants and birds, which respectively compete with and prey on them. Research has also found interactions between birds and ants, potentially resulting in non-additive effects of both groups on arthropod assemblages, although studies of their combined impacts with tests for multiple-predator effects are scarce. We thus aimed to discern the relative effect of ants and birds on the spider assemblage of a citrus grove. We used a split-plot design to factorially exclude these groups over 2 years, preventing ants reaching the canopies by placing sticky bands around tree trunks, and birds by enclosing groups of trees in cages. We sampled spiders from the canopies (beating) and the ground (pitfalls) every 3 months, and we identified them to species. We found a strong influence of ants on the canopy spider assemblage, mainly through a negative effect on the families Araneidae and Theridiidae. Since spiders’ weights from ant-excluded and control trees were similar, these results suggest interference competition of ants on spiders rather than competitive exploitation. Bird exclusion did not affect the spider assemblage, contrasting with other studies reporting a marked predatory pressure of birds on spiders; nor were there any non-additive effects of ants and birds. Our findings show that spider assemblages are not uniformly affected by intraguild competitors.     

Scorpions,spiders and solpugids: predation and competition among distantly related taxa

Summary Scorpions, spiders and solpugids are generalist predators on the same types of arthropod prey. However, these potential competitors also frequently eat one another (=intraguild predation). In a 29 mo. experiment, >6,000 scorpions were removed from 300 (10x10m) quadrats. Significantly more spiders (but not solpugids) occurred in removal versus control quadrats. Two alternate hypotheses potentially explain this result: exploitation competition for jointly exploited prey or intraguild predation. There was no evidence of exploitation competition: no differences existed between removal and control quadrats in prey abundance or spider size and reproductive characteristics. It appears that the release from predation pressure in areas from which scorpions were removed produced the observed increase in spider abundance. Current ecological theory does not fully apply to situations whereby species at the same trophic level interact as both predators and potential competitors.     

Resolving the predator first paradox: Arthropod predator food webs in pioneer sites of glacier forelands

Primary succession on bare ground surrounded by intact ecosystems is, during its first stages, characterized by predator‐dominated arthropod communities. However, little is known on what prey sustains these predators at the start of succession and which factors drive the structure of these food webs. As prey availability can be extremely patchy and episodic in pioneer stages, trophic networks might be highly variable. Moreover, the importance of allochthonous versus autochthonous food sources for these pioneer predators is mostly unknown. To answer these questions, the gut content of 1,832 arthropod predators, including four species of carabid beetles, two lycosid and several linyphiid spider species caught in early and late pioneer stages of three glacier forelands, was screened molecularly to track intraguild and extraguild trophic interactions among all major prey groups occurring in these systems. Two‐thirds of the 2,310 identified food detections were collembolans and intraguild prey, while one‐third were allochthonous flying insects. Predator identity and not successional stage or valley had by far the strongest impact on the trophic interaction patterns. Still, the variability of prey spectra increased significantly from early to late pioneer stage, as did the niche width of the predators. As such the structure of pioneer arthropod food webs in recently deglaciated Alpine habitats seems to be driven foremost by predator identity while site and early successional effects contribute to a lesser extent to food web variability. Our findings also suggest that in these pioneer sites, predatory arthropods depend less on allochthonous aeolian prey but are mainly sustained by prey of local production.     

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.     

Tree Species Richness Strengthens Relationships between Ants and the Functional Composition of Spider Assemblages in a Highly Diverse Forest

In species‐rich ecosystems, such as subtropical and tropical forests, higher trophic level interactions are key mediators of ecosystem functioning. Plant species loss may alter these interactions, but the effects of plant diversity might be modified by intraguild interactions, particularly among predators. We analyzed the relationships between spiders and ants, two dominant predatory arthropod taxa, on tree saplings across a gradient from medium to high woody plant species richness in a subtropical forest in Southeast China. Neither ant nor spider total biomass was significantly related to plant species richness. By contrast, the biomass distribution of web‐building and hunting spiders changed and spider family richness increased in the presence of ants, resulting in more web builder‐dominated assemblages. However, these relationships depended on the plant communities, and were stronger in plots with higher plant species richness. Our results indicate that in addition to potential effects of ants on hunting spiders in particular, ants could indirectly influence intraguild interactions within spider assemblages. The observed shifts in the spider assemblages with increasing ant presence and plant species richness may have functional consequences, as web‐building and hunting spiders have distinct prey spectra. The relationships among ants, spiders, and plant species richness might contribute to explaining the non‐significant relationship between the overall effects of predators and plant diversity previously observed in the same forest plots. Our findings thus give insight into the complexity of biotic interactions in such species‐rich ecosystems.     

More intraguild prey than pest species in arachnid diets may compromise biological control in apple orchards

Understanding the full diet of natural enemies is necessary for evaluating their role as biocontrol agents, because many enemy species do not only feed on pests but also on other natural enemies. Such intraguild predation can compromise pest control if the consumed enemies are actually better for pest control than their predators. In this study, we used gut metabarcoding to quantify diets of all common arachnid species in Swedish and Spanish apple orchards. For this purpose, we designed new primers that reduce amplification of arachnid predators while retaining high amplification of all prey groups. Results suggest that most arachnids consume a large range of putative pest species on apple but also a high proportion of other natural enemies, where the latter constitute almost a third of all prey sequences. Intraguild predation also varied between regions, with a larger content of heteropteran bugs in arachnid guts from Spanish orchards, but not between orchard types. There was also a tendency for cursorial spiders to have more intraguild prey in the gut than web spiders. Two groups that may be overlooked as important biocontrol agents in apple orchards seem to be theridiid web spiders and opilionids, where the latter had several small-bodied pest species in the gut. These results thus provide important guidance for what arachnid groups should be targets of management actions, even though additional information is needed to quantify all direct and indirect interactions occurring in the complex arthropod food webs in fruit orchards.     

Spatial refuge from intraguild predation: implications for prey suppression and trophic cascades

The ability of predators to elicit a trophic cascade with positive impacts on primary productivity may depend on the complexity of the habitat where the players interact. In structurally-simple habitats, trophic interactions among predators, such as intraguild predation, can diminish the cascading effects of a predator community on herbivore suppression and plant biomass. However, complex habitats may provide a spatial refuge for predators from intraguild predation, enhance the collective ability of multiple predator species to limit herbivore populations, and thus increase the overall strength of a trophic cascade on plant productivity. Using the community of terrestrial arthropods inhabiting Atlantic coastal salt marshes, this study examined the impact of predation by an assemblage of predators containing Pardosa wolf spiders, Grammonota web-building spiders, and Tytthus mirid bugs on herbivore populations (Prokelisia planthoppers) and on the biomass of Spartina cordgrass in simple (thatch-free) and complex (thatch-rich) vegetation. We found that complex-structured habitats enhanced planthopper suppression by the predator assemblage because habitats with thatch provided a refuge for predators from intraguild predation including cannibalism. The ultimate result of reduced antagonistic interactions among predator species and increased prey suppression was enhanced conductance of predator effects through the food web to positively impact primary producers. Behavioral observations in the laboratory confirmed that intraguild predation occurred in the simple, thatch-free habitat, and that the encounter and capture rates of intraguild prey by intraguild predators was diminished in the presence of thatch. On the other hand, there was no effect of thatch on the encounter and capture rates of herbivores by predators. The differential impact of thatch on the susceptibility of intraguild and herbivorous prey resulted in enhanced top-down effects in the thatch-rich habitat. Therefore, changes in habitat complexity can enhance trophic cascades by predator communities and positively impact productivity by moderating negative interactions among predators.     

Caught in the web: Spider web architecture affects prey specialization and spider–prey stoichiometric relationships

Quantitative approaches to predator–prey interactions are central to understanding the structure of food webs and their dynamics. Different predatory strategies may influence the occurrence and strength of trophic interactions likely affecting the rates and magnitudes of energy and nutrient transfer between trophic levels and stoichiometry of predator–prey interactions. Here, we used spider–prey interactions as a model system to investigate whether different spider web architectures—orb, tangle, and sheet‐tangle—affect the composition and diet breadth of spiders and whether these, in turn, influence stoichiometric relationships between spiders and their prey. Our results showed that web architecture partially affects the richness and composition of the prey captured by spiders. Tangle‐web spiders were specialists, capturing a restricted subset of the prey community (primarily Diptera), whereas orb and sheet‐tangle web spiders were generalists, capturing a broader range of prey types. We also observed elemental imbalances between spiders and their prey. In general, spiders had higher requirements for both nitrogen (N) and phosphorus (P) than those provided by their prey even after accounting for prey biomass. Larger P imbalances for tangle‐web spiders than for orb and sheet‐tangle web spiders suggest that trophic specialization may impose strong elemental constraints for these predators unless they display behavioral or physiological mechanisms to cope with nutrient limitation. Our findings suggest that integrating quantitative analysis of species interactions with elemental stoichiometry can help to better understand the occurrence of stoichiometric imbalances in predator–prey interactions.     

Web orientation and prey resources for web-building spiders in eastern hemlock

We examined the arthropod community on eastern hemlock, Tsuga canadensis (L.) Carr, in the context of its role in providing potential prey items for hemlock-associated web-weaving spiders. Using sticky traps simulating spider webs, we evaluated what prey items are available to web-weaving spiders in eastern hemlock based on web orientation (horizontal versus vertical) and cardinal direction. We found that the overwhelming majority (>70%) of prey items available to spiders in hemlock canopies were Diptera. Psocoptera, Hymenoptera, and Hemiptera comprised most of the remaining potential prey. A significant direction × orientation interaction, and greater trap capture in some direction-orientation combinations, suggests that spiders might locate their webs in eastern hemlock canopies for thermoregulatory purposes, ultimately optimizing prey capture. We also evaluated these findings in the context of hemlock infestation by the invasive hemlock woolly adelgid, Adelges tsugae Annand. The adelgid is a sedentary insect with a mobile crawler stage that provides a readily available, easily obtained food source for predators in hemlock canopies. However, an abundance of alternative prey will affect within canopy spider distribution and the potential intensity with which spiders consume these prey. Understanding the response of spiders to potential prey availability is essential to understanding the trophic interactions involving these predators and their potential for influencing herbivore populations.     

Evaluating the effects of trophic complexity on a keystone predator by disassembling a partial intraguild predation food web

1. Many taxa can be found in food webs that differ in trophic complexity, but it is unclear how trophic complexity affects the performance of particular taxa. In pond food webs, larvae of the salamander Ambystoma opacum occupy the intermediate predator trophic position in a partial intraguild predation (IGP) food web and can function as keystone predators. Larval A. opacum are also found in simpler food webs lacking either top predators or shared prey. 2. We conducted an experiment where a partial IGP food web was simplified, and we measured the growth and survival of larval A. opacum in each set of food webs. Partial IGP food webs that had either a low abundance or high abundance of total prey were also simplified by independently removing top predators and/or shared prey. 3. Removing top predators always increased A. opacum survival, but removal of shared prey had no effect on A. opacum survival, regardless of total prey abundance. 4. Surprisingly, food web simplification had no effect on the growth of A. opacum when present in food webs with a low abundance of prey but had important effects on A. opacum growth in food webs with a high abundance of prey. Simplifying a partial IGP food web with a high abundance of prey reduced A. opacum growth when either top predators or shared prey were removed from the food web and the loss of top predators and shared prey influenced A. opacum growth in a non-additive fashion. 5. The non-additive response in A. opacum growth appears to be the result of supplemental prey availability augmenting the beneficial effects of top predators. Top predators had a beneficial effect on A. opacum populations by reducing the abundance of A. opacum present and thereby reducing the intensity of intraspecific competition. 6. Our study indicates that the effects of food web simplification on the performance of A. opacum are complex and depend on both how a partial IGP food web is simplified and how abundant prey are in the food web. These findings are important because they demonstrate how trophic complexity can create variation in the performance of intermediate predators that play important roles in temporary pond food webs.     

Intraguild interactions among generalist predator functional groups drive impact on herbivore and decomposer prey

Different functional groups of generalist predators may complement each other in controlling prey populations; but intraguild interactions, common among generalist predators, may also reduce the strength of top–down control. In natural communities greater alterations to ecosystem function are expected if a whole functional group declines in abundance or is lost. Therefore studying functional group diversity is important for predicting effects of predator loss. We studied the top–down impact of web‐building spiders, hunting spiders and ants, which are highly abundant generalist predators in most terrestrial ecosystems, on prey from the herbivore and decomposer system of a grassland food web. The density of the three predator groups was manipulated by continuous removal in a three‐factorial designed field experiment, which was carried out for two years. We found no positive effect of increasing predator functional group richness on prey control. However there was evidence for strong composition effects between the functional groups. The presence of ants in predator assemblages reduced the prey suppression through mostly trait‐mediated intraguild interactions, while hunting and web‐building spiders contributed additively to prey suppression and reduced the density of herbivore and decomposer prey by 50–60%. A trophic cascade on plant biomass triggered by web‐builders and hunting spiders was diminished at levels of higher predator group diversity. In conclusion, our experiments showed that intraguild interactions strongly influence the strength of top–down control by generalist predators. Among spiders there was evidence for a positive relation between functional group richness and prey suppression but the overall outcome strongly depended on the occurrence of interference, driven by trait‐mediated indirect interactions.     

Contributions of detrital subsidies to aboveground spiders during secondary succession, revealed by radiocarbon and stable isotope signatures

Prey subsidies originating from detritus add nutrients and energy to arboreal communities. Measurement of this subsidy is required in the understanding of how food web dynamics respond to changes in surrounding environments. Shrub spiders are one of the key predators involved in food web coupling. We evaluate the effects of potential changes in prey availabilities during secondary succession on the contribution of subsidy from detrital food webs to shrub spiders and how different spider feeding guilds used the subsidy of prey from detrital food webs. We measured the relative importance of the subsidy for the spider feeding guilds, using the ratios of stable isotopes of C (δ¹³C), and N (δ¹⁵N) and C isotope discrimination (Δ¹⁴C). Diet age was calculated from Δ¹⁴C values, because old diet ages of spiders indicate that the spiders consume prey from detrital food sources. Dominant aerial prey (Diptera) had a distinctively old diet age compared with arboreal prey, which indicates that aerial prey were subsidized from detrital food webs. Sit-and-wait spiders tended to have an older diet age than active hunting spiders, which indicates that sit-and-wait spiders depended more on subsidies. Diet age varied only slightly for spiders in stands of different ages, indicating that rates at which spiders use grazing and detrital prey are probably determined more by foraging strategies and not by stand age. A dominance of sit-and-wait predators will lead to higher detrital subsidy inputs in shrub habitats. This study highlights the effect of shrub spider community structure (feeding guild composition) on the volume of the subsidy received from the detrital food web.     

Intraguild interactions between spiders and ants and top-down control in a grassland food web

In most terrestrial ecosystems ants (Formicidae) as eusocial insects and spiders (Araneida) as solitary trappers and hunters are key predators. To study the role of predation by these generalist predators in a dry grassland, we manipulated densities of ants and spiders (natural and low density) in a two-factorial field experiment using fenced plots. The experiment revealed strong intraguild interactions between ants and spiders. Higher densities of ants negatively affected the abundance and biomass of web-building spiders. The density of Linyphiidae was threefold higher in plots without ant colonies. The abundance of Formica cunicularia workers was significantly higher in spider-removal plots. Also, population size of springtails (Collembola) was negatively affected by the presence of wandering spiders. Ants reduced the density of Lepidoptera larvae. In contrast, the abundance of coccids (Ortheziidae) was positively correlated with densities of ants. To gain a better understanding of the position of spiders, ants and other dominant invertebrate groups in the studied food web and important trophic links, we used a stable isotope analysis (¹⁵N and ¹³C). Adult wandering spiders were more enriched in ¹⁵N relative to ¹⁴N than juveniles, indicating a shift to predatory prey groups. Juvenile wandering and web-building spiders showed δ¹⁵N ratios just one trophic level above those of Collembola, and they had similar δ¹³C values, indicating that Collembola are an important prey group for ground living spiders. The effects of spiders demonstrated in the field experiment support this result. We conclude that the food resource of spiders in our study system is largely based on the detrital food web and that their effects on herbivores are weak. The effects of ants are not clear-cut and include predation as well as mutualism with herbivores. Within this diverse predator guild, intraguild interactions are important structuring forces.     

A test of the hypothesis that a pathway of intraguild predation limits densities of a wolf spider

Abstract.  1. The high number of potential predatory interactions among the many arthropod generalist predators in terrestrial food webs makes exhaustive testing of interaction strengths by field experiments unfeasible. Thus, correlative patterns and laboratory observations of behaviour often form the basis of inferences about the strength of interaction pathways involving generalist predators (intraguild predation).
2. Previous research has revealed a negative correlation between survival of juvenile wolf spiders of the genus Schizocosa (Lycosidae) and densities of another abundant spider family, the Gnaphosidae.
3. Feeding trials in laboratory microcosms with a leaf-litter substrate revealed that gnaphosids prey on juvenile Schizocosa in a structurally complex habitat.
4. Gnaphosid densities were manipulated in two different field experiments, each conducted in a different year, in order to test directly the hypothesis that intraguild predation by gnaphosids limits densities of juvenile Schizocosa .
5. Reducing numbers of gnaphosids, and doubling their numbers to two times the mean natural density, had no impact on the survival of juvenile Schizocosa in either field experiment. This finding suggests that correlative patterns in nature and feeding trials in the laboratory may at times provide deceptively simple and potentially misleading generalisations about the strengths of interaction pathways in complex networks of generalist predators.     

Management intensity and vegetation complexity affect web-building spiders and their prey

Agricultural management and vegetation complexity affect arthropod diversity and may alter trophic interactions between predators and their prey. Web-building spiders are abundant generalist predators and important natural enemies of pests. We analyzed how management intensity (tillage, cutting of the vegetation, grazing by cattle, and synthetic and organic inputs) and vegetation complexity (plant species richness, vegetation height, coverage, and density) affect rarefied richness and composition of web-building spiders and their prey with respect to prey availability and aphid predation in 12 habitats, ranging from an uncut fallow to a conventionally managed maize field. Spiders and prey from webs were collected manually and the potential prey were quantified using sticky traps. The species richness of web-building spiders and the order richness of prey increased with plant diversity and vegetation coverage. Prey order richness was lower at tilled compared to no-till sites. Hemipterans (primarily aphids) were overrepresented, while dipterans, hymenopterans, and thysanopterans were underrepresented in webs compared to sticky traps. The per spider capture efficiency for aphids was higher at tilled than at no-till sites and decreased with vegetation complexity. After accounting for local densities, 1.8 times more aphids were captured at uncut compared to cut sites. Our results emphasize the functional role of web-building spiders in aphid predation, but suggest negative effects of cutting or harvesting. We conclude that reduced management intensity and increased vegetation complexity help to conserve local invertebrate diversity, and that web-building spiders at sites under low management intensity (e.g., semi-natural habitats) contribute to aphid suppression at the landscape scale.     

Intraguild predation promotes complex alternative states along a productivity gradient

Intraguild predation is the simplest, ubiquitous form of trophic omnivory, known to greatly influence the structure and functioning of natural and managed food webs. Although alternative states are fundamental to intraguild predation dynamics, only necessary conditions for alternative states have been previously reported. Using simple models, we found complex but systematic patterns in which different alternative states occur along a productivity gradient, and clarified the sufficient conditions to separate these patterns. We found that two quantities known to control the necessary conditions also determine the sufficient conditions: (1) relative energy transfer efficiency through alternative trophic pathways to an intraguild predator, and (2) relative resource exploitation ability between intraguild prey and predator. These governing quantities suggest how body size and stoichiometric relations between intraguild prey and predators can influence the possibility of alternative states. Our results indicate that food webs involving intraguild predation have a high potential of complex alternative states, and their management can be highly precarious.     

Review Behaviour and indirect interactions in food webs of plant-inhabiting arthropods

With the increased use of biological control agents, artificial food webs are created in agricultural crops and the interactions between plants, herbivores and natural enemies change from simple tritrophic interactions to more complex food web interactions. Therefore, herbivore densities will not only be determined by direct predator–prey interactions and direct and indirect defence of plants against herbivores, but also by other direct and indirect interactions such as apparent competition, intraguild predation, resource competition, etc. Although these interactions have received considerable attention in theory and experiments, little is known about their impact on biological control. In this paper, we first present a review of indirect food web interactions in biological control systems. We propose to distinguish between numerical indirect interactions, which are interactions where one species affects densities of another species through an effect on the numbers of an intermediate species and functional indirect interactions, defined as changes in the way that two species interact through the presence of a third species. It is argued that functional indirect interactions are important in food webs and deserve more attention. Subsequently, we discuss experimental results on interactions in an artificial food web consisting of pests and natural enemies on greenhouse cucumber. The two pest species are the two-spotted spider mite Tetranychus urticae and the western flower thrips, Frankliniella occidentalis. Their natural enemies are the predatory mite Phytoseiulus persimilis, which is commonly used for spider mite control and the predatory mites Neoseiulus cucumeris and Iphiseius degenerans and the predatory bug Orius laevigatus, all natural enemies of thrips. First, we analyse the possible interactions between these seven species and we continue by discussing how functional indirect interactions, particularly the behaviour of arthropods, may change the significance and impact of direct interactions and numerical indirect interactions. It was found that a simple food web of only four species already gives rise to some quite complicated combinations of interactions. Spider mites and thrips interact indirectly through resource competition, but thrips larvae are intraguild predators of spider mites. Some of the natural enemies used for control of the two herbivore species are also intraguild predators. Moreover, spider mites produce a web that is subsequently used by thrips to hide from their predators. We discuss these and other results obtained so far and we conclude with a discussion of the potential impact of functional indirect and direct interactions on food webs and their significance for biological control.     

Small-scale spatial pattern of web-building spiders (Araneae) in alfalfa: relationship to disturbance from cutting, prey availability, and intraguild interactions

Understanding the development of spatial patterns in generalist predators will improve our ability to incorporate them into biological control programs. We studied the small-scale spatial patterns of spider webs in alfalfa by analyzing the relationship between web locations over distances ranging from 4 to 66 cm. Using a coordinate-based spatial statistic (O-ring) and assuming a heterogeneous distribution of suitable web sites, we analyzed the impact of cutting and changes in spider abundance on web distribution. We analyzed the influence of small-scale variation in prey availability by comparing web distributions to the pattern of sticky-trap captures of Aphididae and Diptera described by a count-based spatial statistic (SADIE). Cutting of alfalfa reduced the overall density of web-building spiders but had no immediate impact on the spatial distribution of their webs. Availability of aphids was highest before the alfalfa was cut and was clumped at a scale of 66 cm. Spider webs, however, were not clumped at any scale or date. In contrast, webs were regularly distributed at smaller distances (<20 cm) immediately before and after cutting. Because cursorial and web-building spiders were most active during this period, we hypothesize that the development of small-scale regularity in web locations was driven by intraguild interactions. Our results suggest that intraguild interactions contribute to the development of small-scale spatial patterns of spider webs in alfalfa. Variation in prey availability may have more of an influence on web distribution in crops with a different vegetation structure or if patterns are studied at larger spatial scales.     

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.     

Intraguild predation,invertebrate predators,and trophic cascades in lake food webs

The top-down and bottom-up properties of model food webs that include intraguild predation and self-limiting factors such as cannibalism are investigated. Intraguild predation can dampen or even reverse the top-down effects predicted by food chain theory. The degree of self-limitation among the intraguild prey is a key factor in determining the direction and strength of the top-down response. Intraguild predation and self-limiting factors can also substantially alter the bottom-up effects of enrichment. These results can help explain the disparate results of trophic cascade experiments in lakes, where cascades are usually seen when large Daphnia are the primary herbivores, but not when smaller-bodied herbivores are dominant. Top-down manipulations should cascade at least modestly to phytoplankton in those lakes whose food web can be reasonably approximated by a chain (typically, those where Daphnia is the dominant herbivore), as predicted by food chain theory. On the other hand, smaller-bodied zooplankton are often preyed upon heavily by invertebrate predators as well as by planktivorous fish, thereby introducing elements of intraguild predation into these food webs. In this case, conventional food chain theory is likely to give incorrect predictions. Very large cascade effects may be due primarily to regime shifts between intraguild predation-dominated food webs and those that more resemble food chains, rather than due to the simple food chain cascade usually considered.