Importance of habitat structure to the arthropod food-web in Douglas-fir canopies

This study tested the hypothesis that habitat structure dictates the distribution and community composition of arboreal arthropods. A diverse arthropod assemblage of Douglas-fir canopies, which included Araneae, Psocoptera, Collembola and Homoptera, was chosen as a model system. Habitat structural diversity, defined as needle density and branching complexity of Douglas-fir branches, was manipulated in a four-month experiment by needle removal, thinning and tying of branches. Abundance of canopy spiders declined significantly following needle density reduction and branch thinning, branch tying significantly increased spider abundance. Distinct habitat utilization patterns were found among individual spider guilds. Orb weavers (Araneidae) dominated spider assemblages in structurally simple habitats, whereas tied branches were colonized primarily by sheet-web weavers (Linyphiidae) and nocturnal hunting spiders (Anyphaenidae and Clubionidae). Spider species richness and average body size of several spider species increased in structurally more complex habitats. Arboreal spiders appeared to be limited by strong bottom-up effects in the form of habitat quality and, to a lesser degree, prey availability. Habitat manipulations did not affect densities or biomass of flying arthropod colonists in the branch vicinity. Needle removal and branch thinning led to a significant decline in the abundance of Psocoptera and Collembola. Tying of branches resulted in an eight-fold increase in Collembola numbers, organisms most sensitive to habitat alterations. Canopy habitat structure modified vertical dispersal of Collembola from forest litter, which may have significant implications for arboreal consumers. Our results lend strong support to the importance of habitat structural diversity in explaining general patterns of arthropod abundance and diversity on plants.     

Bird predation affects diurnal and nocturnal web-building spiders in a Mediterranean citrus grove

Spiders and birds can greatly decrease insect populations, but birds also limit spider densities in some habitats. Bird predation is thought to be one of the causes behind nocturnal activity in spiders, so night-active spiders that hide in retreats during the day should be less affected by bird foraging than day-active spiders. However, this hypothesis has not yet been tested. We investigated the importance of bird predation on the spider community of a Mediterranean organic citrus grove. We excluded birds by placing net cages over the trees and we conducted visual searches in the canopies to sample web-building spiders. As there are many nocturnal species in the family Araneidae, we conducted searches both by day and by night to compare the abundance of active araneids in these two time periods. We sampled the tree trunks with cardboard bands to collect hunting spiders. In bird-excluded canopies there were more spiders of the families Araneidae and Theridiidae. There were higher numbers of active Araneidae at night, but these were just as negatively affected by bird predation as day-active Araneidae, so there was no evidence of nocturnal activity serving as an anti-predator strategy. We did not find any negative impact of birds on hunting spiders. Our results contrast with other studies reporting a negative effect of birds on hunting but not on web-building spiders.     

Kleptoparasites influence foraging behaviour of the spider <Emphasis Type="Italic">Stegodyphus lineatus</Emphasis> (Araneae,Eresidae)

Prey captured by a predator may attract kleptoparasites which could significantly reduce the amount of food consumed. Stegodyphus lineatus, a cribellate spider, builds an energetically costly web. Ants raid the webs of S. lineatus to steal prey and behave as kleptoparasites. We investigated ant raids in a natural population of S. lineatus and their influence on the spider’s foraging behaviour. Considering spiders that had captured a prey, 31.2% suffered an ant raid within 24 h after the prey capture. Experimental tests showed that the response to ant raid is to delay web rebuilding and this was independent of a spider’s previous foraging success. There was a tendency for spiders that were exposed to ants to build larger webs. Neither prey-handling duration nor prey consumption was modified after exposure to ants. These results suggest that Stegodyphus lineatus adapt its web-building behaviour in response to the risk of kleptoparasitism.     

Poor Display Repertoire, Tolerance and Kleptobiosis: Results of Specialization in an Ant-Eating Spider (Araneae, Zodariidae)

The agonistic display repertoire of myrmecophagous Zodarion rubidum has five displays. This is fewer than in other spiders, which is a result of the short time spent in contests (4 s). Such a short duration seems to be an adaptation to living among foraging ants, which are dangerous to spiders. The interaction procedure was markedly affected by the presence of preyimmobilized ant. Contests between individuals without prey, or each holding prey, were usually resolved by leg waving. But contests between an individual without and an individual with prey escalated to more aggressive levels. Nevertheless, spiders were never observed to harm or cannibalize one another. Absence of cannibalism is explained as a result of diet specialization: only ants elicit a predatory behavior and provide Zodarion spiders with optimal nutrients. Some spiders used kleptobiosis to gain ants. They first tried to gain immobilized prey aggressively and if failed they adopted a stealthy tactic and shared the prey with the owner. Kleptobiosis is an alternative foraging strategy for Zodarion spiders as it reduces risks associated with hunting dangerous ants.     

Lack of diet segregation during breeding by male and female Northern Flickers foraging on ants

Sexual size dimorphism can result in reduced competition if it leads males and females to use different foraging techniques or consume different prey items. Among woodpeckers, differences between males and females in bill length are common and may explain foraging differences in this family of birds. Northern Flickers (Colaptes auratus) are ground‐foraging woodpeckers that specialize on ants. However, the overall contribution of ants to their diet and the proportions of particular ant genera in their diet are not well known. To understand the relationship between bill morphology and the consumption of prey items, we compared the bill length and bill width of male and female flickers. We then collected and analyzed fecal samples from breeding flickers (N = 40 males, 33 females) at a study site in central British Columbia, Canada. Bills of male flickers were significantly longer (4%) and wider (5%) than those of females. Of 11 prey types identified, ants made up over 99% of their diet, and the abundance and composition of ant taxa in the diet did not differ between the sexes. We found significant year and time of season effects, with the abundance of Tapinoma sessile and Lasius spp. increasing from May to the end of June and differing between years. This difference in diet composition between years may have been due to changes in the abundance or accessibility of certain ant taxa related to differences in vegetation structure or weather. Nine ant taxa were consumed by flickers and the four most common were T. sessile, Lasius spp.,Myrmica spp., and the Formica fusca species group. The degree of dimorphism in bill size of male and female Northern Flickers in our study was smaller than reported for several species of arboreal‐foraging woodpeckers, suggesting that bill size of ground‐foraging woodpeckers may not be strongly linked to niche separation at the level of prey selection.     

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.     

Ant versus bird exclusion effects on the arthropod assemblage of an organic citrus grove

1. Predation‐exclusion experiments have highlighted that top‐down control is pervasive in terrestrial communities, but most of these experiments are simplistic in that they only excluded a single group of predators and the effect of removal was evaluated on a few species from the community. The main goal of our study was to experimentally establish the relative effects of ants and birds on the same arthropod assemblage of canopy trees. 2. We conducted 1‐year long manipulative experiments in an organic citrus grove intended to quantify the independent effects of bird and ant predators on the abundance of arthropods. Birds were excluded with plastic nets whereas ants were excluded with sticky barriers on the trunks. The sticky barrier also excluded other ground dwelling insects, like the European earwig Forficula auricularia L. 3. Both the exclusion of ants and birds affected the arthropod community of the citrus canopies, but the exclusion of ants was far more important than the exclusion of birds. Indeed, almost all groups of arthropods had higher abundance in ant‐excluded than in control trees, whereas only dermapterans were more abundant in bird‐excluded than in control trees. A more detailed analysis conducted on spiders also showed that the effect of ant exclusion was limited to a few families rather than being widespread over the entire diverse spectrum of spiders. 4. Our results suggest that the relative importance of vertebrate and invertebrate predators in regulating arthropod populations largely depends on the nature of the predator–prey system.     

Effects of ants on the foraging of birds in spruce trees

I experimentally excluded ants from randomly selected spruce trees Picea abies near colonies of the wood ant Formica aquilonia. Foraging activity of birds in these trees was then compared to the foraging activity of birds in neighboring spruce trees, where ants were allowed to continue foraging. Birds which foraged in the foliage showed the effects of competition with ants: they visited the trees without ants more frequently, and for longer periods. In addition, the insects and spiders that they utilized as food were more abundant in the foliage of trees without ants. Cone-foraging birds, however, which fed on seeds in cones at the tops of the trees, did not show a preference for trees without ants. The differences of tree usage between foliage-gleaning and coneforaging birds can be explained by alteration of the birds' food supply by wood ants: ants did not feed on seeds in cones, and so did not compete with cone-foraging birds. However, foraging wood ants did feed on arthropods living in the foliage, thus reducing the amount of food available to birds there.     

Response of the foraging behaviour of red wood ants (Formica rufa group) to exclusion from trees

Abstract 1 Because of the large numbers within a colony and their aggressive nature, red wood ants (Formica rufa group) have a potential to greatly influence the cold‐temperate forest ecosystem. Wood ants are omnivorous and hunt in trees as well as on the forest floor. 2 A field experiment in a mixed forest in central Sweden was carried out to examine (i) the foraging behaviour of wood ants on the forest floor and (ii) the impact of increased numbers of wood ants on the soil fauna. The foraging behaviour of wood ants was manipulated by excluding the ants from their food resources in the tree canopy, with the intention to increase ant activity on the forest floor. To estimate this activity, the number of trees with foraging ants, the numbers of ants going to and from their nests and the prey carried by home‐running wood ants were determined during the summer period. Pitfall traps were placed in the soil to determine effects on mobile soil invertebrates. 3 When excluded from local trees, wood ants searched other trees further away from the nests rather than searching more intensively for prey on the forest floor. By contrast to the initial hypothesis, more soil‐living prey were caught by ants in the control plots than in the plots where the local trees were not accessible to the wood ants. The proportion of soil‐living to tree‐living prey tended to be greater in the control plots. 4 In the treated plots (no access to the trees), wood ants had a negative effect on the activity of Linyphiidae spiders. There was little effect of wood ants on other soil invertebrates. 5 This study suggests that the role of wood ants as top predators in the forest soil food‐web in central Sweden is limited.     

Predation by an exotic lizard,Anolis sagrei,alters the ant community structure in betelnut palm plantations in southern Taiwan

Abstract 1. Predators can affect prey directly by reducing prey abundance and indirectly by altering behavioural patterns of prey. From previous studies, there is little evidence that ant community structure is affected by vertebrate predation. 2. Researchers tend to consider the interactions between vertebrate predators and ants to be weak. The present study examined the impact of the exotic invasive lizard, Anolis sagrei, on the ant community structure by manipulating the density of lizards within enclosures. The natural density of A. sagrei in the field was surveyed and used as the stocking density rate in the lizard‐present sub‐enclosures. 3. Before the lizard density was manipulated, there was no difference in the ant diversity between sub‐enclosures. After the lizard density manipulation, the ant diversity in sub‐enclosures with A. sagrei present was significantly different from that of enclosures where the lizards were absent, although the overall ant abundance did not differ significantly. 4. The ant diversity difference was generated by a significant reduction of the ant species Pheidole fervens in sub‐enclosures with A. sagrei present. Such an abundance change might be the result of direct predation by the lizards, or it might be generated by a foraging site shift by this ant. 5. The results of this study thus demonstrated that the invasion of an exotic vertebrate can significantly alter the community structure of ants, perhaps through the combined direct and indirect effects of lizards on ants.     

Carbon dioxide effects on stomatal responses to the environment and water use by crops under field conditions

The structural complexity of habitats has been espoused as an important factor influencing natural-enemy abundance and food-web dynamics in invertebrate-based communities, but a rigorous synthesis of published studies has not heretofore been conducted. We performed a meta-analytical synthesis of the density response of natural enemies (invertebrate predators and parasitoids) to experimental increases and decreases in the structural complexity of their habitats using data from 43 published studies, reporting 62 independent taxa. Studies varied in structural complexity at two spatial scales (habitat and within-plant architecture) and comprised a diverse array of natural-enemy taxa (natural-enemy assemblage at large, the entire spider assemblage, hunting spiders, web-building spiders, mites, hemipterans, coccinellid beetles, carabid beetles, ants, and parasitoids). For all taxa combined, increasing habitat structure resulted in a large and significant increase in natural enemy abundance. Similarly, decreasing habitat structure significantly diminished natural enemy abundance. Separate meta-analyses at two spatial scales (habitat and within-plant architecture) found that increasing habitat complexity resulted in significant increases in abundance. In particular, manipulating levels of detritus at the habitat spatial scale had the strongest effect on natural enemy abundance. In general, most guilds of natural enemies were significantly affected when the structural complexity of the habitat was altered. Seven of nine natural enemy guilds were more abundant under conditions of increased habitat complexity, with hunting spiders and web-building spiders showing the strongest response followed by hemipterans, mites, and parasitoids. Spiders in particular were negatively affected when habitat structure was simplified. The mechanisms underlying the accumulation of natural enemies in complex-structured habitats are poorly known. However, refuge from intraguild predation, more effective prey capture, and access to alternative resources (alternative prey, pollen, or nectar), are possible candidates. Our analysis was unable to confirm that predators aggregate in complex-structured habitats because prey (mostly herbivores) are more abundant there. The results of this meta-analysis support the view that basal resources mediate top-down impacts on herbivores, and provide encouragement that manipulations of habitat complexity can be made in agroecosystems that will enhance the effectiveness of the natural enemy complex for more effective pest suppression.Electronic Supplementary Material Supplementary material is available in the online version of this article at     

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.     

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.     

Disentangling the Diversity of Arboreal Ant Communities in Tropical Forest Trees

Tropical canopies are known for their high abundance and diversity of ants. However, the factors which enable coexistence of so many species in trees, and in particular, the role of foragers in determining local diversity, are not well understood. We censused nesting and foraging arboreal ant communities in two 0.32 ha plots of primary and secondary lowland rainforest in New Guinea and explored their species diversity and composition. Null models were used to test if the records of species foraging (but not nesting) in a tree were dependent on the spatial distribution of nests in surrounding trees. In total, 102 ant species from 389 trees occurred in the primary plot compared with only 50 species from 295 trees in the secondary forest plot. However, there was only a small difference in mean ant richness per tree between primary and secondary forest (3.8 and 3.3 sp. respectively) and considerably lower richness per tree was found only when nests were considered (1.5 sp. in both forests). About half of foraging individuals collected in a tree belonged to species which were not nesting in that tree. Null models showed that the ants foraging but not nesting in a tree are more likely to nest in nearby trees than would be expected at random. The effects of both forest stage and tree size traits were similar regardless of whether only foragers, only nests, or both datasets combined were considered. However, relative abundance distributions of species differed between foraging and nesting communities. The primary forest plot was dominated by native ant species, whereas invasive species were common in secondary forest. This study demonstrates the high contribution of foragers to arboreal ant diversity, indicating an important role of connectivity between trees, and also highlights the importance of primary vegetation for the conservation of native ant communities.     

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.     

Within-tree distribution of nest sites and foraging areas of ants on canopy trees in a tropical rainforest in Borneo

It has been argued that canopy trees in tropical rainforests harbor species-rich ant assemblages; however, how ants partition the space on trees has not been adequately elucidated. Therefore, we investigated within-tree distributions of nest sites and foraging areas of individual ant colonies on canopy trees in a tropical lowland rainforest in Southeast Asia. The species diversity and colony abundance of ants were both significantly greater in crowns than on trunks. The concentration of ant species and colonies in the tree crown seemed to be associated with greater variation in nest cavity type in the crown, compared to the trunk. For ants nesting on canopy trees, the numbers of colonies and species were both higher for ants foraging only during the daytime than for those foraging at night. Similarly, for ants foraging on canopy trees, both values were higher for ants foraging only during the daytime than for those foraging at night. For most ant colonies nesting on canopy trees, foraging areas were limited to nearby nests and within the same type of microhabitat (within-tree position). All ants foraging on canopy trees in the daytime nested on canopy trees, whereas some ants foraging on the canopy trees at night nested on the ground. These results suggest that spatial partitioning by ant assemblages on canopy trees in tropical rainforests is affected by microenvironmental heterogeneity generated by three-dimensional structures (e.g., trees, epiphytes, lianas, and aerial soils) in the crowns of canopy trees. Furthermore, ant diversity appears to be enriched by both temporal (diel) and fine-scale spatial partitioning of foraging activity.     

Genetically-based variation between two spider populations in foraging behavior

Summary Optimal foraging theory is based on the assumption that at least some aspects of foraging behavior are genetically determined (Pyke et al. 1977; Kamil and Sargent 1980; Pyke 1984). Nonetheless, very few studies have examined the role of genetics in foraging behavior. Here, we report on geographical differences in the foraging behavior of a spider (Agelenopsis aperta) and investigate whether these differences are genetically determined. Field studies were conducted on two different populations of A. aperta: one residing in a desert riparian habitat, and the other in a desert grassland habitat. Data from the spiders' natural encounters with prey demonstrated that grassland spiders exhibited a higher frequency of attack than riparian spiders towards 13 of 15 prey types, including crickets and ants. Grassland spiders also had shorter latencies to attack 12 of 15 prey types, including crickets and ants, than riparian spiders. Subsequently, we reared grassland and riparian spiders under controlled conditions in the laboratory and observed their interactions with prey to determine whether the populational differences we found in the field could be genetic. Again, grassland spiders showed a shorter latency to attack prey (crickets, ants) than riparian spiders. These latencies were not significantly affected by the hunger state or age of the spiders. Finally, we reared a second generation (F2) of grassland and riparian spiders in the laboratory and observed their interactions with prey to determine whether the populational differences in the previous generation were due to genetic effects or maternal effects. As before, grassland spiders exhibited a shorter latency to attack prey (crickets) than riparian spiders. We conclude that the foraging differences we observed between these two populations of A. aperta are genetically determined. These differences probably have resulted from either natural selection acting directly on attack frequency and the latency to attack prey, or natural selection acting on traits which are genetically correlated with these aspects of foraging behavior.     

Risk Sensitivity and the Paradox of Colonial Web-Building in Spiders

Group foraging is rare in spiders, occurring only where preyavailability is high. If colonial web-building increases individualprey capture rates as shown, why does group foraging not occurmore often where prey are scarce? Risk sensitivity may explainthis paradox, as variance in prey capture is reduced in groups;risk-averse spiders should join groups only when prey exceeda threshold level. Field studies show that group foraging variesas predicted between species, between populations of a singlespecies, and between sites within a population. However, recentmodels suggest the necessity of examining variance within individualsover time rather than between individuals within populations.Additionally, mechanisms responsible for variance reductionin colonial webs may be less effective than previously assumed.New field data suggest that while prey variance over time maybe somewhat less for individual spiders in groups than for solitaries,the relationship between colonial web-building and variancein prey capture is far more complex than originally thought.The influence of risk sensitivity on reproductive success andthe evolution of colonial web-building is discussed.     

Alternative predatory tactics of an araneophagic assassin bug (Stenolemus bituberus)

Predators of dangerous prey risk being injured or killed in counter-attacks and hence may use risk-reducing predatory tactics. Spiders are often dangerous predators to insects, but for a few, including Stenolemus bituberus assassin bugs, web-building spiders are prey. Despite the dangers of counter-attack when hunting spiders, there has been surprisingly little investigation of the predatory tactics used by araneophagic (spider-eating) insects. Here, we compare the pursuit tendency, outcome and predatory tactics of S. bituberus against five species of web-building spider. We found that S. bituberus were most likely to hunt and capture spiders from the genus Achaearanea, a particularly common prey in nature. Capture of Achaearanea sp. was more likely if the prey spider was relatively small, or if S. bituberus was in poor condition. S. bituberus used two distinct predatory tactics, ‘stalking’, in which they slowly approached the prey, and ‘luring’, in which they attracted spiders by manipulating the web to generate vibrations. Tactics were tailored to the prey species, with luring used more often against spiders from the genus Achaearanea, and stalking used more often against Pholcus phalangioides. The choice of hunting tactic used by S. bituberus may reduce the risk posed by the prey spider.