Stingless bee response to spider webs is dependent on the context of encounter

In the course of their foraging bouts, bees frequently encounter spider webs among the vegetation. The ability to see and avoid these webs is vital for the success of the individual bee’s foraging bout. In this study, we report on the response of stingless bees (Trigona carbonaria) towards the webs of the St. Andrew’s Cross spider (Argiope keyserlingi). We studied the ability of bees to avoid webs in different contexts: when bees were on their foraging path or when they were returning to the hive as well as when they were flying North or South. We show that the probability of a bee being able to avoid a web depends on the context of the bee’s flight rather than the visual appearance of the web. Furthermore, the presence of the spider seems to alert the bee to the web, resulting in bees being more able to avoid capture. We show, specifically, that the probability of being captured is higher when the bee is returning to the hive compared with when the bee is foraging. The likelihood of avoiding a web is also influenced by the compass direction of the flight, although to a lesser extent. Our results indicate that the context of the predator–prey encounter has a significant influence on a bee’s ability to escape interception by a spider web.     

Signals for damage control: web decorations in <Emphasis Type="Italic">Argiope keyserlingi</Emphasis> (Araneae: Araneidae)

Orb web spiders of the genus Argiope are permanently located at the hub of the orb web and are thus vulnerable to changing environmental conditions. Severe damage to the web by non-prey animals can have a significant impact on survival, through the cost of producing expensive silk and the loss of foraging opportunities. Thus, selection should favor web protection mechanisms, and the conspicuous web decorations, typical of Argiope spiders, may play a role. Decorated webs suffer less damage than undecorated webs, consistent with the view that they advertise the presence of the web to non-prey animals that may damage the web. However, whether spiders respond to web damage by increasing investment in web decorations has not been investigated. We subjected adult St. Andrew's Cross spider (Argiope keyserlingi) females to three levels of web damage and recorded their subsequent web-decorating behavior. Mild damage, similar to that caused by impacting prey, did not affect either web building or decorating behavior. However, spiders subjected to substantial web damage both reduced the size of subsequent webs and increased investment in web decoration size. These data are consistent with an advertising role of web decorations.     

Why do nocturnal orb-web spiders (Araneidae) search for light?

The nocturnal orb-web spider Larinioides sclopetarius lives near water and frequently builds webs on bridges. In Vienna, Austria, this species is particularly abundant along the artificially lit handrails of a footbridge. Fewer individuals placed their webs on structurally identical but unlit handrails of the same footbridge. A census of the potential prey available to the spiders and the actual prey captured in the webs revealed that insect activity was significantly greater and consequently webs captured significantly more prey in the lit habitat compared to the unlit habitat. A laboratory experiment showed that adult female spiders actively choose artificially lit sites for web construction. Furthermore, this behaviour appears to be genetically predetermined rather than learned, as laboratory-reared individuals which had previously never foraged in artificial light exhibited the same preference. This orb-web spider seems to have evolved a foraging behaviour that exploits the attraction of insects to artificial lights. Received: 8 June 1998 / Received in revised form: 18 January 1999 / Accepted: 19 January 1999     

Parent-offspring aggression in moorhens

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

Effects of predator visibility on prey encounter: a case study on aerial web weaving spiders

Summary Perhaps the most important factor affecting predator-prey interactions is their encounter probability. Predators must either locate sites where prey are active or attract prey to them, and prey must be able to recognize potential predators and flee before capture. In this study we manipulate and describe three components of the foraging system of predatory, web-weaving spiders, the presence of viscid droplets, silk brightness (achromatic surface reflectance), and visibility of the orb pattern, to determine their effect on insect attraction, recognition, and web avoidance. We found that webs with viscid droplets were more visible to prey at close range, but at greater distances the sparkling droplets lured insects to the web area and hence increased insect capture probability. Although the size of viscid droplets and silk brightness are closely correlated (Table 2, Fig. 3), the relationships among droplet size, spider size, and the visual environments in which webs are found are more complicated (Fig. 2, Tables 2, 3). In environments with predictable light exposure, droplet size and hence silk visibility correlate with spider size, and spiders that forage at night produce relatively more visible silks then spiders that forage during the day (Table 3, Fig. 4). In habitats in which light levels are not predictable, silk surface reflectance and spider size are not closely correlated, suggesting that the complexity of the light environment, as well as the visual and foraging behaviors of insects found there, has played an important role in the evolution of spider-insect interactions.Offprint requests to: C.L. Craig     

Colour-independent shape recognition of cryptic predators by bumblebees

Predators hunting for cryptic prey use search images, but how do prey search for cryptic predators? We address this question using the interaction between bumblebees and the colour-changing crab spider Misumena vatia which can camouflage itself on some flowers. In laboratory experiments, we exposed bumblebees to an array of flowers concealing robotic predators (a trapping mechanism combined with a 3D life-sized model of a crab spider or a circle). Groups of bees were trained to avoid either cryptic yellow spiders or yellow circles (equal area to the spiders) or remained predator naive. The bees were then exposed to a new patch of white flowers containing some cryptic predators (either white spiders, white circles or a mixture of both). We monitored individual foraging choices and used a 3D video tracking system to quantify the bees’ flight behaviour. The bees trained to avoid cryptic spiders, chose 40% fewer spider-harbouring flowers than expected by chance, but were indifferent to cryptic circles. They also aborted a higher proportion of landings on flowers harbouring spiders, ultimately feeding from half as many ‘dangerous’ flowers as naive bees. Previous encounters with cryptic spiders also influenced the flight behaviour of bees in the new flower patch. Experienced bees spent more time inspecting the flowers they chose to reject (both with and without concealed spiders) and scanned from side to side more in front of the flowers to facilitate predator detection. We conclude that bees disentangle shape from colour cues and thus can form a generalised search image for spider shapes, independent of colour.     

Do social spiders cooperate in predator defense and foraging without a web?

Nearly all social spiders spin prey-capture webs, and many of the benefits proposed for sociality in spiders, such as cooperative prey capture and reduced silk costs, appear to depend on a mutually shared web. The social huntsman spider, Delena cancerides (Sparassidae), forms colonies under bark with no capture web, yet these spiders remain in tightly associated, long-lasting groups. To investigate how the absence of the web may or may not constrain social evolution in spiders, we observed D. cancerides colonies in the field and laboratory for possible cooperative defense and foraging benefits. We observed spiders’ responses to three types of potential predators and to prey that were introduced into retreats. We recorded all natural prey capture over 447 h both inside and outside the retreats of field colonies. The colony’s sole adult female was the primary defender of the colony and captured most prey introduced into the retreat. She shared prey with younger juveniles about half the time but never with older subadults. Spiders of all ages individually captured and consumed the vast majority of prey outside the retreat. Young spiders benefited directly from maternal defense and prey sharing in the retreat. However, active cooperation was rare, and older spiders gained no foraging benefit by remaining in their natal colony. D. cancerides does not share many of the benefits of group living described in other web-building social spiders. We discuss other reasons why this species has evolved group living.     

A predator’s body coloration enhances its foraging profitability by day and night

Few predators forage by both day and night. It remains unknown, however, how the costs and benefits of foraging or signaling are partitioned in animals that forage at all times. The orb-web spider Cyrtophora moluccensis is brightly colored and forages by day and night. We determined the benefits reaped when it forages by both day and night by estimating the biomass of prey caught in their webs. Additionally, we quantified whether the spider’s presence influences the number of prey caught by day and night and whether its colorful body is visible to diurnal and/or nocturnal insects using diurnal and nocturnal insect vision models. We found that approximately five times the biomass of prey was caught in C. moluccensis’ webs by night than by day. Hemipterans, hymenopterans, and dipterans were predominantly caught by day, while lepidopterans (moths) were predominately caught by night. Accordingly, we concluded that foraging by night is more profitable than foraging by day. We predicted that other benefits, for example, energetic advantages or enhanced fecundity, may promote its daytime activity. Foraging success was greater by day and night when the spider was present in the web than when the spider was absent. We also found that parts of the spider’s body were conspicuous to diurnal and nocturnal insects, possibly through different visual channels. The colorful body of C. moluccensis, accordingly, appears to influence its foraging success by attracting prey during both the day and night.     

Social spider defense against kleptoparasitism

Summary Because of the large amount of webbing they provide, social spider colonies often host other satellite spider species referred to as kleptoparasites or food stealers. Such kleptoparasites may take advantage of increased prey capture rates associated with large spider aggregations. This study investigated the relationship between a cooperatively social spider species, Anelosimus eximius (Araneae: Theridiidae), which lives in the undergrowth of tropical rainforests in Peru, and its kleptoparasite, Argyrodes ululans (Araneae: Theridiidae), which specializes in foraging in An. eximius webs. Although large aggregates of spiders may be more attractive to kleptoparasites, the benefits of group defense may offset this cost. Natural colonies were observed, and enclosed field colonies containing fixed numbers of host spiders were manipulated in order to determine whether kleptoparasite success is affected by the number of social spiders that are available for defense. Prey was less likely to be stolen by Ar. ululans when a greater number of host An. eximius spiders were involved in prey capture. When hosts detected a kleptoparasite earlier and chased it more often, prey was more likely to be successfully defended. Ar. ululans was more successful in stealing small prey items in all colonies and gave up more readily on very large prey (> 11 mm). I conclude that communal living and group defense in An. eximius confer protection from the kleptoparasite Ar. ululans.     

Trap barricading and decorating by a well-armored sit-and-wait predator: extra protection or prey attraction?

Animals may build multiple structures to provide benefits to counter the costs of building. Many orb web spiders add multiple structures, e.g., barricading barrier webs and silk decorations, to their webs and these structures have been hypothesized to function to deter predators or attract prey. The heavily armored spiny spiders construct barrier webs around their orb webs and decorate them with conspicuous silk tufts. Why these organisms, already well protected by a thick cuticle and spines, make the extra investment of building barrier webs and adding conspicuous silk decorations is not known. We predicted that these structures function to both attract prey and deter predators. Field experiments were conducted in two consecutive years using orb webs built by the East Asian spiny spider Thelacantha brevispina. We either (1) concealed the decoration, (2) removed the barrier webs, or (3) left the decorations and barrier webs intact. We found year and treatment to interactively influence prey interception rates. In 2010, but not in 2009, we found prey interception with T. brevispina webs to be greater when the decorations were conspicuous than when they were concealed suggesting that the decorations may lure prey. Prey interception was lower when the barrier webs were present without decorations compared to when they were absent without decorations. The prey-attracting function of the decorations thus may counter the reduction in prey interception incurred by adding a barrier web. Predatory wasp interactions were not influenced by any of our treatments, probably because the spiders’ thick cuticle is the primary means of protection from wasps. Bird predation events, while rare, occurred only when decorations were concealed or the barrier webs were removed. It is therefore plausible that the tuft decorations both lure prey and deter birds.     

Kleptoparasites: a twofold cost of group living for the colonial spider, Metepeira incrassata (Araneae, Araneidae)

Several species of kleptoparasitic and araneophagic spiders (Araneae: Family Theridiidae, Subfamily Argyrodinae) are found in colonial webs of the orb-weaving spider Metepeira incrassata (Araneae, Araneidae) from Mexico, where they steal food and/or prey upon their spider hosts. Census data from natural M. incrassata colonies reveal that the incidence of these species increases with colony size. This pattern may reflect the presence of several other orb-weaving spiders, each with their own kleptoparasitic species, invading larger M. incrassata colonies. As the number of these associated spiders increases, so does the density and number of Argyrodinae species in M. incrassata colonies, suggesting that associated spiders might reduce their own kleptoparasite load by building their webs within M. incrassata colonies. This represents a twofold cost to M. incrassata, as a field enclosure experiment revealed that a primarily kleptoparasitic species (Argyrodes elevatus) may reduce prey available to their hosts, but a kleptoparasitic/araneophagic species (Neospintharus concisus) inflicts high mortality upon M. incrassata. However, the cost of kleptoparasitism and predation by these species may be offset in part for M. incrassata individuals in large colonies by certain defensive mechanisms inherent in groups, i.e., “attack-abatement” and “selfish herd” effects. We conclude that increased occurrence of kleptoparasitic and/or predatory Argyrodinae spiders is a consequence of colonial web building and is an important potential cost of group living for colonial web-building spiders.     

Predation risk makes bees reject rewarding flowers and reduce foraging activity

In the absence of predators, pollinators can often maximize their foraging success by visiting the most rewarding flowers. However, if predators use those highly rewarding flowers to locate their prey, pollinators may benefit from changing their foraging preferences to accept less rewarding flowers. Previous studies have shown that some predators, such as crab spiders, indeed hunt preferentially on the most pollinator-attractive flowers. In order to determine whether predation risk can alter pollinator preferences, we conducted laboratory experiments on the foraging behavior of bumble bees (Bombus impatiens) when predation risk was associated with a particular reward level (measured here as sugar concentration). Bees foraged in arenas containing a choice of a high-reward and a low-reward artificial flower. On a bee’s first foraging trip, it was either lightly squeezed with forceps, to simulate a crab spider attack, or was allowed to forage safely. The foragers’ subsequent visits were recorded for between 1 and 4 h without any further simulated attacks. Compared to bees that foraged safely, bees that experienced a simulated attack on a low-reward artificial flower had reduced foraging activity. However, bees attacked on a high-reward artificial flower were more likely to visit low-reward artificial flowers on subsequent foraging trips. Forager body size, which is thought to affect vulnerability to capture by predators, did not have an effect on response to an attack. Predation risk can thus alter pollinator foraging behavior in ways that influence the number and reward level of flowers that are visited.     

Trait-mediated effects on flowers: artificial spiders deceive pollinators and decrease plant fitness

Although predators can affect foraging behaviors of floral visitors, rarely is it known if these top-down effects of predators may cascade to plant fitness through trait-mediated interactions. In this study we manipulated artificial crab spiders on flowers of Rubus rosifolius to test the effects of predation risk on flower-visiting insects and strength of trait-mediated indirect effects to plant fitness. In addition, we tested which predator traits (e.g., forelimbs, abdomen) are recognized and avoided by pollinators. Total visitation rate was higher for control flowers than for flowers with an artificial crab spider. In addition, flowers with a sphere (simulating a spider abdomen) were more frequently visited than those with forelimbs or the entire spider model. Furthermore, the presence of artificial spiders decreased individual seed set by 42% and fruit biomass by 50%. Our findings indicate that pollinators, mostly bees, recognize and avoid flowers with predation risk; forelimbs seem to be the predator trait recognized and avoided by hymenopterans. Additionally, predator avoidance by pollinators resulted in pollen limitation, thereby affecting some components of plant fitness (fruit biomass and seed number). Because most pollinator species that recognized predation risk visited many other plant species, trait-mediated indirect effects of spiders cascading down to plant fitness may be a common phenomenon in the Atlantic rainforest ecosystem.     

Material investment and prey capture potential of reduced spider webs

Summary The webs of Miagrammopes animotus have a simple structure and variable form. However, both the length of their lines and the total surface area of their capture threads are closely associated with spider size. These spiders' ability to deposit both linear and looped cribellar capture threads along a web's diverging capture lines plays an important role in establishing these relationships. Looped capture threads have the greater surface area and are more prominent in the webs of older spiders where they increase a web's surface area and enhance its ability to retain prey. The predicted performance of these webs is supported by comparisons of the stickiness of their threads and a survey of the prey their owners capture. Cribellar thread stickiness increases with spider size, and larger spiders capture prey that have greater masses.     

Influences of aggregative behaviour on space occupation in the spider Zygiella x-notata (Clerck)

Summary The distribution of the spider Zygiella x-notata was examined using field populations of adult females occupying the outside frames of windows. The structure of the populations was aggregative, and the distribution of individuals on the window frames and the size of the webs were density dependent. Also, the sizes of the webs of neighbouring spiders on the same window alternated. This spatial organization involves interactions between neighbours. If one spider out of two is removed, and if all the webs are destroyed, remaining spiders that previously had small webs significantly increase the size of their construction. In contrast, individuals that previously had large webs do not modify the size of their construction. This shows that individuals of Z. x-notata respond to the presence of neighbours. The influence of intraspecific interactions in such a population is discussed.     

Stabilimenta characteristics of the spider Argiope argentata on small islands: support of the predator-defense hypothesis

Summary A variety of orb-weaving spider species construct stabilimenta, patterned areas of dense silk, typically near the hub of the orb. The adaptive significance of this construction, along with associated behaviors such as shuttling and vibrating, is much debated. Arigiope argentata on small islands of the Bahamas frequently possess stabilimenta; we studied 397 individuals of this species to investigate possible functions of their stabilimenta, paying particular attention to predator-defense hypotheses. Cruciform stabilimenta were commoner in all size classes of spiders than discoid stabilimenta or no stabilimentum at all; discoid stabilimenta occurred mostly among intermediate size classes. Within the cruciform type, two-segmented stabilimenta were especially common among the very smallest spiders. Size of cruciform stabilimenta showed a curvilinear relation to spider body length; the fitted curve for total segment length had a maximum at an intermediate spider length. We argue that this relationship (among other phenomena) supports an apparently-larger-size hypothesis, whereby intermediate-sized spiders in particular appear much larger than they actually are. This could discourage predators, including those that are gape-limited such as lizards. We argue that stabilimenta in the smallest spiders, in which typically two segments are opposed, so that they more or less line up, serve as camouflage. When disturbed experimentally, spiders with discoid stabilimenta shuttle to the opposite side of the centrally located stabilimentum. This seems an obvious defensive behavior and occurs less frequently among spiders with cruciform stabilimenta. Large spiders vibrate more frequently than small ones, but no relation exists between vibration frequency and stabilimentum type. We argue (see also Tolbert 1975) that vibrating behavior, in which the spider can become a blur, renders its location more difficult to discern and the spider more difficult to grasp, rather than increasing apparent size. Correspondence to: T.W. Schoener     

Thermal constraints on prey-capture behavior of a burrowing spider in a hot environment

Summary Seothyra henscheli (Eresidae) is a burrowing spider that lives in the dune sea of the southern Namib Desert, Namibia. Prey capture by these spiders involves a foray from a cool subterranean retreat to the undersurface of a capture web that can be lethally hot. Striking, disentangling and retrieving prey from the capture web typically involves several short trips to the capture web, alternating with retreats to the cool burrow. It has been suggested that this behavior limits the increase of body temperature a spider must experience while working at the hot capture web. We used biophysical models in conjunction with direct observations of prey-capture behavior and distributions of sand temperature to estimate body temperatures experienced by S. henscheli during prey capture. In the circumstances we observed, only the relatively long post-strike retreat from the capture web is important in keeping spiders' body temperatures from exceeding their lethal limits. After the post-strike retreat, shuttling appreciably limits the increase in body temperature of small individuals, but may have little effect on body temperature increase in larger spiders. Correspondence to: J.S. Turner at the present address     

Experimental manipulations of dispersal in the subsocial spider,Theridion pictum

Summary Theridion pictum (Walckenaer) is a subsocial spider, that is, the mother actively takes care of her offspring for an extended period of time after they emerge. It is demonstrated that spiderling dispersal is affected by the availability of food in the maternal web. Experimental manipulations indicate that in those webs where less food was provided, spiderlings dispersed significantly earlier and in significantly greater numbers. These results illustrate the importance of food availability as a permissive factor in the evolution of social spiders from subsocial ancestors. Food availability also affects the pattern of sex-biased philopatry and dispersal. Although both sexes are more likely to remain in the maternal web when more food is available, females are relatively more likely to do so. When less food is available, females are relatively more likely to disperse than are males.     

Comparisons of forager distributions from matched honey bee colonies in suburban environments

We conducted experiments designed to examine the distribution of foraging honey bees (Apis mellifera) in suburban environments with rich floras and to compare spatial patterns of foraging sites used by colonies located in the same environment. The patterns we observed in resource visitation suggest a reduced role of the recruitment system as part of the overall colony foraging strategy in habitats with abundant, small patches of flowers. We simultaneously sampled recruitment dances of bees inside observation hives in two colonies over 4 days in Miami, Florida (1989) and from two other colonies over five days in Riverside, California (1991). Information encoded in the dance was used to determine the distance and direction that bees flew from the hive for pollen and nectar and to construct foraging maps for each colony. The foraging maps showed that bees from the two colonies in each location usually foraged at different sites, but occasionally they visited the same patches of flowers. Each colony shifted foraging effort among sites on different days. In both locations, the mean flight distances differed between colonies and among days within colonies. The flight distances observed in our study are generally shorter than those reported in a similar study conducted in a temperate deciduous forest where resources were less dense and floral patches were smaller.     

Ecological costs and benefits of communal behavior in a presocial spider

Summary Philoponella oweni (Uloboridae) is a facultatively communal spider found in the southwestern United States. Solitary females and communal groups of females coexist in the same habitats. Communal females are known to produce more eggs than solitary females. Communal females might acquire the energy for the production of additional eggs by one of three means: they might have a longer life span, and thus more time in which to acquire energy; web maintenance might be more efficient in a communal group; or communal females might spend more time feeding. Field data from a marked population in Arizona showed the third alternative was true.Two possible causes for the greater time spent feeding by communal females were investigated; either the communal web or group is a more efficient trap for insects, or the communal groups are located at sites where insects are more abundant. Analysis of feeding rates and sticky trap data confirmed the second alternative.Potential web attachment sites are limited in some habitats, and mortality during dispersal is very high. When potential web attachment sites are scarce more of the population is found in communal groups; when they are abundant more of the population is solitary. The pattern of dispersal and overwintering is this species suggests that females renewing a colony in the spring could be related.