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.     

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.     

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.     

Thermal behaviour of a small spider (Araneae : Araneidae : Araneinae) on horizontal webs in semi-arid Western Australia

Summary The solar orientation response of a small spider (gen. nov.: Araneae: Araneidae: Araneinae) is examined at a semi-arid location in tropical Western Australia. The spiders rest beneath horizontal dome-shaped webs close to the ground surface on spinifex (Triodia sp.). The spiders are small (c. 5 mm long) and robust, being only twice as long as wide. Spider orientation, elevation, wind velocity, radiation and wet and dry bulb temperatures together with spider and egg-sac temperatures were measured throughout the day. The spiders orientated their long axis to the solar azimuth through much of the day and postured to the solar elevation; in so doing they tracked the apparent movement of the sun throughout the day with considerable precision, far exceeding that of other spiders studied. Stepwise regression indicated that the solar position was the most important environmental factor associated with the posturing and orientation. The variance in the orientation of the population was associated with a suite of environmental variables related to wind velocity and temperature. Spider (abdominal) temperature had a curvilinear relationship with ambient temperature, with a plateau at c. 35° C, whereas egg-sac temperature had a strictly linear relationship with ambient temperature. Stepwise regression indicates that both the spider and egg-sac temperatures are related mainly to net radiation and ambient temperature. The spiders can alter the incident radiation to which they are exposed by c. 2.2 times which represent the limits of the exposed silhouette area between full dorsal sunlight and the posterior (or anterior) aspect. Under hot conditions the spiders posture and orientate such that the long axis of the body is orientated abdomen towards the sun.     

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.     

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.     

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     

Exploring the effects of individual traits and within-colony variation on task differentiation and collective behavior in a desert social spider

Social animals are extraordinarily diverse and ecologically abundant. In understanding the success of complex animal societies, task differentiation has been identified as a central mechanism underlying the emergence and performance of adaptive collective behaviors. In this study, we explore how individual differences in behavior and body size determine task allocation in the social spider Stegodyphus dumicola. We found that individuals with high body condition indices were less likely to participate in prey capture, and individuals’ tendency to engage in prey capture was not associated with either their behavioral traits or body size. No traits were associated with individuals’ propensity to participation in web repair, but small individuals were more likely to engage in standard web-building. We also discovered consistent, differences among colonies in their collective behavior (i.e., colony-level personality). At the colony level, within-colony variation in behavior (aggressiveness) and body size were positively associated with aggressive foraging behavior. Together, our findings reveal a subtly complex relationship between individual variation and collective behavior in this species. We close by comparing the relationship between individual variation and social organization in nine species of social spider. We conclude that intraspecific variation is a major force behind the social organization of multiple independently derived lineages of social spider.     

Cooperative prey capture by young subsocial spiders: II. Behavioral mechanism

Subsocial spiders demonstrate an intermediate stage in the evolution of permanent sociality. Cooperative hunting is an important attribute of their sociality, but has not been documented in subsocial arthropods. After cannibalizing their mother, young of the subsocial spider Amaurobius ferox (Araneae, Amaurobiidae) remain together for several instars and feed communally. We monitored the collective prey capture behavior of the spiderlings. All the clutches showed collective capturing sequence (latency–orientation–moving–touching–seizing–feeding) toward the prey that was 10 times more massive than each individual. The first three individuals that exhibited attacking behavior were responsible for 90% of the total number of attacks, while 68% of the individuals within the group never exhibited attacks during the first 10 min following the introduction of prey into the communal web. First arriving individuals at the prey most often seized the antennae and legs of the prey, which probably facilitate access to the prey for subsequent individuals. The spiderlings that arrived later occupied more likely the abdomen and thorax, which contain more nutrition than the extremities occupied earlier. The individual apportionment of collective hunting behavior suggests a coordinated teamwork among individuals.     

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     

Aggressive chemical mimicry of moth pheromones by a bolas spider: how does this specialist predator attract more than one species of prey?

Summary. The bolas spider, Mastophora hutchinsoni, attracts Lacinipolia renigera and Tetanolita mynesalis males by mimicking the female moth sex pheromones. However, as the prey species use completely different pheromone blends we conducted experiments to determine how this is accomplished by the predator. The periodicity of L. renigera mate-seeking activities occurs early in the scotophase, whereas male T. mynesalis are active late at night, corresponding with periods when these moths are captured by the spider. The pheromone blend of early-flying L. renigera interferes with attraction of late-flying T. mynesalis to its pheromone in a dose-dependent manner, suggesting the spider must always produce a single sub-optimal “compromise” blend for both species or that it adjusts its allomonal blend to optimize capture of the respective prey species at different times during the night. We delayed (L. renigera) or advanced (T. mynesalis) the periodicity of male activity through photoperiodic manipulation and found that the bolas spider attracted both prey species outside their normal activity windows. These results support the idea that bolas spiders produce components of both species at all times rather than producing the pheromone of each prey species at different times of the night. However, using coupled gas chromatography-electroantennography, we also demonstrated that the spider decreases its emission of the L. renigera pheromone over the course of the night. This modification should reduce the behavioral antagonism of the L. renigera pheromone on T. mynesalis males and increase the predator's success of attracting T. mynesalis during this prey's normal activity window late at night. Received 13 October 2001; accepted 28 December 2001.     

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.     

The spider and fly revisited: ploy–counterploy behavior in a unique predator–prey system

Earlier studies have shown that the sarcophagid fly Arachnidomyia lindae is the principal egg-sac predator of the colonial orb-weaving spider Metepeira incrassata, and that risk of egg-sac loss increases with group size. Observations of specialized behaviors for attack (flies) and defense (spiders) suggest that this predator-prey relationship may incorporate elements of ploy and counterploy behavior. Here we explore this relationship in detail and test hypotheses regarding efficacy of attack and defense behaviors. Egg-sac guarding by the spider includes defensive behaviors specific to this fly predator, which were observed during experimental "attacks" with tethered A. lindae, but not with similar presentations of non-predatory Musca domestica. Experimental studies also show that Metepeira incrassata recognizes this predatory fly by airborne cues (i.e., the signature frequency of wing-beats), and can distinguish between this predator and other flies (potential prey) on the basis of wing-beat frequency differences. Removal of female spiders results in a significantly higher probability of unguarded egg-sacs being parasitized, demonstrating the adaptive value of spider defensive behaviors. We also present evidence that A. lindae utilizes a behavioral ploy for circumventing spider guarding behavior (aggressive mimicry - producing vibrations of captured prey in the web), and that Metepeira incrassata, in turn, exhibits a counter-ploy behavior (signal thread cutting) to eliminate this potentially distracting vibratory information. While previous studies have shown that this colonial web-building spider uses a number of general attack-abatement mechanisms against a diversity of predators and parasitoids, results of this study suggest that selection pressures from a highly specialized predator may also result in evolution of predator-specific prey responses.     

Defensive behavior of the black widow spider Latrodectus hesperus (Araneae: Theridiidae)

Summary When threatened, the black widow spider, Latrodectus hesperus, emits an adhesive, viscous web, pulls the strand from its spinnerets with its fourth pair of legs, and spreads its appendages. This behavior positions the web over the delicate abdomen, increases the area of protection, and enables the spider to place the web onto the offender, if necessary.In laboratory interactions, the viscous web protected black widows from mice (Peromyscus spp.). Mature female spiders, which had their spinnerets blocked and hence could not discharge the viscid silk, escaped less often than did black widows that were not experimentally altered.The viscid silk is palatable to mice and it appears that the deterrent effect of the web is due solely to mechanical irritation.The defensive behavior is elicited most often from mature females, which may suffer greater predation than other age groups. Males lose the ability to produce the defensive web at maturity and may shift their energy resources totally into reproductive effort.     

Predator foraging behavior in response to perception and learning by its prey: interactions between orb-spinning spiders and stingless bees

Although rewarded bees learn and remember colors and patterns, they have difficulty in learning to avoid negative stimuli such as decorated spider webs spun by Argiope argentata. A. argentata decorates its web with silk patterns that vary unpredictably (Fig. 1) and thus foraging insects that return to sites where spiders are found encounter new visual cues daily. Stingless bees can learn to avoid spider webs but avoidance-learning is slowed or inhibited by daily variation in web decorations (Figs. 3,4; Tables 1,2). In addition, even if bees learn to avoid decorated webs found in one location, they are unable to generalize learned-avoidance responses to similarly decorated webs found at other sites. A. argentata seems to have evolved a foraging behavior that is tied to the ways insects perceive and process information about their environment. Because of the evolutionary importance of bee-flower interdependence, the predatory behavior of web-decorating spiders may be difficult for natural selection to act against.     

Within-group behavioral variation promotes biased task performance and the emergence of a defensive caste in a social spider

The social spider Anelosimus studiosus exhibits a behavioral polymorphism where colony members express either a passive, tolerant behavioral tendency (social) or an aggressive, intolerant behavioral tendency (asocial). Here we test whether asocial individuals act as colony defenders by deflecting the suite of foreign (i.e., heterospecific) spider species that commonly exploit multi-female colonies. We (1) determined whether the phenotypic composition of colonies is associated with foreign spider abundance, (2) tested whether heterospecific spider abundance and diversity affect colony survival in the field, and (3) performed staged encounters between groups of A. studiosus and their colony-level predator Agelenopsis emertoni (A. emertoni)to determine whether asocial females exhibit more defensive behavior. We found that larger colonies harbor more foreign spiders, and the number of asocial colony members was negatively associated with foreign spider abundance. Additionally, colony persistence was negatively associated with the abundance and diversity of foreign spiders within colonies. In encounters with a colony-level predator, asocial females were more likely to exhibit escalatory behavior, and this might explain the negative association between the frequency of asocial females and the presence of foreign spider associates. Together, our results indicate that foreign spiders are detrimental to colony survival, and that asocial females play a defensive role in multi-female colonies.     

Three-dimensional barricading of a predatory trap reduces predation and enhances prey capture

Animal structures come at material, energetic, time, and expression costs. Some orb-web spiders add three-dimensional barrier structures to their webs, but many do not. Predator protection is considered to be the principal benefit of adding these structures. Accordingly, it remains paradoxical why some orb-web spiders might construct the barriers while others do not. Here, we experimentally determined whether the barrier structure added to the horizontal orb web of the spider Cyrtophora moluccensis deters predators at the cost of reducing the amount of prey captured in the field. We conducted experiments by day and night to assess whether the effects vary with the time of day. We found that the three-dimensional barriers not only offered protection from predatory wasps by day but also enhanced the amount of prey captured by day and night. Moreover, the barrier structure appears particularly useful at catching moths, the largest and most energetically profitable prey that it encounters. We, therefore, concluded that reducing the energetic and time costs associated with producing and depositing extra silk threads is the principal reason why barrier structures are used intermittently among orb-web spiders.     

The role of thermoregulation in lizard biology: Predatory efficiency in a temperate diurnal basker

Summary Decreasing levels of simulated solar radiation have the following effects in the diurnal basking lizard Lacerta vivipara: (i) increase in time spent basking with a consequent decrease in time available for foraging (ii) decrease in speed of movement whilst foraging (iii) decrease in total foraging distance, and hence contact with potential prey (iv) decrease in searching efficiency in an experimental arena. Complete absence of simulated solar radiation accentuates these effects, and reduces the proportion of faster-moving prey in the diet. Time taken to swallow prey (handling time) increases exponentially with decreasing body temperature. It is concluded from these results that maintaining relatively high activity temperatures (30–36°C) is adaptive for the species because the loss of potential foraging time caused by lengthy periods of basking is offset by the following advantages: increased contact with and capture of prey, increased efficiency of prey handling, and availability of a wider range of prey types.     

Using stable isotopes to reveal shifts in prey consumption by generalist predators.

The effectiveness of generalist predators in biological control may be diminished if increased availability of alternative prey causes individual predators to decrease their consumption of crop pests. Farming practices that enhance densities of microbidetritivores in the detrital food web can lead to increased densities of generalist predators that feed on pest species. The ability to predict the net biocontrol impact of increased predator densities depends upon knowing the extent to which individual predators may shift to detrital prey and feed less on crop pests when prey of the detritus-based food web are more abundant. We addressed this question by comparing ratios of stable isotopes of carbon (delta13C) and nitrogen (delta15N) in generalist ground predators and two types of prey (crop pests and microbidetritivores) in replicated 8 x 8 m cucurbit gardens subjected to one of two treatments: a detrital subsidy or no addition of detritus (control). Small sheet-web spiders (Linyphiidae) and small wolf spiders (Lycosidae) had delta13C values similar to those of Collembola in both the detrital and control treatments, indicating that small spiders belong primarily to the detrital food web. In control plots the larger generalist predators had delta13C values similar to those of the major insect pests, consistent with their known effectiveness as biocontrol agents. Adding detritus may have caused delta13C of one species of large wolf spider to shift toward that of the microbi-detritivores, although evidence is equivocal. In contrast, another large wolf spider displayed no shift in delta13C in the detrital treatment. Thus, stable isotopes revealed which generalist predators will likely continue to feed on pest species in the presence of greater densities of alternative prey.     

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.