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.     

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.     

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.     

Trade-offs in foraging success and predation risk with spatial position in colonial spiders

Summary Colonial web-building spiders respond to trade-offs between selective forces relative to spatial position within colonies and thus provide support for the selfish herd theory. The size distribution of spiders within colonies of Metepeira incrassata, a colonial orb-weaver (Araneae: Araneidae) from tropical Mexico is nonrandom; larger (mature) spiders and females guarding eggsacs are more prevalent in the center, whereas more small (immature) spiders are found on the periphery. Experimental field studies with spiders of selected size classes show that larger spiders actively and aggressively seek protected positions in the center of the colony webbing, even though prey availability and capture rates are significantly higher on the periphery. Attacks by predatory wasps, other spiders, and hummingbirds are more frequent on the periphery than in the core of the colony. Reproductive females on the periphery are at greater risk because they are captured more often than smaller spiders, and if their egg sacs consequently remain unguarded, chances of cocoon parasitism are increased. As a result, spiders in the core of the colony have greater reproductive success, producing more egg sacs with greater hatching frequency. Colonial spiders thus appear to be making a trade-off between foraging and protection from predation and show a spatial organization predicted by the selfish herd theory. The influence of such trade-offs on individual fitness and the structure of colonies is discussed. Offprint requests to: G.W. Uetz     

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.     

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     

Functional values of stabilimenta in a wasp spider, Argiope bruennichi: support for the prey-attraction hypothesis

Many orb-weaving spiders decorate their webs with conspicuous ultraviolet (UV)-reflective stabilimenta. The prey-attraction hypothesis suggests that stabilimenta are visually attractive to prey and thus may increase the spiders’ foraging success. However, previous studies on the function of stabilimenta have produced conflicting results in Argiope species. Using a combination of field and laboratory studies, we examined whether the linear stabilimentum of Argiope bruennichi contributes to prey interception. We recorded prey interceptions in 53 webs with stabilimenta and 37 equally-sized webs without stabilimenta, classifying captured prey according to their taxonomical group and size. On average, 6.2?±?4.7 prey items were intercepted in webs with stabilimenta, while 3.2?±?2.9 items were intercepted in webs without stabilimenta. The effects of stabilimenta on foraging success appear to be due to increased interception of UV-sensitive insect pollinators, including 20 families of Diptera, Hymenoptera, Coleoptera, and Lepidoptera. The mean number of UV-sensitive prey was 4.4?±?3.6 in webs with stabilimenta compared with 1.8?±?2.1 in webs without stabilimenta. Webs with and without stabilimenta did not differ in the mean number of UV-nonsensitive prey captured. The linear stabilimentum showed strong positive effects on the interception of large prey: webs with stabilimenta captured more than twice as many large prey (≥5?mm) than webs without stabilimenta, whereas there was only a slight difference in the interception rates for small prey (<5?mm). Comparisons among different Argiope species suggest that the stabilimentum may have different adaptive functions in different species or ecological contexts.     

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.     

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.     

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.     

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.     

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.     

Sexual cohabitation as mate-guarding in the leaf-curling spider Phonognatha graeffei Keyserling (Araneoidea, Araneae)

The leaf-curling spider Phonognatha graeffei incorporates a twisted leaf into the central hub of its orb-web that is used as a retreat. This species is unusual among orb-weaving spiders because males cohabit in the leaf retreat with both immature and mature females, mating with the former shortly after the female molts. Cohabitation appears to be a form of mate-guarding because cohabiting males respond agonistically to rival males that venture onto the web, and their behaviour depends upon the reproductive status of the female; males defending immature females are more aggressive than those defending virgin, adult females. Males copulate with previously mated females for significantly longer than with virgin females. Females may cannibalise cohabiting males, which occurs independently of whether the female has been deprived of food. Females that cannibalise a single male do not have a higher fecundity than non-cannibalistic females. Received: 2 February 1996 / Accepted after revision: 27 October 1996     

Defensive chemistry of lycid beetles and of mimetic cerambycid beetles that feed on them

Summary.  Beetles of the family Lycidae have long been known to be chemically protected. We present evidence that North American species of the lycid genera Calopteron and Lycus are rejected by thrushes, wolf spiders, and orb-weaving spiders, and that they contain a systemic compound that could account, at least in part, for this unacceptability. This compound, a novel acetylenic acid that we named lycidic acid, proved actively deterrent in feeding tests with wolf spiders and coccinellid beetles. Species of Lycuscommonly figure as models of mimetic associations. Among their mimics are species of the cerambycid beetle genus Elytroleptus, remarkable because they prey upon the model lycids. We postulated that by doing so Elytroleptus might incorporate the lycidic acid from their prey for their own defense. However, judging from analytical data, the beetles practice no such sequestration, explaining why they remain relatively palatable (in tests with wolf spiders) even after having fed on lycids. Chemical analyses also showed the lycids to contain pyrazines, such as were already known from other Lycidae, potent odorants that could serve in an aposematic capacity to forestall predatory attacks. David Utterback: Deceased     

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.     

Community composition and nearest neighbor relationships in orb-weaving spiders: the product of aggression?

Summary Nearest-neighbor distances and species' densities were calculated for 949 individuals of 8 species of orb-weaving spiders from eight edge and meacow sites in Panama. These data were used to test the hypothesis that interspecific aggressive interactions influenced spider community structure by: (1) eliminating or decreasing the densities of sympatric, less aggressive species, or, (2) affecting nearest-neighbor relationships so that large or aggressive individuals had smaller, less aggressive or more distant neighbors. Although there was a negative relationship between the densities of two species pairs (Leucauge sp. and Mangora pia; Eriophora fuliginea and Cyclosa caroli), overall spider densities could not be predicted from indices of spider aggression obtained by introduction experiments. The spatial distribution of spiders in three dimensions was not significantly different than random. I attributed the lack of clumping to the relative vegetational homogeneity of my sites; the lack of overdispersion may reflect the fact that websites are not limiting. Relationships between nearest neighbors generally supported the aggression hypothesis, however, comparisons of the actual data with equivalent randomly-assorted data indicate that a hypothesis of randomly chosen neighborsis equally applicable. This result suggests that, unless websites differ significantly in their quality, with high quality websites in short supply, spider aggression may not affect community structure.     

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.     

A predator’s preference for egg-carrying prey: a novel cost of parental care

Using a subsocial spitting spider (Scytodes pallida) as the prey and a spider-eating jumping spider (Portia labiata) as the predator, the cost of parental care is investigated. Our findings suggest that being singled out as preferred prey by P. labiata is, for egg-carrying females of S. pallida, an important cost of parental care. In survival tests, during which P. labiata was given access to egg-carrying and eggless S. pallida females, egg-carrying females were preyed on more often than eggless females. In preference tests, motionless lures instead of living S. pallida were used. The lures were made by mounting dead egg-carrying and dead eggless S. pallida females in lifelike posture in webs. In these tests, P. labiata detected and identified, by vision alone, both kinds of prey (egg-carrying and eggless), and singled out egg-carrying females as preferred prey.Communicated by M.A. Elgar     

The concentration of Cu and Pb in the funnel spider Eratigena atrica (C. L. Koch 1843) (Araneae: Agelenidae) and its web

Studies on the heavy metal concentrations on spider webs in relation to the pathways of pollution penetration (external and internal) have yet to be performed. This work assesses the concentration levels of two heavy metals: essential copper (Cu) and toxic lead (Pb) in spider webs and spiders (females, males and juveniles). Spiders divided into three treatment groups were exposed to the heavy metals in their diet as prey (fruit flies and mealworms larvae) were artificially contaminated with Cu and Pb. In general, we found higher rates of Cu compared to Pb concentrations in spiders and their webs. A positive correlation between levels of Cu and Pb in webs and spiders was found. Males had higher concentrations of both metals Cu and Pb in their bodies and webs compared to females. In an additional experiment, washed webs had significantly less metals than unwashed suggesting the dominance of external pollution in the contamination pathway.     

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.