Selectivity underlies the dissociation between seasonal prey availability and prey consumption in a generalist predator

Generalist predators are capable of selective foraging, but are predicted to feed in close proportion to prey availability to maximize energetic intake especially when overall prey availability is low. By extension, they are also expected to feed in a more frequency‐dependent manner during winter compared to the more favourable foraging conditions during spring, summer and fall seasons. For 18 months, we observed the foraging patterns of forest‐dwelling wolf spiders from the genus Schizocosa (Araneae: Lycosidae) using PCR‐based gut‐content analysis and simultaneously monitored the activity densities of two common prey: springtails (Collembola) and flies (Diptera). Rates of prey detection within spider guts relative to rates of prey collected in traps were estimated using Roualdes’ c_st model and compared using various linear contrasts to make inferences pertaining to seasonal prey selectivity. Results indicated spiders foraged selectively over the course of the study, contrary to predictions derived from optimal foraging theory. Even during winter, with overall low prey densities, the relative rates of predation compared to available prey differed significantly over time and by prey group. Moreover, these spiders appeared to diversify their diets; the least abundant prey group was consistently overrepresented in the diet within a given season. We suggest that foraging in generalist predators is not necessarily restricted to frequency dependency during winter. In fact, foraging motives other than energy maximization, such as a more nutrient‐focused strategy, may also be optimal for generalist predators during prey‐scarce winters.     

Secondary predation: quantification of food chain errors in an aphid-spider-carabid system using monoclonal antibodies

"Secondary predation" occurs when one predator feeds on a second predator, which has in turn eaten a target prey. Detection of prey remains within predators using monoclonal antibodies cannot distinguish between primary and secondary predation, potentially leading to quantitative and qualitative food chain errors. We report the first fully replicated experiments to measure secondary predation effects, using an aphid-spider-carabid system. Aphids, Sitobion avenae, were fed to spiders, Lepthyphantes tenuis, which were allowed to digest their prey for a range of time intervals. The spiders were then fed to carabids, Poecilus (=Pterostichus) cupreus, which were again allowed to digest their prey for set periods. The anti-aphid monoclonal antibody used to identify S. avenae remains in P. cupreus was one that detected an epitope that increased in availability over the first few hours of digestion, amplifying the signal, extending detection periods and thus increasing the chances of detecting secondary predation. Despite this, and the fact that spiders are known to digest their prey more slowly than many other predators, detection of secondary predation was only possible if the carabids were killed immediately after consuming at least two spiders which were, in turn, eaten immediately after consuming aphids. As this scenario is unlikely to occur frequently in the field it was concluded that secondary predation is unlikely to be a serious source of error during field studies.     

Spiders are special: fear and disgust evoked by pictures of arthropods

Because all spiders are predators and most subdue their prey with poison, it has been suggested that fear of spiders is an evolutionary adaptation. However, it has not been sufficiently examined whether other arthropods similarly elicit fear or disgust. Our aim was to examine if all arthropods are rated similarly, if only potentially dangerous arthropods (spiders, bees/wasps) elicit comparable responses, or if spiders are rated in a unique way. We presented pictures of arthropods (15 spiders, 15 beetles, 15 bees/wasps, and 15 butterflies/moths) to 76 students who rated each picture for fear, disgust, and how dangerous they thought the animal is. They also categorized each animal into one of the four animal groups. In addition, we assessed the participants' fear of spiders and estimates for trait anxiety. The ratings showed that spiders elicit significantly greater fear and disgust than any other arthropod group, and spiders were rated as more dangerous. Fear and disgust ratings of spider pictures significantly predicted the questionnaire scores for fear of spiders, whereas dangerousness ratings of spiders and ratings of other arthropods do not provide any predictive power. Thus, spider fear is in fact spider specific. Our results demonstrate that potential harmfulness alone cannot explain why spiders are feared so frequently.     

Perception of ultraviolet light by crab spiders and its role in selection of hunting sites

The perception of ultraviolet (UV) light by spiders has so far been only demonstrated in salticids. Crab spiders (Thomisidae) hunt mostly on flowers and need to find appropriate hunting sites. Previous studies have shown that some crab spiders that reflect UV light use UV contrast to enhance prey capture. The high UV contrast can be obtained either by modulation of body colouration or active selection of appropriate backgrounds for foraging. We show that crab spiders (Thomisus sp.) hunting on Spathiphyllum plants use chromatic contrast, especially UV contrast, to make themselves attractive to hymenopteran prey. Apart from that, they are able to achieve high UV contrast by active selection of non-UV reflecting surfaces when given a choice of UV-reflecting and non-UV reflecting surfaces in the absence of odour cues. Honeybees (Apis cerana) approached Spathiphyllum plants bearing crab spiders on which the spiders were high UV-contrast targets with greater frequency than those plants on which the UV contrast of the spiders was low. Thus, crab spiders can perceive UV and may use it to choose appropriate backgrounds to enhance prey capture, by exploiting the attraction of prey such as honeybees to UV.     

Comparative Growth and Development of Spiders Reared on Live and Dead Prey

Scavenging (feeding on dead prey) has been demonstrated across a number of spider families, yet the implications of feeding on dead prey for the growth and development of individuals and population is unknown. In this study we compare the growth, development, and predatory activity of two species of spiders that were fed on live and dead prey. Pardosa astrigera (Lycosidae) and Hylyphantes graminicola (Lyniphiidae) were fed live or dead fruit flies, Drosophila melanogaster. The survival of P. astrigera and H. graminicola was not affected by prey type. The duration of late instars of P. astrigera fed dead prey were longer and mature spiders had less protein content than those fed live prey, whereas there were no differences in the rate of H. graminicola development, but the mass of mature spiders fed dead prey was greater than those fed live prey. Predation rates by P. astrigera did not differ between the two prey types, but H. graminicola had a higher rate of predation on dead than alive prey, presumably because the dead flies were easier to catch and handle. Overall, the growth, development and reproduction of H. graminicola reared with dead flies was better than those reared on live flies, yet for the larger P. astrigera, dead prey may suit smaller instars but mature spiders may be best maintained with live prey. We have clearly demonstrated that dead prey may be suitable for rearing spiders, although the success of the spiders fed such prey appears size- and species specific.     

Resource Management in the Orb-Weaving Spider Zygiella x-notata II. Ability to Manage a Set of Various Prey

Management of a set of various prey received ‘simultaneously’ by the orb-weaving spider Zygiella x-notata is investigated in relation to prey species, weight, order of arrival in the web, and factors concerning the spider's state of nourishment. When the spiders encountered several different prey ‘simultaneously’, they subdued all of them, took one of them to begin consumption, and stored the others in the web. The first item for consumption was chosen according to the characteristics of the available prey. Factors concerning previous feeding influenced the spiders's choice of the first item to be ingested. Exploitation of the first item, estimated by the quantity of food extracted and the time necessary for this extraction, depended on the type and the quantity of prey at the spider's disposal. When the spiders received several prey, they spent less time feeding on the first cricket compared with spiders receiving only one cricket, while no difference was found when the first prey item exploited was a fly.     

Scavenging by spiders (Araneae) and its relationship to pest management of the brown recluse spider

Experiments reported in Sandidge (2003; Nature 426: 30) indicated that the brown recluse spider, Loxosceles reclusa Gertsch & Mulaik, preferred to scavenge dead prey over live prey and that the spiders were not detrimentally affected when fed insecticide-killed crickets. Extrapolations made in subsequent media coverage disseminating the results of this research made counter-intuitive statements that pesticide treatment in houses would increase brown recluse populations in homes. This information was presented as if the scavenging behavior was specialized in the brown recluse; however, it was more likely that this behavior has not been well studied in other species. To provide a comparison, the current laboratory study examined the likelihood of non-Loxosceles spiders to scavenge dead prey. Of 100 non-Loxosceles spiders that were tested (from 11 families, 24 genera, and at least 29 species from a variety of spider hunting guilds), 99 scavenged dead crickets when offered in petri dishes. Some of the spiders were webspinners in which real-world scavenging of dead prey is virtually impossible, yet they scavenge when given the opportunity. Therefore, scavenging is a flexible opportunistic predatory behavior that is spread across a variety of taxa and is not a unique behavior in brown recluses. These findings are discussed in relation to pest management practices.     

The non-filter function of orb webs in spiders

Summary Experiments with artificial webs show that there is no correlation between prey size and mesh width. The number of prey items per thread increases with mesh width. Spiders with narrow-meshed webs catch more prey, whilst spiders with wide-meshed webs save more spinning material.     

Influence of prey availability and conspecifics on patch quality for a cannibalistic forager: laboratory experiments with the wolf spider Schizocosa

 Because cannibals are potentially both predator and prey, the presence of conspecifics and alternative prey may act together to influence the rate at which cannibals prey upon each other or emigrate from a habitat patch. Wolf spiders (Lycosidae) are cannibalistic-generalist predators that hunt for prey with a sit-and-wait strategy characterized by changes in foraging site. Little information is available on how both prey abundance and the presence of conspecifics influence patch quality for these cursorial, non-web-building spiders. To address this question, laboratory experiments were conducted with spiderlings and older juveniles of the lycosid genus Schizocosa. The presence of insect prey consistently reduced rates of spider emigration when spiders were housed either alone or in groups. Solitary juvenile Schizocosa that had been recently collected from the field exhibited a median giving-up time (GUT) of 10 h in the absence of prey (Collembola); providing Collembola increased the median GUT to 64 h. For solitary spiders, the absence of prey increased by about fourfold the rate of emigration during the first 24 h. In contrast, for spiders in patches with a high density of conspecifics, the absence of prey increased the 24-h emigration rate by only 1.6-fold. For successful cannibals in the no-prey patches, the presence of conspecifics improved patch quality by providing a source of food. Mortality by cannibalism was affected by both prey availability and openness of the patch to net emigration. In patches with no net emigration, the presence of prey reduced rates of cannibalism from 79% to 57%. Spiders in patches open to emigration but not immigration experienced a rate of cannibalism (16%) that was independent of prey availability. The results of these experiments indicate that for a cannibalistic forager such as the wolf spider Schizocosa, (1) the presence of conspecifics can improve average patch quality when prey are absent, and (2) cannibalism has the potential to be a significant mortality factor under natural field conditions because cannibalism persisted in prey patches that were open to emigration. Received: 12 April 1996 / Accepted: 14 August 1996     

The effect of hunger level on predation dynamics in the spider Nesticodes rufipes: a functional response study

It is well known that a predator has the potential to regulate a prey population only if the predator responds to increases in prey density and inflicts greater mortality rates. Predators may cause such density-dependent mortality depending on the nature of the functional and numerical responses. As spiders are usually faced with a shortage of prey, the killing behavior of the spider Nesticodes rufipes at varying densities of Musca domestica was examined here through laboratory functional response experiments where spiders were deprived of food for 5 (well-fed) or 20 days (hungry). An additional laboratory experiment was also carried out to assess handling time of spiders. The number of prey killed by spiders over 24- and 168-h periods of predator–prey interaction was recorded. Logistic regression analyses revealed the type II functional response for both well-fed and hungry spiders. We found that the lower predation of hungry spiders during the first hours of experimentation was offset later by an increase in predation (explained by estimated handling times), resulting in similarity of functional response curves for well-fed and hungry spiders. It was also observed that the higher number of prey killed by well-fed spiders over a 24-h period of spider–prey interaction probably occurred due to their greater weights than hungry spiders. We concluded that hungry spiders may be more voracious than well-fed spiders only over longer time periods, since hungry spiders may spend more time handling their first prey items than well-fed spiders.     

Predatory behaviour of an araneophagic assassin bug

Assassin bugs from the genus Stenolemus (Heteroptera, Reduviidae) are predators of web-building spiders. However, despite their fascinating lifestyle, little is known about how these insects hunt and catch their dangerous prey. Here we characterise in detail the behaviour adopted by Stenolemus bituberus (Stål) during encounters with web-building spiders, this being an important step toward understanding this species’ predatory strategy. These bugs employed two distinct predatory tactics, “stalking” and “luring”. When stalking their prey, bugs slowly approached the prey spider until within striking range, severing and stretching threads of silk that were in the way. When luring their prey, bugs attracted the resident spider by plucking and stretching the silk with their legs, generating vibrations in the web. Spiders approached the luring bug and were attacked when within range. The luring tactic of S. bituberus appears to exploit the tendency of spiders to approach the source of vibrations in the web, such as might be generated by struggling prey.     

Temperature and prey capture: opposite relationships in two predator taxa

Abstract 1. All other things equal, predator capture rates are expected to depend on encounter rate with prey, prey escape capability (including prey defences), and on predator agility. Ectotherm predators and their prey both respond to increasing temperature by increased activity, i.e. predators increase their search area and prey may enhance their escape capability. This means that, as temperature changes, the ability of a predator to catch prey will decrease, increase, or remain unchanged depending on the relative effect of temperature on predator and prey. Their responses may further be differentially moulded by light conditions depending on whether the predator is diurnally or nocturnally active. It was hypothesised that flying Diptera are vulnerable to carabid beetles only at low temperatures and over the full temperature range for spiders because carabids, in contrast to spiders, are not built to catch swiftly moving prey. 2. The first experiment examined the spontaneous locomotor activity of the predators and of fruit flies at different temperatures (5, 10, 15, 20, 25, and 30 °C) and light conditions (light, dark). A second experiment examined the effect of temperature and light on the predation rate of two carabid beetles (Pterostichus versicolor and Calathus fuscipes) and two spiders (Clubiona phragmitis and Pardosa prativaga) using fruit flies (Drosophila melanogaster) as prey. 3. All four predators and the fruit fly increased their locomotory activity at higher temperatures. Activity of the carabid beetles peaked at intermediate temperatures; spiders and fruit flies were most active at the highest temperatures. Predation rate of the spiders increased with temperature whereas the beetles caught flies only at low temperatures (5 and 10 °C). 4. Diurnal variation in temperature may bring different prey groups within the set of potential prey at different times of the day or at different seasons. The ability of many carabid beetles to forage at low temperatures may have nutritional benefits and increases the diversity of interactions in terrestrial food webs.     

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.     

Ontogenetic approach reveals how cognitive capability and motivation shape prey‐searching behavior in Pholcus phalangioides cellar spiders

An important part of understanding the evolution of behavior is understanding how and why behavior develops and changes throughout ontogeny. Patterns of behavior are shaped by an animal's capabilities as well as its motivations, both of which are subject to selection. We ran an experiment to see how spiders' efforts to recover lost prey change with age and to determine the relative contributions of shifts in capability and motivation. We found that as spiders mature, they spend less time searching to recover lost prey, and they discriminate less between prey of different sizes. We also found that even the youngest, least experienced spiders are cognitively equipped to search for lost prey. Thus, predatory behavior in spiders fluctuated primarily with each age group's motivations to capture and consume prey, and did not seem to be hindered by behavioral or cognitive limitations at young ages.     

Foraging behavior of the communal spider,Philoponella republicana (Araneae: Uloboridae)

The communal orb-weaving spider, Philoponella republicana,was observed in the subtropical moist forest of Southeast Peru. These spiders live in colonies of conspecifics whose individual orbs are connected by silk. The wrapping of a prey prior to feeding is a large component of the prey capture process because P. republicanahas no venom with which to kill an insect. Wrapping time was the only aspect of prey capture that was strongly correlated with the size of the insect captured. Occasionally we observed several individuals working together to wrap a prey item. These joint efforts were more frequent on prey larger than the capturing spider. Although group captures accounted for only 5.5% of captures, they represented 14.7% of the biomass obtained. A comparison of the relationship between wrapping time and prey size for solitary and group efforts suggested that, by working together, the spiders reduced their total handling time. In most cases only one spider fed on the captured prey.     

Effects of prey quality and availability on the life history of a trap-building predator

Although generalist predators catch a great diversity of prey species, foraging theory has mostly been concerned with quantitative aspects and neglected questions about the nutrient quality of prey. Here, we test the hypothesis that the life history of a trap-building predator is affected by both prey availability and by the nutrient quality of prey. Under controlled laboratory conditions, orb-weaving spiders ( Zygiella x-notata ) were raised from hatchlings to maturity on prey of different nutrient quality and in different amounts. Both prey nutrient quality and availability had significant but different effects on many important life history traits, such as instar duration, number of instars used in the development, body weight at maturation and development time. Prey availability was especially important for growth rates whereas prey nutrient quality had the most severe effects on juvenile survivorship and female fecundity. Furthermore, while prey of low quality tended to reduce the number of instars used in the development, prey availability induced sex-specific responses in instar numbers. Thus, both prey nutrient quality and availability may be important factors shaping the evolution of life history traits in generalist predators.     

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.     

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

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

Impact of predation by spiders on a territorial damselfly (Odonata: Calopterygidae)

Summary Predation by orb-weaving spiders and crab spiders on the damselfly Calopteryx haemorrhoidalis was studied at a small stream in Southern France. One species of orb-weaving spider, Larinioides folium, caught 76% of the damselflies which fell prey to spiders. Displacement experiments on spiders on sections of bank and the positioning of webs in male territories show that the density and distribution of damselflies is not influenced by orbweb density or by the position of webs. Predation rates corresponded to orb-web density, but neither for sex nor for stage was there a relationship with damselfly density. Mean daily predation rates ranged between 0.9% for females and 4.1% for adult males. Predation risk to adult damselflies by orb-weaving spiders was male biased, whereas among tenerals there was no bias. Males were captured more frequently at territories near the water. Captures show a maximum at noon when territorial disputes of adult males were most frequent. After orbwebs were placed within territories predation rate of males was strongly increased. Predation risk to adult females in the direct vicinity of the stream was less than in the bank vegetation where they perch close to orbwebs. The risk of predation by crab spiders, which catch damselflies at their perching sites, was not sex-biased.     

蛛网结构性能及其适应性

蛛网是蜘蛛的捕食工具,蛛网的结构性能不仅影响蜘蛛的捕食效率,也关系着蜘蛛的捕食投入。在不同的内外环境条件影响下,蜘蛛会通过蛛网结构性能上的相应变化来调整捕食策略和维持网结构的稳定性。本文主要综述了蛛网的结构性能以及蜘蛛通过蛛网结构性能表现出的对环境因子的适应性。