Anti-predator defences of Pholcus phalangioides (Araneae, Pholcidae), a web-building and web-invading spider

The behaviours used by Pholcus phalangioides (Fuesslin) (Araneae, Pholcidae) to evade its predators were studied with particular attention being given to a special defence behaviour, whirling. To whirl, this long-legged web-building spider swings its body around in a circle, with its legs remaining on the silk. Experiments were carried out to determine the types of stimuli that elicited whirling. Touching the spider or its web elicited whirling, as did air movement over the spider, but there was no evidence that chemical stimuli from potential predators were important. Small juveniles differed from adult females and larger juveniles by more often dropping from the web instead of whirling when confronted by a potential predator. Besides catching prey on its own web P. phalangioides invades other spiders' webs to catch the other spiders. By whirling in alien webs, P. phalangioides could deter attacks by the resident spider, but P. phalangioides was less inclined to whirl when in an alien than when in its own web.     

Anti-predator defences of a web-building spider, Holocnemus pluchei (Araneae, Pholcidae)

Anti-predator defence behaviour of Holocnemus pluchei (Araneae, Pholcidae) was studied in the laboratory and the field. Bouncing, in which this web-building spider moves its body rapidly up and down while keeping its legs on the silk, was the most frequent response to disturbance. Less often, H. pluchei left the web when disturbed. Experiments were carried out to determine the types of stimuli that elicited bouncing and leaving the web: tactile stimulation (striking the spider with a brush) and air movement (blowing on the spider) elicited bouncing and leaving the web, but there was no evidence that chemical stimuli from potential predators were important. Tactile stimulation elicited stronger responses than air movement. There were no evident differences in how H. pluchei responded to tactile stimulation in the laboratory and the field.     

Predator-prey interactions between web-invading jumping spiders and Argiope appensa (Araneae, Araneidae), a tropical orb-weaving spider

A range of web-invading jumping spiders with different predatory strategies was tested with A. appensa in the laboratory: Mimetus maculosus (Mimetidae), Pholcus phalangioides (Pholcidae), Taieria erebus (Gnaphosidae), and 11 species of salticids. Spiders that are known to specialize at web-invading, either by leaping into webs or by walking slowly into webs and practising aggressive mimicry, captured A. appensa ; three salticid species not known to be web-invaders never did. Web-invaders that practised aggressive mimicry were more efficient than were species that only leapt into webs. Portia fimbriata from Queensland was the most consistent at using aggressive mimicry and was also the most efficient at catching A. appensa . Web-invaders that were more efficient at catching A. appensa were also better able to avoid setting off pumping, a special defence behaviour used by A. appensa . Portia fimbriata from Queensland was especially efficient at avoiding setting off pumping: P. fimbriata more consistently than other Portia made its final approach toward A. appensa by coming down from above the web on a dragline and making minimal contact with the web. An experiment, in which A. appensa was artificially induced to pump whenever the predator was near, provided additional evidence that pumping is effective in defending A. appensa against web-invaders.     

Predator-induced plasticity in web-building behaviour

Many orb-web weaving spiders add conspicuous silken structures, called stabilimenta, to the hub of their webs, which are hypothesized to attract more prey. However, they may also attract predators. Orb spiders should therefore alter their web-building behaviour to minimize predation risk. We tested this hypothesis by experimentally examining web-building responses of the St Andrew cross spider, Argiope versicolor, to predation risk from one of its natural predators, the jumping spider Portia labiata. We randomly assigned A. versicolor juveniles to one of three treatments: (1) blank control (clean blotting paper: no odour from the predator or nonpredator); (2) predator odour cues from P. labiata; and (3) nonpredator control (odour cues from Leucauge decorata). Each individual of A. versicolor was monitored until it had built five consecutive webs (two webs before and three webs after the introduction of predator cues). When exposed to predator cues, the juveniles not only decreased the frequency of stabilimentum building but also refrained from increasing stabilimentum area, capture area and capture silk thread with subsequent webs compared with the blank control and the nonpredator control. Web-building traits, however, were not significantly different between the blank control and the nonpredator control. One plausible explanation is that A. versicolor juveniles can detect and discriminate between predators and nonpredators through olfactory cues and alter stabilimentum building and other web traits in response to the risk of predation. This is the first demonstration of an adaptive, plastic web-building behavioural response induced by chemical cues from a predator.     

Anti-predator defences of Psilochorus sphaeroides and Smeringopus pallidus (Araneae, Pholcidae), tropical web-building spiders

Anti-predator defence behaviours of Psilochorus sphaeroides from Queensland and Smeringopus pallidus from Sri Lanka were studied in the laboratory. Whirling, in which these web-building pholcid spiders move their bodies rapidly around in circles, with legs remaining on the silk, is the most frequent response of adults to disturbance. Experiments were carried out to determine what types of stimuli elicited whirling. Forceful tactile and air movement stimuli elicited whirling, but there was no evidence that chemical stimuli from potential predators were important. Tactile stimulation was more effective than air movement. The behaviour of these pholcids was compared to that of a cosmopolitan pholcid, Pholcus phalangioides .     

Wasp Attacks and Spider Defence in the Orb Weaving Species <Emphasis Type="Italic">Zygiella x-notata</Emphasis>

Predator–prey relationships are generally based on arm-race. Wasps and spiders are both predators, which could be potential prey for each other. The orb weaver spider Zygiella x-notata is sometimes a prey for the wasp Vespula germanica. We observed the wasp hunting behaviour under natural conditions, and we tested the influence of the spider’s behaviour on the wasp attack success. Wasps were active predators during the reproductive period of the spider. Results showed that wasps located more easily male spiders than females particularly when they were engaged in mate guarding. Female location depended on the presence of a web, but also of prey or prey remains in the web. On the other hand, their location depend neither on the characteristics and the position of the retreat in the environment nor on the size of the web. After location, males were more often captured than females whatever their behaviour (mate guarding or not). Presence of prey remains or prey in the web did not increase the risk for the spider to be captured. There was also no influence of the retreat’s characteristics or of its position in the habitat on the risk for the spider to be captured; but wasp successful attacks were less numerous when silk was present around the entrance of the retreat or when the spider was completely inside. As prey and prey remains favoured location of spiders by the wasps, we tested spider web cleaning behaviour as a response to wasp predatory pressure. By throwing small polystyrene pellets in the webs, we observed that more 80% of the spiders rejected the pellets in less than one minute. Our data indicated that wasps were significant predators of Z. x-notata and wasp attack could have been a selective pressure that had favoured spider defensive behaviours such as web cleaning.     

Asymmetry in spider orb webs: a result of physical constraints?

A typical feature of most vertical orb webs is that the upper web region is smaller and contains less silk than the lower web region, creating an asymmetrical web. The degree of web asymmetry changes during the spider's development: small juveniles construct more symmetrical webs, but older and larger individuals decrease the upper web region. This implies that weight may control the extent of web asymmetry. Using two species, Argiope keyserlingi and Larinioides sclopetarius, we tested the effect of weight increase on web asymmetry by naturally increasing weight through feeding and by artificially adding lead weights to the abdomen of the spiders. Weight increase (natural or artificial) resulted in more asymmetric webs through a reduction of the upper web region. Added weight may interfere with spiral placement in the upper region, because the spider has to lift its abdomen above the carapace during the process. In the lower region, however, the position of the spider is mostly head up during spiral placement. Therefore, amongst other factors, weight and gravitational forces may be physical constraints during web construction. Copyright 1999 The Association for the Study of Animal Behaviour.     

Effect of abiotic factors on the foraging strategy of the orb‐web spider Argiope keyserlingi (Araneae: Araneidae)

Abstract Environmental conditions such as light level, background contrast and temperature might influence a spider's prey capture success and risk of predation. Thus it may often be advantageous for spiders to adjust web‐building behaviour in response to variation in these environmental conditions. This hypothesis was examined in a study of the construction of webs and web decorations (conspicuous strands of silk at the hub of the web) of the orb‐web spider Argiope keyserlingi. Web decorations are thought to have one or more separate functions. They may attract prey, deter predators or advertise the web to oncoming birds, thus preventing web damage. In this series of experiments, relationships between weather parameters and the construction of webs and web decorations were considered. In complementary laboratory experiments, A. keyserlingi spiders were exposed to two different light levels (700 and 90 lx), background contrasts (black and white) and temperature conditions (20 and 26°C). Of the available weather parameters, only temperature was significantly related to web decorating behaviour but not to web size. In the laboratory, temperature also influenced web‐decorating behaviour, and spiders in dim light (700 lx) constructed larger webs and longer decorations. Background contrast did not significantly alter web size or web decorations. These data suggest that when prey availability is reduced at low temperatures, spiders may use web decorations to attract prey to the web. Similarly, in dim light, spiders may build more and larger decorations to increase the visual signal to approaching prey or to advertise the web to oncoming birds.     

Review Behaviour and indirect interactions in food webs of plant-inhabiting arthropods

With the increased use of biological control agents, artificial food webs are created in agricultural crops and the interactions between plants, herbivores and natural enemies change from simple tritrophic interactions to more complex food web interactions. Therefore, herbivore densities will not only be determined by direct predator–prey interactions and direct and indirect defence of plants against herbivores, but also by other direct and indirect interactions such as apparent competition, intraguild predation, resource competition, etc. Although these interactions have received considerable attention in theory and experiments, little is known about their impact on biological control. In this paper, we first present a review of indirect food web interactions in biological control systems. We propose to distinguish between numerical indirect interactions, which are interactions where one species affects densities of another species through an effect on the numbers of an intermediate species and functional indirect interactions, defined as changes in the way that two species interact through the presence of a third species. It is argued that functional indirect interactions are important in food webs and deserve more attention. Subsequently, we discuss experimental results on interactions in an artificial food web consisting of pests and natural enemies on greenhouse cucumber. The two pest species are the two-spotted spider mite Tetranychus urticae and the western flower thrips, Frankliniella occidentalis. Their natural enemies are the predatory mite Phytoseiulus persimilis, which is commonly used for spider mite control and the predatory mites Neoseiulus cucumeris and Iphiseius degenerans and the predatory bug Orius laevigatus, all natural enemies of thrips. First, we analyse the possible interactions between these seven species and we continue by discussing how functional indirect interactions, particularly the behaviour of arthropods, may change the significance and impact of direct interactions and numerical indirect interactions. It was found that a simple food web of only four species already gives rise to some quite complicated combinations of interactions. Spider mites and thrips interact indirectly through resource competition, but thrips larvae are intraguild predators of spider mites. Some of the natural enemies used for control of the two herbivore species are also intraguild predators. Moreover, spider mites produce a web that is subsequently used by thrips to hide from their predators. We discuss these and other results obtained so far and we conclude with a discussion of the potential impact of functional indirect and direct interactions on food webs and their significance for biological control.     

Predators induce interspecific herbivore competition for food in refuge space

Resource competition among herbivorous arthropods has long been viewed as unimportant because herbivore populations are controlled by predators. Although recently resurrected as an organizing force in arthropod communities on plants, there is still general agreement that resource competition among herbivores is reduced by predators. Here we show the reverse: predators induce interspecific resource competi-tion among herbivores. We found that thrips larvae ( Frankliniella occidentalis ) use the web produced and inhabited by the spider mite Tetranychus urticae as a refuge from predation by the phytoseiid mite Neoseiulus cucumeris . Thrips larvae prefer clean plant parts, but move into the web upon perceiving volatile cues associated with thrips and predators. This behaviour leads to lower predation risk, but also to reduced developmental rate and lower production of thrips larvae due to competition with spider mites. In addition, thrips larvae consume spider-mite eggs. Thus, predators induce interspecific competition and intraguild predation among herbivores within refuge space, even when host plants have an overall green appearance.     

Spider mite web mediates anti-predator behaviour

Herbivores suffer significant mortality from predation and are therefore subject to natural selection on traits promoting predator avoidance and resistance. They can employ an array of strategies to reduce predation, for example through changes in behaviour, morphology and life history. So far, the anti-predator response studied most intensively in spider mites has been the avoidance of patches with high predation risk. Less attention has been given to the dense web produced by spider mites, which is a complex structure of silken threads that is thought to hinder predators. Here, we investigate the effects of the web produced by the red spider mite, Tetranychus evansi Baker & Pritchard, on its interactions with the predatory mite, Phytoseiulus longipes Evans. We tested whether female spider mites recognize predator cues and whether these can induce the spider mites to produce denser web. We found that the prey did not produce denser web in response to such cues, but laid more eggs suspended in the web, away from the leaf surface. These suspended eggs suffered less from predation by P. longipes than eggs that were laid on the leaf surface under the web. Thus, by altering their oviposition behaviour in response to predator cues, females of T. evansi protect their offspring.     

Habitat selection in a large orb‐weaving spider: vegetational complexity determines site selection and distribution

1. The distribution of the large orb‐weaving spider Argiope trifasciata in old field habitats of North America and the habitat selection process this species used was studied for 2 years. 2. Because web spiders have limited dispersal abilities and an energetically costly prey capture device, they do not have the ability to sample potential foraging sites. Structural complexity of the vegetation to which the web must be attached is relatively easy to assess. The hypothesis that the structural complexity is a primary factor in determining initial web site selection was tested both by relating the natural distribution of the spiders across habitats to vegetational complexity and by manipulating the complexity of the habitats in a series of experiments. 3. Argiope trifasciata was not distributed evenly among three old field vegetation types. Habitat complexity was related to spider density in both years although no measure of insect activity, prey capture, or prey consumption was correlated with spider distribution. 4. Three experimental manipulations were conducted to test the impact of habitat structure on spider establishment: (1) the amount of natural vegetation was reduced, (2) structures were added to a simple habitat, and (3) the complexity of the structures added was varied. In each case, spiders were introduced and establishment of webs was monitored. In all manipulations, spider establishment was related to the complexity of the substrate available. 5. These results are important for understanding the cues that influence foraging site selection and therefore provide insight into the distribution of species with limited dispersal abilities and high site investment requirements.     

Allocating time between feeding, resting and moving by the two-spotted spider mite, Tetranychus urticae and its predator Phytoseiulus persimilis

This study characterizes the timing of feeding, moving and resting for the two-spotted spider mite, Tetranychus urticae Koch and a phytoseiid predator, Phytoseiulus persimilis Athias-Henriot. Feeding is the interaction between T. urticae and plants, and between P. persimilis and T. urticae. Movement plays a key role in locating new food resources. Both activities are closely related to survival and reproduction. We measured the time allocated to these behaviours at four ages of the spider mite (juveniles, adult females immediately after moult and adult females 1 and 3 days after moult) and two ages of the predatory mite (juveniles and adult females). We also examined the effect of previous spider mite-inflicted leaf damage on the spider mite behaviour. Juveniles of both the spider mite and the predatory mite moved around less than their adult counterparts. Newly emerged adult female spider mites spent most of their time moving, stopping only to feed. This represents the teneral phase, during which adult female spider mites are most likely to disperse. With the exception of this age group, spider mites moved more and fed less on previously damaged than on clean leaves. Because of this, the spider mite behaviour was initially more variable on damaged leaves. Phytoseiulus persimilis rested at all stages for a much larger percentage of the time and spent less time feeding than did T. urticae; the predators invariably rested in close proximity to the prey. Compared to adult predators, juveniles spent approximately four times as long handling a prey egg. The predator-prey interaction is dependent upon the local movement of both the predators and prey. These details of individual behaviours in a multispecies environment can provide an understanding of population dynamics.     

Habitat selection and web plasticity by the orb spider Argiope keyserlingi (Argiopidae): Do they compromise foraging success for predator avoidance?

Abstract Orb web spiders face a dilemma: forage in open habitats and risk predation or forage in closed habitats to minimize risk but at reduced foraging profitability. We tested whether Argiope keyserlingi opts for safer habitats at the expense of foraging success by (i) determining habitat selection indices in open and closed habitats; (ii) marking and releasing individual juvenile, subadult and adults over two 4‐week periods to determine if life‐history stage influences habitat selection; and (iii) determining the biotic and abiotic environmental parameters that relate to A. keyserlingi abundance. We found that A. keyserlingi selected closed habitats. Sedge and anthropogenic structures were selected and trees were avoided. Juveniles were never found in open habitats, most likely because of high postdispersal mortality. Subadults and adults may shift from closed to open habitats while juveniles never shifted habitat. Foliage density, plant height, potential prey abundance, and mantid and bird abundance were correlated with A. keyserlingi abundance, with only bird abundance explaining habitat selection. We measured web capture area, spiral distance (distance between spiral threads) and the number of decoration arms (0, 1, 2, 3 or 4) in the field and did laboratory experiments to test the influence of (i) space and vegetation; (ii) prey abundance; and (iii) web damage, on web architecture. Argiope keyserlingi webs exhibited geometric plasticity by having larger prey capture areas and spiral distances in open habitats. Decoration design did not differ between habitats however. Variation in space availability, air temperature, prey abundance and web damage explained the variations in web architecture. Potential prey size and diversity differed between habitats but prey abundance did not. As large prey may be important for spider survivorship, foraging success appears to be compromised by occupying closed habitats.     

Habitat structure mediates top–down effects of spiders and ants on herbivores

Differences in structural complexity of habitats have been suggested to modify the extent of top–down forces in terrestrial food webs. In order to test this hypothesis, we manipulated densities of generalist invertebrate predators and the complexity of habitat structure in a two-factorial design. We conducted two field experiments in order to study predation effects of ants and spiders and, in particular, of the wasp spider Argiope bruennichi on herbivorous arthropods such as grasshoppers, plant- and leafhoppers in a grassland. Predator densities were manipulated by removal in habitats of higher and lower structural diversity, and the effects on herbivore densities were assessed by suction sampling. Habitat structure was changed by cutting the vegetation to half its height and removing leaf litter.We found a significant negative effect of this assemblage of generalist predators on plant- and leafhoppers, which were 1.6 times more abundant in predator removal plots. This effect was stronger in low-structured (cut) than in uncut vegetation. Densities of the most abundant planthopper Ribautodelphax pungens (Delphacidae) were 2.2 times higher in predator removal plots. Furthermore, adult plant- and leafhoppers responded more strongly than juveniles and epigeic species more strongly than hypergeic species. The presence of predators had a positive effect on plant- and leafhopper species diversity. In a second field experiment, we tested the exclusive impact of Argiope bruennichi on its prey, and found that its effect was also significant, although weaker than the effect of the predator assemblage. This effect was stronger in grass-dominated vegetation compared to structurally more complex mixed vegetation of grasses and herbs. We conclude that habitat structure and in particular vegetation height and architectural complexity strongly modify the strength of top–down forces and indirectly affect the diversity of herbivorous arthropods.     

Does individual internal state affect the presence of a barrier web in Argiope bruennichii (Araneae: Araneidae)?

A barrier web created by some orb-web spiders has a three-dimensional web structure made of nonsticky threads, which is believed to provide a defensive function against predators. To test for a possible antipredatory function, we investigated relationships between the presence of a barrier web and an individuals internal state in the orb-web spider Argiope bruennichii, which builds orb-webs with and without a barrier web. Hunger level and molting as measures of internal state were represented by relative abdominal width and relative web size, respectively. Field survey showed that individuals with a wider abdomen and a smaller web tended to construct a barrier web. The positive relationship between body width and the presence of a barrier web supports the hypothesis of an antipredatory function as well-fed individuals are likely to expend more effort for defense. The small web size, however, may not be representative of molting because a correlation existed between web size and potential web substrates; therefore, the effect of molting on the presence of barrier web remains unclear.     

Anemotactic responses of the predatory mite,Phytoseiulus persimilis Athias-Henriot,and their role in prey finding

Although it is well established that the predatory mite Phytoseiulus persimilis Athias-Henriot responds to odour emanating from leaves infested by its phytophagous prey, the two-spotted spider mite Tetranychus urticae Koch, little is known of the behavioural mechanisms elicited by odour perception and how they contribute to finding the prey. In this paper the influence of prey-related odour on orientation to wind direction is discussed. It was analysed by observing the predator's walking paths in still air and in an air stream uniformly permeated either with or without prey-related odour stimuli. The results show that well-fed predator females move upwind in presence of these stimuli, but downwind otherwise. Starved predators always move upwind. The anemotactic responses observed are therefore both odour-conditioned and (feeding) state-dependent.In an attempt to explain these responses it is argued that the anemotactic responses of well-fed predators may contribute to arrestment within the area marked by a cluster of prey-colonized leaves. The anemotactic responses of starved predators may help them to find clusters of spider mite colonies located upwind. Why predatory mites also move upwind in absence of prey-related odour stimuli, is a question that remains to be answered.     

Absence of odour-mediated avoidance of heterospecific competitors by the predatory mite Phytoseiulus persimilis

Arthropods use odours associated with the presence of their food, enemies and competitors when searching for patches. Responses to these odours therefore determine the spatial distribution of animals, and are decisive for the occurrence and strength of interactions among species. Therefore, a logical first step in studying food web interactions is the analysis of behaviour of individuals that are searching for patches of food. We followed this approach when studying interactions in an artificial food web occurring on greenhouse cucumber in the Netherlands. In an earlier paper we found that one of the predators of the food web, the predatory mite Phytoseiulus persimilis Athias-Henriot, used to control spider mites, discriminates between odours from plants with spider mites, Tetranychus urticae Koch, and plants with spider mites plus conspecific predators. The odours used for discrimination are produced by adult prey in response to the presence of predators, and probably serve as an alarm pheromone to warn related spider mites. Other predator species may also trigger production of this alarm pheromone, which P. persimilis could use in turn to avoid plants with heterospecific predators. We therefore studied the response of the latter to odours from plants with spider mites and 3 other predator species, i.e. the generalist predatory bug Orius laevigatus (Fieber), the polyphagous thrips Frankliniella occidentalis and the spider-mite predator Neoseiulus californicus (McGregor). Both olfactometer and greenhouse release experiments yielded no evidence that P. persimilis avoids plants with any of the 3 heterospecific predators. This suggests that these predators do not elicit production of alarm pheromones in spider mites, and we argue that this is caused by a lack of coevolutionary history. The consequences of the lack of avoidance of heterospecific predators for interactions in food webs and biological control are discussed.     

The sterile male technique: Irradiation negatively affects male fertility but not male courtship

The sterile male technique is a common method to assign paternity, widely adopted due to its relative simplicity and low cost. Male sterility is induced by exposure to sub lethal doses of chemosterilants or irradiation, the dosage of which has to be calibrated for every species to provide successful male sterilisation, without affecting male physiology and behaviour. While the physiological effects of sterilisation are usually assessed for each study, the behavioural ones are rarely analysed in detail. Using the orb web spider Argiope keyserlingi as a model we first tested (1) the validity of the thread assay, which simulates male courtship behaviour in a standardised context, as a proxy representing courtship on a female web. We then investigated (2) the effectiveness of male sterilisation via irradiation and (3) its consequences on male courtship behaviour. Our results validate the thread assay and the sterile male technique as legitimate tools for the study of male courtship behaviour and fertilisation success. We show that these techniques are time and cost effective and reduce undesirable variation, thereby creating opportunities to study and understand the mechanisms underlying sexual selection.     

Responses of the spider Argiope aurantia to low frequency phasic and continuous vibrations

I maintained female orb-weaving spiders, Argiope aurantia, in the laboratory and subjected them to low frequency phasic and continuous transverse vibrations of known frequency, amplitude, and position on the web. Stimuli were administered for periods of 1 min and all behaviour recorded. Pumping was associated with tactile contact to the spider's abdomen. Pumping increased with increasing frequency of phasic stimuli delivered to the web surface. Attack increased linearly with the log of stimulus frequencies between 0 and 14 Hz for stimuli delivered to the catching spiral. Dropping occurred only when high frequencies and amplitudes of stimuli were administered to the spider's abdomen and the hub of the web.