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.     

Predator perception of detritus and eggsac decorations spun by orb-web spiders Cyclosa octotuberculata: Do they function to camouflage the sniders?

Camouflage is one of the most widespread and powerful strategies that animals use to make detection/recognition more difficult. Many orb-web spiders of the genus Cyclosa add prey remains, plant debris, moults, and/or eggsacs to their webs called web decorations. Web decorations resembling spider body colour pattern have been considered to camouflage the spider from predators. While this camouflage is obvious from a human's perspective, it has rarely been investigated from a predator's perspective. In this study, we tested the visibility of web decorations by calculating chromatic and achromatic contrasts of detritus and eggsac decorations built by Cyclosa octotuberculata, against four different backgrounds viewed by both bird (e.g., blue tits) and hymenopteran (e.g. Wasps) predators. We showed that both juvenile and adult spiders on webs with detritus or egg-sac deco-rations were undetectable by both hymenopteran and bird predators over short and long distances. Our results thus suggest that decorating webs with detritus or eggsacs by C. Octotuberculata may camouflage the spiders from both hymenopteran and bird predators in their common habitats.     

Predator hunting mode influences patterns of prey use from grazing and epigeic food webs

Multichannel omnivory by generalist predators, especially the use of both grazing and epigeic prey, has the potential to increase predator abundance and decrease herbivore populations. However, predator use of the epigeic web (soil surface detritus/microbe/algae consumers) varies considerably for reasons that are poorly understood. We therefore used a stable isotope approach to determine whether prey availability and predator hunting style (active hunting vs. passive web-building) impacted the degree of multichannel omnivory by the two most abundant predators on an intertidal salt marsh, both spiders. We found that carbon isotopic values of herbivores remained constant during the growing season, while values for epigeic feeders became dramatically more enriched such that values for the two webs converged in August. Carbon isotopic values for both spider species remained midway between the two webs as values for epigeic feeders shifted, indicating substantial use of prey from both food webs by both spider species. As the season progressed, prey abundance in the grazing food web increased while prey abundance in the epigeic web remained constant or declined. In response, prey consumption by the web-building spider shifted toward the grazing web to a much greater extent than did consumption by the hunting spider, possibly because passive web-capture is more responsive to changes in prey availability. Although both generalist predator species engaged in multichannel omnivory, hunting mode influenced the extent to which these predators used prey from the grazing and epigeic food webs, and could thereby influence the strength of trophic cascades in both food webs.     

Intra-guild predation relaxes natural enemy impacts on herbivore populations

Abstract. 1. To investigate the role of intra-guild predation in mediating the impact of the natural enemy complex on herbivore populations, a manipulative field experiment was conducted using uncaged plots (islets of Spartina cordgrass) on a North American salt marsh. The densities (moderate or low) of two invertebrate predators, the generalist wolf spider Pardosa littoralis and the specialist mirid bug Tytthus vagus , were manipulated in a 2 × 2 factorial design, and the resulting treatment effects on the population growth of their herbivorous prey, Prokelisia planthoppers, were assessed.
2. The abundance of wolf spiders on experimental islets was unaffected by the presence of mirid bugs, however the density of mirid bugs was influenced very negatively by the presence of the wolf spider.
3. The negative effect of the wolf spider on mirid bugs most probably resulted from the intra-guild predation of mirids by spiders because planthopper limitation by the wolf spider alone was significantly greater than when both predators were present.
4. As a result of intra-guild predation, planthopper population growth was positive in the presence of both predators, despite the fact that each predator alone promoted a decrease in planthopper population growth.
5. Notably, the occurrence of intra-guild predation diminished top-down impacts on planthopper populations in a relatively simple food web where strong top-down effects were expected. This result, however, was limited to habitats on the marsh with simply structured vegetation lacking leaf litter.     

Conspicuous colouration attracts prey to a stationary predator

Abstract 1. Conspicuous body colouration is counter‐intuitive in stationary predators because sit‐and‐wait tactics frequently rely on concealed traps to capture prey. Consequently, bright colours and contrasting patterns should be rare in predators using traps as they may alert potential prey. Yet, some orb‐weaving spiders are brightly coloured and contrastingly patterned. How can conspicuousness of trap‐building sit‐and‐wait predators be favoured by natural selection? 2. Observations of spiny spiders Gasteracantha fornicata in north‐eastern Australia showed that the size of spiders relative to their orb webs correlated positively with relative prey numbers already captured in their webs. A possible explanation is that the relatively larger appearance of the yellow–black striped dorsal surface of this spider attracts more visually oriented prey items. Prey attracted to webs may get trapped, thereby increasing the spiders' foraging success. 3. To test this hypothesis for the function of conspicuous body colouration, a field experiment was conducted that documented the prey capture rates of spiny spiders after manipulating or sham‐manipulating their appearance. 4. As predicted, spiders that were dyed black on their striped dorsal surface caught relatively fewer prey items than did control spiders. Thus, conspicuous dorsal body colouration may be adaptive in spiny spiders because it increases foraging success and, presumably, survival rates and reproductive outputs. Overall, these data support the colour‐as‐prey‐attractant hypothesis in a stationary, trap‐building predator.     

Small-scale spatial pattern of web-building spiders (Araneae) in alfalfa: relationship to disturbance from cutting, prey availability, and intraguild interactions

Understanding the development of spatial patterns in generalist predators will improve our ability to incorporate them into biological control programs. We studied the small-scale spatial patterns of spider webs in alfalfa by analyzing the relationship between web locations over distances ranging from 4 to 66 cm. Using a coordinate-based spatial statistic (O-ring) and assuming a heterogeneous distribution of suitable web sites, we analyzed the impact of cutting and changes in spider abundance on web distribution. We analyzed the influence of small-scale variation in prey availability by comparing web distributions to the pattern of sticky-trap captures of Aphididae and Diptera described by a count-based spatial statistic (SADIE). Cutting of alfalfa reduced the overall density of web-building spiders but had no immediate impact on the spatial distribution of their webs. Availability of aphids was highest before the alfalfa was cut and was clumped at a scale of 66 cm. Spider webs, however, were not clumped at any scale or date. In contrast, webs were regularly distributed at smaller distances (<20 cm) immediately before and after cutting. Because cursorial and web-building spiders were most active during this period, we hypothesize that the development of small-scale regularity in web locations was driven by intraguild interactions. Our results suggest that intraguild interactions contribute to the development of small-scale spatial patterns of spider webs in alfalfa. Variation in prey availability may have more of an influence on web distribution in crops with a different vegetation structure or if patterns are studied at larger spatial scales.     

Previous foraging success influences web building in the spider Stegodyphus lineatus (Eresidae)

Stegodyphus lineatus (Eresidae) is a desert spider that buildsan aerial capture web on bushes in the Negev desert in southernIsrael. Web building for spiders is costly in energy, time,and risk of predation. Spiders should trade-off these costswith the benefits in terms of prey capture. We tested the hypothesisthat the previous foraging success of the spider influencesthe effort invested in foraging. Specifically, we asked whetheran increase in food intake causes spiders to reduce web renewalactivity and web size. Alternatively, time constraints on foragingand development, resulting from a short growing season, couldinduce spiders to continue foraging even when supplemented withprey. The cost of web building was measured as time and massloss. To build an average size web (about 150 cm²), we calculatedthat a spider requires 6 h and that spiders lose 3%-7% of their weight.In field experiments, spiders responded differently to food supplementationin 2 different years. In 1994, they improved their condition comparedto individuals whose webs were removed to reduce foraging opportunitiesand compared to control spiders. In 1995, spiders tested earlier inthe season than the previous year did not improve their conditionin response to prey supplementation. Nonetheless, in both years, food-supplementedspiders built significantly smaller webs than food-deprived andcontrol spiders. This result was confirmed in a laboratory experiment whereprey intake was controlled. We conclude that for S. lineatus immediateforaging risks outweigh the potential time constraints on foraging.     

Web tuning of an orb-web spider, Octonoba sybotides, regulates prey-catching behaviour

An uloborid spider (Oclonoba sybotides constructs two types of web which are distinguished by linear or spiral stabilimenta. Food-deprived spiders tend to construct webs with spiral stabilimenta and food-satiated spiders tend to construct webs with linear stabilimenta. I experimentally examined the influence of web type on the speed of a spider's response to small and large flies. The results indicated that web type rather than the spiders' energetic condition influences the response speed to small or large Drosophila flies. I also examined whether thread tension affects the response speed of spiders by increasing the tension of the radial threads. The results showed that spiders on an expanded web responded to small prey as quickly as spiders on webs with spiral stabilimenta. The tension of the radial threads may be regulated by the degree of distortion of the radial threads at the hub. O. sybotides seems to construct orb webs which induce different responses for smaller, less-profitable prey according to its energetic state. The spider appears to increase the tension of the radial threads so that it can sense smaller prey better when hungry.     

SIGNAL POLYMORPHISM IN THE WEB-DECORATING SPIDER ARGIOPE ARGENTATA IS CORRELATED WITH REDUCED SURVIVORSHIP AND THE PRESENCE OF STINGLESS BEES, ITS PRIMARY PREY

Abstract.— Many spiders, and in particular those in the genus Argiope, spin highly visible web decorations whose function and significance are the subject of spirited debate. In this work, we present data to address two of the competing hypotheses that fuel this controversy. In particular, we examine the relationship between the presence of web decorations and spider survivorship (predator‐protection hypothesis) and the relationship between the presence of prey and spider decorating behavior (the prey‐attraction hypothesis). Our laboratory studies reveal that the decorating behavior of the spider A. argentata has a genetic component but that the expression of decorating behavior tends to be elicited only when a spider is well fed. Furthermore, our field studies show that in the presence of abundant stingless bees, spider decorating behavior is induced. Nevertheless, our field surveys also suggest that spiders that decorate their webs show reduced survivorship. We propose that the high correlation between web decorating in the presence of stingless bees supports the hypothesis that A. argentata engage in decorating behavior when attracting or targeting specific prey types. However, we also propose that web decorations attract the predators of A. argentata because high‐frequency decorators suffer lower survivorship than spiders that decorate moderately or rarely. These findings suggest that spider web decorating behavior is affected by conflicting selection pressures: the positive effect of prey attraction versus the negative effect of predator attraction. Due to the heritable component of decorating behavior, web decorating among A. argentata is likely to be particularly sensitive to the spider's local ecology as well as local patterns of gene flow.     

Local adaptation to temperature conserves top-down control in a grassland food web

A fundamental limitation in many climate change experiments is that tests represent relatively short-term 'shock' experiments and so do not incorporate the phenotypic plasticity or evolutionary change that may occur during the gradual process of climate change. However, capturing this aspect of climate change effects in an experimental design is a difficult challenge that few studies have accomplished. I examined the effect of temperature and predator climate history in food webs composed of herbaceous plants, generalist grasshopper herbivores and spider predators across a natural 4.8°C temperature gradient spanning 500 km in northeastern USA. In these grasslands, the effects of rising temperatures on the plant community are indirect and arise via altered predator-herbivore interactions. Experimental warming had no direct effect on grasshoppers, but reduced predation risk effects by causing spiders from all study sites to seek thermal refuge lower in the plant canopy. However, spider thermal tolerance corresponded to spider origin such that spiders from warmer study sites tolerated higher temperatures than spiders from cooler study sites. As a consequence, the magnitude of the indirect effect of spiders on plants did not differ along the temperature gradient, although a reciprocal transplant experiment revealed significantly different effects of spider origin on the magnitude of top-down control. These results suggest that variation in predator response to warming may maintain species interactions and associated food web processes when faced with long term, chronic climate warming.     

The multiple disguises of spiders: web colour and decorations, body colour and movement

Diverse functions have been assigned to the visual appearance of webs, spiders and web decorations, including prey attraction, predator deterrence and camouflage. Here, we review the pertinent literature, focusing on potential camouflage and mimicry. Webs are often difficult to detect in a heterogeneous visual environment. Static and dynamic web distortions are used to escape visual detection by prey, although particular silk may also attract prey. Recent work using physiological models of vision taking into account visual environments rarely supports the hypothesis of spider camouflage by decorations, but most often the prey attraction and predator confusion hypotheses. Similarly, visual modelling shows that spider coloration is effective in attracting prey but not in conveying camouflage. Camouflage through colour change might be used by particular crab spiders to hide from predator or prey on flowers of different coloration. However, results obtained on a non-cryptic crab spider suggest that an alternative function of pigmentation may be to avoid UV photodamage through the transparent cuticle. Numerous species are clearly efficient locomotory mimics of ants, particularly in the eyes of their predators. We close our paper by highlighting gaps in our knowledge.     

Effects of subsidized spiders on coastal food webs in the Baltic Sea area

The aim of this study was to examine top-down effects of cursorial spiders in subsidized coastal food webs. Top-down effects were examined by selectively removing cursorial spiders, mainly wolf spiders, from small islands (26–1834 m²) during 2004–2007. The removal success varied among islands and years, and spider densities were reduced by 30–65%. To examine treatment effects, arthropods were sampled using a vacuum sampling device at three occasions each summer. The densities of other arthropod predators, especially web spiders and carabids, were higher on islands where cursorial spiders had been removed compared to control islands. This treatment effect probably occurred through a combination of competitive release and reduced intraguild predation from cursorial spiders. No treatment effects were found on herbivore or detritivore densities and plant biomass. This lack of effect may either be because spiders indeed have fairly weak effects on herbivore and detritivore densities on Baltic shorelines or that the removal success of spiders was insufficient for observing such effects. Treatment effects may also be weak because negative effects exerted by spiders on herbivore and detritivore populations were balanced by increased predation by insect predators.     

Chemical Mediation of Prey Recognition by Spider-Hunting Wasps

Predator–prey interactions are important in maintaining the structure and dynamics of ecological communities. Both predators and prey use cues from a range of sensory modalities to detect and assess one another; identification of these cues is necessary to understand how selection operates to shape predator–prey interactions. Mud-dauber wasps (Sphecidae) provision their larval nests with paralyzed spiders, and different genera of wasps specialize on particular spider taxa. Sceliphron caementarium (Drury 1773) wasps preferentially capture spiders that build two-dimensional (2D) webs, rather than those that construct three-dimensional (3D) webs, but the basis of this preference is not clear. Wasps may choose spiders based on an assessment of their web architecture, as 3D webs may provide better defenses against wasp predation than do 2D webs. However, because many hymenopterans use chemical cues to locate and recognize prey, it is also possible that mud-dauber wasps rely on chemical cues associated with the spider and/or the web to assess prey suitability. When we offered foraging S. caementarium wasps 2D and 3D spiders both on and off their webs, we found that in both cases the wasps took 2D spiders and avoided 3D spiders, demonstrating that the web itself is not the impediment. Results of a series of behavioral choice assays involving filter paper discs containing spider cues and chemically manipulated spiders or spider dummies corroborated the importance of spider chemical cues in mediation of prey recognition by mud-dauber wasps. We also discuss the relative importance of visual and chemical cues for prey recognition by wasps, examine the anti-predator behaviors of 2D and 3D spiders, and consider the role of wasp predation in spider diversification.     

Impact of avian and arthropod predation on lepidopteran caterpillar densities and plant productivity in an ephemeral agroecosystem

Abstract.  1. Most studies evaluating the combined impact of spiders and other predators on herbivore densities in agroecosystems have focused primarily on their trophic connections with invertebrate predators (e.g. carabids, chrysopids); however linkages among spiders and vertebrate predators may also help structure the population dynamics of insect herbivores. A field experiment was conducted to examine the impact of avian and spider predation on lepidopteran caterpillar densities and plant productivity within a Brassica agroecosystem.
2. Arthropod abundance, leaf-chewing damage, and final plant productivity associated with broccoli, Brassica oleracea L. (var. italica ), were recorded for four treatments: (1) bird present but spiders removed; (2) both birds and spiders present; (3) birds excluded, spiders present; and (4) birds and spiders both excluded.
3. Densities of Artogeia rapae L. (Lepidoptera: Pieridae) and Trichoplusia ni Hübner (Lepidoptera: Noctuidae) large caterpillars and post feeding stages were reduced significantly by bird predation. The abundance of large caterpillars was also reduced on spider-inhabited plants during early plant growth; however the assemblage of birds and spiders did not suppress caterpillar densities more significantly than either predator alone.
4. Plants protected by birds, spiders, and birds plus spiders sustained less folivory attributable to leaf chewing caterpillars than check plants. Plant productivity was also greater for predator-protected plants than check plants.
5. Although spiders and parasitoids were responsible for some of the mortality inflicted upon lepidopteran caterpillars, it was concluded that in this study system, birds are the most important natural enemies of folivores.     

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.     

Effects of ant competition and bird predation on the spider assemblage of a citrus grove

Characterizing intraguild interactions is key to improving understanding of food webs because they are major forces in the structuring of communities. Spiders are generalist predators with intermediate positions in the food web that establish intraguild interactions with ants and birds, which respectively compete with and prey on them. Research has also found interactions between birds and ants, potentially resulting in non-additive effects of both groups on arthropod assemblages, although studies of their combined impacts with tests for multiple-predator effects are scarce. We thus aimed to discern the relative effect of ants and birds on the spider assemblage of a citrus grove. We used a split-plot design to factorially exclude these groups over 2 years, preventing ants reaching the canopies by placing sticky bands around tree trunks, and birds by enclosing groups of trees in cages. We sampled spiders from the canopies (beating) and the ground (pitfalls) every 3 months, and we identified them to species. We found a strong influence of ants on the canopy spider assemblage, mainly through a negative effect on the families Araneidae and Theridiidae. Since spiders’ weights from ant-excluded and control trees were similar, these results suggest interference competition of ants on spiders rather than competitive exploitation. Bird exclusion did not affect the spider assemblage, contrasting with other studies reporting a marked predatory pressure of birds on spiders; nor were there any non-additive effects of ants and birds. Our findings show that spider assemblages are not uniformly affected by intraguild competitors.     

Behavioural and biomaterial coevolution in spider orb webs

Mechanical performance of biological structures, such as tendons, byssal threads, muscles, and spider webs, is determined by a complex interplay between material quality (intrinsic material properties, larger scale morphology) and proximate behaviour. Spider orb webs are a system in which fibrous biomaterials—silks—are arranged in a complex design resulting from stereotypical behavioural patterns, to produce effective energy absorbing traps for flying prey. Orb webs show an impressive range of designs, some effective at capturing tiny insects such as midges, others that can occasionally stop even small birds. Here, we test whether material quality and behaviour (web design) co‐evolve to fine‐tune web function. We quantify the intrinsic material properties of the sticky capture silk and radial support threads, as well as their architectural arrangement in webs, across diverse species of orb‐weaving spiders to estimate the maximum potential performance of orb webs as energy absorbing traps. We find a dominant pattern of material and behavioural coevolution where evolutionary shifts to larger body sizes, a common result of fecundity selection in spiders, is repeatedly accompanied by improved web performance because of changes in both silk material and web spinning behaviours. Large spiders produce silk with improved material properties, and also use more silk, to make webs with superior stopping potential. After controlling for spider size, spiders spinning higher quality silk used it more sparsely in webs. This implies that improvements in silk quality enable ‘sparser’ architectural designs, or alternatively that spiders spinning lower quality silk compensate architecturally for the inferior material quality of their silk. In summary, spider silk material properties are fine‐tuned to the architectures of webs across millions of years of diversification, a coevolutionary pattern not yet clearly demonstrated for other important biomaterials such as tendon, mollusc byssal threads, and keratin.     

Carcass Decoration Changes Web Structure and Prey Capture Rate in an Orb-Web Spider, Cyclosa mulmeinensis (Araneae, Araneidae)

Cyclosa spiders attach prey carcasses as decorations to their webs, but the functions of the carcasses are unclear and controversial. We investigated distinctive features of these webs in the field and conducted prey-capture experiments in the lab using the orb-web spider Cyclosa mulmeinensis. Webs with attached decoration had a significantly narrower mesh width than those without decoration and a higher degree of vertical asymmetry in the web’s shape. In the laboratory, webs without decorations trapped significantly more prey, even though other features of the webs were nearly identical. These results suggest that web decorations do not attract prey in this species, but might play other roles such as blinding predators to the spider’s presence.     

Web orientation and prey resources for web-building spiders in eastern hemlock

We examined the arthropod community on eastern hemlock, Tsuga canadensis (L.) Carr, in the context of its role in providing potential prey items for hemlock-associated web-weaving spiders. Using sticky traps simulating spider webs, we evaluated what prey items are available to web-weaving spiders in eastern hemlock based on web orientation (horizontal versus vertical) and cardinal direction. We found that the overwhelming majority (>70%) of prey items available to spiders in hemlock canopies were Diptera. Psocoptera, Hymenoptera, and Hemiptera comprised most of the remaining potential prey. A significant direction × orientation interaction, and greater trap capture in some direction-orientation combinations, suggests that spiders might locate their webs in eastern hemlock canopies for thermoregulatory purposes, ultimately optimizing prey capture. We also evaluated these findings in the context of hemlock infestation by the invasive hemlock woolly adelgid, Adelges tsugae Annand. The adelgid is a sedentary insect with a mobile crawler stage that provides a readily available, easily obtained food source for predators in hemlock canopies. However, an abundance of alternative prey will affect within canopy spider distribution and the potential intensity with which spiders consume these prey. Understanding the response of spiders to potential prey availability is essential to understanding the trophic interactions involving these predators and their potential for influencing herbivore populations.     

Foraging Benefits in a Colour Polymorphic Neotropical Orb Web Spider

Conspicuous body colouration in sedentary predators such as orb web spiders seems paradoxical as potential prey can see and avoid the webs. Several studies have demonstrated that rather than deterring prey, the colours act as sensory traps for flower‐seeking insects. In chromatically polymorphic species, the existence of more than one colour morph may lead to differing levels of prey attraction. To explore these issues, we studied a neotropical orb web spider, Verrucosa arenata, which shows colour polymorphism with predominantly white or yellow abdomen colours. We asked whether a particular morph is dominant in the population, and whether a particular morph is associated with enhanced foraging success and body condition. Here we showed that although yellow morphs attracted more prey, white morphs were in better body condition. We showed that model prey such as honeybees are able to discriminate between the morphs. We discuss these findings in relation to the functional significance of bright body colouration and colour polymorphism in spiders.