Resource Management in the Orb-Weaving Spider Zygiella x-notata: I. Influence of Spider Behaviour and Available Prey

Management of resource exploitation by the orb-weaving spider Zygiella x-notata was investigated in relation to available prey. Prey capture is studied according to prey type (cricket or fly), and the spider's behaviour (waiting without prey or consuming other prey), at the moment when a prey item was placed onto the web. Characteristics of the spider's exploitation of the previous item (feeding time, and quantities ingested) were analysed according to the type and quantity of prey presented, (1 or 2 items at 1-h intervals). Capture time did not vary in relation to prey type, nor spider activity. However, duration and quantities of certain behavioural components (biting and wrapping) did vary according to prey type and the spider's behaviour, at the moment of presentation of prey. Whatever the type of prey received, if the first item consumed was a cricket, the spiders reduced time spent feeding on this prey without any decrease in the quantities ingested.     

Are you what you eat? A highly transient and prey‐influenced gut microbiome in the grey house spider Badumna longinqua

Stable core microbial communities have been described in numerous animal species and are commonly associated with fitness benefits for their hosts. Recent research, however, highlights examples of species whose microbiota are transient and environmentally derived. Here, we test the effect of diet on gut microbial community assembly in the spider Badumna longinqua. Using 16S rRNA gene amplicon sequencing combined with quantitative PCR, we analyzed diversity and abundance of the spider's gut microbes, and simultaneously characterized its prey communities using nuclear rRNA markers. We found a clear correlation between community similarity of the spider's insect prey and gut microbial DNA, suggesting that microbiome assembly is primarily diet‐driven. This assumption is supported by a feeding experiment, in which two types of prey—crickets and fruit flies—both substantially altered microbial diversity and community similarity between spiders, but did so in different ways. After cricket consumption, numerous cricket‐derived microbes appeared in the spider's gut, resulting in a rapid homogenization of microbial communities among spiders. In contrast, few prey‐associated bacteria were detected after consumption of fruit flies; instead, the microbial community was remodelled by environmentally sourced microbes, or abundance shifts of rare taxa in the spider's gut. The reshaping of the microbiota by both prey taxa mimicked a stable core microbiome in the spiders for several weeks post feeding. Our results suggest that the spider's gut microbiome undergoes pronounced temporal fluctuations, that its assembly is dictated by the consumed prey, and that different prey taxa may remodel the microbiota in drastically different ways.     

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.     

Repeatability but no short-term plasticity in the web architecture of Latrodectus hesperus western black widow spiders (Araneae: Theridiidae)

A spider's web is an extended phenotype and a direct product of behavior. Variation in web architecture may be the result of individual differences in genotype, environment, or experience. We analyzed variation in the web architecture of western black widow spiders, Latrodectus hesperus, by testing two hypotheses: (a) intrinsic individual differences and (b) plasticity according to experience of the site of prey capture. We used treatments manipulating the site of prey capture experienced by the spiders: sheet only, gum-footed lines only, and alternating patterns between sheet and gum-footed lines equally. We found considerable consistent individual variation (repeatability of r = .56) in one web component (gum-footed lines) and negligible plasticity in either component. We discuss our results in terms of the potential for the design of these web components to respond to selection, and of potential additional causes of variation in extended phenotypes.     

Upside-down spiders build upside-down orb webs: web asymmetry,spider orientation and running speed in Cyclosa

Almost all spiders building vertical orb webs face downwards when sitting on the hubs of their webs, and their webs exhibit an up–down size asymmetry, with the lower part of the capture area being larger than the upper. However, spiders of the genus Cyclosa, which all build vertical orb webs, exhibit inter- and intraspecific variation in orientation. In particular, Cyclosa ginnaga and C. argenteoalba always face upwards, and C. octotuberculata always face downwards, whereas some C. confusa face upwards and others face downwards or even sideways. These spiders provide a unique opportunity to examine why most spiders face downwards and have asymmetrical webs. We found that upward-facing spiders had upside-down webs with larger upper parts, downward-facing spiders had normal webs with larger lower parts and sideways-facing spiders had more symmetrical webs. Downward-facing C. confusa spiders were larger than upward- and sideways-facing individuals. We also found that during prey attacks, downward-facing spiders ran significantly faster downwards than upwards, which was not the case in upward-facing spiders. These results suggest that the spider''s orientation at the hub and web asymmetry enhance its foraging efficiency by minimizing the time to reach prey trapped in the web.     

Bt maize pollen exposure and impact on the garden spider, Araneus diadematus

Concerns have been raised that Bt maize pollen may have adverse effects on non‐target organisms; consequently, there is a general call for Bt maize risk assessment evaluating lethal and sublethal side effects. Spiders play an important economic and ecological role as pest predators in various crops, including maize. Web‐building spiders, especially, may be exposed to the Cry1Ab toxin of Bt maize by the ingestion of pollen via ‘recycling’ of pollen‐dusted webs and intentional pollen feeding. In this study, the potential Bt maize pollen exposure of orb‐web spiders was quantified in maize fields and adjacent field margins, and laboratory experiments were conducted to evaluate the possible effects of Bt maize pollen consumption on juvenile garden spiders, Araneus diadematus (Clerck) (Araneae: Araneidae). In maize fields and neighbouring field margins, web‐building spiders were exposed to high amounts of Bt maize pollen. However, a laboratory bioassay showed no effects of Bt maize pollen on weight increase, survival, moult frequency, reaction time, and various web variables of A. diadematus. A pyrethroid insecticide (Baythroid) application affected weight increase, survival, and reaction time of spiders negatively. In conclusion, the insecticide tested showed adverse effects on the garden spider, whereas the consumption of Bt maize pollen did not. This study is the first one on Bt maize effects on orb‐web spiders, and additional research is recommended in order to account for further spider species, relative fitness parameters, prey‐mediated effects, and possible long‐term chronic consequences of Bt exposure.     

Body size and tree species composition determine variation in prey consumption in a forest‐inhabiting generalist predator

Trophic interactions may strongly depend on body size and environmental variation, but this prediction has been seldom tested in nature. Many spiders are generalist predators that use webs to intercept flying prey. The size and mesh of orb webs increases with spider size, allowing a more efficient predation on larger prey. We studied to this extent the orb‐weaving spider Araneus diadematus inhabiting forest fragments differing in edge distance, tree diversity, and tree species. These environmental variables are known to correlate with insect composition, richness, and abundance. We anticipated these forest characteristics to be a principle driver of prey consumption. We additionally hypothesized them to impact spider size at maturity and expect shifts toward larger prey size distributions in larger individuals independently from the environmental context. We quantified spider diet by means of metabarcoding of nearly 1,000 A. diadematus from a total of 53 forest plots. This approach allowed a massive screening of consumption dynamics in nature, though at the cost of identifying the exact prey identity, as well as their abundance and putative intraspecific variation. Our study confirmed A. diadematus as a generalist predator, with more than 300 prey ZOTUs detected in total. At the individual level, we found large spiders to consume fewer different species, but adding larger species to their diet. Tree species composition affected both prey species richness and size in the spider''s diet, although tree diversity per se had no influence on the consumed prey. Edges had an indirect effect on the spider diet as spiders closer to the forest edge were larger and therefore consumed larger prey. We conclude that both intraspecific size variation and tree species composition shape the consumed prey of this generalist predator.     

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.     

Web-building management in an orb-weaving spider, Zygiella x-notata: influence of prey and conspecifics

According to optimal foraging theory, spiders should adapt their web building to environmental variations. Until now, there was no data on the influence of simultaneous information coming from different environmental factors on web building behaviour. Under laboratory conditions, we studied the behaviour of Zygiella x-notata in the presence of prey, conspecifics, or both simultaneously. There was a stimulating effect of prey, but web building was not affected by the presence of conspecifics. When spiders and prey were present simultaneously, the effect was similar to that of prey alone; it seemed that there was no interactive influence of both factors. We discussed about the use of environmental information by spiders in foraging behaviour.     

Prey Luring Coloration of A Nocturnal Semi‐Aquatic Predator

Body coloration serves a variety of purposes in animals. Diurnal and nocturnal predators such as spiders may use their body coloration to lure prey. We predicted here that the white patches on the forelegs on females of the nocturnal semi‐aquatic spider Dolomedes raptor lure prey, explaining why they are primarily displayed when the spider forages along the water edge. To test our prediction, we developed a color vision model assessing whether the patches are visible to pygmy grasshoppers, the spider's primary prey. We conducted a field experiment using cardboard dummies that resemble D. raptor in size, shape, and color, but with half of them lacking leg patches, and we staged interactions between pygmy grasshoppers and D. raptor with and without leg patches in a greenhouse. We found the white patches to be visible to grasshoppers. The dummies with white patches attracted more grasshopper prey than the dummies without the patches. Moreover, grasshoppers were more attracted to spiders when their white patches were present. Our results supported the hypothesis that the white patches of D. raptor lure prey. Our findings, nevertheless, could not be explained as the spider's body coloration acting as a sensory trap but it should not be ruled out. More studies on a wider range of predators and prey will give more meaningful insights into the co‐evolution of predatory lures and prey sensory modalities.     

Prey capture tactics of spiders: An analysis based on a simulation model for spider's growth

The prey capture tactics of spiders was analyzed, considering the energy gained by the capture of prey and that required for it. For the purpose of it, a growth model of spiders was constructed, expressing the flow rate of prey biomass to the spider's body by differential equations. Solving these equations under the differing values of three parameters, growth curves of spiders was obtained. These three parameters are the amount of prey biomass supplied daily to spiders, x₀, the rate of prey capture of spiders, α, and a coefficient of the respiration rate required for the capture of prey, k. When the value of k increased, spiders could grow only at high value of x₀. These results suggest that habitats with small prey biomass are preferred by spiders adopting a sit-and-wait tactics for prey capture, which requires small values of k. Wolf spiders are one of these spiders showing that tactics. On the other hand, web-builders which require large amount of energy for spinning webs (namely, take large value of k), are able to grow only in the habitats with large prey biomass. Each species of spiders are considered to locate in a certain point between both extremes of these tactics for the capture of prey.     

Contributions of detrital subsidies to aboveground spiders during secondary succession, revealed by radiocarbon and stable isotope signatures

Prey subsidies originating from detritus add nutrients and energy to arboreal communities. Measurement of this subsidy is required in the understanding of how food web dynamics respond to changes in surrounding environments. Shrub spiders are one of the key predators involved in food web coupling. We evaluate the effects of potential changes in prey availabilities during secondary succession on the contribution of subsidy from detrital food webs to shrub spiders and how different spider feeding guilds used the subsidy of prey from detrital food webs. We measured the relative importance of the subsidy for the spider feeding guilds, using the ratios of stable isotopes of C (δ¹³C), and N (δ¹⁵N) and C isotope discrimination (Δ¹⁴C). Diet age was calculated from Δ¹⁴C values, because old diet ages of spiders indicate that the spiders consume prey from detrital food sources. Dominant aerial prey (Diptera) had a distinctively old diet age compared with arboreal prey, which indicates that aerial prey were subsidized from detrital food webs. Sit-and-wait spiders tended to have an older diet age than active hunting spiders, which indicates that sit-and-wait spiders depended more on subsidies. Diet age varied only slightly for spiders in stands of different ages, indicating that rates at which spiders use grazing and detrital prey are probably determined more by foraging strategies and not by stand age. A dominance of sit-and-wait predators will lead to higher detrital subsidy inputs in shrub habitats. This study highlights the effect of shrub spider community structure (feeding guild composition) on the volume of the subsidy received from the detrital food web.     

An order-specific monoclonal antibody to Diptera reveals the impact of alternative prey on spider feeding behavior in a complex food web

Generalist predators have the capacity to restrict pest population growth, especially early in the season before densities increase. However, their polyphagous feeding habits sometimes translate into reduced pest consumption when they target alternative prey. An order-specific monoclonal antibody was developed to examine the strength of trophic connections between Diptera, a major category of non-pest prey, and linyphiid spiders in alfalfa. We report the development and characterization of a monoclonal antibody with order-level specificity to Diptera. This antibody elicited strong absorbance to 22 Diptera from 13 families, no false-positive reactivity to non-dipteran invertebrates, and antigen detection periods following prey consumption that were comparable between spiders. Over 900 field-collected females of the linyphiid spiders Erigone autumnalis and Bathyphantes pallidus were screened for Diptera antigen. Significantly more B. pallidus screened positive for Diptera (40%) compared to E. autumnalis (16%), indicating differential reliance on these prey. In parallel with the collection of spiders for gut-content analysis, prey availability was estimated at web sites. The two spiders exhibited different feeding responses to prey availability. Consumption of Diptera by B. pallidus was strongly correlated with Diptera abundance whilst the availability of other potential prey did not influence predation rates. Conversely, E. autumnalis did not prey upon Diptera in proportion to availability, but increased Collembola activity-density reduced dipteran consumption. Integration of molecular gut-content analysis with precise sampling of prey demonstrated how two closely related linyphiid spiders exhibit different feeding responses to the availability of prey under natural field conditions. Elucidating the feeding preferences of natural enemies is critical to effective incorporation of biological control by generalist predators in the management of agricultural pests.     

Emergent properties in the behaviour of a virtual spider robot

Using a virtual spider robot, we studied hypotheses about the weaving behaviour of orb spiders. Our model spiders built virtual webs that mimicked perfectly the visual architecture of real webs of the garden cross spider Araneus diadematus. The matching of capture spiral and auxiliary spiral pitch was an apparently emergent property in both types of web. This validated our interpretation of the garden spider''s web-building decision rules, which use strictly local interactions with previously placed threads to generate global architecture.     

Correction to: Testing traditional hypotheses about prey capture efficiency in orb-web spiders

To capture prey, orb-web spiders create complex traps whose efficiency is contingent on a variety of factors that are not yet completely understood, including web size, competition for food, sun exposure, presence of web decorations and web orientation. Here we evaluate such factors in the field and ask which of them are the most influential variables affecting the quantity of prey captured in Argiope argentata webs. Webs were observed during the morning and the number of prey attached to each web was counted. Using the approach of information criteria based on the Akaike information criterion (AIC) values of each candidate model, we averaged the parameters of a global model, finding that the only predictor which 95% confidence interval did not include zero, was exposure to sunlight (whether the web is continuously shaded or continuously exposed to sunlight). All other variables did not explain variation in prey capture. We conclude that only sun exposure has an important effect on orb-web spiders’ prey capture efficiency in A. argentata. We additionally argue that silk decorations have different functions depending on the habitat and the species.

     

Web-building behaviour in the orb-weaving spider Zygiella x-notata: influence of experience

Zygiella x-notata is an orb-weaving spider that often renews its trap daily. Web building has associated costs and benefits, and building successive webs may have consequences for lifetime reproductive success. In the laboratory, we tested the ability of Z. x-notata to modify its building behaviour in response to various stages in predation (prey detection, capture and ingestion) experienced with a previous web. We determined which stages provided information for the spiders. Spiders that detected, captured and ingested prey and then rebuilt their web used less silk and made a smaller capture area than in the previous web. There was no effect of prey detection alone on the next web. Capture without feeding gave the same results as capture followed by feeding. The spiders that ate prey without detection and capture (feeding by hand) had the same energetic gains as spiders that caught prey but delayed building a new web. The spiders thus showed plasticity in web-building behaviour and in the amount of silk used (energetic investment) in the short term (from one web to the next). Changes in body condition may therefore influence web construction. Moreover, information gained during prey capture appeared to influence the size and structure of the next web. This ability should enable spiders to adapt their web building to maximize their fitness. Copyright 2000 The Association for the Study of Animal Behaviour.     

Multichannel feeding by spider functional groups is driven by feeding strategies and resource availability

Multichannel feeding, whereby consumers feed across resource channels such as upon herbivore and detritivore resources, acts to link discrete compartments of a food web with implications for ecosystem functioning and stability. Currently however, we have little understanding which feeding strategies of consumers underlie multichannel feeding. We therefore link spider functional group and resource density‐dependent or density‐independent feeding strategies to multichannel feeding by quantifying not only consumer diet, but also the relative availability of resources. Here we analysed herbivore (green) and detritivore (brown) prey use by spider communities in grasslands, and tested if available prey biomass proportions were linked to observed spider dietary proportions. Different spider functional groups each linked green and brown resource channels, but while green prey were always consumed in proportion to their relative biomass, brown prey were consumed independently of proportion by some functional groups. Additionally, we found greater intraguild predation by cursorial spiders when green resources were relatively scarcer, suggesting green prey was preferred, and needed to be compensated for when rare. Overall, we observed a stronger consumer connection to the green than brown resource channel, yet this green connection was more variable due to greater range in green resource availability across grasslands and density‐dependent consumption on green prey. Consequently, multichannel feeding by spiders was determined by density‐dependent and density‐independent feeding strategies that varied by spider functional group and across resources channels. Our results demonstrate that the role of multichannel feeding by spiders in linking separate food web compartments is a dynamic component of food web structure in these wild grasslands.     

Experimental Evidence that a Desert Parasitoid Keeps its Host Cool

The wasp parasitoid Pseudopompilus humboldti (Hymenoptera; Pompilidae) in the Negev desert of Israel stores its paralyzed host, the spider Stegodyphus lineatus (Araneae; Eresidae), at the entrance of the spider's nest. The spider is moved by the wasp from the depth of the nest to the entrance in spite of increased exposure to visually-searching predators, such as birds. We examined the hypothesis that this behaviour has evolved to prevent the wasp's host from overheating in this hot desert. Experimental manipulations of the position of the parasitized spiders demonstrated that spiders and wasp larvae could not survive the heat experienced deep in the spider's nest during summer. By contrast, in the cooler nest entrance, spiders and larvae survived, if they were overlooked by predators. This host-storing behaviour is an adaptive trade-off between thermoregulatory requirements and predation risk. This appears to be the first evidence that a parasitoid manipulates the temperature of its host, albeit for its own ultimate gain.     

Does the Giant Wood Spider Nephila pilipes Respond to Prey Variation by Altering Web or Silk Properties?

Recent studies demonstrated that orb‐weaving spiders may alter web architectures, the amount of silk in webs, or the protein composition of silks in response to variation in amount or type of prey. In this study, we conducted food manipulations to examine three mechanisms by which orb‐weaving spiders may adjust the performance of webs to variation in prey by altering the architectures of webs, making structural changes to the diameters of silk threads, and manipulating the material properties or amino acid composition of silk fibers. We fed Nephila pilipes two different types of prey, crickets or flies, and then compared orb structure and the chemical and physical properties of major ampullate (MA) silk between groups. Prey type did not affect orb structures in N. pilipes, except for mesh size. However, MA silk diameter and the stiffness of orbs constructed by spiders fed crickets were significantly greater than for the fly group. MA fibers forcibly silked from N. pilipes fed crickets was significantly thicker, but less stiff, than silk from spiders fed flies. Spiders in the cricket treatment also produced MA silk with slightly, but statistically significantly, more serine than silk from spiders in the fly treatment. Percentages of other major amino acids (proline, glycine, and glutamine) did not differ between treatments. This study demonstrated that orb‐weaving spiders can simultaneously alter some structural and material properties of MA silk, as well as the physical characteristics of webs, in response to different types of prey.     

Web damage and feeding experience influence web site tenacity in the orb-web spider Argiope keyserlingi Karsch

For orb-web spiders, the decision to remain at a particular site or to relocate elsewhere will ultimately depend on the quality of the site. In the past, the quality of the site has been measured in terms of prey availability; spiders experiencing large numbers of prey items are less likely to relocate. However, regular web damage caused by larger nonprey animals may also contribute to the quality of a particular site. Laboratory experiments revealed that the frequency and extent of web relocation by the orb-web spider Argiope keyserlingi was influenced by the feeding regime and the rate of nonprey web damage. Daily movement patterns were influenced by web damage, and these movements were in the direction away from the source of damage. However, the cumulative distance moved during the 7 days of the experiment was influenced by the frequency with which spiders were fed. Spiders that were not given prey moved further than spiders that obtained prey. These data indicate that spiders respond to web damage on a daily basis, but the cumulative movement of spiders over a longer period is influenced mostly by the history of prey ingestion rate. Copyright 2000 The Association for the Study of Animal Behaviour.