Ontogenetic Changes in Web Design in Two Orb‐Web Spiders

The first orb web built by newly hatched spiders resembles the adult web in its overall form and structure. However, many details show ontogenetic changes. One possible explanation for these changes is that the tiny early‐instar spiders with their minute brains will make more mistakes and build less ‘perfect’ orb webs than older and larger juveniles and adults. To test this hypothesis, known as the size limitation hypothesis, I analysed orb webs from three developmental stages, spiderlings, juveniles and adult females, in two neotropical orb‐web spiders, the araneid Eustala illicita and the nephilid Nephila clavipes. Neither species showed clear signs of being behaviourally limited or more prone to committing errors as spiderlings than were older juveniles or adults. These findings therefore do not support the size limitation hypothesis in either species. Finally, I looked for evidence of the ‘biogenetic law’, which predicts that juveniles should build less derived orb webs than the adults. Evidence for this was found in E. illicita, but not in N. clavipes.     

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.     

The Effect of Wind Exposure on the Web Characteristics of a Tetragnathid Orb Spider

Studies on spiders in their natural habitats are necessary for determining the full range of plasticity in their web-building behaviour. Plasticity in web design is hypothesised to be important for spiders building in habitats where environmental conditions cause considerable web damage. Here we compared web characteristics of the orb spider Metellina mengei (Araneae, Tetragnathidae) in two different forest habitats differing in their wind exposure. We found a notable lack of differences in web geometry, orientation and inclination between webs built along an exposed forest edge and those built inside the forest, despite marked differences in wind speed. This suggests that M. mengei did not exhibit web-building plasticity in response to wind in the field, contrasting with the findings of laboratory studies on other species of orb spiders. Instead, differences in prey capture and wind damage trade-offs between habitats may provide an explanation for our results, indicating that different species employ different strategies to cope with environmental constraints.     

Age-related sequential web building in the colonial spider Metepeira incrassata (Araneidae): an adaptive spacing strategy

Colonial orb-weaving spiders provide insight into the proximate mechanisms by which social animals space themselves within a group. We examined mechanisms for the temporal patterns of web building that determine individual positions in Metepeira incrassata (Araneidae) colonies. The spiders display a characteristic age-related sequence of daily web building, with larger spiders completing their webs significantly earlier than smaller ones. We used data on behavioural interactions, web building, prey capture and predator attacks to evaluate four hypotheses. (1) Larger spiders are better competitors and pre-empt optimal spatial positions. (2) Smaller spiders reduce competition with larger individuals by building webs later. (3) Prey captured by different size classes is available at different times. (4) Differential predation risk determines web-building times. Large individuals dominated behavioural interactions. Disturbances by larger spiders during web construction significantly delayed the completion of smaller individuals' webs and precipitated movements to new web sites. One prediction of the first hypothesis, that spatial needs translate into earlier building, was confirmed by significantly earlier web building by mature females with egg sacs (which are unable to move their egg sacs) compared with same-sized females without eggs (which can change locations freely). Experiments to determine whether the presence of large spiders inhibited the web building of smaller individuals were equivocal. Prey availability and risk of predation are not factors affecting web-building patterns. Sequential web building appears to be a result of both larger spiders competing to pre-empt space from one another and smaller individuals attempting to reduce conflict during web construction. Sequential web building is a proximate mechanism that influences spacing among colonial orb-weaving spiders and helps shape the typical hierarchical size distribution of spiders within the colony. Similar spacing mechanisms may be seen in colonial birds and marine invertebrates. Copyright 2000 The Association for the Study of Animal Behaviour.     

Predatory behaviour of an araneophagic assassin bug

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

Conspecific and Heterospecific Attraction: A Mechanism of Web-site Selection Leading to Aggregation Formation by Web-building Spiders

This study investigated whether conspecific and/or heterospecific attraction to silk is a mechanism of web-site selection leading to aggregation formation by two species of web-building spiders, Hypochilus thorelli Marx (Araneae: Hypochilidae) and Achaearanea tepidariorum (C.L. Koch) (Araneac: Theridiidae). We determined that the spatial distribution of these two spiders was clumped and that H. thorelli had a greater tendency to aggregate than did A. tepidariorum. To determine the mechanism responsible for this spatial pattern, we conducted three field experiments. We examined web-site selection by H. thorelli in three contexts: no spiders or webs present (cue removed), vacant webs present, and occupied webs present. In the case where no webs were present, there was no tendency for spiders to choose previously occupied sites as web sites. When vacant webs were present, spiders chose to occupy the vacant webs. When occupied webs were present, spiders either invaded webs and evicted the owners, or settled adjacent to and attached their webs to those of residents. Various microhabitat variables (height, angle, temperature, humidity, and substrate character of the rock surface) of randomly selected unoccupied sites and the web sites chosen by spiders were compared. There were no detectable differences between web sites and unoccupied sites with respect to any of the variables measured. This leads us to conclude that web-site choice by immigrating spiders was based on the presence of silk rather than other features of the site.     

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.     

Aging and foraging efficiency in an orb-web spider

Aging is often associated with reduced behavioral performance such as decreased locomotion or food consumption, related to a deterioration in physiological functions. In orb-web spiders, webs are used to capture prey and aging can affect web-building behavior and web structure. Here, we investigated the effect of aging on prey capture in the orb-web spider Zygiella x-notata. The ability of adult females to capture flies was examined at different ages. The rate of prey capture did not change with age, but older spiders took more time to subdue and capture the prey. Alterations which appeared in web structure with age (increase in the number of anomalies affecting radii and capture spiral) affected prey capture behavior. Furthermore, the analysis of individual performance (carried out on 17 spiders at two different ages) showed that older females spent more time handling the prey and finding it in the web. Our results suggest that, in the laboratory, age does not affect prey capture rates but it influences prey capture behavior by affecting web structure or/and spider motor functions.     

蛛网结构性能及其适应性

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

Contrasting responses of web-building spiders to deer browsing among habitats and feeding guilds

We examined web-building spider species richness and abundance in forests across a deer density gradient to determine the effects of sika deer browsing on spiders among habitats and feeding guilds. Deer decreased the abundance of web-building spiders in understory vegetation but increased their abundance in the litter layer. Deer seemed to affect web-building spiders in the understory vegetation by reducing the number of sites for webs because vegetation complexity was positively correlated with spider density and negatively correlated with deer density. In contrast, the presence of vegetation just above the litter layer decreased the spider density, and deer exerted a negative effect on this vegetation, possibly resulting in an indirect positive effect on spider density. The vegetation just above the litter layer may be unsuitable as a scaffold for building webs if it is too flexible to serve as a reliable web support, and may even hinder spiders from building webs on litter. Alternatively, the negative effect of this vegetation on spiders in the litter may be as a result of reduced local prey availability under the leaves because of the reduced accessibility of aerial insects. The response to deer browsing on web-building spiders that inhabit the understory vegetation varied with feeding guild. Deer tended to affect web-invading spiders, which inhabit the webs of other spiders and steal prey, more heavily than other web-building spiders, probably because of the accumulated effects of habitat fragmentation through the trophic levels. Thus, the treatment of a particular higher-order taxon as a homogeneous group could result in misleading conclusions about the effects of mammalian herbivores.     

Prey preferences ofPortia fimbriata,an araneophagic,web-building jumping spider (Araneae: Salticidae) from Queensland

Portia fimbriata from Queensland, a previously studied jumping spider (Salticidae), routinely includes web-building spiders and cursorial salticids in its diet, both of these types of prey being dangerous and unusual prey for a salticid. The present paper is the first detailed study ofP. fimbriata's prey preferences. Three basic types of tests of prey preference were used, providing evidence that (1)P. fimbriata males and females prefer spiders (both web-building spiders in webs and salticids away from webs) to insects; (2)P. fimbriata males and females prefer salticids to web-building spiders; (3)P. fimbriata males and females prefer larger spiders to smaller spiders; (4) there are intersexual differences in the preferences ofP. fimbriata for prey size, females preferring larger prey and males preferring smaller prey; and (5)P. fimbriata's prey preferences are not affected by a prior period without food of 2 weeks. When preferences were tested for by using both living, active prey and dead, motionless lures, the same preferences were expressed, indicating thatP. fimbriata can distinguish among different types of prey independent of the different movement patterns of different prey.     

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.     

The biology of New Zealand and Queensland pirate spiders (Araneae, Mimetidae): aggressive mimicry, araneophagy and prey specialization

Mimetus sp. indet. and Mimetus maculosus , from New Zealand and Australia, respectively, were studied in the laboratory and in nature. Behaviourally, the two species were very similar. Each was found to be primarily an araneophagic spider which invaded alien webs, acted as an aggressive mimic by performing a variety of vibratory behaviours to which the prey-spider responded as it normally would to its own prey, and attacked by lunging at close range, subduing its victim with a strong, apparently spider-specific venom while holding the spider in a 'basket' formed by its spine-covered legs. In nature, these mimetids were observed to feed on a restricted range of spiders: orb web-building araneids and space web-building theridiids. Sometimes, they occupied other types of webs, but in the laboratory they captured only araneids and theridiids efficiently. They captured non-cribellate amaurobiids considerably less efficiently, and never captured other types of spiders. Occasionally, the mimetids fed on insects ensnared in araneid and theridiid webs and on eggs of theridiids. Experimental evidence indicated that vision was of little or no importance in the predatory behaviour of these mimetids. The behaviour of the mimetids is compared to that of Portia , an araneophagic web-invading salticid, and the results of this study are discussed in relation to hypotheses concerning salticid evolution.     

The Trochidae and Turbinidae of the Kermadec Ridge (Mollusca: Gastropoda)

Abstract

Taieria erebus (Gnaphosidae) was found to be a versatile predator: it captured insects both cursorially (away from webs) and kleptopar-asitically (on alien webs); it captured spiders in both the presence and absence of webs; and it also ate the eggs of host spiders (oophagy). When T. erebus invaded webs, it was as an aggressive mimic — it performed a repertoire of vibratory behaviours to lure the host spider. Although T. erebus pursued and captured spiders on diverse web-types, it was more effective as a predator when invading densely (rather than sparsely) woven cribellate and non-sticky webs, and was especially effective on non-cribellate sticky webs. Gnaphosids are traditionally referred to as hunting spiders, but T. erebus built a small prey-capture web. T. erebus also preyed on segestriid spiders, then used their webs to catch more prey, this being an unusual example of a spider using, as a tool for predation, the spinning-work of another species from an unrelated family. T. erebus used specialised behaviours to prey on nesting cursorial spiders. Prey was either grasped or stabbed; the venom of T. erebus was highly potent against spiders. Experiments indicated that vision was of little or no importance in the predatory behaviour of T. erebus. The behaviour of T. erebus is compared to that of Portia, a web-building salticid spider which is very versatile in its predatory behaviour and has acute vision. T. erebus is discussed in relation to hypotheses concerning gnaphosid and salticid evolution.     

Risk Sensitivity and the Paradox of Colonial Web-Building in Spiders

Group foraging is rare in spiders, occurring only where preyavailability is high. If colonial web-building increases individualprey capture rates as shown, why does group foraging not occurmore often where prey are scarce? Risk sensitivity may explainthis paradox, as variance in prey capture is reduced in groups;risk-averse spiders should join groups only when prey exceeda threshold level. Field studies show that group foraging variesas predicted between species, between populations of a singlespecies, and between sites within a population. However, recentmodels suggest the necessity of examining variance within individualsover time rather than between individuals within populations.Additionally, mechanisms responsible for variance reductionin colonial webs may be less effective than previously assumed.New field data suggest that while prey variance over time maybe somewhat less for individual spiders in groups than for solitaries,the relationship between colonial web-building and variancein prey capture is far more complex than originally thought.The influence of risk sensitivity on reproductive success andthe evolution of colonial web-building is discussed.     

Competition for a limited space in kleptoparasitic Argyrodes spiders revealed by field experiments

Most kleptoparasitic Argyrodes spiders rely exclusively on host spider webs for obtaining their food. Because their densities occasionally reach high levels within a restricted area, competitive interactions may be important for determining the density of these unique spiders. Here I used two Argyrodes species commonly found on webs of the large orb-web spider Nephila clavata to clarify whether inter- and intraspecific competition influences abundance and within-web distribution by using observational data and field experiment. Removing Argyrodes flavescens from the host webs induced a remarkably high immigration of that species while density on control webs remained almost at the same level, which is evidence for strong intraspecific competition. Larger individuals of A. flavescens were located more frequently at the capture area of the host webs where it is easy to access prey ignored by the host spider, and spiders immigrating into webs from which that species had been removed were smaller in body size, suggesting interference competition for space among conspecific kleptoparasites. Argyrodes bonadea increased in number on webs from which A. flavescens had been removed, and the increase was correlated with the number of A. flavescens removed. This finding is evidence for interspecific competition that is rarely reported in spiders. A multiple regression model including numbers of a conspecific parasite as well as web and body sizes of the host spider could not detect competitive interactions between species, suggesting the importance of experimental approaches. Received: May 22, 2000 / Accepted: December 1, 2000     

Behavioral Consequences of Laser Lesions in the Central Nervous System of Araneus diadematus Cl.

A laser beam of 6.5-1 joule is directed for 1 msec at dorsalsurfaces of adult female Araneus diadematus Cl. spiders. Thelesion is located by means of histological methods, and relationshipof its position to behavioral consequences—as measuredby changes in the web-pattern—are studied. Twelve animalswith lesions in the area of the supra-esophageal ganglion showdifferent degrees of extensive web-pattern disturbances consistingof smaller and rounder webs with less detail and irregular spacingof threads. Limitations of the method and the role of the supra-esophagealganglion in Araneus as an overall integrating center of sensorymotor coordination in web-building are discussed.     

Interactions between citrus psylla,Trioza erytreae (Hem. Triozidae), and spiders in an unsprayed citrus orchard in the transvaal lowveld

During a two year survey, a total of 3,054 spiders represented by 21 families were sampled in an unsprayed citrus orchard in the Transvaal Lowveld. Numerically the Salticidae was the dominant family (34.4%) followed by the Theridiidae (21.9%), Thomisidae (11.9%), Araneidae (7.9%), Clubionidae (7.0%) and the Tetragnathidae (3.7%). Eighteen species of spiders were observed to prey on citrus psylla,Trioza erytreae (Del Guercio) (Hemiptera: Triozidae), while six species trap nymphs and adults under their retreats and webs. There were significant positive correlations between the weekly psylla populations and the weekly populations of web-building spiders and wandering spiders present one to four weeks later but no significant correlation between the weekly spider populations and the weekly psylla populations present one to five weeks later. This seems to indicate that while spiders are unable to keep citrus psylla populations at acceptable low levels, they may contribute in reducing their numbers.      

Individual decisions and group dynamics: why pholcid spiders join and leave groups

To examine the role of individual variation in the dynamics of group formation, I conducted a mark-recapture study and a series of laboratory and field experiments with Holocnemus pluchei spiders (Araneae: Pholcidae). These spiders can either share webs or live alone, and individuals shift frequently between these strategies. Spiders' decisions were influenced by size and recent feeding success. In the laboratory, small hungry spiders introduced into a web that held a larger conspecific resident were more likely than small well-fed spiders to abandon the web and build their own web. This behaviour pattern gradually reversed as spiders grew: large hungry spiders were more likely than large well-fed spiders to stay in the shared web. When I introduced spiders into empty webs, they were more likely to stay compared with spiders tested with conspecifics. However, hunger level also influenced behaviour even when conspecifics were not present. Food-deprived spiders were more likely to abandon webs and build their own, consistent with the idea that spiders were following a win-stay/lose-shift strategy. In the field, spiders were more likely to stay in webs overnight when they were given supplemental food. In another experiment, spiders that were found building webs in cleared areas were smaller and thinner than average. Finally, I tested whether the size of the intruder or the resident affected whether spiders joined webs. Large intruders were more successful at remaining in webs than smaller intruders, although spiders of all sizes had some success in joining groups. Additional synthetic theoretical work is needed to integrate the complex processes underlying the formation and persistence of groups.     

Do stabilimenta in orb webs attract prey or defend spiders?

Orb-weaving spiders are ideal organisms for the study of conflictbetween behavioral investments in foraging and defense becausetheir webs provide physical manifestations of those investments.We examined the impact of including stabilimenta, designs ofbright-white noncapture silk, at the center of orb webs forforaging and defense in Argiope aurantia. Our findings suggestthat stabilimentum building is a defensive behavior, supportingthe "web advertisement" hypothesis that the high visibilityof stabilimenta can prevent birds from flying through webs.Yet, spiders often do not include stabilimenta in their webs,indicating that a serious cost is associated with them. We alsoshow, through comparison of paired webs with and without stabilimenta,that stabilimenta reduce the prey capture success of spidersby almost 30%. This demonstrates the potential impact that defensivebehaviors of spiders can have on their foraging success andsuggests that much of the variation in stabilimenta may be accountedfor by a cost—benefit trade-off made when including stabilimentain webs.