Function of bright coloration in the wasp spider Argiope bruennichi (Araneae: Araneidae)

There are two major competing explanations for the counter-intuitive presence of bright coloration in certain orb-web spiders. Bright coloration could lure insect prey to the web vicinity, increasing the spider's foraging success. Alternatively, the markings could function as disruptive camouflage, making it difficult for the insect prey to distinguish spiders from background colour variation. We measured the prey capture rates of wasp spiders, Argiope bruennichi, that were blacked out, shielded from view using a leaf fragment, or left naturally coloured. Naturally coloured spiders caught over twice the number of prey as did either blacked-out or leaf-shielded spiders, and almost three times as many orthopteran prey. Spectrophotometer measurements suggest that the bright yellow bands on the spider's abdomen are visible to insect prey, but not the banding on the legs, which could disguise the spider's outline. Thus, our results provide strong support for the hypothesis that bright coloration in the wasp spider acts as a visual lure for insect prey and weak support for the hypothesis that the arrangement of the banding pattern across the spider's body disguises the presence of the spider on the web.     

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

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

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.     

蜘蛛位置对成功捕获猎物和球型网图案的影响

静坐在球型网的中心,蜘蛛可能遭受天敌的攻击并暴露在不利的天气条件下,如风和雨。然而,栖居于网的中心使蜘蛛比隐藏在隐蔽场所中的蜘蛛能更迅速地察觉并捕获猎物,这是因为猎物的位置仅能被位于网中心的蜘蛛所确定。对在隐蔽场所中的蜘蛛而言,提高对猎物捕获率的方式之一是尽量减少隐蔽所与网中心的距离。而且,网中心与隐蔽所之间较短的距离使蜘蛛能更迅速地逃离危险境况。我使用既在网中心、又在隐蔽场所的硬类肥蛛(Larinioides sclopetarius Clerck),来检验这两种行为如何影响对猎物的捕获成功率。隐藏在隐蔽场所中的蜘蛛更经常忽略猎物,使猎物也有比较多的逃离机会,这样,与在网中心的蜘蛛相比,猎物的损失率就更高。另外,研究了隐蔽场所的位置对球型网图案的影响。在大多数球型网中,网中心上方的区域比网下方小,丝也比较少,形成了结构不对称的网;隐蔽场所通常在网的上方。当隐蔽场所的位置在实验中被倒转时,就形成了非典型的球型网。最后,L．sclopetarius建造的网有很突出的边缘非对称性,与隐蔽场所相邻的区域面积较小,而远离隐蔽场所的区域面积较大,这也可解释为减少了隐蔽场所和网中心之间的距离[动物学报50(4)：559-565．2004]。     

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.     

Assassin bug uses aggressive mimicry to lure spider prey

Assassin bugs (Stenolemus bituberus) hunt web-building spiders by invading the web and plucking the silk to generate vibrations that lure the resident spider into striking range. To test whether vibrations generated by bugs aggressively mimic the vibrations generated by insect prey, we compared the responses of spiders to bugs with how they responded to prey, courting male spiders and leaves falling into the web. We also analysed the associated vibrations. Similar spider orientation and approach behaviours were observed in response to vibrations from bugs and prey, whereas different behaviours were observed in response to vibrations from male spiders and leaves. Peak frequency and duration of vibrations generated by bugs were similar to those generated by prey and courting males. Further, vibrations from bugs had a temporal structure and amplitude that were similar to vibrations generated by leg and body movements of prey and distinctly different to vibrations from courting males or leaves, or prey beating their wings. To be an effective predator, bugs do not need to mimic the full range of prey vibrations. Instead bugs are general mimics of a subset of prey vibrations that fall within the range of vibrations classified by spiders as 'prey'.     

The ecological and evolutionary interdependence between web architecture and web silk spun by orb web weaving spiders

Spider orb webs are dynamic, energy absorbing nets whose ability to intercept prey is dependent on both the mechnical properties of web design and the material properties of web silks. Variation in web designs reflects variation in spider web spinning behaviours and variation in web silks reflects variation in spider metabolic processes. Therefore, natural selection may affect web function (or prey capture) through two independent and alternative pathways. In this paper, I examine the ways in which architectural and material properties, singly and in concert, influence the ability of webs to absorb insect impact energy. These findings are evaluated in the context of the evolution of diverse aerial webs. Orb webs range along a continuum from high to low energy absorbing. No single feature of web architecture characterizes the amount of energy webs can absorb, but suites of characters indicate web function. In general, webs that intercept heavy and fast flying prey (high energy absorbing webs) are large, built by large spiders, suspended under high tension and characterized by a ratio of radii to spiral turns per web greater than one. In contrast, webs that intercept light and slow flying prey (low energy absorbing webs) are suspended under low tension, are small and are characterized by radial to spiral turn ratios that are less than one. The data suggest that for spiders building high energy absorbing webs, the orb architecture contributes much to web energy absorption. In contrast, for spiders that build low energy absorbing webs, orb architecture contributes little to enhance web energy absorption. Small or slow flying insects can be intercepted by web silks regardless of web design. Although there exists variation in the material properties of silk collected from high and low energy absorbing webs, only the diameter of web fibres varies predictably with silk energy absorption. Web fibre diameter and hence the amount of energy absorbed by web silks is an isometric function of spider size. The significance of these results lies in the apparent absence of selective advantage of orb architecture to low energy absorbing webs and the evolutionary trend to small spiders that build them. Where high energy absorption is not an exacting feature of web design, web architecture should not be tightly constrained to the orb. Assuming the primitive araneoid web design is the orb web, I propose that the evolution of alternative web building behaviours is a consequence of the general, phyletic trend to small size among araneoids. Araneoids that build webs of other than orb designs are able to use new habitats and resources not available to their ancestors.     

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.     

Spinning an elastic ribbon of spider silk

The Sicarid spider Loxosceles laeta spins broad but very thin ribbons of elastic silk that it uses to form a retreat and to capture prey. A structural investigation into this spider's silk and spinning apparatus shows that these ribbons are spun from a gland homologous to the major ampullate gland of orb web spiders. The Loxosceles gland is constructed from the same basic parts (separate transverse zones in the gland, a duct and spigot) as other spider silk glands but construction details are highly specialized. These differences are thought to relate to different ways of spinning silk in the two groups of spiders. Loxosceles uses conventional die extrusion, feeding a liquid dope (spinning solution) to the slit-like die to form a flat ribbon, while orb web spiders use an extrusion process in which the silk dope is processed in an elongated duct to produce a cylindrical thread. This is achieved by the combination of an initial internal draw down, well inside the duct, and a final draw down, after the silk has left the spigot. The spinning mechanism in Loxosceles may be more ancestral.     

The escape strategy of green lacewings from orb webs

When green lacewings (Neuroptera: Chrysopidae) fly into spider orb webs, they often simply reverse their flight direction and pull away (Table I). If a lacewing is trapped, it uses a specialized escape behavior. It first cuts away the sticky strands entangling head, feet, and antennae. If an antenna cannot be freed by tugging, it uses an antenna climb (Fig. 5A). After its body is free, the lacewing remains suspended by its hair-covered wings, which are held in a characteristic cruciform position (Fig. 5B). Orb web sticky strands adhere poorly to the hairy wings (Fig. 7), so the chrysopid may just wait until the strands slide off and it falls free. If placed in an orb web when the spider is at the web hub and ready to attack, a lacewing usually does not have time to escape (Fig. 1). When the spider is at the hub but eating, the chances of escape improve, and when the spider is away from the hub attacking other prey, nearly all lacewings in our experiment were able to escape. This finding emphasizes the importance of the spider's activity in its capture success.Paper No. 88 of the series Defense Mechanisms of Arthropods.     

Design features of the orb web of the spider, Araneus diadematus

Analysis of orb webs of the garden cross spider (Araneus diadematus)showed that these vertical webs have a significant up/down asymmetry.Experiments demonstrated that the spider runs down faster thanup, and thus confers a relatively higher foraging value to sectionsbelow the hub. Simulations suggested that the density of capturespiral spacing, prey size, and the density of prey should allaffect the capture efficiency of a web. Webs lose effectivecapture area because of overlap of the capture zone around eachthread; the smaller the prey, the finer the mesh can be withoutlosing effective area. Lower sectors of the web have a particularmesh size (height and length of capture spiral segments) throughout,whereas in the upper sectors the mesh size changes, wideningfrom the hub towards the periphery.     

The biology of Pholcus phalangioides (Araneae, Pholcidae): predatory versatility, araneophagy and aggressive mimicry

Predatory versatility occurs in Pholcus phalangioides (Fuesslin). In addition to building prey-catching space webs, P. phalangioides invades webs of other spiders and feeds on the occupants. It acts as an aggressive mimic by performing specialized vibratory behaviours to which the prey-spider responds as it normally would to its own prey. Prey (spiders and insects) is attacked by wrapping. Prey that trips over lines at the edge of a web of P. phalangioides , but fails to enter the web, is successfully attacked: P. phalangioides leans out of its web to throw silk over the prey, keeping as few as two legs on the silk. However, P. phalangioides does not attack prey that is completely away from webs. Occasionally, P. phalangioides feeds on eggs of other spiders and on ensnared insects it encounters in alien webs. Experimental evidence indicates that vision is of little or no importance in the predatory behaviour of P. phalangioides . Although P. phalangioides invades diverse types of webs, in addition to using its own web, its efficiency as a predator varies with web-type. It is most efficient as a predator of spiders and, especially, insects on its own web, and least efficient as a predator of amaurobiids on their cribellate sheet webs. Sensory, locomotory and other factors which influence differential predatory efficiency are discussed. The behaviour of P. phalangioides 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.     

Transmission of vibrations in funnel and sheet spider webs.

Behavior of the funnel web building spider, Hololena curta, was observed. The spider is dependent on vibratory signals transmitted through the web. Prey localization usually necessitates two to four pauses and reorientation, and cannot proceed if the prey ceases to struggle. Treating the web as a stretched membrane, we calculated fundamental and overtone normal frequencies and transmission velocities. Unlike the results reported for orb webs, resonance does appear to be a significant factor in the funnel web.     

Prey analysis of four species of tropical orb-weaving spiders (Araneae: Araneidae) and a comparison with araneids of the temperate zone

Summary The actual prey in the orb webs of four araneid spiders (Nephila clavipes, Eriophora fuliginea, Argiope argentata, and A. savignyi) and the relative abundance of their potential prey (pitfall traps, yellow traps, and sweep-netting) was investigated over 1 year at different locations in Panama. The relative abundance of insects and spiders depends on seasonal fluctuations (Fig. 2) which are reflected by corresponding variations in the effectiveness of the webs. The main prey groups are Nematocera (50%–68%), winged Formicoidea (6%–15%) and Hymenoptera, Coleoptera, and Brachycera (4%–10% each) (Fig. 4-6). The remaining 10%–17% of the prey comes from up to 26 other groups (Table 2). Differences in prey size and prey composition between the spider species are small (Fig. 7). Most prey items are 1–2 mm long: only a few insects exceed 30 mm body length (Figs. 9–12). Relative to the available prey, some groups (e.g. Nematocera, Aphidoidea, Psocoptera) are caught selectively, while other groups (e.g. Heteroptera, Coleoptera, Brachycera, Orthoptera) are underrepresented in the prey spectrum and obviously avoid orb webs (Table 7). The differences in prey composition between araneids of the tropics and of the temperate zone are discussed (Table 8) and compared to those recorded in other studies (Table 9, 10). Most of these report large numbers of big prey items (Odonata, Lepidoptera, wasps/bees). It is pointed out that those studies do not take into account the total available prey in a spider's web but only that part which the spider selects from the web (mainly according to size). The importance of small prey items even for large spiders is explained and an obvious lack of niche partitioning among coexisting araneids is discussed (Table 11).     

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.     

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.     

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.     

Discrimination and release of unpalatable butterflies by Nephila clavipes, a neotropical orb-weaving spider

Abstract. 1. Nephila clavipes (L.), a common spider in neotropical forests, discriminates some unpalatable prey and releases them unharmed from its web. Release is not accidental but results from a specific behavioural sequence.
2. Field trials with twenty-seven butterfly species showed that spiders respond consistently to butterfly species and higher taxa. Ithomiinae and some Danainae are almost always released while Heliconiinae, Nymphalinae, Acraeinae, Pieridae and Papilionidae are usually eaten.
3. Paired tests showed that an immediately preceding experience with a different butterfly did not reveal any change in the spider's usual response to a particular butterfly.
4. Warning coloration is not involved in spider response. Spiders rejected the models but ate the mimics of two different butterfly species pairs. Distastefulness is probably signalled by chemical cues.
5. Some unpalatable butterflies stay motionless when entangled and while the spiders release them. Motionlessness in webs seems to be a requisite to allow recognition of their distastefulness without being bitten by the spider.
6. Spiders differ from vertebrates in the prey accepted and in rejection mechanics. Although there is no indication of learning, spiders may select butterflies for distastefulness, chemical signals and motionlessness, at the individual level.     

Color polymorphic lures target different visual channels in prey

Selection for signal efficacy in variable environments may favor color polymorphism, but little is known about this possibility outside of sexual systems. Here we used the color polymorphic orb‐web spider Gasteracantha fornicata, whose yellow‐ or white‐banded dorsal signal attracts dipteran prey, to test the hypothesis that morphs may be tuned to optimize either chromatic or achromatic conspicuousness in their visually noisy forest environments. We used data from extensive observations of naturally existing spiders and precise assessments of visual environments to model signal conspicuousness according to dipteran vision. Modeling supported a distinct bias in the chromatic (yellow morph) or achromatic (white morph) contrast presented by spiders at the times when they caught prey, as opposed to all other times at which they may be viewed. Hence, yellow spiders were most successful when their signal produced maximum color contrast against viewing backgrounds, whereas white spiders were most successful when they presented relatively greatest luminance contrast. Further modeling across a hypothetical range of lure variation confirmed that yellow versus white signals should, respectively, enhance chromatic versus achromatic conspicuousness to flies, in G. fornicata's visual environments. These findings suggest that color polymorphism may be adaptively maintained by selection for conspicuousness within different visual channels in receivers.