Ladder webs in orb‐web spiders: ontogenetic and evolutionary patterns in Nephilidae

Spider web research bridges ethology, ecology, functional morphology, material science, development, genetics, and evolution. Recent work proposes the aerial orb web as a one‐time key evolutionary innovation that has freed spider‐web architecture from substrate constraints. However, the orb has repeatedly been modified or lost within araneoid spiders. Modifications include not only sheet‐ and cobwebs, but also ladder webs, which secondarily utilize the substrate. A recent nephilid species level phylogeny suggests that the ancestral nephilid web architecture was an arboricolous ladder and that round aerial webs were derived. Because the web biology of the basalmost Clitaetra and the derived Nephila are well understood, the present study focuses on the webs of the two phylogenetically intervening genera, Herennia and Nephilengys, to establish ontogenetic and macroevolutionary patterns across the nephilid tree. We compared juvenile and adult webs of 95 Herennia multipuncta and 143 Nephilengys malabarensis for two measures of ontogenetic allometric web changes: web asymmetry quantified by the ladder index, and hub asymmetry quantified by the hub displacement index. We define a ‘ladder web’ as a vertically elongated orb exceeding twice the length over width (ladder index ≥ 2) and possessing (sub)parallel rather than round side frames. Webs in both genera allometrically grew from orbs to ladders, more so in Herennia. Such allometric web growth enables the spider to maintain its arboricolous web site. Unexpectedly, hub asymmetry only increased significantly in heavy‐bodied Nephilengys females, and not in Herennia, challenging the commonly invoked gravity hypothesis. The findings obtained in the present study support the intrageneric uniformness of nephilid webs, with Herennia etruscilla webs being identical to H. multipuncta. The nephilid web evolution suggests that the ancestor of Nephila reinvented the aerial orb web because the orb arises at a much more inclusive phylogenetic level, and all intervening nephilids retained the secondarily acquired substrate‐dependent ladder web. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 849–866.     

Host behavior modification of Achaearanea tingo (Araneae: Theridiidae) induced by the parasitoid wasp Zatypota alborhombarta (Hymenoptera: Ichneumonidae)

Behavioral manipulation involving Zatypota (Ichneumonidae: Pimplinae) parasitoids and their spider hosts is usually associated with an increase in web complexity at the location where the parasitoid larva builds its cocoon. A higher number of web threads at this location may improve stability and provide a physical barrier against potential predators. However, we observed that parasitized individuals of Achaearanea tingo attacked by Z. alborhombarta change the three‐dimensional structure of their webs to a very simple and strong structure composed of two cables attached to the surrounding vegetation. This structure holds the curled leaf formerly used by the spider as a shelter. The parasitoid larva remains protected within this shelter after killing the host. The architectural pattern of the cocoon webs of A. tingo indicates that host manipulation is characterized by the repetition of one specific subroutine involved in web construction. Similar alterations have been previously described for cocoon webs constructed by parasitized orb‐weavers, but not for the three‐dimensional webs of theridiids.     

Cues that Spiders (Araneae: Araneidae,Tetragnathidae) Use to Build Orbs: Lapses in Attention to One Set of Cues because of Dissonance with Others?

Even for small animals such as spiders, behavioral decisions are sometimes influenced by multiple cues. Orb webs constitute exquisitely precise records of the stimuli the spider experienced and the decisions that it made while building its web. In addition, because spiders appear to sense their webs largely by touch, direct behavioral observations can determine which stimuli they probably sense. Previous studies have shown that when an orb‐weaving spider decides how far apart to space successive sticky lines during orb construction, it responds to at least five different kinds of stimuli, all of which apparently use a cue from the web, the location of the previous, inner loop of sticky spiral (IL location), as a point of reference. Here we show that two additional cues from the web, which are related to the position of the temporary spiral (TS), also influence sticky spiral spacing. A combination of direct observations of spider movements, analyses of complete and partially complete webs, and responses to experimental modifications of the web of two species in different families, Micrathena duodecimspinosa (Araneidae) and Leucauge mariana (Tetragnathidae), indicate that both the TS‐IL distance itself and the short‐term memory of the change in TS‐IL distance compared with that on other recently encountered radii correlate with sticky spiral spacing. When the TS‐IL distance was large, the spiders apparently ceased to attend to other cues. Thus, even the relatively stereotyped behavior of orb construction includes variation that stems from attention‐like mental processes.     

Environmentally induced post‐spin property changes in spider silks: influences of web type,spidroin composition and ecology

Many spiders use silk to construct webs that must function for days at a time, whereas many other species renew their webs daily. The mechanical properties of spider silk can change after spinning under environmental stress, which could influence web function. We hypothesize that spiders spinning longer‐lasting webs produce silks composed of proteins that are more resistant to environmental stresses. The major ampullate (MA) silks of orb web spiders are principally composed of a combination of two proteins (spidroins) called MaSp1 and MaSp2. We expected spider MA silks dominated by MaSp1 to have the greatest resistance to post‐spin property change because they have high concentrations of stable crystalline β‐sheets. Some orb web spiders that spin three‐dimensional orb webs, such as Cyrtophora, have MA silks that are predominantly composed of MaSp1. Hence, we expected that the construction of three‐dimensional orb webs might also coincide with MA silk resistance to post‐spin property change. Alternatively, the degree of post‐spin mechanical property changes in different spider silks may be explained by factors within the spider's ecosystem, such as exposure to solar radiation. We exposed the MA silks of ten spider species from five genera (Nephila, Cyclosa, Leucauge, Cyrtophora, and Argiope) to ecologically high temperatures and low humidity for 4 weeks, and compared the mechanical properties of these silks with unexposed silks. Using species pairs enabled us to assess the influence of web dimensionality and MaSp composition both with and without phylogenetic influences being accounted for. We found neither the MaSp composition nor the three‐dimensionality of the orb web to be associated with the degree of post‐spin mechanical property changes in MA silk. The MA silks in Leucauge spp. are dominated by MaSp2, which we found to have the least resistance to post‐spin property change. The MA silk in Argiope spp. is also dominated by MaSp2, but has high resistance to post‐spin property change. The ancestry of Argiope is unresolved, but it is largely a tropical genus inhabiting hot, open regions that present similar stressors to silk as those of our experiment. Ecological factors thus appear to influence the vulnerability of orb web spider MA silks to post‐spin property change. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 580–588.     

An organic coating keeps orb‐weaving spiders (Araneae,Araneoidea, Araneidae) from sticking to their own capture threads

More than 95% of orb‐weaving spider species ensure prey capture success by producing viscous threads equipped with gluey droplets. However, this trap may bear serious risks for the web‐inhabiting spider as well. The obvious question, how a spider avoids getting stuck in its own capture spiral, has gained little scientific attention up till now. In 1905, the French naturalist Jean‐Henry Fabre concluded from anecdotal observation that orb‐weaving spiders protect themselves by a fatty surface coating. Here, we test this hypothesis by indirectly measuring the force necessary to detach an autotomized spider’s leg from the capture spiral of its own web (here called ‘index of adhesion’, IOA). Three groups of legs, each of the species Araneus diadematus Clerck, 1757 and Larinioides sclopetarius (Clerck, 1757), were tested. One was left untreated, one was washed with distilled water (H₂O), and one was washed with the organic solvent carbon disulphide (CS₂). In both species, we found a weak IOA between the spider leg and the gluey capture spiral in untreated and water‐washed legs without significant differences between the two. The IOA approximately doubled, when spider legs had been washed with carbon disulphide prior to measurement, that is, the CS₂‐washed legs stuck significantly more strongly than the untreated and water‐washed legs. These results provide indirect evidence for a protective anti‐adhesive organic coating on the spider’s body surface and so support Fabre’s hypothesis.     

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]。     

The phylogenetic placement of Psechridae within Entelegynae and the convergent origin of orb‐like spider webs

Evolutionary convergence of phenotypic traits provides evidence for their functional success. The origin of the orb web was a critical event in the diversification of spiders that facilitated a spectacular radiation of approximately 12 000 species and promoted the evolution of novel web types. How the orb web evolved from ancestral web types, and how many times orb‐like architectures evolved in spiders, has been debated for a long time. The little known spider genus Fecenia (Psechridae) constructs a web that resembles the archetypical orb web, but morphological data suggest that Psechridae (Psechrus + Fecenia) does not belong in Orbiculariae, the ‘true orb weavers’, but to the ‘retrolateral tibial apophysis (RTA) clade’ consisting mostly of wandering spiders, but also including spiders building less regular webs. Yet, the data are sparse and no molecular phylogenetic study has estimated Fecenia's exact position in the tree of life. Adding new data to sequences pulled from GenBank, we reconstruct a phylogeny of Entelegynae and phylogenetically test the monophyly and placement of Psechridae, and in doing so, the alternative hypotheses of monophyletic origin of the orb web and the pseudo‐orb versus their independent origins, a potentially spectacular case of behavioural convergence. We also discuss the implications of our results for Entelegynae systematics. Our results firmly place a monophyletic Psechridae within the RTA clade, phylogenetically distant from true orb weavers. The architectural similarities of the orb and the pseudo‐orb are therefore clearly convergent, as also suggested by detailed comparisons of these two web types, as well as the spiders' web‐building behaviours and ontogenetic development. The convergence of Fecenia webs with true orbs provides a remarkable opportunity to investigate how these complex sets of traits may have interacted during the evolution of the orb.     

Signals and Signal Choices made by the Araneophagic Jumping Spider Portia fimbriata while Hunting the Orb-Weaving Web spiders Zygiella x-notata and Zosis geniculatus

Portia fimbriata is a web-invading araneophagic jumping spider (Salticidae). The use of signal-generating behaviours is characteristic of how P. fimbriata captures its prey, with three basic categories of signal-generating behaviours being prevalent when the prey spider is in an orb web. The predatory behaviour of P. fimbriata has been referred to as aggressive mimicry, but no previous studies have provided details concerning the characteristics of P. fimbriata 's signals. We attempt to determine the model signals for P. fimbriata 's 'aggressive mimicry' signals. Using laser Doppler vibrometer and the orb webs of Zygiella x-notata and Zosis geniculatus , P. fimbriata 's signals are compared with signals from other sources. Each of P. fimbriata 's three categories of behaviour makes a signal that resembles one of three signals from other sources: prey of the web spider (insects) ensnared in the capture zone of the web, prey making faint contact with the periphery of the web and large-scale disturbance of the web (jarring the spider's cage). Experimental evidence from testing P. fimbriata with two sizes of lure made from Zosis (dead, mounted in a lifelike posture in standard-size orb web) clarifies P. fimbriata 's signal-use strategy: (1) when the resident spider is small, begin by simulating signals from an insect ensnared in the capture zone (attempt to lure in the resident spider); (2) when the resident spider is large, start by simulating signals from an insect brushing against the periphery of the web (keep the resident spider out in the web, but avoid provoking from it a full-scale predatory attack); (3) when walking in the resident spider's web, regardless of the resident spider's size, step toward the spider while making a signal that simulates a large-scale disturbance of the web (mask footsteps with a self-made vibratory smokescreen).     

Behavioral manipulation of the orb‐weaver spider Argiope argentata (Araneae: Araneidae) by Acrotaphus chedelae (Hymenoptera: Ichneumonidae)

We observed the first case of host‐behavioral manipulation of an orb‐weaver spider Argiope argentata induced by a parasitoid wasp of the genus Acrotaphus. The modified web is similar of those constructed by other orb weavers attacked by wasps of the close related genus Hymenoepimecis. The stick spirals and radii are absent and the web is composed of a three‐dimensional structure of non adhesive threads. The discovery of the ability to induce changes in host's web‐building behavior in Acrotaphus is indicative that this trait may be primitively present in the clade that includes the genus Hymenoepimecis.     

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.     

The spinning apparatus of Polenecia producta (Araneae,Uloboridae): Structure and histochemistry

Summary The spinning apparatus of the uloborid spider Polenecia producta was studied to complete previous studies on the same family of spiders. The structure of spinnerets and spigots, under scanning electron microscopy, and the main anatomical and histochemical characteristics of the spinning glands of adult females and males are described. In addition some observations on the spinning apparatus at three successive stages of development are made. There are nine kinds of silk glands in Polenecia, i.e. one more (aciniform — B glands) than found in other uloborids. The spinning apparatus of Polenecia is, therefore, the most complex so far known. It is also more complex than that presently known of Araneoidea. The characteristics of the spinning glands of Polenecia are compared with those of other uloborids. Present knowledge of the spinning apparatus of uloborids leads to a renewed discussion of the origin of the orb web in this family and in araneids. It is concluded that these two types of orb webs emerged from independent evolutionary processes.     

Different positions of males approaching towards females and male choices between webs with and without stabilimentum in a sexually cannibalistic orb‐web spider,Argiope bruennichi

Males of the orb‐weaving spider species Argiope bruennichi (Araneidae) are frequently victims of sexual cannibalism. Therefore, a male spider approaching a female should have strategies to avoid being killed before copulation. Our present field study detected six types of A. bruennichi male positions vis‐à‐vis the female web. In 78% of situations (39/50), only one male attached to a female. Two males attached to the same female in 11 cases. We observed no cases of three or more males on the same female web. We most commonly observed the situation of a male staying in its own web and connecting to a female's web with its silk thread (46% of cases). Of the female webs chosen by males, 68% were decorated with both an upper and lower portion of stabilimentum – a conspicuous white silk structure that reflects much more ultraviolet light than other spider silks in the web. Only 14% (7/50) of the selected webs lacked stabilimentum. Therefore, we conducted an experiment to investigate the males' choice between females' webs with and without stabilimentum. Of the 24 males used in the experiment, 10 chose webs with stabilimentum. This result did not show a strong preference of the male for stabilimentum between equally sized webs, and thus did not support an earlier suggestion that stabilimentum in A. bruennichi might function to guide males to females for mating.     

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.     

How Did the Spider Cross the River? Behavioral Adaptations for River-Bridging Webs in Caerostris darwini (Araneae: Araneidae)

Background

Interspecific coevolution is well described, but we know significantly less about how multiple traits coevolve within a species, particularly between behavioral traits and biomechanical properties of animals'' “extended phenotypes”. In orb weaving spiders, coevolution of spider behavior with ecological and physical traits of their webs is expected. Darwin''s bark spider (Caerostris darwini) bridges large water bodies, building the largest known orb webs utilizing the toughest known silk. Here, we examine C. darwini web building behaviors to establish how bridge lines are formed over water. We also test the prediction that this spider''s unique web ecology and architecture coevolved with new web building behaviors.

Methodology

We observed C. darwini in its natural habitat and filmed web building. We observed 90 web building events, and compared web building behaviors to other species of orb web spiders.

Conclusions

Caerostris darwini uses a unique set of behaviors, some unknown in other spiders, to construct its enormous webs. First, the spiders release unusually large amounts of bridging silk into the air, which is then carried downwind, across the water body, establishing bridge lines. Second, the spiders perform almost no web site exploration. Third, they construct the orb capture area below the initial bridge line. In contrast to all known orb-weavers, the web hub is therefore not part of the initial bridge line but is instead built de novo. Fourth, the orb contains two types of radial threads, with those in the upper half of the web doubled. These unique behaviors result in a giant, yet rather simplified web. Our results continue to build evidence for the coevolution of behavioral (web building), ecological (web microhabitat) and biomaterial (silk biomechanics) traits that combined allow C. darwini to occupy a unique niche among spiders.     

Body‐colour variation in an orb‐web spider and its effect on predation success

Animal body coloration serves several functions such as thermoregulation, camouflage, aposematism, and intraspecific communication. In some orb‐web spiders, bright and conspicuous body colours are used to attract prey. On the other hand, there are other species whose body colour does not attract prey. Using a spider species showing individual body‐colour variation, the present study aimed to determine whether or not the variation in body colour shows a correlation with predation rates. We studied the orb‐web spider (Cyclosa argenteoalba) using both field observations and T‐maze experiments, in which the prey were exposed to differently coloured spiders. Cyclosa argenteoalba has silver‐ and black‐coloured areas on its dorsal abdomen, with the ratio of these two colours varying continuously among individuals. The bright and conspicuous silver area reflects ultraviolet light. Results of both field observations and colour choice experiments using Drosophila flies as prey showed that darker spiders have a greater chance of capturing prey than silver spiders. This indicates that body‐colour variation affects predation success among individuals and that the bright silver colour does not function to attract prey in C. argenteoalba.     

Forces in the spider orb web

Summary Pretensile forces were measured in individual threads of intact spider webs. In the orb web of Araneus diadematus forces decrease from mooring threads to frame threads and radii, a typical ratio being 1071. The smaller number of radii in the upper than in the lower half of the orb is paralleled by force ratios of 21 to 31. A similar difference between radii built first during web construction and radii added after completion of the frame underlines the importance of the former as part of the scaffolding. High tensions in the auxiliary spiral stabilize the radii in addition to providing a pathway for the spider when inserting the sticky spiral. Radial pretension (F) changes with spider mass (m). F/m is similar for different animals indicating an adaptation of radial forces to those resulting from spider mass. Several observations suggest tension control by the spider. When forced to anchor its web to thin flexible rods tension in the threads remains in the normal range. Tension values are similar in the webs of A. diadematus, Zygiella x-notata, Nuctenea umbratica, and Nephila clavipes indicating independence from details of web geometry. Only the mooring threads of Nephila show unusually large forces suggesting a narrower working range of tensions for the catching area than for the scaffolding.     

Screening of Necrotizing Spiders in Korea Using Nonradioactive Sphingomyelinase Assay(II) Web-building Spiders

Spider bites cause a range of symptoms from simple swellings to disfiguring necrotic lesions, and occasionally death. While spider bites are not a major medical problem in Korea, it would be of great value to know which species of spiders pose a threat to human health. A middle molecular weight protein, sphingomyelinase D, has been identified in the venom of the brown recluse spider and strong evidence suggests that they have a major role in spider bite necrosis. For the identification of necrotizing species, we have investigated using recently developed non‐radioactive assay of sphingomyelinase for rapidly screening the necrotizing venoms. Here, we demonstrate the fetal toxicity of total 65 species (24 genera, 9 families) of the web‐building spiders among 622 identified spider species in Korea. It has been revealed that four species of the orb‐weaving spider, Araneus ventricosus (family Araneidae, 0.3509), Dipoena castrata (0.2413, family Theridiidae), Argiope minuta (0.1836, family Araneidae), and Paracoelotes spinivulva (0.1760, family Amaurobiidae) have relatively strong activities among themselves. However, comparing to that of the brown recluse spider, Loxosceles recluse (1.814) in North America, the necrotizing shpingomyelinase activities of these Korean web‐building spiders are still very low. Based on our results, we may thus conclude that there would be little possibilities in South Korea to create serious medical problems caused by necrotizing arachnidism.     

Spider web guilds in cacao agroforestry – comparing tree,plot and landscape‐scale management

Aim Owing to their role as insect predators, web‐building spiders can be important biological control agents within agricultural systems. In complex tropical agroecosystems such as agroforests, management determines plant architecture, vegetation composition and associated ant density, but little is known on how these attributes, together with landscape context, determine spider web density. We hypothesized that all three spatial scales and the presence of Philidris ants significantly contribute to the explanation of spider web density with web types being differently affected. Location In 42 differently managed cacao agroforestry systems in Sulawesi, Indonesia. Methods We surveyed the distribution of five spider‐web types on 420 cacao trees to determine how these relate to habitat variables and a numerically dominant ant species at three different spatial scales, comparing tree, plot and landscape features. We fitted linear mixed‐effects model, selected the best model subset using information‐theoretic criteria and calculated the model‐averaged estimates. We used non‐metric multidimensional scaling (NMDS) to determine and visualize guild level responses to the effects of the tree, plot and landscape‐scale variables. Results The five spider guilds preferred different features of cacao tree architecture. Most frequently recorded webs belonged to the line‐ and orb‐web type. At the tree scale, overall web density was positively related to canopy openness. At the plot scale, a higher number of shade trees was related to a higher web density. At the landscape scale, the altitude determined the distribution patterns of web‐building spiders. Presence of Philidris ants was positively associated with density of orb webs, while no pattern was found for other web types. Main conclusions Results suggest spider web density could be increased by pruning of cacao trees while keeping shade trees at high density in cacao plots. The results emphasize the need to consider scale dependency of crop management and web‐guild‐specific responses that may be related to different functional roles of spiders as a high‐density predator group in agroforestry.     

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

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