M. E. Herberstein

The functional significance of silk decorations of orb-web spiders: a critical review of the empirical evidence

A number of taxonomically diverse species of araneoid spiders adorn their orb‐webs with conspicuous silk structures, called decorations or stabilimenta. The function of these decorations remains controversial and several explanations have been suggested. These include:(1)stabilising and strengthening the web;(2) hiding and concealing the spider from predators;(3) preventing web damage by larger animals, such as birds;(4) increasing foraging success; or (5) providing a sunshield. Additionally, they may have no specific function and are a consequence of stress or silk regulation. This review evaluates the strength of these explanations based on the evidence. The foraging function has received most supporting evidence, derived from both correlative field studies and experimental manipulations. This contrasts with the evidence provided for other functional explanations, which have not been tested as extensively. A phylogenetic analysis of the different decoration patterns suggests that the different types of decorations are as evolutionary labile as the decorations themselves: the analysis shows little homology and numerous convergences and independent gains. Therefore, it is possible that different types of decorations have different functions, and this can only be resolved by improved species phylogenies, and a combination of experimental and ultimately comparative analyses.

Sperm competition and small size advantage for males of the golden orb-web spider Nephila edulis.

Sexual selection, through female choice and/or male–male competition, has influenced the nature and direction of sexual size dimorphism in numerous species. However, few studies have examined the influence of sperm competition on size dimorphism. The orb‐web spider Nephila edulis has a polygamous mating system and extreme size dimorphism. Additionally, the frequency distribution of male body size is extremely skewed with most males being small and few large. The duration of copulation, male size and sexual cannibalism have been identified as the significant factors determining patterns of sperm precedence in spiders. In double mating trials, females were assigned to three treatments: either they mated once with both males or the first or the second male was allowed to mate twice. Paternity was strongly associated with the duration of copulation, independent of mating order. Males that were allowed to mate twice not only doubled the duration of copulation but also their paternity. Small males had a clear mating advantage, they copulated longer than large males and fertilized more eggs. Males of different sizes used different tactics to mate. Large males were more likely to mate through a hole they cut into the web, whereas small males approached the female directly. Furthermore, small males usually mated at their first attempt but large males required several attempts before mating took place. There was no obvious female reaction towards males of different sizes.

Costs of courtship and mating in a sexually cannibalistic orb-web spider: female mating strategies and their consequences for males

Abstract. The costs of courtship and mating may include increased risks of predation, the transmission of pathogens, and a loss of foraging opportunities. Thus, a females decision to tolerate a courting male will depend upon how these costs offset the benefits of mating, which will depend on her reproductive and nutritional status. While these costs may be similar for mated and unmated females, the benefits of mating will be less for mated than virgin females. However, the cost of lost foraging opportunities may be higher for females with fewer nutritional reserves necessary for forming eggs. We examined how these costs and benefits influence the courtship and mating behaviour of male and female orb-web spiders, Argiope keyserlingi. In the field, females on webs that also contained a courting male intercepted fewer prey items per hour than females on webs without a male. In the laboratory, the presence of a courting male at the hub also attracted mantid predators to the web, increasing the risk of predation for both male and female. Staged mating experiments in the laboratory revealed that the frequency of female attacks and pre-copulatory cannibalism was greater among mated than virgin females. Feeding history did not affect aggression in virgin females but, among mated females, food-deprived spiders attacked and cannibalized males more frequently than sated females and only the latter ever remated. These differences in female behaviour influenced male mating strategies. Choice experiments demonstrated that males preferred to venture onto the silk threads of virgin rather than those of mated females. Similar patterns of mate selectivity were observed in the field; females with narrow abdomens attracted more males to the webs than females with broad abdomens, and copulations were observed more frequently among females with narrow abdomens. These smaller females are likely to be virgins that have recently molted. Males that preferentially mate with virgin females will not only avoid potentially fatal attacks but also obtain, on average, a higher fertilization success.

Signalling conflict between prey and predator attraction

Predators may utilize signals to exploit the sensory biases of their prey or their predators. The inclusion of conspicuous silk structures called decorations or stabilimenta in the webs of some orb‐web spiders (Araneae: Araneidae, Tetragnathidae, Uloboridae) appears to be an example of a sensory exploitation system. The function of these structures is controversial but they may signal to attract prey and/or deter predators. Here, we test these predictions, using a combination of field manipulations and laboratory experiments. In the field, decorations influenced the foraging success of adult female St. Andrew’s Cross spiders, Argiope keyserlingi: inclusion of decorations increased prey capture rates as the available prey also increased. In contrast, when decorations were removed, prey capture rates were low and unrelated to the amount of available prey. Laboratory choice experiments showed that significantly more flies (Chrysomya varipes; Diptera: Calliphoridae) were attracted to decorated webs. However, decorations also attracted predators (adult and juvenile praying mantids, Archimantis latistylus; Mantodea: Mantidae) to the web. St. Andrew’s Cross spiders apparently resolve the conflicting nature of a prey‐ and predator‐attracting signal by varying their decorating behaviour according to the risk of predation: spiders spun fewer decorations if their webs were located in dense vegetation where predators had greater access, than if the webs were located in sparse vegetation.

EVALUATION OF FORMULAE TO ESTIMATE THE CAPTURE AREA AND MESH HEIGHT OF ORB WEBS (ARANEOIDEA, ARANEAE)

Abstract We evaluated several formulae to estimate the capture area (the area of the web covered by capture spirals) and the mesh height (the distance between capture spirals) of orb webs constructed by Argiope keyserlingi Karsch. The accuracy of the various formulae was estimated through regression analyses. Accordingly, we propose two new formulae specifically suited for asymmetric orb webs, which provide accurate estimates of capture area and mesh height.

The role of experience in web-building spiders (Araneidae)

Abstract A typical feature of vertical orb-webs is the ‘top/bottom’ asymmetry, where the lower web region is larger than the upper web region. This asymmetry may improve prey capture success, because, sitting in the hub of the web, a spider can reach prey entangled below the hub faster than prey entangled in the area above the hub. While web asymmetry is known to vary intraspecifically, we tested if this variation also exists at the individual level and whether it is the result of experience, using two orb-web spider species, Argiope keyserlingi and Larinioides sclopetarius. The results reveal that experienced web-building spiders constructed more asymmetric webs than conspecifics deprived of any prior building experience over a period of several months. Experienced individuals invested more silk material into the web region below the hub, which covered a larger area. Moreover, web asymmetry was also influenced by previous prey capture experiences, as spiders increased the lower region of the web if it intercepted the most prey over a period of 6 days. Consequently, spiders may be able to use long-term web-building experience as well as short-term prey capture experience to build better traps. In contrast to previous views of spiders, experience can contribute to intraspecific as well as to individual variations in web design.

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 spiders 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.

LABORATORY METHODS FOR MAINTAINING AND STUDYING WEB-BUILDING SPIDERS

Abstract Web-building spiders are an important model system to address questions in a variety of biological fields. They are attractive because of their intriguing biology and because they can be fairly easily collected and maintained in the laboratory. However, the only published instructions for working with web-building spiders are somewhat outdated and not easily accessible. This paper aims to provide an up-to-date guide on how to best collect, keep and study web-building spiders. In particular, it describes how to obtain spiders by capturing them or by raising them from cocoons, how to keep and feed spiders in the laboratory and how to encourage them to build webs. Finally it describes how to document and analyze web building and web structure.

Food caching in orb-web spiders (Araneae: Araneoidea)

Abstract. Caching or storing surplus prey may reduce the risk of starvation during periods of food deprivation. While this behaviour occurs in a variety of birds and mammals, it is infrequent among invertebrates. However, golden orb-web spiders, Nephila edulis, incorporate a prey cache in their relatively permanent web, which they feed on during periods of food shortage. Heavier spiders significantly reduced weight loss if they were able to access a cache, but lost weight if the cache was removed. The presence or absence of stored prey had no effect on the weight loss of lighter spiders. Furthermore, N. edulis always attacked new prey, irrespective of the number of unprocessed prey in the web. In contrast, females of Argiope keyserlingi, who build a new web every day and do not cache prey, attacked fewer new prey items if some had already been caught. Thus, a necessary pre-adaptation to the evolution of prey caching in orb-web spiders may be a durable or permanent web, such as that constructed by Nephila.

DOES THE PRESENCE OF POTENTIAL PREY AFFECT WEB DESIGN IN ARGIOPE KEYSERLINGI (ARANEAE, ARANEIDAE)?

Abstract Orb-web spiders may anticipate their future prey environment by detecting the presence of prey and adjusting their web building behavior accordingly. Here we investigate the effect of different prey sizes and density on the web size and mesh height of the orb webs constructed by Argiope keyserlingi. The experimental design allowed the transmission of prey vibrations but prevented any capture. We found that A. keyserlingi constructed webs more frequently in the presence of prey, but did not alter the web size or mesh height of their webs.