SciCombinator

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Concept: Spider web

169

Correlated evolution of traits can act synergistically to facilitate organism function. But, what happens when constraints exist on the evolvability of some traits, but not others? The orb web was a key innovation in the origin of >12,000 species of spiders. Orb evolution hinged upon the origin of novel spinning behaviors and innovations in silk material properties. In particular, a new major ampullate spidroin protein (MaSp2) increased silk extensibility and toughness, playing a critical role in how orb webs stop flying insects. Here, we show convergence between pseudo-orb-weaving Fecenia and true orb spiders. As in the origin of true orbs, Fecenia dragline silk improved significantly compared to relatives. But, Fecenia silk lacks the high compliance and extensibility found in true orb spiders, likely due in part to the absence of MaSp2. Our results suggest how constraints limit convergent evolution and provide insight into the evolution of nature’s toughest fibers.

Concepts: Evolution, Organism, Insect, Materials science, Spider silk, Spider, Spider web, Orb-weaver spider

34

Natural materials are renowned for exquisite designs that optimize function, as illustrated by the elasticity of blood vessels, the toughness of bone and the protection offered by nacre. Particularly intriguing are spider silks, with studies having explored properties ranging from their protein sequence to the geometry of a web. This material system, highly adapted to meet a spider’s many needs, has superior mechanical properties. In spite of much research into the molecular design underpinning the outstanding performance of silk fibres, and into the mechanical characteristics of web-like structures, it remains unknown how the mechanical characteristics of spider silk contribute to the integrity and performance of a spider web. Here we report web deformation experiments and simulations that identify the nonlinear response of silk threads to stress–involving softening at a yield point and substantial stiffening at large strain until failure–as being crucial to localize load-induced deformation and resulting in mechanically robust spider webs. Control simulations confirmed that a nonlinear stress response results in superior resistance to structural defects in the web compared to linear elastic or elastic-plastic (softening) material behaviour. We also show that under distributed loads, such as those exerted by wind, the stiff behaviour of silk under small deformation, before the yield point, is essential in maintaining the web’s structural integrity. The superior performance of silk in webs is therefore not due merely to its exceptional ultimate strength and strain, but arises from the nonlinear response of silk threads to strain and their geometrical arrangement in a web.

Concepts: Tensile strength, Elasticity, Linear elasticity, Silk, Spider silk, Spider, Spider web, Spinneret

28

Although it is well known that spatial learning can be important in the biology of predators that actively move around in search for food, comparatively little is known about ways in which spatial learning might function in the strategies of sit-and-wait predators. In this study, Cyclosa octotuberculata, an orb-web spider that uses its legs to contract radial threads of its web to increase thread tension, was trained to capture prey in limited web sectors. After training, spiders that had captured prey in horizontal web sectors applied more tension on radial threads connected to horizontal sectors than spiders that had captured prey in vertical sectors. This result suggests that the effect of experience on C. octotuberculata’s behaviour is not expressed in the way the trained spider responds to prey-derived stimuli and is instead expressed in behaviour by which the spider anticipates the likely direction from which prey will arrive in the future. This illustrates that learning can be important even when the predator remains in one location during foraging bouts.

Concepts: Predation, Knowledge, Optimal foraging theory, Spider, Spider web, Orb-weaver spider, Araneomorphae, Ambush predator

28

Among a myriad of spider web geometries, the orb web presents a fascinating, exquisite example in architecture and evolution. Orb webs can be divided into two categories according to the capture silk used in construction: cribellate orb webs (composed of pseudoflagelliform silk) coated with dry cribellate threads and ecribellate orb webs (composed of flagelliform silk fibres) coated by adhesive glue droplets. Cribellate capture silk is generally stronger but less-extensible than viscid capture silk, and a body of phylogenic evidence suggests that cribellate capture silk is more closely related to the ancestral form of capture spiral silk. Here, we use a coarse-grained web model to investigate how the mechanical properties of spiral capture silk affect the behaviour of the whole web, illustrating that more elastic capture spiral silk yields a decrease in web system energy absorption, suggesting that the function of the capture spiral shifted from prey capture to other structural roles. Additionally, we observe that in webs with more extensible capture silk, the effect of thread strength on web performance is reduced, indicating that thread elasticity is a dominant driving factor in web diversification.

Concepts: World Wide Web, Web 2.0, Silk, Adhesive, Spider silk, Spider, Spider web, Spider anatomy

24

Tephritid pests controlled through the sterile insect technique (SIT) are mass-reared and subsequently released in affected areas. Several quality parameters are currently used to test adults, but none take into account interactions with a predator. When sterile males are released in the field, they will need to avoid predators until they reach sexual maturity and survive long enough to mate with wild females. Spiders are one of the most common predators that flies may encounter in release sites. In this study, we evaluated the antipredator behavior of a mass-reared sterile unisexual strain (‘Tapachula-7’) of the Mexican fruit fly Anastrepha ludens (Diptera: Tephritidae) against their spider predators. We sampled spiders in citrus trees to determine which families could be more common. We established the baseline activity rates of sterile Tapachula-7 (Tap-7) flies in comparison with wild flies. We also tested the behavior of the fertile and sterile bisexual strain and wild flies against hunting spiders (Family Salticidae) and orb building spiders (Family Tetragnathidae). We recorded 18 spider families, with Salticidae being the most dominant. Tap-7 flies diminished their activity in comparison with wild males at 1800┬áh but showed similar activity levels earlier in the day. When exposed to orb-web spiders (Leucauge venusta), Tap-7, fertile and sterile males from the bisexual strain had similar rates of survival, but Tap-7 males showed lower survival than wild males. Against hunting spiders (Phidippus audax), wild males had higher probability of defensive wing displays, but there was no difference in spider attack rates. In general, sterile Tap -7 males performed as well as males from the bisexual strain, although they had lower survival than wild males. This could be due to either mass-rearing and/or irradiation effects. We recommend the use of the defensive wing display behavior as a quality parameter and propose a rapid and effective method to evaluate fly activity. The efficiency of SIT will be improved if released sterile males have the same antipredator repertoire as their wild counterparts.

Concepts: Insect, Predation, Antipredator adaptation, Fly, Spider, Spider web, Salticidae, Phidippus

11

Capture success of spider webs has been associated with their microstructure, ornamentation, and wind-induced vibrations. Indirect evidence suggests that statically charged objects can attract silk thread, but web deformations induced by charged insects have not yet been described. Here, we show under laboratory conditions that electrostatically charged honeybees, green bottle flies, fruit flies, aphids, and also water drops falling near webs of cross-spiders (Araneus diadematus) induce rapid thread deformation that enhances the likelihood of physical contact, and thus of prey capture.

Concepts: Insect, Spider silk, Spider, Spider web, Orb-weaver spider, Spinneret, European garden spider, Araneus

5

Spider orb webs are multifunctional, acting to absorb prey impact energy and transmit vibratory information to the spider. This paper explores the links between silk material properties, propagation of vibrations within webs and the ability of the spider to control and balance web function. Combining experimental and modelling approaches, we contrast transverse and longitudinal wave propagation in the web. It emerged that both transverse and longitudinal wave amplitude in the web can be adjusted through changes in web tension and dragline silk stiffness, i.e. properties that can be controlled by the spider. In particular, we propose that dragline silk supercontraction may have evolved as a control mechanism for these multifunctional fibres. The various degrees of active influence on web engineering reveals the extraordinary ability of spiders to shape the physical properties of their self-made materials and architectures to affect biological functionality, balancing trade-offs between structural and sensory functions.

Concepts: Function, Physical quantities, World Wide Web, Web 2.0, Spider silk, Spider, Transverse wave, Spider web

5

Spiders constitute one of the most successful clades of terrestrial predators [1]. Their extraordinary diversity, paralleled only by some insects and mites [2], is often attributed to the use of silk, and, in one of the largest lineages, to stereotyped behaviors for building foraging webs of remarkable biomechanical properties [1]. However, our understanding of higher-level spider relationships is poor and is largely based on morphology [2-4]. Prior molecular efforts have focused on a handful of genes [5, 6] but have provided little resolution to key questions such as the origin of the orb weavers [1]. We apply a next-generation sequencing approach to resolve spider phylogeny, examining the relationships among its major lineages. We further explore possible pitfalls in phylogenomic reconstruction, including missing data, unequal rates of evolution, and others. Analyses of multiple data sets all agree on the basic structure of the spider tree and all reject the long-accepted monophyly of Orbiculariae, by placing the cribellate orb weavers (Deinopoidea) with other groups and not with the ecribellate orb weavers (Araneoidea). These results imply independent origins for the two types of orb webs (cribellate and ecribellate) or a much more ancestral origin of the orb web with subsequent loss in the so-called RTA clade. Either alternative demands a major reevaluation of our current understanding of the spider evolutionary chronicle.

Concepts: Evolution, Insect, Phylogenetics, Polyphyly, Spider, Mygalomorphae, Spider web, Cribellum

5

Spider webs are made of silk, the properties of which ensure remarkable efficiency at capturing prey. However, remaining on, or near, the web exposes the resident spiders to many potential predators, such as ants. Surprisingly, ants are rarely reported foraging on the webs of orb-weaving spiders, despite the formidable capacity of ants to subdue prey and repel enemies, the diversity and abundance of orb-web spiders, and the nutritional value of the web and resident spider. We explain this paradox by reporting a novel property of the silk produced by the orb-web spider Nephila antipodiana (Walckenaer). These spiders deposit on the silk a pyrrolidine alkaloid (2-pyrrolidinone) that provides protection from ant invasion. Furthermore, the ontogenetic change in the production of 2-pyrrolidinone suggests that this compound represents an adaptive response to the threat of natural enemies, rather than a simple by-product of silk synthesis: while 2-pyrrolidinone occurs on the silk threads produced by adult and large juvenile spiders, it is absent on threads produced by small juvenile spiders, whose threads are sufficiently thin to be inaccessible to ants.

Concepts: Predation, Ecology, Silk, Spider silk, Spider, Spider web, Orb-weaver spider, Spinneret

3

Modern orb-weaving spiders use micron-sized glue droplets on their viscid silk to retain prey in webs. A combination of low molecular weight salts and proteins makes the glue viscoelastic and humidity responsive in a way not easily achieved by synthetic adhesives. Optically, the glue droplet shows a heterogeneous structure, but the spatial arrangement of its chemical components is poorly understood. Here, we use optical and confocal Raman microscopy to show that salts and proteins are present ubiquitously throughout the droplet. The distribution of adhesive proteins in the peripheral region explains the superior prey capture performance of orb webs as it enables the entire surface area of the glue droplet to act as a site for prey capture. The presence of salts throughout the droplet explains the recent Solid-State NMR results that show salts directly facilitate protein mobility. Understanding the function of individual glue components and the role of the droplet’s macro-structure can help in designing better synthetic adhesives for humid environments.

Concepts: Understanding, Molecule, Liquid, Silk, Adhesive, Spider silk, Spider, Spider web