Jonas O. Wolff

Surface roughness effects on attachment ability of the spider Philodromus dispar (Araneae, Philodromidae)

SUMMARY The morphology of the tarsal attachment system of the running spider Philodromus dispar Walckenaer 1826 (Araneae, Philodomidae) was studied using scanning electron microscopy and its performance was experimentally tested in traction force measurements. Each pretarsus bears a hierarchically built hairy adhesive pad that consists of a dense array of flattened setae covered with numerous microtrichia on the substrate-facing side. Microtrichia carry spatulate end tips that allow close contact with the substrate. Forces were estimated on tethered living specimens on rough epoxy resin surfaces (asperity size 0.3, 1, 3, 9 and 12 μm) and on a smooth surface as a control. A strong reduction in adhesion was observed for substrates with an asperity size of 0.3 and 1 μm. Comparison of the present data with previous results of different organisms demonstrates that the difference in force reduction on rough substrata depends on the dimensions of terminal contact elements (spatulae).

The influence of humidity on the attachment ability of the spider Philodromus dispar (Araneae, Philodromidae)

Hairy attachment devices that are not supplemented with fluid secretion have evolved independently in lizards and spiders. van der Waals forces have previously been shown to be responsible for excellent adhesive properties of these structures, but it has recently been reported that wetting phenomena also play an important role in such ‘dry adhesives’. To investigate the effect of ambient humidity on the attachment of the living spider Philodromus dispar, traction force was measured on a smooth epoxy resin surface at relative humidities (RHs) of 15, 50, 70, 80 and 99 per cent. The results show that attachment ability is significantly higher at an intermediate humidity compared with that in a dry atmosphere and at high humidity. Water condensation on the substrate surface almost completely abolishes adhesion. Experimental results obtained may be explained by an increase in capillarity or changes in mechanical properties of setae and spatulae owing to water absorption by the cuticle at an intermediate RH. The results obtained show dry adhesion limits under different environmental conditions and are important for understanding spider biology.

The whole is more than the sum of all its parts: collective effect of spider attachment organs

Dynamic attachment is the key to moving safely and fast in a three-dimensional environment. Among lizards, hexapods and arachnids, several lineages have evolved hairy foot pads that can generate strong friction and adhesion on both smooth and rough surfaces. A strongly expressed directionality of attachment structures results in an anisotropy of frictional properties, which might be crucial for attachment control. In a natural situation, more than one leg is usually in contact with the substrate. In order to understand the collective effect of hairy foot pads in the hunting spider Cupiennius salei (Arachnida, Ctenidae), we performed vertical pulling experiments combined with stepwise disabling of the pads. We found the attachment force of the spider to be not simply the sum of single leg forces because with leg pair deactivation a much greater decrease in attachment forces was found than was predicted by just the loss of available adhesive pad area. This indicates that overall adhesion ability of the spider is strongly dependent on the antagonistic work of opposing legs, and the apparent contact area plays only a minor role. It is concluded that the coordinated action of the legs is crucial for adhesion control and for fast and easy detachment. The cumulative effect of anisotropic fibrillar adhesive structures could be potentially interesting for biomimetic applications, such as novel gripping devices.

The Great Silk Alternative: Multiple Co-Evolution of Web Loss and Sticky Hairs in Spiders

Spiders are the most important terrestrial predators among arthropods. Their ecological success is reflected by a high biodiversity and the conquest of nearly every terrestrial habitat. Spiders are closely associated with silk, a material, often seen to be responsible for their great ecological success and gaining high attention in life sciences. However, it is often overlooked that more than half of all Recent spider species have abandoned web building or never developed such an adaptation. These species must have found other, more economic solutions for prey capture and retention, compensating the higher energy costs of increased locomotion activity. Here we show that hairy adhesive pads (scopulae) are closely associated with the convergent evolution of a vagrant life style, resulting in highly diversified lineages of at least, equal importance as the derived web building taxa. Previous studies often highlighted the idea that scopulae have the primary function of assisting locomotion, neglecting the fact that only the distal most pads (claw tufts) are suitable for those purposes. The former observations, that scopulae are used in prey capture, are largely overlooked. Our results suggest the scopulae evolved as a substitute for silk in controlling prey and that the claw tufts are, in most cases, a secondary development. Evolutionary trends towards specialized claw tufts and their composition from a low number of enlarged setae to a dense array of slender ones, as well as the secondary loss of those pads are discussed further. Hypotheses about the origin of the adhesive setae and their diversification throughout evolution are provided.

Radial arrangement of Janus-like setae permits friction control in spiders

Dynamic attachment is the key to move on steep surfaces, with mechanisms being still not well understood. The hunting spider Cupiennius salei (Arachnida, Ctenidae) possesses hairy attachment pads (claw tufts) at its distal legs, consisting of directional branched setae. The morphological investigation revealed that adhesive setae are arranged in a radial manner within the distal tarsus. Friction of claw tufts on smooth glass was measured to reveal the functional effect of seta arrangement within the pad. Measurements revealed frictional anisotropy in both longitudinal and transversal directions. Contact behaviour of adhesive setae was investigated in a reflection interference contrast microscope (RICM). Observations on living spiders showed, that only a small part of the hairy pads is in contact at the same time. Thus the direction of frictional forces is depending on leg placement and rotation. This may aid controlling the attachment to the substrate.

Comparative morphology of pretarsal scopulae in eleven spider families.

Many wandering spiders bear attachment pads (scopulae) on their tarsi, consisting of hierarchically-branching adhesive setae. Amongst spider families and even species, these show remarkable differences in morphology. Using scanning electron microscopy, the scopula microstructure of sixteen spider species was described, with the focus on pretarsal scopulae (claw tufts). Area and shape of the claw tuft, seta and setule density, as well as seta and spatula dimensions were analysed and compared. Claw tufts of the majority of species studied show a similar gradient in size and shape from anterior to posterior legs: the dimension of pads increases, while setal density decreases. Commonly, there is also a gradient of both the seta and spatula size within the claw tuft: Setae become larger from the proximal to the distal part of the pad, and spatulae size increases in the same direction at the level of individual seta. Often, different hierarchical levels of claw tuft organisation are differently expressed in different species: Species with lower setal density usually have broader setae. Smaller spatula size often implicates higher setule density. Evolutionary and ecological aspects of the scopula origin are discussed.

Composition and substrate-dependent strength of the silken attachment discs in spiders

Araneomorph spiders have evolved different silks with dissimilar material properties, serving different purposes. The two-compound pyriform secretion is used to glue silk threads to substrates or to other threads. It is applied in distinct patterns, called attachment discs. Although ubiquitously found in spider silk applications and hypothesized to be strong and versatile at low material consumption, the performance of attachment discs on different substrates remains unknown. Here, we analyse the detachment forces and fracture mechanics of the attachment discs spun by five different species on three different substrates, by pulling on the upstream part of the attached thread. Results show that although the adhesion of the pyriform glue is heavily affected by the substrate, even on Teflon it is frequently strong enough to hold the spiders weight. As plant surfaces are often difficult to wet, they are hypothesized to be the major driving force for evolution of the pyriform secretion.

How to stay on mummy's back: Morphological and functional changes of the pretarsus in arachnid postembryonic stages.

A specific type of maternal care occurs in several groups of Arachnida: mothers carry their offspring on their back (pulli-carrying behaviour). In scorpions, whip scorpions and whip spiders it is the prenymphal stage that settles on the mother. The prenymph is not yet fully developed for a free life and very limited in its mobility, but its feet are equipped with special adhesive organs (arolia) that become lost at the nymphal stage. Here we study the morphology, ultrastructure and mechanical function of the arolia. In scorpions (Scorpiones) the contact area between arolia and substrate and thus adhesion of the pad is controlled by the antagonistic work of hydrostatic pressure and muscular retraction. Arolia of whip scorpions (Thelyphonida) do not require muscular action for strong attachment. Arrays of long, branching fibres in the mesocuticle lead to high compliancy of the pad. In whip spiders (Amblypygi) the prenymphal pretarsus is already equipped with sclerites and claws. Its arolium is retained in nymphs and adults in some taxa, but acquires a more complex structure. These results contribute to our knowledge on the postembryonic development of arachnids and to the understanding of attachment pad evolution among arthropods. Some of the described developmental, structural, and mechanical phenomena are not known from other animals and might be of potential interest for further biomimetic developments.

Gluing the ‘unwettable’: soil-dwelling harvestmen use viscoelastic fluids for capturing springtails

Gluing can be a highly efficient mechanism of prey capture, as it should require less complex sensory–muscular feedback. Whereas it is well known in insects, this mechanism is much less studied in arachnids, except spiders. Soil-dwelling harvestmen (Opiliones, Nemastomatidae) bear drumstick-like glandular hairs (clavate setae) at their pedipalps, which were previously hypothesized to be sticky and used in prey capture. However, clear evidence for this was lacking to date. Using high-speed videography, we found that the harvestman Mitostoma chrysomelas was able to capture fast-moving springtails (Collembola) just by a slight touch of the pedipalp. Adhesion of single clavate setae increased proportionally with pull-off velocity, from 1 μN at 1 μm s−1 up to 7 μN at 1 mm s−1, which corresponds to the typical weight of springtails. Stretched glue droplets exhibited characteristics of a viscoelastic fluid forming beads-on-a-string morphology over time, similar to spider capture threads and the sticky tentacles of carnivorous plants. These analogies indicate that viscoelasticity is a highly efficient mechanism for prey capture, as it holds stronger the faster the struggling prey moves. Cryo-scanning electron microscopy of snap-frozen harvestmen with glued springtails revealed that the gluey secretions have a high affinity to wet the microstructured cuticle of collembolans, which was previously reported to be barely wettable for both polar and non-polar liquids. Glue droplets can be contaminated with the detached scaly setae of collembolans, which may represent a counter-adaptation against entrapment by the glue, similar to the scaly surfaces of Lepidoptera and Trichoptera (Insecta) facilitating escape from spider webs.

Adhesive foot pads: an adaptation to climbing? An ecological survey in hunting spiders.

Hairy pads relying on dry adhesion are fascinating structures that convergently evolved among spiders and lizards. Numerous studies underline the functional aspects leading to their strong adhesion to smooth surfaces, but rarely has their role been studied in the context of natural habitats and surfaces that animals are faced with. In hunting spiders, the hairy foot pads (claw tufts) underneath the paired claws are assumed to be an adaptation to a climbing lifestyle, particularly on smooth plant surfaces. However, surfaces that are too smooth for claws to generate a sufficient grip are rather rare in natural habitats and above-ground habitats are occupied by hunting spiders both with and without claw tufts. In this study we estimated the proportion of claw tuft-bearing hunting spiders (ct+ ratio) among microhabitat-specific assemblages by conducting both a field study and a meta-analysis approach. The effect of surface characteristics, structure fragmentation and altitude of the microhabitat niche on the ct+ ratio was analyzed. We hypothesized that the ct+ ratio will be higher in (i) hunting spider assemblages obtained from microhabitats above the ground than from those at the ground and (ii) in hunting spider assemblages obtained from microhabitats with smoother surfaces (tree foliage) than those with rougher surfaces (barks, stones), and lower in (iii) hunting spider assemblages obtained from microhabitats with more fragmented structures (small leaves) than in those with comparable but less fragmented structures (large leaves). We found the ct+ ratio to be significantly affected by the microhabitats distance from the ground, whereas surface characteristics and fragmentation of the substrates were of minor importance. This suggests that claw tufts are highly beneficial when the microhabitats height exceeds a value where the additional pad-related costs are exceeded by the costs of dropping. We assume the benefit to be mainly due to gaining a high safety factor at a lower energy demand if compared to alternative attachment devices (claws, silk). The previously presumed enhanced access to new microhabitat sites may play only a minor role as hunting spiders without claw tufts are present in most microhabitats.