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Secure landing is indispensable for both leaping animals and robotics. Tree frogs, renowned for their adhesive capabilities, can effectively jump across intricate 3D terrain and land safely. Compared with jumping, the mechanisms underlying their landing technique, particularly in arboreal environments, have remained largely unknown. In this study, we focused on the landing patterns of the tree frog Polypedates dennysi on horizontally placed perches, explicitly emphasizing the influence of perch diameters. Tree frogs demonstrated diverse landing postures, including the utilization of: (1) single front foot, (2) double front feet, (3) anterior bellies, (4) middle bellies, (5) posterior bellies, (6) single hind foot, or (5) double hind feet. Generally, tree frogs favoured bellies on slimmer targets but double front feet on large perches. Analysis of limb-trunk relationships revealed their adaptability to modify postures, including body positions and limb orientations, for successful landing. The variations in the initial landing postures affected the subsequent landing procedures and, consequently, the dynamics. As the initial contact position switched from front foot back to the hind foot, the stabilization time decreased at first, reaching a minimum in middle belly landings, and then increased again. The maximum vertical forces showed an inverse trend, whereas the maximum fore-aft forces continuously increased as the initial contact position switched. As the perch diameter increased, the time expended dropped, whereas the maximum impact force increased. These findings not only add to our understanding of frog landings but also highlight the necessity of considering perch diameters and landing styles when studying the biomechanics of arboreal locomotion., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2024. Published by The Company of Biologists Ltd.)

The most effective way to avoid intense inter- and intra-specific competition at the dung source, and to increase the distance to the other competitors, is to follow a single straight bearing. While ball-rolling dung beetles manage to roll their dung balls along nearly perfect straight paths when traversing flat terrain, the paths that they take when traversing more complex (natural) terrain are not well understood. In this study, we investigate the effect of complex surface topographies on the ball-rolling ability of Kheper lamarcki. Our results reveal that ball-rolling trajectories are strongly influenced by the characteristic scale of the surface structure. Surfaces with an increasing similarity between the average distance of elevations and the ball radius cause progressively more difficulties during ball transportation. The most important factor causing difficulties in ball transportation appears to be the slope of the substrate. Our results show that, on surfaces with a slope of 7.5 deg, more than 60% of the dung beetles lose control of their ball. Although dung beetles still successfully roll their dung ball against the slope on such inclinations, their ability to roll the dung ball sideways diminishes. However, dung beetles do not seem to adapt their path on inclines such that they roll their ball in the direction against the slope. We conclude that dung beetles strive for a straight trajectory away from the dung pile, and that their actual path is the result of adaptations to particular surface topographies., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2024. Published by The Company of Biologists Ltd.)

Asteraceae, one of the largest flowering plant families, are adapted to a vast range of ecological niches. Their adaptability is partially based on their strong ability to reproduce. The initial, yet challenging, step for the reproduction of animal-pollinated plants is to transport pollen to flower-visiting pollinators. We adopted Hypochaeris radicata as a model species to investigate the functional morphology of the typical floral feature of Asteraceae, a pollen-bearing style. Using quantitative experiments and numerical simulations, here we show that the pollen-bearing style can serve as a ballistic lever for catapulting pollen grains to pollinators. This can potentially be a pollen dispersal strategy to propel pollen to safe sites on pollinators' bodies, which are beyond the physical reach of the styles. Our results suggest that the specific morphology of the floret and the pollen adhesion avoid pollen waste by catapulting pollen within a specific range equal to the size of a flowerhead. The insights into the functional floral oscillation may shed light on the superficially unremarkable, but ubiquitous functional floral design of Asteraceae., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2023. Published by The Company of Biologists Ltd.)

Stick and leaf insects (Phasmatodea) are exclusively herbivores. As they settle in a broad range of habitats, they need to attach to and walk on a wide variety of plant substrates, which can vary in their surface free energy (SFE). The adhesive microstructures (AMs) on the euplantulae of phasmids are assumed to be adapted to such substrate properties. Moreover, the natural substrates can often be covered with water as a result of high relative humidity or rain. Although considerable experimental research has been carried out on different aspects of stick insect attachment, the adaptations to cope with the influence of flooded water on attachment performance remain unclear. To elucidate the role of AMs in this context, we here measured attachment forces in three species of stick insects with different AMs. The results show that attachment forces of the three species studied were influenced by the SFE and the presence of water: they all showed higher pull-off (vertical) and traction (horizontal) forces on dry surfaces, compared with when the surfaces were covered with a water film. However, the extent to which the surface properties influenced attachment differed depending on the species and its AMs. All three species showed approximately the same attachment performance on dry surfaces with different surface free energy but maintained attachment underwater to different extents., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2023. Published by The Company of Biologists Ltd.)

Animals that habitually cross the boundary between water and land face specific challenges with respect to locomotion, respiration, insulation, fouling and waterproofing. Many semi-aquatic invertebrates and plants have developed complex surface microstructures with water-repellent properties to overcome these problems, but equivalent adaptations of the skin have not been reported for vertebrates that encounter similar environmental challenges. Here, we document the first evidence of evolutionary convergence of hydrophobic structured skin in a group of semi-aquatic tetrapods. We show that the skin surface of semi-aquatic species of Anolis lizards is characterized by a more elaborate microstructural architecture (i.e. longer spines and spinules) and a lower wettability relative to closely related terrestrial species. In addition, phylogenetic comparative models reveal repeated independent evolution of enhanced skin hydrophobicity associated with the transition to a semi-aquatic lifestyle, providing evidence of adaptation. Our findings invite a new and exciting line of inquiry into the ecological significance, evolutionary origin and developmental basis of hydrophobic skin surfaces in semi-aquatic lizards, which is essential for understanding why and how the observed skin adaptations evolved in some and not other semi-aquatic tetrapod lineages., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2021. Published by The Company of Biologists Ltd.)

Intraoral food processing mechanisms are known for all major vertebrate groups, but the form and function of systems used to crush, grind or puncture food items can differ substantially between and within groups. Most vertebrates display flexible mechanisms of intraoral food processing with respect to different environmental conditions or food types. It has recently been shown that newts use cyclical loop-motions of the tongue to rasp prey against the palatal dentition. However, it remains unknown whether newts can adjust their food processing behavior in response to different food types or environmental conditions. Newts are interesting models for studying the functional adaptation to different conditions because of their unique and flexible lifestyle: they seasonally change between aquatic and terrestrial habitats, adapt their prey-capture mode to the respective environment, and consume diverse food types with different mechanical properties. Using X-ray high-speed recordings, anatomical investigations, behavioral analyses and mechanical property measurements, we tested the effects of the medium in which feeding occurs (water/air) and the food type (maggot, earthworm, cricket) on the processing behavior in Triturus carnifex We discovered that food processing, by contrast to prey capture, differed only slightly between aquatic and terrestrial habitats. However, newts adjusted the number of processing cycles to different prey types: while maggots were processed extensively, earthworm pieces were barely processed at all. We conclude that, in addition to food mechanical properties, sensory feedback such as smell and taste appear to induce flexible processing responses, while the medium in which feeding occurs appears to have less of an effect., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2020. Published by The Company of Biologists Ltd.)

The aim of this study was to find out how strongly the parasitic insect Stylops ovinae , which has tarsi equipped with tenent hairs and lacking claws, attaches to different substrates. We investigated adhesion of male S. ovinae to the abdomen of its hymenopteran host ( Andrena vaga ), the hairier abdomen of a Bombus sp. and two artificial smooth reference surfaces with different degrees of hydrophilicity. In our experiments, the male S. ovinae developed significantly higher forces on smooth surfaces. However, the forces were significantly lower on all the hymenopteran surfaces used in the experiment. The absence of anisotropy in the force grip in cranial/caudal direction relative to the host might indirectly indicate that S. ovinae generate forces by adhesion rather than mechanical interlocking with the host hairs. The tolerance of the attachment system of S. ovinae to the substrate chemistry might be explained by the primary contribution of van der Waals interactions and not capillary forces to adhesion in S. ovinae., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2020. Published by The Company of Biologists Ltd.)

Phasmatodea (stick and leaf insects) are herbivorous insects well camouflaged on plant substrates as a result of cryptic masquerade. Also, their close association with plants has allowed them to adapt to different substrate geometries and surface topographies of the plants they imitate. Stick insects are gaining increasing attention in attachment- and locomotion-focused research. However, most studies experimentally investigating stick insect attachment have been performed either on single attachment pads or on flat surfaces. In contrast, curved surfaces, especially twigs or stems of plants, are dominant substrates for phytophagous insects, but not much is known about the influence of curvature on their attachment. In this study, by combining analysis of tarsal usage with mechanical traction and pull-off force measurements, we investigated the attachment performance on curved substrates with different diameters in two species of stick insects with different tarsal lengths. We provide the first quantitative data for forces generated by stick insects on convex curved substrates and show that the curvature significantly influences attachment ability in both species. Within the studied range of substrate curvatures, traction force decreases and pull-off force increases with increasing curvature. Shorter tarsi demonstrate reduced forces; however, tarsus length only has an influence for diameters thinner than the tarsal length. The attachment force generally depends on the number of tarsi/tarsomeres in contact, tarsus/leg orientation and body posture on the surface. Pull-off force is also influenced by the tibiotarsal angle, with higher pull-off force for lower angles, while traction force is mainly influenced by load, i.e. adduction force., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2020. Published by The Company of Biologists Ltd.)

The evolution of wings has played a key role in the success of insect species, allowing them to diversify to fill many niches. Insect wings are complex multifunctional structures, which not only have to withstand aerodynamic forces but also need to resist excessive stresses caused by accidental collisions. This Commentary provides a summary of the literature on damage-reducing morphological adaptations in wings, covering natural causes of wing collisions, their impact on the structural integrity of wings and associated consequences for both insect flight performance and life expectancy. Data from the literature and our own observations suggest that insects have evolved strategies that (i) reduce the likelihood of wing damage and (ii) allow them to cope with damage when it occurs: damage-related fractures are minimized because wings evolved to be damage tolerant and, in the case of wing damage, insects compensate for the reduced aerodynamic efficiency with dedicated changes in flight kinematics., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2020. Published by The Company of Biologists Ltd.)

Stick insects are well adapted in their locomotion to various surfaces and topographies of natural substrates. Single pad measurements characterised the pretarsal arolia of these insects as shear-sensitive adhesive pads and the tarsal euplantulae as load-sensitive friction pads. Different attachment microstructures on the euplantulae reveal an adaptation of smooth euplantulae to smooth surfaces and nubby eupantulae to a broader range of surface roughness. However, how different attachment pads and claws work in concert and how strong the contribution of different structures is to the overall attachment performance remains unclear. We therefore assessed combinatory effects in the attachment system of two stick insect species with different types of euplantular microstructures by analysing their usage in various posture situations and the performance on different levels of substrate roughness. For comparison, we provide attachment force data of the whole attachment system. The combination of claws, arolia and euplantulae provides mechanical interlocking on rough surfaces, adhesion and friction on smooth surfaces in different directions, and facilitates attachment on different inclines and on a broad range of surface roughness, with the least performance in the range 0.3-1.0 µm. On smooth surfaces, stick insects use arolia always, but employ euplantulae if the body weight can generate load on them (upright, wall). On structured surfaces, claws enable mechanical interlocking at roughnesses higher than 12 µm. On less-structured surfaces, the attachment strength depends on the use of pads and, corroborating earlier studies, favours smooth pads on smooth surfaces, but nubby euplantulae on micro-rough surfaces., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2019. Published by The Company of Biologists Ltd.)

The shell of Mytilus edulis is a multilayered system for protecting this bivalve. In contrast to well-developed research on the nacre materials, the protective function of the complete M. edulis shell has not been widely studied. In particular, the question of why nacre is situated on the inner side of the shell rather than on the outer side remains unclear. Herein, the acid resistance of different shell layers was compared using etching tests and the mechanical protection performance of the shell was tested using three-point bending. Two bending loads, including static and dynamic, were applied on the shell samples from outside in (i.e. out-in bending) and from inside out (i.e. in-out bending), respectively. Our etching results show that the external prismatic calcite endows M. edulis with stronger acid resistance than if nacre was on the outside. In contrast, the static out-in and in-out bending tests reveal that a better mechanical protection of the shell against slow mechanical attacks is achieved if the nacre is on the outside. However, the shell has the same mechanical properties against dynamic mechanical attacks regardless of nacre location. Briefly, the nacre should be on the outside of the shell for better mechanical protection while the outside location of the prismatic layer offers a stronger resistance against etching. The inside natural location of nacre is a compromise between mechanical and chemical protection mechanisms against a complex survival environment. This strongly contributes to our understanding of biological design principles and further development of shell-inspired protective materials., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2019. Published by The Company of Biologists Ltd.)

Attachment to surfaces is a major aspect of an animal's interaction with the environment. Consequently, shaping of the attachment system in relation to weight load and substrate is considered to have occurred mainly by natural selection. However, sexual selection may also be important because many animals attach to their partner during mating. The two hypotheses generate opposing predictions in species where males are smaller than females. Natural selection predicts that attachment ability will scale positively with load, and hence body size, and so will be larger in females than males. Sexual selection predicts attachment forces in males will be larger than those in females, despite the males' smaller size because males benefit from uninterrupted copulation by stronger attachment to the female. We tested these predictions in the common bedbug Cimex lectularius , a species in which both sexes, as well as nymphs, regularly carry large loads: blood meals of up to 3 times their body weight. By measuring attachment forces to smooth surfaces and analysing in situ fixed copulating pairs and the morphology of attachment devices, we show that: (i) males generate twice the attachment force of females, despite weighing 15% less; (ii) males adhere to females during copulation using hairy tibial adhesive pads; (iii) there are more setae, and more setae per unit area, in the pads of males than in those of females but there is no difference in the shape of the tarsal setae; and (iv) there is an absence of hairy tibial attachment pads and a low attachment force in nymphs. These results are consistent with a sexually selected function of attachment in bedbugs. Controlling sperm transfer and mate guarding by attaching to females during copulation may also shape the evolution of male attachment structures in other species. More generally, we hypothesise the existence of an arms race in terms of male attachment structures and female counterparts to impede attachment, which may result in a similar evolutionary diversification to male genitalia., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2019. Published by The Company of Biologists Ltd.)

Many parasitic wasps use slender and steerable ovipositors to lay eggs in hosts hidden in substrates, but it is currently unknown how steering is achieved. The ovipositors generally consist of three longitudinally connected elements, one dorsal and two ventral valves that can slide along each other. For the parasitic wasp Diachasmimorpha longicaudata , it has been shown that protraction of the ventral valves causes incurving of the ventral valves towards the dorsal one, which results in a change in probing direction. We hypothesize that this shape change is due to differences in bending stiffness along the ovipositor. Alignment of the stiff tip of the dorsal valve with a more flexible ventral S-shaped region situated just behind the tip straightens this S-bend and results in upwards rotation of the ventral tip. We show that the S-shaped region of the ventral valves has a low bending stiffness because it contains soft materials such as resilin. In contrast, the large cross-sectional area of the dorsal valve tip area probably results in a high bending stiffness. Elsewhere, the dorsal valve is less stiff than the ventral valves. Our results support the hypothesis that the interaction between the stiff dorsal valve portion and the more flexible S-shaped region co-determines the configurational tip changes required for steering the ovipositor in any desired direction along curved paths in the substrate. This provides novel insights in the understanding of steering mechanisms of the hymenopteran ovipositor, and for application in man-made probes., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2019. Published by The Company of Biologists Ltd.)

Prehensile and gripping organs are recurring structures in different organisms that enhance friction by the reinforcement and redirection of normal forces. The relationship between organ structure and biomechanical performance is poorly understood, despite a broad relevance for microhabitat choice, movement ecology and biomimetics. Here, we present the first study of the biomechanics of prehensile feet in long-legged harvestmen. These arachnids exhibit the strongest sub-division of legs among arthropods, permitting extreme hyperflexion (i.e. curling up the foot tip). We found that despite the lack of adhesive foot pads, these moderately sized arthropods are able to scale vertical smooth surfaces, if the surface is curved. Comparison of three species of harvestmen differing in leg morphology shows that traction reinforcement by foot wrapping depends on the degree of leg sub-division, not leg length. Differences are explained by adaptation to different microhabitats on trees. The exponential increase of foot section length from distal to proximal introduces a gradient of flexibility that permits adaptation to a wide range of surface curvature while maintaining integrity at strong flexion. A pulley system of the claw depressor tendon ensures the controlled flexion of the high number of adesmatic joints in the harvestman foot. These results contribute to the general understanding of foot function in arthropods and showcase an interesting model for the biomimetic engineering of novel transportation systems and surgical probes., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2019. Published by The Company of Biologists Ltd.)

The louse fly Crataerina pallida is an obligate blood-sucking ectoparasite of the common swift Apus apus As a result of reduction of the wings, C. pallida is unable to fly; thus, an effective and reliable attachment to their host's plumage is of utmost importance. The attachment system of C. pallida shows several modifications in comparison to that of other calyptrate flies, notably the large tridentate claws and the dichotomously shaped setae located on the pulvilli. Based on data from morphological analysis, confocal laser scanning microscopy, cryo-scanning electron microscopy and attachment force experiments performed on native (feathers) as well as artificial substrates (glass, epoxy resin and silicone rubber), we showed that the entire attachment system is highly adapted to the fly's lifestyle as an ectoparasite. The claws in particular are the main contributor to strong attachment to the host. Resulting attachment forces on feathers make it impossible to detach C. pallida without damage to the feathers or to the legs of the louse fly itself. Well-developed pulvilli are responsible for the attachment to smooth surfaces. Both dichotomously shaped setae and high setal density explain high attachment forces observed on smooth substrates. For the first time, we demonstrate a material gradient within the setae, with soft, resilin-dominated apical tips and stiff, more sclerotized bases in Diptera. The empodium seems not to be directly involved in the attachment process, but it might operate as a cleaning device and may be essential to maintain the functionality of the entire attachment system., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2018. Published by The Company of Biologists Ltd.)

During the lifespan of the stick insect Carausius morosus , their long and narrow tibiae experience substantial compressive loads. The mechanical load on the tibiae increases as the weight of the insect rises. The increase in body weight is accompanied by a notable increase in the insect's body size and, accordingly, by an increase in the length of the tibiae. Both of these changes can raise the risk of buckling of the tibiae. In this study, we tracked changes in the material and geometric properties of the hindleg tibia of C. morosus during growth. The results show that although buckling (either by Euler buckling or local buckling) is the dominant failure mode under compression, the tibia is very capable of maintaining its buckling resistance in each postembryonic developmental stage. This is essentially the result of a compromise between the increasing slenderness of the tibia and its increasing material stiffness. The use of an optimal radius to thickness ratio, a soft resilin-dominated core, and chitin fibres oriented in both longitudinal and circumferential directions are presumably additional strategies preventing buckling of the tibia. This study, providing the first quantitative data on changes in the biomechanical properties of cuticle during the entire life of an insect, is expected to shed more light on the structure-property-function relationship in this complex biological composite., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2018. Published by The Company of Biologists Ltd.)

Foraging is one of the main evolutionary driving forces shaping the phenotype of organisms. In predators, a significant, though understudied, cost of foraging is the risk of being injured by struggling prey. Hunting spiders that feed on dangerous prey like ants or other spiders are an extreme example of dangerous feeding, risking their own life over a meal. Here, we describe an intriguing example of the use of attachment silk (piriform silk) for prey immobilization that comes with the costs of reduced silk anchorage function, increased piriform silk production and additional modifications of the extrusion structures (spigots) to prevent their clogging. We show that the piriform silk of gnaphosids is very stretchy and tough, which is an outstanding feat for a functional glue. This is gained by the combination of an elastic central fibre and a bi-layered glue coat consisting of aligned nanofibrils. This represents the first tensile test data on the ubiquitous piriform gland silk, adding an important puzzle piece to the mechanical catalogue of silken products in spiders., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2017. Published by The Company of Biologists Ltd.)

The ability of many insects to walk on vertical smooth surfaces such as glass or even on the ceiling has fascinated biologists for a long time, and has led to the discovery of highly specialized adhesive organs located at the distal end of the animals' legs. So far, research has primarily focused on structural and ultrastructural investigations leading to a deeper understanding of adhesive organ functionality and to the development of new bioinspired materials. Genetic approaches, e.g. the analysis of mutants, to achieve a better understanding of adhesive organ differentiation have not been used so far. Here, we describe the first Drosophila melanogaster mutant that develops malformed adhesive organs, resulting in a complete loss of climbing ability on vertical smooth surfaces. Interestingly, these mutants fail to make close contact between the setal tips and the smooth surface, a crucial condition for wet adhesion mediated by capillary forces. Instead, these flies walk solely on their claws. Moreover, we were able to show that the mutation is caused by a P-element insertion into the Su(z)2 gene locus. Remobilization of the P-element restores climbing ability. Furthermore, we provide evidence that the P-element insertion results in an artificial Su(z)2 transcript, which most likely causes a gain-of-function mutation. We presume that this transcript causes deregulation of yet unknown target genes involved in pulvilli differentiation. Our results nicely demonstrate that the genetically treatable model organism Drosophila is highly suitable for future investigations on adhesive organ differentiation., (© 2015. Published by The Company of Biologists Ltd.)

Numerous studies deal with insect attachment onto surfaces with different roughness; however, little is known about insect attachment onto surfaces that have different chemistry. In the present study, we describe the attachment structures of the water-lily leaf beetle Galerucella nymphaeae and test the hypothesis that the larval and adult stages generate the strongest attachment on surfaces with contact angles that are similar to those of leaves of their host plants. The larvae bear a smooth attachment system with arolium-like structures at their legs and a pygopodium at the abdomen tip. Adults have pointed setae on the ventral side of the two proximal tarsomeres and densely arranged spatula-shaped ones on their third tarsomere. In a centrifugal force tester, larvae and adults attained the highest friction forces and safety factors on surfaces with a water contact angle of 83 deg compared to those of 6, 26 and 109 deg. This comes close to the contact angle of their host plant Nuphar lutea (86 deg). The similarity in larval and adult performances might be a result of the similar chemical composition of their attachment fluid. We compare our findings with previous studies on the forces that insects generate on surfaces with different surface energies., (© 2014. Published by The Company of Biologists Ltd.)

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., (© 2014. Published by The Company of Biologists Ltd.)

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.

Monogeneans, which are mainly fish ectoparasites, use various types of haptoral (posterior) attachment apparatus to secure their attachment onto their hosts. However, it remains unclear how strongly a monogenean can attach onto its host. In the present study, we aimed for the first time to (1) measure pull-off forces required to detach a pair of clamp-bearing monogeneans, Diplozoon paradoxum, from gills of Abramis brama and (2) determine the contribution of muscles to the clamp movements. A mean force of 6.1±2.7 mN (~246 times the animals' weight) was required to dislodge a paired D. paradoxum vertically from the gills. There were significant differences (P<0.05, Tukey test) between the widths of clamp openings in D. paradoxum treated in three different solutions: the widest clamp openings were observed in the monogeneans treated in 100 mmol l(-1) potassium chloride solution (58.26±13.44 μm), followed by those treated in 20 mmol l(-1) magnesium chloride solution (37.91±7.58 μm), and finally those treated in filtered lake water (20.16±8.63 μm). This suggests that the closing of the clamps is probably not due to the continuous contraction of extrinsic muscles but is caused by the elasticity of the clamp material and that muscle activity is required for clamp opening.

The activity pattern of the claw retractor muscle of Pachnoda marginata peregrina beetles was examined in this study. We found this muscle to be located in the tibia, without a femoral part, as is the case in other insects. Electromyograms of the muscle revealed a rather similar activity pattern during beetle locomotion on rough and smooth substrates. We recorded units with small and large amplitude, with the smaller unit being active during almost the entire stance phase, and the larger unit active roughly in the first half of stance. Small but significant differences were found in the precise onset and end of activity. Both small and large units began their activity earlier on the rough surface. Although there was no difference at the end of activity in the small unit between surfaces, the large unit ended its activity significantly earlier on the rough substrate. The spike frequencies on both surfaces were also significantly different for small and large units. The small unit showed a higher spike frequency on the smooth surface, while the large unit had a higher spike frequency on the rough surface. From our experiments, we conclude that the muscle is controlled by the same basic activity pattern on different surfaces, with some adjustments that are due to sensory feedback. The adjustments cause differences in onset and end of activity, as well as in spike frequency of the involved muscle units.

Insect tarsal adhesive structures secrete a thin layer of fluid into the contact area. It was previously reported that the presence of this fluid significantly increases adhesion on various substrata. Previous data obtained from representatives of different insect groups suggest a difference not only in the chemical composition of the fluid, but also in its physical properties. In the present study, we have measured for the first time changes in the droplet geometry over time and the evaporation rate of the fluid in flies (Calliphora vicina) and beetles (Coccinella septempunctata) by the use of atomic force microscopy. Flattened droplets of the beetle had lower evaporation rates than hemispherical footprints of the fly. Within 1 h, the droplet volume reduced to 21% of the initial volume for the fly, and to 65% for the beetle, suggesting a larger fraction of volatile compounds in the fly fluid. It was revealed that drop geometry changes significantly during evaporation and shows pinning effects for the fly footprints due to an assumed self-organizing oil layer on top of the water fraction of the micro-emulsion. The data obtained suggest that the adhesion strength in capillarity-based switchable adhesive systems must be time-dependent because of the specific evaporation rate of the adhesive fluid. These results are important for our understanding of the functional mechanism of insect adhesive systems and also for biomimetics of artificial capillarity-based adhesive systems.

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 activity of the middle part of the claw retractor muscle was examined in two species of stick insects (Carausius morosus and Cuniculina impigra). We performed electromyographic recordings while the animals were standing on a smooth or a rough surface of a platform in horizontal, vertical or inverted positions, as well as during rotations of the platform. We recorded tonic and phasic motor units. The tonic units were active all the time without significant differences in spike frequency, regardless of the position of the animals (although there was a tendency for higher discharge frequencies to occur during platform rotations). The phasic units were active almost exclusively during platform movement. In contrast to the tonic units, we detected significant differences in the activities of the phasic units; namely, higher spike frequencies during rotations compared with the stationary phases, especially for rotations into 'more awkward' positions. A comparison of the two species revealed no difference in muscle activity, despite differences in the animals' tarsal attachment structures. The same was true when comparing the muscle activity of the two species on both the smooth and the rough surfaces.

The surface of some insect eyes consists of arrays of cuticular protuberances, which are 50-300 nm in diameter, and are termed corneal nipples or ommatidia gratings. They were widely reported to reduce the reflectance for normally incident light, contributing to camouflage by reducing glare to predators, while furthermore enhancing the intake of light, which is especially important for nocturnal insects. Our preliminary observations suggest a third function: in contrast to the rest of the body, ommatidia of various insects remain clean, even in a heavy contaminated environment. In order to prove such an anti-contamination hypothesis of these structures, we measured the adhesive properties of polymer moulds of insect ommatidia, and compared these data with control surfaces having the same curvature radii but lacking such a nanostructure. A scanning electron microscope (SEM) study and force measurements using an atomic force microscope (AFM) on the eye surfaces of three different insect species, dragonfly Aeshna mixta (Odonata), moth Laothoe populi (Lepidoptera) and fly Volucella pellucens (Diptera), were undertaken. We revealed that adhesion is greatly reduced by corneal grating in L. populi and V. pellucens when compared with their smooth controls. The smooth cornea of A. mixta showed no statistically significant difference to its control. We assume that this anti-adhesive phenomenon is due to a decrease in the real contact area between contaminating particles and the eye's surface. Such a combination of three functions in one nanostructure can be interesting for the development of industrial multifunctional surfaces capable of enhancing light harvesting while reducing light reflection and adhesion.

Setose pads of aquatic Epeorus assimilis larvae are specialised structures located ventrally on the part of the gill lamella contacting the substrate and were suggested to have an attachment function in strong currents. In order to test the role of these setose pads in underwater attachment for the first time, we measured friction (shear) forces generated by the gill lamellae on solid substrates. Moreover, the influence of a different kind of surface roughness on attachment was investigated. Scanning electron microscopy showed that four different seta types can be found on the pads. Our results revealed that the pads significantly contributed to friction force generated on smooth and on some rough substrates but not on certain surfaces of intermediate roughness. The contribution of pads to the friction coefficient in experiments was lower than expected under natural conditions, which may be caused by a smaller contact area between the pads and the substrate (changes in material properties, lack of the active control of body positioning of the larva). The friction coefficient of the gill lamellae with the substrate depended on the surface roughness of the substrate and on the pulling direction. These results suggest that interlocking between structures of the insect cuticle and substrate irregularities, as well as molecular adhesion, contribute to friction.

For the first time, the terminal elements (spatulae) of setal (hairy) attachment devices of the beetle Gastrophysa viridula (Coleoptera, Chrysomelidae) and the fly Lucilia caesar (Diptera, Calliphoridae) were studied using transmission X-ray microscopy (TXM) with a lateral resolution of about 30 nm. Since images are taken under ambient conditions, we demonstrate here that this method can be applied to study the contact behaviour of biological systems, including animal tenent setae, in a fresh state. We observed that the attached spatulae show a viscoelastic behavior increasing the contact area and providing improved adaptability to the local topography of the surface. The technique can be extended to TXM tomography, which would provide three-dimensional information and a deeper insight into the details of insect attachment structures.

Each tarsus of Coreus marginatus L. (Coreidae) bears a pair of smooth flexible pulvilli adapted for attachment to relatively smooth surfaces, such as their host plant Rumex crispus L. (Polygonaceae). This account quantifies insect attachment abilities on smooth surfaces at various stages of ontogenesis. Friction (shear) force (FF) of adults and juvenile insects was measured by the use of a computer controlled centrifugal force tester equipped with a fibre optical sensor. Pad area, body size and body mass were determined individually for each experimental insect. Light microscopy revealed no difference in pulvilli area between different leg pairs. Pulvilli area demonstrated a stronger increase with increasing linear dimensions, as predicted by scaling laws. Since friction coefficient (relationship between FF and body weight) (FC) was always higher than 1, it was concluded that adhesion has strongly contributed to the measured friction. The frictional properties of pulvilli do not change during ontogenesis. Thus, only the growth of pulvilli and, therefore, the increased contact area, contribute to the increasing attachment ability in insects at later larval stages. Due to different scaling of the body mass and area of attachment organs, smaller insects attach relatively more strongly. Both FF and FC were higher in experiments in which higher angular acceleration (AC) was applied. Lateral tenacity determined individually for experimental insects and pooled for all animals and accelerations is 0.097 N m(-2). These data led us to suggest that viscosity of the pad secretion and/or visco-elastic properties of the foam-like material of pulvilli play an important role in the attachment ability of insects.

This paper studies slide-resisting forces generated by claws in the free-walking beetle Pachnoda marginata (Coleoptera, Scarabaeoidea) with emphasis on the relationship between the dimension of the claw tip and the substrate texture. To evaluate the force range by which the claw can interact with a substrate, forces generated by the freely moving legs were measured using a load cell force transducer. To obtain information about material properties of the claw, its mechanical strength was tested in a fracture experiment, and the internal structure of the fractured claw material was studied by scanning electron microscopy. The bending stress of the claw was evaluated as 143.4-684.2 MPa, depending on the cross-section model selected. Data from these different approaches led us to propose a model explaining the saturation of friction force with increased texture roughness. The forces are determined by the relative size of the surface roughness R(a) (or an average particle diameter) and the diameter of the claw tip. When surface roughness is much bigger than the claw tip diameter, the beetle can grasp surface irregularities and generate a high degree of attachment due to mechanical interlocking with substrate texture. When R(a) is lower than or comparable to the claw tip diameter, the frictional properties of the contact between claw and substrate particles play a key role in the generation of the friction force.