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Abstract Background Heelwalkers possess a highly modified tarsal attachment system. All extant species lift the distalmost tarsomere permanently off the substrate and primarily use their euplantulae for locomotion. The combination of a smooth adhesive pad (arolium) on the pretarsus and fibrillary attachment pads on the euplantulae offers valuable insights for translational approaches, but its infra-order diversity remains unexplored. Results We explored the morphology of the tarsal attachment apparatus of Mantophasmatodea based on a representative taxon sampling spanning a large fraction of species of this group and compared morphological differences in the specialized morphology of this system across species and sexes. Our scanning electron microscope investigation of the tarsi of 11 species (52% of all described extant species) revealed an overall very consistent ground pattern and almost no specific adaptations. There are only minor, but mostly clade-specific differences in the shape of the adhesive setae on the tarsal euplantulae and in the morphology and density of the acanthae on the pretarsal arolium. Both features differ primarily between Austrophasmatidae in comparison to the remaining Mantophasmatodea taxa. Conclusion We conclude that the strong specialization of the mantophasmatodean tarsal attachment sufficiently copes with the diversity of substrates the insects are exposed to.

Abstract WingAnalogy is a computer tool for automated insect wing morphology and asymmetry analysis. It facilitates project management, enabling users to import pairs of wing images obtained from individual insects, such as left and right, fore- and hindwings. WingAnalogy employs image processing and computer vision to segment wing structures and extract cell boundaries, and junctions. It quantifies essential metrics encompassing cell and wing characteristics, including area, length, width, circularity, and centroid positions. It enables users to scale and superimpose wing images utilizing Particle Swarm Optimization (PSO). WingAnalogy computes regression, Normalized Root Mean Square Error (NRMSE), various cell-based parameters, and distances between cell centroids and junctions. The software generates informative visualizations, aiding researchers in comprehending and interpreting asymmetry patterns. WingAnalogy allows for dividing wings into up to five distinct wing cell sets, facilitating localized comparisons. The software excels in report generation, providing detailed asymmetry measurements in PDF, CSV, and TXT formats.

Abstract Background Centipedes are terrestrial and predatory arthropods that possess an evolutionary transformed pair of appendages used for venom injection—the forcipules. Many arthropods incorporate reinforcing elements into the cuticle of their piercing or biting structures to enhance hardness, elasticity or resistance to wear and structural failure. Given their frequent exposure to high mechanical stress, we hypothesise that the cuticle of the centipede forcipule might be mechanically reinforced. With a combination of imaging, analytical techniques and mechanical testing, we explore the centipede forcipule in detail to shed light on its morphology and performance. Additionally, we compare these data to characteristics of the locomotory leg to infer evolutionary processes. Results We examined sclerotization patterns using confocal laser-scanning microscopy based on autofluorescence properties of the cuticle (forcipule and leg) and elemental composition by energy-dispersive X-ray spectroscopy in representative species from all five centipede lineages. These experiments revealed gradually increasing sclerotization towards the forcipular tarsungulum and a stronger sclerotization of joints in taxa with condensed podomeres. Depending on the species, calcium, zinc or chlorine are present with a higher concentration towards the distal tarsungulum. Interestingly, these characteristics are more or less mirrored in the locomotory leg’s pretarsal claw in Epimorpha. To understand how incorporated elements affect mechanical properties, we tested resistance to structural failure, hardness (H) and Young’s modulus (E) in two representative species, one with high zinc and one with high calcium content. Both species, however, exhibit similar properties and no differences in mechanical stress the forcipule can withstand. Conclusions Our study reveals similarities in the material composition and properties of the forcipules in centipedes. The forcipules transformed from an elongated leg-like appearance into rigid piercing structures. Our data supports their serial homology to the locomotory leg and that the forcipule’s tarsungulum is a fusion of tarsus and pretarsal claw. Calcium or zinc incorporation leads to comparable mechanical properties like in piercing structures of chelicerates and insects, but the elemental incorporation does not increase H and E in centipedes, suggesting that centipedes followed their own pathways in the evolutionary transformation of piercing tools.

Abstract In many insects the surface of the eye is nanostructured by arrays of protuberances termed ommatidial gratings which provide the cuticle with anti-reflective, anti-wetting and self-cleaning properties. The hypothesised anti-contamination role of the gratings against dust and pollen results from theoretical predictions on grating geometry and experiments on synthetic replicas of ommatidia surfaces but has not yet been proven in an animal. Whiteflies are biological test beds for anti-contamination surfaces because they deliberately distribute wax particles extruded from abdominal plates over their entire bodies. The numerous particles protect the animal against water evaporation and radiation, but may severely impair vision. Using scanning electron microscopy (SEM) and CryoSEM, we here show that the cornea of whiteflies exhibits ~ 220 nm wide mesh-like structures forming hexagonal gratings with thin ~ 40 nm connecting walls. Quantitative measurements of wax particles on the eye show that the nanostructures reduce particle contamination by more than ~ 96% compared to other areas of the cuticle. Altogether, our study is the first description of a predicted optimized grating geometry for anti-contamination in an arthropod. The findings serve as evidence of the high effectiveness of nanostructured surfaces for reducing contact area and thus adhesion forces between biological surfaces and contaminating particles.

Many insect species have found their way into ageing research as small and easy-to-keep model organisms. A major sign of ageing is the loss of locomotory functions due to neuronal disorders or tissue wear. Soft and pliable attachment pads on the tarsi of insects adapt to the substrate texture to maximize their real contact area and, thereby, generate attachment during locomotion. In the majority of stick insects, adhesive microstructures covering those pads support attachment. Stick insects do not molt again after reaching the imaginal stage; hence, the cuticle of their pads is subject to continuous ageing. This study aims to quantify how attachment ability changes with age in the stick insect Sungaya aeta Hennemann, 2023 and elucidate the age effects on the material and microstructure of the attachment apparatus. Attachment performance (adhesion and friction forces) on substrates with different roughnesses was compared between two different age groups, and the change of attachment performance was monitored extending over a larger time frame. Ageing effects on the morphology of the attachment pads and the autofluorescence of the cuticle were documented using light, scanning electron, and confocal laser scanning microscopy. The results show that both adhesion and friction forces decline with age. Deflation of the pads, scarring of the cuticle, and alteration of the autofluorescence, likely indicating stiffening of the cuticle, were observed to accumulate over time. This would reduce the attachment ability of the insect, as pads lose their pliant properties and cannot properly maintain sufficient contact area with the substrate.

The mechanism by which insects achieve attachment and locomotion across diverse substrates has long intrigued scientists, prompting extensive research on the functional morphology of attachment pads. In stick insects, attachment and locomotion are facilitated by two distinct types of smooth cuticular attachment pads: the primary adhesion force-generating arolium and the friction force-generating euplantulae. They are both supported by an adhesive secretion delivered into the interspace between the attachment pads and the substrate. In this study, we analysed and compared internal morphology, material composition and ultrastructure, as well as the transportation pathways in both adhesive organs in the stick insect Medauroidea extradentata using scanning electron microscopy, micro-computed tomography, light microscopy, and confocal laser scanning microscopy. Our observations revealed structural differences between both attachment pads, reflecting their distinct functionality. Furthermore, our results delineate a potential pathway for adhesive secretions, originating from exocrine epidermal cells and traversing various layers before reaching the surface. Within the attachment pad, the fluid may influence the viscoelastic properties of the pad and control the attachment/detachment process. Understanding the material composition of attachment pads and the distribution process of the adhesive secretion can potentially aid in the development of more effective artificial attachment systems.

This study focuses on experimental testing of the contamination hypothesis and examines how the contamination of insect adhesive pads with three-dimensional epicuticular waxes of different plant species contributes to the reduction of insect attachment. We measured traction forces of tethered Chrysolina fastuosa male beetles having hairy adhesive pads on nine wax-bearing plant surfaces differing in both shape and dimensions of the wax structures and examined insect adhesive organs after they have contacted waxy substrates. For comparison, we performed the experiments with the same beetle individuals on a clean glass sample just before (gl1) and immediately after (gl2) the test on a plant surface. The tested insects showed a strong reduction of the maximum traction force on all waxy plant surfaces compared to the reference experiment on glass (gl1). After beetles have walked on waxy plant substrates, their adhesive pads were contaminated with wax material, however, to different extents depending on the plant species. The insects demonstrated significantly lower values of both the maximum traction force and the first peak of the traction force and needed significantly longer time to reach the maximum force value in the gl2 test than in the gl1 test. These effects were especially pronounced in cases of the plant surfaces covered with wax projections having higher aspect ratios. The data obtained clearly indicated the impact of waxy plant surfaces on the insect ability to subsequently attach to the clean smooth surface. This effect is caused by the contamination of adhesive pads and experimentally supports the contamination hypothesis.

Abstract We explore the use of movable automata in numerical modelling of male competition for territory. We used territorial dragonflies as our biological inspiration for the model, assuming two types of competing males: (a) faster and larger males that adopt a face‐off strategy and repulse other males; (b) slower and smaller males that adopt a non‐aggressive strategy. The faster and larger males have higher noise intensity, leading to faster motion and longer conservation of motion direction. The velocity distributions resemble the Maxwell distributions of velocity, expected in Brownian dynamics, with two probable velocities and distribution widths for the two animal subpopulations. The fast animals' trajectories move between visually fixed density folds of the slower animal subpopulation. A correlation is found between individual velocity and individual area distribution, with smaller animals concentrated in a region of small velocities and areas. Attraction between animals results in a modification of the system behaviour, with larger animals spending more time being surrounded by smaller animals and being slowed down by their interaction with the surroundings. Overall, the study provides insights into the dynamics of animal competition for territory and the impact of attraction between animals.

Abstract Scaling in insect wings is a complex phenomenon that seems pivotal in maintaining wing functionality. In this study, the relationship between wing size and the size, location, and shape of wing cells in dragonflies and damselflies (Odonata) is investigated, aiming to address the question of how these factors are interconnected. To this end, WingGram, the recently developed computer‐vision‐based software, is used to extract the geometric features of wing cells of 389 dragonflies and damselfly wings from 197 species and 16 families. It has been found that the cell length of the wings does not depend on the wing size. Despite the wide variation in wing length (8.42 to 56.5 mm) and cell length (0.1 to 8.5 mm), over 80% of the cells had a length ranging from 0.5 to 1.5 mm, which was previously identified as the critical crack length of the membrane of locust wings. An isometric scaling of cells is also observed with maximum size in each wing, which increased as the size increased. Smaller cells tended to be more circular than larger cells. The results have implications for bio‐mimetics, inspiring new materials and designs for artificial wings with potential applications in aerospace engineering and robotics.

Abstract Insect feeding structures, such as mandibles, interact with the ingesta (food or/and substrate) and can be adapted in morphology, composition of material and mechanical properties. The foraging on abrasive ingesta, as on algae covering rocks, is particularly challenging because the mandibles will be prone to wear and structural failure, thus suggesting the presence of mandibular adaptations to accompany this feeding behavior. Adaptations to this are well studied in the mouthparts of molluscs and sea urchins, but for insects there are large gaps in our knowledge. In this study, we investigated the mandibles of a grazing insect, the larvae of the trichopteran Glossosoma boltoni. Using scanning electron microscopy, wear was documented on the mandibles. The highest degree was identified on the medial surface of the sharp mandible tip. Using nanoindentation, the mechanical properties, such as hardness and Young’s modulus, of the medial and lateral mandible cuticles were tested. We found, that the medial cuticle of the tip was significantly softer and more flexible than the lateral one. These findings indicate that a self-sharpening mechanism is present in the mandibles of this species, since the softer medial cuticle is probably abraded faster than the harder lateral one, leading to sharp mandible tips. To investigate the origins of these properties, we visualized the degree of tanning by confocal laser scanning microscopy. The autofluorescence signal related to the mechanical property gradients. The presence of transition and alkaline earth metals by energy dispersive X-ray spectroscopy was also tested. We found Ca, Cl, Cu, Fe, K, Mg, Mn, P, S, Si, and Zn in the cuticle, but the content was very low and did not correlate with the mechanical property values.

Abstract Marine mammals host a great variety of parasites, which usually co-evolved in evolutionary arms races. However, little is known about the biology of marine mammal insect parasites, and even less about physical aspects of their life in such a challenging environment. One of 13 insect species that manage to endure long diving periods in the open sea is the seal louse, Echinophthirius horridus, parasitising true seals. Its survival depends on its specialised adaptations for enduring extreme conditions such as hypoxia, temperature changes, hydrostatic pressure, and strong drag forces during host dives. To maintain a grip on the seal fur, the louse’s leg morphology is equipped with modified snap hook claws and soft pad-like structures that enhance friction. Through techniques including CLSM, SEM, and histological staining, we have examined the attachment system’s detailed structure. Remarkably, the seal louse achieves exceptional attachment forces on seal fur, with safety factors (force per body weight) reaching 4500 in average measurements and up to 18000 in peak values, indicating superior attachment performance compared to other insect attachment systems. These findings underscore the louse’s remarkable adaptations for life in a challenging marine environment, shedding light on the relationship between structure and function in extreme ecological niches.

Abstract Suckermouth armoured catfish (Loricariidae) are a highly speciose and diverse freshwater fish family, which bear upper and lower lips forming an oral disc. Its hierarchical organisation allows the attachment to various natural surfaces. The discs can possess papillae of different shapes, which are supplemented, in many taxa, by small horny projections, i.e. unculi. Although these attachment structures and their working mechanisms, which include adhesion and interlocking, are rather well investigated in some selected species, the loricariid oral disc is unfortunately understudied in the majority of species, especially with regard to comparative aspects of the diverse oral structures and their relationship to the ecology of different species. In the present paper, we investigated the papilla and unculi morphologies in 67 loricariid species, which inhabit different currents and substrates. We determined four papilla types and eight unculi types differing by forms and sizes. Ancestral state reconstructions strongly suggest convergent evolution of traits. There is no obvious correlation between habitat shifts and the evolution of specific character states. From handling the structures and from drying artefacts we could infer some information about their material properties. This, together with their shape, enabled us to carefully propose hypotheses about mechanisms of interactions of oral disc structures with natural substrates typical for respective fish species.

Abstract To adhere their silk threads for the construction of webs and to fix the dragline, spiders produce attachment discs of piriform silk. Uniquely, the aquatic spider Argyroneta aquatica spends its entire life cycle underwater. Therefore, it has to glue its attachment discs to substrates underwater. Here we show that Argyroneta aquatica applies its thread anchors within an air layer around the spinnerets maintained by superhydrophobic setae. During spinning, symmetric movements of the spinnerets ensure retaining air in the contact area. The flat structure of the attachment discs is thought to facilitate fast curing of the piriform adhesive cement and improves the resistance against drag forces. Pull-off tests on draglines connected with attachment discs on different hydrophilic substrates point to dragline rupture as the failure mode. The Young´s modulus of the dragline (8.3 GPa) is within the range as in terrestrial spiders. The shown structural and behavioral adaptations can be the model for new artificial underwater gluing devices.

This article introduces WingSegment, a MATLAB app‐designed tool employing a hybrid approach of computer vision and graph theory for precise insect wing image segmentation. WingSegment detects cells, junctions, Pterostigma, and venation patterns, measuring geometric features and generating Voronoi patterns. The tool utilizes region‐growing, thinning, and Dijkstra's algorithms for boundary detection, junction identification, and vein path extraction. It provides histograms and box plots of geometric features, facilitating comprehensive wing analysis. WingSegment's efficiency is validated through comparisons with established tools and manual measurements, demonstrating accurate results. The tool further enables exporting detected boundaries as FreeCAD macro files for 3D modeling and printing, supporting finite element analysis. Beyond advancing insect wing morphology understanding, WingSegment holds broader implications for diverse planar structures, including leaves and geocells. This tool not only enhances automated geometric analysis and 3D model generation in insect wing studies but also contributes to the broader advancement of analysis, 3D printing, and modeling technologies across various planar structures.

The ectoparasitic seal louse, Echinophthirius horridus infects harbour (Phoca vitulina) and grey seals (Halichoerus grypus) in the North and Baltic Sea. The endoparasitic heartworm Acanthocheilonema spirocauda parasitizes the right heart and blood vessels of harbour seals. The complete lifecycle of the heartworm is not entirely understood although the seal louse is assumed to serve as vector for its transmission. Knowledge about the impact of both parasite species on host health are scarce. In this study, necropsy data and archived parasites of harbour and grey seals in German waters were analysed to determine long-term seal louse (SLP) and heartworm prevalence (HWP) from 2014 to 2021. Histology, microbiology and scanning electron microscopy (SEM) were applied on seal louse infected and uninfected skin to investigate associated lesions and the health impact. During the study period, HWP in harbour seals was 13%, the SLP in harbour seals was 4% and in grey seals 10%. HWP of harbour seals was significantly higher during the winter months compared to the summer. SLP in adults was significantly higher in comparison to juvenile harbour seals. SLP varied significantly between grey seals from the North and Baltic Sea. Filarial nematodes were detected in the haemocoel, pharynx, and intestine of E. horridus highlighting the seal louse as vector for heartworms. Alopecia and folliculitis were associated with the attachment posture of E. horridus and microbiological investigations isolated bacteria commonly associated with folliculitis.

The ability of many insects to adhere vertically or even upside down to smooth substrates is closely related to the morphology and distribution of the adhesive structures on their legs. During locomotion, the legs are in direct contact with different substrates, and it is hypothesized that the adhesive structures have been evolved as an adaption to smooth substrates in specific environments. To investigate whether there is a relationship between the presence of adhesive structures and the combined effects of different environments and mating behavior, we compared five species of tiger beetles belonging to two tribes living in arboreal and non-arboreal environments, respectively. In three non-arboreal species, we found a specific type of adhesive structure consisting of elongated spoon-like setae present on the protarsi of males but absent on the male meso- and metatarsi and on females. In Tricondyla pulchripes, an arboreal species living on stems, we found three types of adhesive setae on male protarsi, while only two types of setae were found on male meso- and metatarsi and on females. In Neocollyris linearis, an arboreal species living on leaves, we found three types of adhesive setae on male pro-, meso- and meta-tarsi but only two types of adhesive setae on females. The adaptive evolution of these adhesive structures was probably driven by the selective pressures of both mating behavior and the presence of smooth substrates in the respective environments. It is discussed that the adhesive structures in tiger beetles may be an adaptive evolutionary response to the plant surfaces and may play an important role in species differentiation.

To reduce negative effects of floral visitation by ants, which do not serve as reliable cross-pollinators, some plants have developed a non-floral, stem-based defense mechanism called greasy pole syndrome. In the present study, we examined the effects of two surface features (trichomes and three-dimensional epicuticular wax coverage) on stems of Alliaria petiolata plants on visiting frequencies, travelled distances, and running velocities of Lasius niger ants. The experiments were performed with stem samples prepared from different (apical and basal) stem portions showing different surface morphologies (smooth control, covered by wax and trichomes + wax, respectively). The control, mechanically wiped stem samples lacking any surface features were significantly more often visited by ants, where they travelled significantly longer distances and moved with significantly higher velocities, compared to the intact stems. The apical and basal stem portions showed no significant differences in the measured parameters. Based on data obtained, we conclude about the main contribution of the wax to the greasy pole function of the A. petiolata stem via reduction of ant adhesion to the wax-bearing stem surface, whereas trichomes presumably serve as the first barrier for ants approaching usually from the ground level and protect the fragile wax coverage from an excessive deterioration.

Insect attachment devices enhance adhesion to complex-geometry substrates by increasing the real contact area. In nature, insects mainly interact with plant surfaces that are often covered by 3D wax structures. Here, we describe, discuss, and give a mechanical interpretation of plant waxes and the possible fracture mechanisms of these wax structures during their interactions with the adhesive pads of insects. It is argued that these plant surface microstructures significantly influence insect adhesion through reducing the contact area and contaminating the insect pads.

Abstract The presence of wear of the joint cuticle of the beetle legs is demonstrated experimentally and is represented by abrasive wear. Presence of the abrasive wear in the natural‐habiting beetles of various families is also established. Abrasive wear comprises of both two‐body and three‐body wear as a result of the action of particles that have entered the joint. The presence of the sliding wear in the absence of abrasive particles is not found. The wear effects are represented by microcuttings, indentations, cracking, delamination, brittle fracturing, and destruction of the surface layer (epicuticle). The correlation of quantitative indicators of wear between the types of experimental substrates (paper, wood, sand) and the length of exposure of beetles on a particular substrate turned out to be very weak. It is hypothesized that the lubricating material in the leg joints of beetles may provide an anti‐wear function by binding particles and facilitating their removal thereby reducing surface damage by sharp asperities, and protecting the damaged surface to prevent further wear.

Suspension feeding via setae collecting particles is common within Crustacea. Even though the mechanisms behind it and the structures themselves have been studied for decades, the interplay between the different setae types and the parameters contributing to their particle collecting capacities remain partly enigmatic. Here, we provide a numerical modeling approach to understand the relationship among the mechanical property gradients, the mechanical behavior and the adhesion of setae, and the feeding efficiency of the system. In this context, we set-up a simple dynamic numerical model that takes all of these parameters into account and describes the interaction with food particles and their delivery into the mouth opening. By altering the parameters, it was unraveled that the system performs best when the long and short setae have different mechanical properties and different degrees of adhesion since the long setae generate the feeding current and the short ones establish the contact with the particle. This protocol can be applied to any system in the future as the parameters (i.e., properties and arrangement of particles and setae) can be easily altered. This will shed light on the biomechanical adaptations of these structures to suspension feeding and provide inspiration for biomimetics in the field of filtration technologies.

Abstract Phylliidae are herbivorous insects exhibiting impressive cryptic masquerade and are colloquially called “walking leaves”. They imitate angiosperm leaves and their eggs often resemble plant seeds structurally and in some cases functionally. Despite overall morphological similarity of adult Phylliidae, their eggs reveal a significant diversity in overall shape and exochorionic surface features. Previous studies have shown that the eggs of most Phylliidae possess a specialised attachment mechanism with hierarchical exochorionic fan-like structures (pinnae), which are mantled by a film of an adhesive secretion (glue). The folded pinnae and glue respond to water contact, with the fibrous pinnae expanding and the glue being capable of reversible liquefaction. In general, the eggs of phylliids appear to exhibit varying structures that were suggested to represent specific adaptations to the different environments the eggs are deposited in. Here, we investigated the diversity of phylliid eggs and the functional morphology of their exochorionic structure. Based on the examination of all phylliid taxa for which the eggs are known, we were able to characterise eleven different morphological types. We explored the adhesiveness of these different egg morphotypes and experimentally compared the attachment performance on a broad range of substrates with different surface roughness, surface chemistry and tested whether the adhesion is replicable after detachment in multiple cycles. Furthermore, we used molecular phylogenetic methods to reconstruct the evolutionary history of different egg types and their adhesive systems within this lineage, based on 53 phylliid taxa. Our results suggest that the egg morphology is congruent with the phylogenetic relationships within Phylliidae. The morphological differences are likely caused by adaptations to the specific environmental requirements for the particular clades, as the egg morphology has an influence on the performance regarding the surface roughness. Furthermore, we show that different pinnae and the adhesive glue evolved convergently in different species. While the evolution of the Phylliidae in general appears to be non-adaptive judging on the strong similarity of the adults and nymphs of most species, the eggs represent a stage with complex and rather diverse functional adaptations including mechanisms for both fixation and dispersal of the eggs.

Micro- and nanostructures of the white and black scales on the tarsi of the mosquito Aedes albopictus are analysed using scanning electron microscopy, transmission electron microscopy, and fluorescence microscopy. Reflectance spectra of the white areas are measured. No clear difference is present in the morphology of micro- and nanostructures of black and white scales in SEM and TEM, but black scales contain a dark pigment. The white colour of the scales has a structural origin. The structural white produced by the micro- and nanostructures of the scales on the tarsi of Ae. albopictus appears bright and is angle-dependent, since the reflected light changes according to the angle detection and according to the tarsus orientation. The optical appearance of the scale system of Ae. albopictus has a complex nature and can be explained by the combination of several effects. Among them, multiple refraction and reflection on the micro- and nanostructures of the scales are mainly responsible for the white appearance. The results suggest that mosquito scales, in addition to their superhydrophobic function, produce structural white. The biological role of white and black patches in mate recognition and defensive behaviour in the mosquitoes of the genus Aedes is hypothesized.

The goal of the research is to describe the aggregation process inside the mucilage produced by plant seeds using molecular dynamics (MD) combined with time series algorithmic analysis based on the recurrence plots. The studied biological molecules model is seed mucilage composed of three main polysaccharides, i.e. pectins, hemicellulose, and cellulose. The modeling of biological molecules is based on the assumption that a classical–quantum passage underlies the aggregation process in the mucilage, resulting from non-covalent interactions, as they affect the macroscopic properties of the system. The applied recurrence plot approach is an important tool for time series analysis and data mining dedicated to analyzing time series data originating from complex, chaotic systems. In the current research, we demonstrated that advanced algorithmic analysis of seed mucilage data can reveal some features of the dynamics of the system, namely temperature-dependent regions with different dynamics of increments of a number of hydrogen bonds and regions of stable oscillation of increments of a number of hydrophobic–polar interactions. Henceforth, we pave the path for automatic data-mining methods for the analysis of biological molecules with the intermediate step of the application of recurrence plot analysis, as the generalization of recurrence plot applications to other (biological molecules) datasets is straightforward.

Abstract The gastric mill of Decapoda is a unique feature, which comprises teeth, stabilizing ossicles, and particle sorting setae. Involved in the fragmentation and sorting of the food, this structure serves as interface between the organism and its environment. As material properties complement morphology and hold information about function and trophic preferences, we here provide a basis for more comparative research on gastric mills. For gastric mill components of the adult red swamp crayfish Procambarus clarkii, we studied (a) the micro-structure via scanning electron microscopy, (b) the elemental composition by energy-dispersive X-ray spectroscopy, (c) the heterogeneities in material properties and degree of tanning (autofluorescence) by confocal laser scanning microscopy, and (d) the mechanical properties hardness and elasticity by nanoindentation technique. The morphology and micro-structure were previously described for this species, but the mechanical properties and the autofluorescence were not studied before. As epicuticle and exocuticle could be analyzed individually, material property gradients, with values decreasing from the interacting surface towards interior, could be determined. Finally, we were able to relate the mechanical property data with the elemental composition and the degree of tanning. We found that the epicuticle of the teeth is among the hardest and stiffest biological materials in invertebrates due to the incorporations of high proportions of silicon.

Abstract The jumping mechanism with supporting morphology and kinematics is described in the marsh beetle Scirtes hemisphaericus (Coleoptera: Scirtidae). In marsh beetles, the jump is performed by the hind legs by the rapid extension of the hind tibia. The kinematic parameters of the jump are: 139–1536 m s−2 (acceleration), 0.4–1.9 m s−1 (velocity), 2.7–8.4 ms (time to take-off), 0.2–5.4 × 10–6 J (kinetic energy) and 14–156 (g-force). The power output of a jumping leg during the jumping movement is 3.5 × 103 to 9.6 × 103 W kg−1. A resilin-bearing elastic extensor ligament is considered to be the structure that accumulates the elastic strain energy. The functional model of the jumping involving an active latching mechanism is proposed. The latching mechanism is represented by the conical projection of the tibial flexor sclerite inserted into the corresponding socket of the tibial base. Unlocking is triggered by the contraction of flexor muscle pulling the tibial flexor sclerite backwards which in turn comes out of the socket. According to the kinematic parameters, the time of full extension of the hind tibia, and the value of the jumping leg power output, this jumping mechanism is supposed to be latch-mediated spring actuation using the contribution of elastically stored strain energy.

Abstract The molluscan feeding structure is the radula, a chitinous membrane with teeth, which are highly adapted to the food and the substrate to which the food is attached. In Polyplacophora and Patellogastropoda, the handling of hard ingesta can be facilitated by high content of chemical compounds containing Fe or Si in the tooth cusps. Other taxa, however, possess teeth that are less mineralized, even though animals have to avoid structural failure or high wear during feeding as well. Here, we investigated the gastropod Gastropteron rubrum, feeding on hard Foraminifera, diatoms and Porifera. Tooth morphologies and wear were documented by scanning electron microscopy and their mechanical properties were tested by nanoindentation. We determined that gradients of hard‐ and stiffness run along each tooth, decreasing from cusp to basis. We also found that inner lateral teeth were harder and stiffer than the outer ones. These findings allowed us to propose hypotheses about the radula‐ingesta interaction. In search for the origins of the gradients, teeth were visualized using confocal laser scanning microscopy, to determine the degree of tanning, and analyzed with energy‐dispersive X‐ray spectroscopy, to test the elemental composition. We found that the mechanical gradients did not have their origins in the elemental content, as the teeth did not contain high proportions of metals or other minerals. This indicates that their origin might be the degree of tanning. However, in the tooth surfaces that interact with the ingesta high Si and Ca contents were determined, which is likely an adaptation to reduce wear.

Gripping, holding, and moving objects are among the main functional purposes of robots. Ever since automation first took hold in society, optimizing these functions has been of high priority, and a multitude of approaches has been taken to enable cheaper, more reliable, and more versatile gripping. Attempts are ongoing to reduce grippers’ weight, energy consumption, and production and maintenance costs while simultaneously improving their reliability, the range of eligible objects, working loads, and environmental independence. While the upper bounds of precision and flexibility have been pushed to an impressive level, the corresponding solutions are often dependent on support systems (e.g., sophisticated sensors and complex actuation machinery), advanced control paradigms (e.g., artificial intelligence and machine learning), and typically require more maintenance owed to their complexity, also increasing their cost. These factors make them unsuited for more modest applications, where moderate to semi-high performance is desired, but simplicity is required. In this paper, we attempt to highlight the potential of the tarsal chain principle on the example of a prototype biomimetic gripping device called the TriTrap gripper, inspired by the eponymous tarsal chain of insects. Insects possess a rigid exoskeleton that receives mobility due to several joints and internally attaching muscles. The tarsus (foot) itself does not contain any major intrinsic muscles but is moved by an extrinsically pulled tendon. Just like its biological counterpart, the TriTrap gripping device utilizes strongly underactuated digits that perform their function using morphological encoding and passive conformation, resulting in a gripper that is versatile, robust, and low cost. Its gripping performance was tested on a variety of everyday objects, each of which represented different size, weight, and shape categories. The TriTrap gripper was able to securely hold most of the tested objects in place while they were lifted, rotated, and transported without further optimization. These results show that the insect tarsus selected approach is viable and warrants further development, particularly in the direction of interface optimization. As such, the main goal of the TriTrap gripper, which was to showcase the tarsal chain principle as a viable approach to gripping in general, was achieved.

Propolis, a naturally sticky substance used by bees to secure their hives and protect the colony from pathogens, presents a fascinating challenge. Despite its adhesive nature, honeybees adeptly handle propolis with their mandibles. Previous research has shown a combination of an anti-adhesive fluid layer and scale-like microstructures on the inner surface of bee mandibles. Our aim was to deepen our understanding of how surface energy and microstructure influence the reduction in adhesion for challenging substances like propolis. To achieve this, we devised surfaces inspired by the intricate microstructure of bee mandibles, employing diverse techniques including roughening steel surfaces, creating lacquer structures using Bénard cells, and moulding resin surfaces with hexagonal patterns. These approaches generated patterns that mimicked the bee mandible structure to varying degrees. Subsequently, we assessed the adhesion of propolis on these bioinspired structured substrates. Our findings revealed that on rough steel and resin surfaces structured with hexagonal dimples, propolis adhesion was significantly reduced by over 40% compared to unstructured control surfaces. However, in the case of the lacquer surface patterned with Bénard cells, we did not observe a significant reduction in adhesion.

Evolutionary and functional analyses alongside new fossil specimens expand our understanding of rolling-into-ball behaviour in pill scarab beetles.

Abstract Despite extensive research on the biomechanics of insect wings over the past years, direct mechanical measurements on sensitive wing specimens remain very challenging. This is especially true for examining delicate museum specimens. This has made the finite element method popular in studies of wing biomechanics. Considering the complexities of insect wings, developing a wing model is usually error-prone and time-consuming. Hence, numerical studies in this area have often accompanied oversimplified models. Here we address this challenge by developing a new tool for fast, precise modelling of insect wings. This application, called WingGram, uses computer vision to detect the boundaries of wings and wing cells from a 2D image. The app can be used to develop wing models that include complex venations, corrugations and camber. WingGram can extract geometric features of the wings, including dimensions of the wing domain and subdomains and the location of vein junctions. Allowing researchers to simply model wings with a variety of forms, shapes and sizes, our application can facilitate studies of insect wing morphology and biomechanics. Being an open-access resource, WingGram has a unique application to expand how scientists, educators, and industry professionals analyse insect wings and similar shell structures in other fields, such as aerospace.

Using cryo scanning electron microscopy, the surface micromorphology of vegetative (leaf blade and ligule) and generative (pedicel and outer glume) organs in Deschampsia antarctica, one of the only two flowering plants native to Antarctica, was examined. Whereas the pedicel and outer glume were wax-free, both leaf sides had a prominent epicuticular wax coverage consisting of two superimposed layers: polygonal rodlets formed by fused irregular platelets (the lower wax layer) and membraneous platelets (the upper wax layer). Although the adaxial (inner) and abaxial (outer) leaf surfaces showed a similar microstructure of the wax coverage, they differed in the thickness ratio between lower and upper wax layer. The ligule bore a very loose wax coverage composed of separate scale-like projections or clusters of them. We suppose that the two-layered wax densely covering both leaf surfaces might contribute to the plant adaptation to severe environmental conditions in Antarctica due to an increase of its resistance against cold temperatures, icing, harmful UV radiation, and dehydration. The presence of the epicuticular wax on the abaxial leaf side and the ligule as well as the hierarchical structure of the wax coverage on both leaf surfaces is described in D. antarctica for the first time.

Abstract Background The radula, a chitinous membrane with embedded teeth, is one important molluscan autapomorphy. In some taxa (Polyplacophora and Patellogastropoda) one tooth type (the dominant lateral tooth) was studied intensively in the last decades with regard to its mechanical properties, chemical and structural composition, and the relationship between these parameters. As the dominant lateral tooth is probably one of the best studied biological materials, it is surprising, that data on elements and mechanical properties of the other tooth types, present on a chiton radula, is lacking. Results We provide data on the elemental distribution and mechanical properties (hardness and elasticity, i.e. Young’s modulus) of all teeth from the Polyplacophora Lepidochitona cinerea (Linnaeus, 1767) [Chitonidae: Ischnochitonidae]. The ontogeny of elements, studied by energy-dispersive X-ray spectroscopy, and of the mechanical properties, determined by nanoindentation, was analysed in every individual tooth type. Additionally, we performed breaking stress experiments with teeth under dry and wet condition, highlighting the high influence of the water content on the mechanical behaviour of the radula. We thereby could determine the forces and stresses, teeth can resist, which were previously not studied in representatives of Polyplacophora. Overall, we were able to relate the mineral (iron, calcium) content with the mechanical parameters (hardness and Young’s modulus) and the breaking force and stress in every tooth type. This led to a better understanding of the relationship between structure, material, and function in radular teeth. Further, we aimed at determining the role of calcium for the mechanical behaviour of the teeth: we decalcified radulae by ethylene diamine tetra acetic acid and performed afterwards elemental analyses, breaking stress experiments, and nanoindentation. Among other things, we detected that wet and decalcified radular teeth could resist highest forces, since teeth have a higher range of bending motion leading to a higher capability of teeth to gain mechanical support from the adjacent tooth row. This indicates, that the tooth material is the result of a compromise between failure reduction and the ability to transfer forces onto the ingesta. Conclusion We present novel data on the elemental composition, mechanical properties, and the mechanical behaviour of chiton teeth, which allows conclusions about tooth function. We could also relate the parameters mentioned, which contributes to our understanding on the origins of mechanical property gradients and the processes reducing structural failure in radular teeth. Additionally, we add more evidence, that the elemental composition of radular is probably species-specific and could be used as taxonomic character.

Abstract The molluscan phylum is the second specious animal group with its taxa feeding on a variety of food sources. This is enabled by the radula, a chitinous membrane with embedded teeth, one important autapomorphy. Between species, radulae can vary in their morphology, mechanical, and chemical properties. With regard to chemical composition, some taxa (Polyplacophora and Patellogastropoda) were studied extensively in the past decades, due to their specificity to incorporate high proportions of iron, calcium, and silicon. There is, however, a huge lack of knowledge about radular composition in other taxa. The work presented aims at shedding light on the chemistry by performing energy-dispersive X-ray spectroscopy analyses on 24 molluscan species, thereof two Polyplacophora, two Cephalopoda, and 20 Gastropoda, which was never done before in such a comprehensiveness. The elements and their proportions were documented for 1448 individual, mature teeth and hypotheses about potential biomineralization types were proposed. The presented work additionally comprises a detailed record on past studies about the chemical composition of molluscan teeth, which is an important basis for further investigation of the radular chemistry. The found disparity in elements detected, in their distribution and proportions highlights the diversity of evolutionary solutions, as it depicts multiple biomineralization types present within Mollusca.

Cuticle is one of the most abundant, but least studied, biological composites. As a result, it has contributed very little to the field of biomimetics. An important step to overcome this problem is to study cuticle biomechanics by means of accurate mechanical measurements. However, due to many reasons, mechanical testing on fresh cuticle specimens is not always possible. Hence, researchers often use stored specimens to measure properties of arthropod cuticle. Our knowledge about the influence of different treatment methods on cuticle properties is currently very limited. In this study, we investigated the effect of freezing, desiccation, and rehydration on the elastic modulus of the hind tibial cuticle of locusts obtained by nanoindentation. We found that all the mentioned treatments significantly influence cuticle properties. This is in contrast to previous reports suggesting that freezing did not significantly influence the elastic modulus of native cuticle specimens tested in bending. In the light of our data, we suggest that changes of the elastic modulus of cuticle are not solely due to changes of the water content. Our results provide a platform for more accurate measurements of cuticle properties.

Abstract Background In most arthropods, adult females are larger than males, and male competition is a race to quickly locate and mate with scattered females (scramble competition polygyny). Variation in body size among males may confer advantages that depend on context. Smaller males may be favored due to more efficient locomotion leading to higher mobility during mate searching. Alternatively, larger males may benefit from increased speed and higher survivorship. While the relationship between male body size and mobility has been investigated in several systems, how different aspects of male body morphology specifically affect their locomotor performance in different contexts is often unclear. Results Using a combination of empirical measures of flight performance and modelling of body aerodynamics, we show that large body size impairs flight performance in male leaf insects (Phyllium philippinicum), a species where relatively small and skinny males fly through the canopy in search of large sedentary females. Smaller males were more agile in the air and ascended more rapidly during flight. Our models further predicted that variation in body shape would affect body lift and drag but suggested that flight costs may not explain the evolution of strong sexual dimorphism in body shape in this species. Finally, empirical measurements of substrate adhesion and subsequent modelling of landing impact forces suggested that smaller males had a lower risk of detaching from the substrates on which they walk and land. Conclusions By showing that male body size impairs their flight and substrate adhesion performance, we provide support to the hypothesis that smaller scrambling males benefit from an increased locomotor performance and shed light on the evolution of sexual dimorphism in scramble competition mating systems.

The stability of the body during locomotion is a fundamental requirement for walking animals. The mechanisms that coordinate leg movement patterns are even more complex at water–air interfaces. Water striders are agile creatures on the water surface, but they can be vulnerable to leg damage, which can impair their movement. One can assume the presence of certain compensatory biomechanical factors that are involved in the maintenance of postural balance lost after an amputation. Here, we studied changes in load distribution among the legs and assessed the effects of amputation on the locomotory behavior and postural defects that may increase the risk of locomotion failure. Apparently, amputees recover a stable posture by applying leg position modifications (e.g., widening the stance) and by load redistribution to the remaining legs. Water striders showed steering failure after amputation in all cases. Amputations affected locomotion by (1) altering motion features (e.g., shorter swing duration of midlegs), (2) functional constraints on legs, (3) shorter travelled distances, and (4) stronger deviations in the locomotion path. The legs functionally interact with each other, and removal of one leg has detrimental effects on the others. This research may assist the bioinspired design of aquatic robots.

In this paper, a new concept of extra-durable and sustainable wind turbine blades is presented. The two critical materials science challenges of the development of wind energy now are the necessity to prevent the degradation of wind turbine blades for several decades, and, on the other side, to provide a solution for the recyclability and sustainability of blades. In preliminary studies by DTU Wind, it was demonstrated that practically all typical wind turbine blade degradation mechanisms (e.g., coating detachment, buckling, spar cap/shell adhesive joint degradation, trailing edge failure, etc.) have their roots in interface degradation. The concept presented in this work includes the development of bio-inspired dual-mechanism-based interface adhesives (combining mechanical interlocking of fibers and chemical adhesion), which ensures, on the one side, extra-strong attachment during the operation time, and on the other side, possible adhesive joint separation for re-use of the blade parts. The general approach and physical mechanisms of adhesive strengthening and separation are described.

Attachment to the substrate is an important phenomenon that determines the survival of many organisms. Most insects utilize wet adhesion to support attachment, which is characterized by fluids that are secreted into the interface between the tarsus and the substrates. Previous research has investigated the composition and function of tarsal secretions of different insect groups, showing that the secretions are likely viscous emulsions that contribute to attachment by generating capillary and viscous adhesion, leveling surface roughness and providing self-cleaning of the adhesive systems. Details of the structural organization of these secretions are, however, largely unknown. Here, we analyzed footprints originating from the arolium and euplantulae of the stick insect Medauroidea extradentata using cryo-scanning electron microscopy (cryo-SEM) and white light interferometry (WLI). The secretion was investigated with cryo-SEM, revealing four morphologically distinguishable components. The 3D WLI measurements of the droplet shapes and volumes over time revealed distinctly different evaporation rates for different types of droplets. Our results indicate that the subfunctionalization of the tarsal secretion is facilitated by morphologically distinct components, which are likely a result of different proportions of components within the emulsion. Understanding these components and their functions may aid in gaining insights for developing adaptive and multifunctional biomimetic adhesive systems.

Abstract The radula, a chitinous membrane with embedded tooth rows, is the molluscan autapomorphy for feeding. The morphologies, arrangements and mechanical properties of teeth can vary between taxa, which is usually interpreted as adaptation to food. In previous studies, we proposed about trophic and other functional specialisations in taenioglossan radulae from species of African paludomid gastropods. These were based on the analysis of shape, material properties, force-resistance, and the mechanical behaviour of teeth, when interacting with an obstacle. The latter was previously simulated for one species (Spekia zonata) by the finite-element-analysis (FEA) and, for more species, observed in experiments. In the here presented work we test the previous hypotheses by applying the FEA on 3D modelled radulae, with incorporated material properties, from three additional paludomid species. These species forage either on algae attached to rocks (Lavigeria grandis), covering sand (Cleopatra johnstoni), or attached to plant surface and covering sand (Bridouxia grandidieriana). Since the analysed radulae vary greatly in their general size (e.g. width) and size of teeth between species, we additionally aimed at relating the simulated stress and strain distributions with the tooth sizes by altering the force/volume. For this purpose, we also included S. zonata again in the present study. Our FEA results show that smaller radulae are more affected by stress and strain than larger ones, when each tooth is loaded with the same force. However, the results are not fully in congruence with results from the previous breaking stress experiments, indicating that besides the parameter size, more mechanisms leading to reduced stress/strain must be present in radulae.

Abstract The water strider group demonstrates a very complex dynamics consisting of competition for the food items, territoriality and aggression to the conspecific individuals, escaping from the predators, etc. The situation is even more complex due to the presence of different instars, which in most water strider species live in the same habitat and occupy the same niche. The presented swarm model of water striders demonstrates the realistic population dynamics. For the swarm formation in the model, attraction and repulsion forces were used. Animal motion in the model takes into account inertia and kinetic energy dissipation effects. The model includes three different rates related to the growth of individuals: food appearance rate, food assimilation rate, and stored energy loss rate. The results of our modeling show that the size distribution of individuals seems to be an adequate measure for population status, and it has a characteristic shape for different model parameter combinations. Distribution of the distances between nearest neighbors is other important measure of the population density and its dynamics. Parameters of the model can be tuned in such a way, that the shape of both distributions in a steady phase coincides with that shape observed in a natural population, which helps to understand the factors leading to particular momentary distribution of both parameters (size and distance) in the population. From this point of view, the model can predict how both distributions can further develop from certain state depending on particular combination of factors.

We are pleased to announce that starting in July 2023, Biomimetics will publish 12 monthly online issues [...]

Biomimetics (bionics, bioinspired technology) refers to research on living systems and attempts to transfer their properties to engineering applications [...]

Adhesive pads are functional systems with specific micro- and nanostructures which evolved as a response to specific environmental conditions and therefore exhibit convergent traits. The functional constraints that shape systems for the attachment to a surface are general requirements. Different strategies to solve similar problems often follow similar physical principles, hence, the morphology of attachment devices is affected by physical constraints. This resulted in two main types of attachment devices in animals: hairy and smooth. They differ in morphology and ultrastructure but achieve mechanical adaptation to substrates with different roughness and maximise the actual contact area with them. Species-specific environmental surface conditions resulted in different solutions for the specific ecological surroundings of different animals. As the conditions are similar in discrete environments unrelated to the group of animals, the micro- and nanostructural adaptations of the attachment systems of different animal groups reveal similar mechanisms. Consequently, similar attachment organs evolved in a convergent manner and different attachment solutions can occur within closely related lineages. In this review, we present a summary of the literature on structural and functional principles of attachment pads with a special focus on insects, describe micro- and nanostructures, surface patterns, origin of different pads and their evolution, discuss the material properties (elasticity, viscoelasticity, adhesion, friction) and basic physical forces contributing to adhesion, show the influence of different factors, such as substrate roughness and pad stiffness, on contact forces, and review the chemical composition of pad fluids, which is an important component of an adhesive function. Attachment systems are omnipresent in animals. We show parallel evolution of attachment structures on micro- and nanoscales at different phylogenetic levels, focus on insects as the largest animal group on earth, and subsequently zoom into the attachment pads of the stick and leaf insects (Phasmatodea) to explore convergent evolution of attachment pads at even smaller scales. Since convergent events might be potentially interesting for engineers as a kind of optimal solution by nature, the biomimetic implications of the discussed results are briefly presented.

Lietz, Rajabi and colleagues investigate the physiological and structural components of damping in insect wings. Notably, flight muscles and hemolyph appear to have no direct involvement in wing damping, compared to the function of the wing hinge.

Traumatic insemination describes an unusual form of mating during which a male penetrates the body wall of its female partner to inject sperm. Females unable to prevent traumatic insemination have been predicted to develop either traits of tolerance or of resistance, both reducing the fitness costs associated with the male-inflicted injury. The evolution of tolerance traits has previously been suggested for the bed bug. Here we present data suggesting that tolerance traits also evolved in females of the twisted-wing parasite species Stylops ovinae and Xenos vesparum. Using micro-indentation experiments and confocal laser scanning microscopy, we found that females of both investigated species possess a uniform resilin-rich integument that is notably thicker at penetration sites than at control sites. As the thickened cuticle does not seem to hamper penetration by males, we hypothesise that thickening of the cuticle resulted in reduced penetration damage and loss of haemolymph and in improved wound sealing. To evaluate the evolutionary relevance of the Stylops-specific paragenital organ and penis shape variation in the context of inter- and intraspecific competition, we conducted attraction and interspecific mating experiments, as well as a geometric-morphometric analysis of S. ovinae and X. vesparum penises. We found that S. ovinae females indeed attract sympatrically distributed congeneric males. However, only conspecific males were able to mate. In contrast, we did not observe any heterospecific male attraction by Xenos females. We therefore hypothesise that the paragenital organ in the genus Stylops represents a prezygotic mating barrier that prevents heterospecific matings.

Egg parasitoids play an important role in biological control of pest species attacking and killing their hosts at an early stage of their development. During the antagonistic coevolution with their hosts, egg parasitoids have developed a great ability to locate their host using chemical cues. A considerable amount of literature is available on this topic, while nothing is known about a possible adaptation of egg parasitoids to topography and mechanical properties of egg surface features and its shape when attaching to the host egg for oviposition. In the present investigation, the attachment ability of adults of both sexes of the egg parasitoid Anastatus bifasciatus (Hymenoptera: Eupelmidae) to artificial (polishing paper, flat glass, glass beads as dummies of the host egg) and natural surfaces (eggs of Halyomorpha halys and Nezara viridula, both Heteroptera: Pentatomidae), with different roughness and wettability, was measured using centrifugal force tester and traction force experiments. The parasitoid attachment devices and the egg surfaces were examined under cryo scanning electron microscope, wettability and roughness of natural and artificial substrates were characterised. We detected differences in the attachment devices and attachment ability of the two sexes. The collected data revealed a special ability of the female to attach to the eggs of the host species, thus suggesting an adaptation of the A. bifasciatus female to the surface features of the eggs during oviposition.

Abstract The radula is the food gathering and processing structure and one important autapomorphy of the Mollusca. It is composed of a chitinous membrane with small, embedded teeth representing the interface between the organism and its ingesta. In the past, various approaches aimed at connecting the tooth morphologies, which can be highly distinct even within single radulae, to their functionality. However, conclusions from the literature were mainly drawn from analyzing mounted radulae, even though the configuration of the radula during foraging is not necessarily the same as in mounted specimens. Thus, the truly interacting radular parts and teeth, including 3D architecture of this complex structure during foraging were not previously determined. Here we present an experimental approach on individuals of Vittina turrita (Neritidae, Gastropoda), which were fed with algae paste attached to different sandpaper types. By comparing these radulae to radulae from control group, sandpaper-induced tooth wear patterns were identified and both area and volume loss could be quantified. In addition to the exact contact area of each tooth, conclusions about the 3D position of teeth and radular bending during feeding motion could be drawn. Furthermore, hypotheses about specific tooth functions could be put forward. These feeding experiments under controlled conditions were introduced for stylommatophoran gastropods with isodont radulae and are now applied to heterodont and complex radulae, which may provide a good basis for future studies on radula functional morphology.

Abstract The forces exerted by the animal’s food processing structures can be important parameters when studying trophic specializations to specific food spectra. Even though molluscs represent the second largest animal phylum, exhibiting an incredible biodiversity accompanied by the establishment of distinct ecological niches including the foraging on a variety of ingesta types, only few studies focused on the biomechanical performance of their feeding organs. To lay a keystone for future research in this direction, we investigated the in vivo forces exerted by the molluscan food gathering and processing structure, the radula, for five stylommatophoran species (Gastropoda). The chosen species and individuals have a similar radular morphology and motion, but as they represent different body mass classes, we were enabled to relate the forces to body mass. Radular forces were measured along two axes using force transducers which allowed us to correlate forces with the distinct phases of radular motion. A radular force quotient, AFQ = mean Absolute Force/bodymass0.67, of 4.3 could be determined which can be used further for the prediction of forces generated in Gastropoda. Additionally, some specimens were dissected and the radular musculature mass as well as the radular mass and dimensions were documented. Our results depict the positive correlation between body mass, radular musculature mass, and exerted force. Additionally, it was clearly observed that the radular motion phases, exerting the highest forces during feeding, changed with regard to the ingesta size: all smaller gastropods rather approached the food by a horizontal, sawing-like radular motion leading to the consumption of rather small food particles, whereas larger gastropods rather pulled the ingesta in vertical direction by radula and jaw resulting in the tearing of larger pieces.

Abstract We unveiled the penile penetration mechanics of two earwig species, Echinosoma horridum, whose intromittent organ, termed virga, is extraordinarily long, and E. denticulatum, whose virga is conversely short. We characterised configuration, geometry, material and bending stiffness for both virga and spermatheca. The short virga of E. denticulatum has a material gradient with the stiffer base, whereas the long virga of E. horridum and the spermathecae of both species are homogeneously sclerotised. The long virga of E. horridum has a lower bending stiffness than the spermatheca. The virga of E. denticulatum is overall less flexible than the spermatheca. We compared our results to a previous study on the penetration mechanics of elongated beetle genitalia. Based on the comparison, we hypothesised that the lower stiffness of the male intromittent organ comparing to the corresponding female structure is a universal prerequisite for the penetration mechanics of the elongated intromittent organ in insects.