Subdivision of the neotropical Prisopodinae Brunner von Wattenwyl, 1893 based on features of tarsal attachment pads (Insecta, Phasmatodea)

The euplantulae of species from all five genera of the Prisopodinae Brunner von Wattenwyl, 1893 were examined using scanning electron microscopy with the aim to reveal the significance of attachment pads regarding their phylogenetic relationships. The split into the conventional two sister groups is supported by the two-lobed structure of the euplantulae with a smooth surface in the Prisopodini and a nubby surface microstructure in the Paraprisopodini. The two lineages are well distinguishable by this feature, as well as by the shape of the euplantulae themselves. The functional importance of the attachment pad surface features is discussed

Adhesion Performance in the Eggs of the Philippine Leaf Insect Phyllium Philippinicum (Phasmatodea: Phylliidae)

Leaf insects (Phasmatodea: Phylliidae) exhibit perfect crypsis imitating leaves. Although the special appearance of the eggs of the species Phyllium philippinicum, which imitate plant seeds, has received attention in different taxonomic studies, the attachment capability of the eggs remains rather anecdotical. We herein elucidate the specialized attachment mechanism of the eggs of this species and provide the first experimental approach to systematically characterize the functional properties of their adhesion by using different microscopy techniques and attachment force measurements on substrates with differing degrees of roughness and surface chemistry, as well as repetitive attachment/detachment cycles while under the influence of water contact. We found that a combination of folded exochorionic structures (pinnae) and a film of adhesive secretion contribute to attachment, which both respond to water. Adhesion is initiated by the glue, which becomes fluid through hydration, enabling adaption to the surface profile. Hierarchically structured pinnae support the spreading of the glue and reinforcement of the film. This combination aids the egg's surface in adapting to the surface roughness, yet the attachment strength is additionally influenced by the egg's surface chemistry, favoring hydrophilic substrates. Repetitive detachment and water-mediated adhesion can optimize the location of the egg to ensure suitable environmental conditions for embryonic development. Furthermore, this repeatable and water-controlled adhesion mechanism can stimulate further research for biomimeticists, ecologists and conservationalists

Gripping performance in the stick insect Sungaya inexpectata in dependence on the pretarsal architecture

Insect attachment devices and capabilities have been subject to research efforts for decades, and even though during that time considerable progress has been made, numerous questions remain. Different types of attachment devices are known, alongside most of their working principles, however, some details have yet to be understood. For instance, it is not clear why insects for the most part developed pairs of claws, instead of either three or a single one. In this paper, we investigated the gripping forces generated by the stick insect Sungaya inexpectata, in dependence on the number of available claws. The gripping force experiments were carried out on multiple, standardized substrates of known roughness, and conducted in directions both perpendicular and parallel to the substrate. This was repeated two times: first with a single claw being amputated from each of the animals' legs, then with both claws removed, prior to the measurement. The adhesive pads (arolia) and frictional pads (euplantulae) remained intact. It was discovered that the removal of claws had a detractive effect on the gripping forces in both directions, and on all substrates. Notably, this also included the control of smooth surfaces on which the claws were unable to find any asperities to grip on. The results show that there is a direct connection between the adhesive performance of the distal adhesive pad (arolium) and the presence of intact claws. These observations show collective effects between different attachment devices that work in concert during locomotion, and grant insight into why most insects possess two claws

Multifunctional Adhesives on the Eggs of the Leaf Insect Phyllium philippinicum (Phasmatodea: Phylliidae): Solvent Influence and Biomimetic Implications

Leaf insects (Phylliidae) are well-camouflaged terrestrial herbivores. They imitate leaves of plants almost perfectly and even their eggs resemble seeds-visually and regarding to dispersal mechanisms. The eggs of the leaf insect Phyllium philippinicum utilize an adhesive system with a combination of glue, which can be reversibly activated through water contact and a water-responding framework of reinforcing fibers that facilitates their adjustment to substrate asperities and real contact area enhancement. So far, the chemical composition of this glue remains unknown. To evaluate functional aspects of the glue-solvent interaction, we tested the effects of a broad array of chemical solvents on the glue activation and measured corresponding adhesive forces. Based on these experiments, our results let us assume a proteinaceous nature of the glue with different functional chemical subunits, which enable bonding of the glue to both the surface of the egg and the unpredictable substrate. Some chemicals inhibited adhesion, but the deactivation was always reversible by water-contact and in some cases yielded even higher adhesive forces. The combination of glue and fibers also enables retaining the adhesive on the egg, even if detached from the egg's surface. The gained insights into this versatile bioadhesive system could hereafter inspire further biomimetic adhesives

The frequency of wing damage in a migrating butterfly

The ability to fly is crucial for migratory insects. Consequently, the accumulation of damage on the wings over time can affect survival, especially for species that travel long distances. We examined the frequency of irreversible wing damage in the migratory butterfly Vanessa cardui to explore the effect of wing structure on wing damage frequency, as well as the mechanisms that might mitigate wing damage. An exceptionally high migration rate driven by high precipitation levels in their larval habitats in the winter of 2018-2019 provided us with an excellent opportunity to collect data on the frequency of naturally occurring wing damage associated with long-distance flights. Digital images of 135 individuals of V. cardui were collected and analyzed in Germany. The results show that the hindwings experienced a greater frequency of damage than the forewings. Moreover, forewings experienced more severe damage on the lateral margin, whereas hindwings experienced more damage on the trailing margin. The frequency of wing margin damage was higher in the painted lady butterfly than in the migrating monarch butterfly and in the butterfly Pontia occidentalis following artificially induced wing collisions. The results of this study could be used in future comparative studies of patterns of wing damage in butterflies and other insects. Additional studies are needed to clarify whether the strategies for coping with wing damage differ between migratory and nonmigratory species

Louse flies holding on mammals' hair: Comparative functional morphology of specialized attachment devices of ectoparasites (Diptera: Hippoboscoidea)

Hippoboscidae and Nycteribiidae of the dipteran superfamily Hippoboscoidea are obligate ectoparasites, which feed on the blood of different mammals. Due to their limited flight capability, the attachment system on all tarsi is of great importance for a secure grasp onto their host and thus for their survival. In this study, the functional morphology of the attachment system of two hippoboscid species and two nycteribiid species was compared in their specificity to the host substrate. Based on data from scanning electron microscopy and confocal laser scanning microscopy, it was shown that the attachment systems of both Hippoboscidae and Nycteribiidae (Hippoboscoidea) differ greatly from that of other calyptrate flies and are uniform within the respective families. All studied species have an attachment system with two monodentate claws and two pulvilli. The claws and pulvilli of the Hippoboscidae are asymmetric, which is an adaptation to the fur of even-toed ungulates (Artiodactyla). The fur of these mammals possesses both, thinner woolen and thicker coat hair; thus, the asymmetry of the attachment system of the hippoboscid species enables a secure attachment to all surfaces of their hosts. The claws and pulvilli of the nyceribiid species do not show an asymmetry, since the fur of their bat (Chiroptera) hosts consists of hairs with the same thickness. The claws are important for the attachment to mammals' fur, because they enable a secure grip by mechanical interlocking of the hairs through the claws. Additionally, well-developed pulvilli are able to attach on thicker hairs of Artiodactyla or on smooth substrates such as the skin

The exceptional attachment ability of the ectoparasitic bee louse Braula coeca (Diptera, Braulidae) on the honeybee

Bee lice (Braulidae) are small parasitic flies, which are adapted to live on their bee host. As such, the wingless Braula coeca is a parasite of the common honey bee Apis mellifera and it is well adapted to attach to its hairy surface. The attachment system of B. coeca provides a secure grip on the fine setae of the bee. This is crucial for the parasite survival, as detachment from the host is fatal for the bee louse. The feet morphology of B. coeca is well adapted to the challenging bee surface, notably by strongly broadened claws, which are split into a high number of comb-like teeth, perfectly matching the diameter of the bee hairs. Based on microscopy observations, both the morphology and material composition of the tarsi of B. coeca are characterized in detail. Using high-speed video analysis, we combine the morphology data on the attachment system with a behavioural context. Furthermore, we directly measured the attachment forces generated by the bee lice in contact with the host. In particular, the claws are involved in attachment to the host, as the interstices between the teeth-like spines allow for the collection of several hairs and generate strong friction, when the hairs slip to the narrow gap between the spines. The overall morphology of the tarsus produces strong attachment, with average safety factors (force per body weight) around 1130, and stabilizes the tarsal chain with lateral stoppers against overflexion, but also allows for the fast detachment by the tarsal chain torsion.Deutsche Forschungsgemeinschaft; Human Frontier Science Program and the National Research Foundation.https://onlinelibrary.wiley.com/journal/13653032hj2021Zoology and Entomolog

Subdivision of the neotropical Prisopodinae Brunner von Wattenwyl, 1893 based on features of tarsal attachment pads (Insecta, Phasmatodea)

The euplantulae of species from all five genera of the Prisopodinae Brunner von Wattenwyl, 1893 were examined using scanning electron microscopy with the aim to reveal the significance of attachment pads regarding their phylogenetic relationships. The split into the conventional two sister groups is supported by the two-lobed structure of the euplantulae with a smooth surface in the Prisopodini and a nubby surface microstructure in the Paraprisopodini. The two lineages are well distinguishable by this feature, as well as by the shape of the euplantulae themselves. The functional importance of the attachment pad surface features is discussed

Subdivision of the neotropical Prisopodinae Brunner von Wattenwyl, 1893 based on features of tarsal attachment pads (Insecta, Phasmatodea)

The euplantulae of species from all five genera of the Prisopodinae Brunner von Wattenwyl, 1893 were examined using scanning electron microscopy with the aim to reveal the significance of attachment pads regarding their phylogenetic relationships. The split into the conventional two sister groups is supported by the two-lobed structure of the euplantulae with a smooth surface in the Prisopodini and a nubby surface microstructure in the Paraprisopodini. The two lineages are well distinguishable by this feature, as well as by the shape of the euplantulae themselves. The functional importance of the attachment pad surface features is discussed

Convergent Evolution of Adhesive Properties in Leaf Insect Eggs and Plant Seeds: Cross-Kingdom Bioinspiration

Plants and animals are often used as a source for inspiration in biomimetic engineering. However, stronger engagement of biologists is often required in the field of biomimetics. The actual strength of using biological systems as a source of inspiration for human problem solving does not lie in a perfect copy of a single system but in the extraction of core principles from similarly functioning systems that have convergently solved the same problem in their evolution. Adhesive systems are an example of such convergent traits that independently evolved in different organisms. We herein compare two analogous adhesive systems, one from plants seeds and one from insect eggs, to test their properties and functional principles for differences and similarities in order to evaluate the input that can be potentially used for biomimetics. Although strikingly similar, the eggs of the leaf insect Phyllium philippinicum and the seeds of the ivy gourd Coccinia grandis make use of different surface structures for the generation of adhesion. Both employ a water-soluble glue that is spread on the surface via reinforcing fibrous surface structures, but the morphology of these structures is different. In addition to microscopic analysis of the two adhesive systems, we mechanically measured the actual adhesion generated by both systems to quantitatively compare their functional differences on various standardized substrates. We found that seeds can generate much stronger adhesion in some cases but overall provided less reliable adherence in comparison to eggs. Furthermore, eggs performed better regarding repetitive attachment. The similarities of these systems, and their differences resulting from their different purposes and different structural/chemical features, can be informative for engineers working on technical adhesive systems