Insects have developed specialized structures on their feet for adhering to surfaces, with stick and leaf insects or Phasmatodea exhibiting an unexpectedly high diversity of these structures. In Phasmatodea, attachment on different substrates is achieved by two types of pads on the legs: the euplantulae on the tarsomeres and the arolium on the pretarsus. The euplantulae are adhesive structures capable of adaptability to the substrate profile and generation of the required attachment strength. The diversity of euplantular microstructures of 56 species that represent all major lineages recognized within Phasmatodea and the whole biogeographical distribution of the group are examined using scanning electron microscopy (SEM). Nine different types of attachment structures can be distinguished whereby one, the nubby type, can be further divided into three different distinct types based on the specific ratio of each conical outgrowth. We mapped the morphological data from the SEM onto a phylogenetic tree we reconstructed based on molecular data. Previously, the evolution of different adhesive microstructures (AMs) on these pads has been suggested to reflect phylogenetic groups. However, different types of AMs are found within monophyletic groups, and our ancestral character state reconstruction suggests smooth euplantulae in the ground pattern of Euphasmatodea and multiple independent origins of other forms. The type of AM appears to be strongly associated with ecomorphs, e.g., smooth euplantular surfaces are more frequently found in tree-dwellers than in ground-dwellers, whilst the attachment pads of ground-dwelling species primarily bear conical cuticular outgrowths (nubby euplantulae).

Stick and leaf insects (Phasmatodea) are large terrestrial herbivorous arthropods known for masquerading as plant parts such as bark, twigs and leaves. Their evolutionary history is largely shaped by convergent evolution associated with adaptive radiations on geographically isolated landmasses that have repeatedly generated ground‐dwelling ecomorphs. The members of one lineage, however, the Oriental Heteropterygidae, are morphologically rather uniform, and have a predominantly ground‐dwelling lifestyle. The phylogeny of Heteropterygidae that comprises approximately 130 described species is controversial and remains uncertain. In particular, the systematic position of the giant Jungle Nymph Heteropteryx dilatata, whose males are capable of flight and exhibit the most plesiomorphic wing morphology among extant phasmatodeans, is of major interest to the scientific community. Here, we analysed a set of seven nuclear and mitochondrial genes to infer the phylogeny of Heteropterygidae covering the group's overall diversity. The divergence time estimation and reconstruction of the historical biogeography resulted in an ancestral distribution across Sundaland with long distance dispersal events to Wallacea, the Philippines and the South Pacific. We were able to resolve the relationships among the three principal subgroups of Heteropterygidae and revealed the Dataminae, which contain entirely wingless small forms, as the sister group of Heteropteryginae + Obriminae. Within Heteropteryginae, Haaniella is recovered as paraphyletic in regard to Heteropteryx . Consequently, Heteropteryx must be considered a subordinate taxon deeply embedded within a flightless clade of stick insects. Within Obriminae, the Bornean Hoploclonia is strongly supported as the earliest diverging lineage. Based on this finding, we recognize only two tribes of equal rank among Obriminae, the Hoplocloniini trib. nov. and Obrimini sensu nov . Within the latter, we demonstrate that previous tribal assignments do not reflect phylogenetic relationships and that a basal splitting event occurred between the wing‐bearing clade Miroceramia + Pterobrimus and the remaining wingless Obrimini. The Philippine genus Tisamenus is paraphyletic with regard to Ilocano hebardi , thus, we transfer the latter species to Tisamenus as Tisamenus hebardi comb. nov. and synonymize Ilocano with Tisamenus . We discuss character transformations in the light of the new phylogenetic results and conclude that the current taxonomic diversity appears to be mainly driven by allopatry and not to be the result of niche differentiation. This radiation is thus best described as a nonadaptive radiation.

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 roughnesses. How different attachment pads and claws work in concert, and how strong the contribution of different structures to the overall attachment performance is, however, 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 roughnesses, with the least performance in a range of 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.

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.

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.

In its evolution, the diverse group of stick and leaf insects (Phasmatodea) has undergone a rapid radiation. These insects evolved specialized structures to adhere to different surfaces typical for their specific ecological environments. The cuticle of their tarsal attachment pads (euplantulae) is known to possess a high diversity of attachment microstructures (AMS) which are suggested to reflect ecological specializations of different groups within phasmids. However, the origin of these microstructures and their developmental background remain largely unknown. Here, based on the detailed scanning electron microscopy study of pad surfaces, we present a theoretical approach to mathematically model an outstanding diversity of phasmid AMS using the reaction–diffusion model by Alan Turing. In general, this model explains pattern formation in nature. For the first time, we were able to identify eight principal patterns and simulate the transitions among these. In addition, intermediate transitional patterns were predicted by the model. The ease of transformation suggests a high adaptability of the microstructures that might explain the rapid evolution of pad characters. We additionally discuss the functional morphology of the different microstructures and their assumed advantages in the context of the ecological background of species.