Complementary effect of attachment devices in stick insects (Phasmatodea).

Stick insects are well adapted in their locomotion to various surfaces and topographies of natural substrates. Single pad measurements characterised the pretarsal arolia of these insects as shear-sensitive adhesive pads and the tarsal euplantulae as load-sensitive friction pads. Different attachment microstructures on the euplantulae reveal an adaptation of smooth euplantulae to smooth surfaces and nubby eupantulae to a broader range of surface roughness. However, how different attachment pads and claws work in concert and how strong the contribution of different structures is to the overall attachment performance remains unclear. We therefore assessed combinatory effects in the attachment system of two stick insect species with different types of euplantular microstructures by analysing their usage in various posture situations and the performance on different levels of substrate roughness. For comparison, we provide attachment force data of the whole attachment system. The combination of claws, arolia and euplantulae provides mechanical interlocking on rough surfaces, adhesion and friction on smooth surfaces in different directions, and facilitates attachment on different inclines and on a broad range of surface roughness, with the least performance in the range 0.3-1.0 µm. On smooth surfaces, stick insects use arolia always, but employ euplantulae if the body weight can generate load on them (upright, wall). On structured surfaces, claws enable mechanical interlocking at roughnesses higher than 12 µm. On less-structured surfaces, the attachment strength depends on the use of pads and, corroborating earlier studies, favours smooth pads on smooth surfaces, but nubby euplantulae on micro-rough surfaces.