Animals with widely varying body weight, such as flies, spiders, and geckos, can adhere to and move along vertical walls and even ceilings. This ability is caused by very efficient attachment mechanisms in which patterned surface structures interact with the profile of the substrate. An extensive microscopic study has shown a strong inverse scaling effect in these attachment devices. Whereas m dimensions of the terminal elements of the setae are sufficient for flies and beetles, geckos must resort to sub-m devices to ensure adhesion. This general trend is quantitatively explained by applying the principles of contact mechanics, according to which splitting up the contact into finer subcontacts increases adhesion. This principle is widely spread in design of natural adhesive systems and may also be transferred into practical applications.walking ͉ adhesion ͉ locomotion ͉ legs ͉ insects A ttachment structures have independently developed several times in animal evolution (1, 2). Setose or hairy systems of various animal groups, such as insects, spiders, and lizards contain surfaces covered by fine patterns of protuberances of different origin. These highly specialized structures are not restricted to one particular area of the leg and may be located on different derivatives of the tarsus and pretarsus (3). Even among insects, the protuberances belong to different types: representatives of the Coleoptera and Dermaptera have setae with sockets providing additional mobility of setae, whereas representatives of Diptera have setae without sockets (acanthae). Setae range in their length from several millimeters to a few micrometers (4).Despite Ͼ300 years of studies on hairy attachment systems, there is still a debate concerning the attachment mechanism of animals walking on smooth walls or ceilings. Different hypotheses have been proposed to explain the mechanism of attachment: sticking fluid, microsuckers, and electrostatic forces (5). Based on experimental data, some of these theories have been rejected, and adhesion has been attributed to a combination of molecular interactions and capillary attractive forces mediated by secretions (6) or purely van der Waals interactions (7). Because some animals produce secretory fluids (insects) (8-10) in the contact area, whereas others do not (spiders, geckos) (11, 12), one can expect different basic physical forces contributing to the overall adhesion. Recently, strong evidence has been presented (13) that the adhesion of gecko setae is caused by van der Waals interaction, rejecting mechanisms relying on capillary adhesion. Elements of contact mechanics have also been applied to this problem (13,14); it was predicted that arrays with smaller setae endings should result in greater adhesive strength. In the present study, we combine an extensive microscopical study ¶ of biological surface devices with the theory of contact mechanics based on molecular adhesion. We will show that the scaling of the surface protuberances, for animals differing in weight by 6 orders of magnitude, ca...

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Attachment devices of representatives of most higher taxa of hexapods were examined. Short descriptions of tibial, tarsal and pretarsal adhesive structures for each order are presented. In their evolution, hexapods have developed two distinctly dierent mechanisms to attach themselves to a variety of substrates: hairy surfaces and smooth¯exible pads. The¯exible properties of pad material guarantees a maximal contact with surfaces, regardless of the microsculpture. These highly specialized structures are not restricted to one particular area of the leg. They may be located on dierent parts, such as claws, derivatives of the pretarsus, tarsal apex, tarsomeres, or tibia. The 10 characters of the two alternative designs of attachment devices ± smooth and hairy ± were coded and analysed together with a data matrix containing 105 additional morphological characters of dierent stages and body parts. The analysis demonstrates, that similar structures (arolium, euplantulae, hairy tarsomeres) have evolved independently in several lineages. Nevertheless, some of them support monophyletic groups (e.g. Embioptera + Dermaptera; Dictyoptera + Phasmatodea + Grylloblattodea + Orthoptera; Dictyoptera + Phasmatodea; Hymenoptera + Mecopterida; Neuropterida + Strepsiptera + Coleoptera). Other structures such as claw pads (Ephemeroptera), balloon-shaped eversible pads (Thysanoptera), or fossulae spongiosae (Reduviidae) are unique for larger or smaller monophyletic units. It is plausible to assume that the evolution of¯ight and the correlated necessity to cling to vegetation or other substrates was a major trigger for the evolution of adhesive structures. Groups with a potential to evolve a great variety of designs of adhesive pads are Hemiptera and Diptera. Even though characters of the adhesive pads are strongly subject to selection, they can provide phylogenetic information. The results of the cladistic analyses are largely congruent with current hypotheses of hexapod phylogeny. A sistergroup relationship between Diplura and Insecta and between Zygentoma (excl. Tricholepidion) and Pterygota is con®rmed. Plecoptera are probably the sistergroup of the remaining Neoptera. Dermaptera are the sistergroup of Embioptera and Dictyoptera the sistergroup of Phasmatodea. Paurometabola excl. Dermaptera + Embioptera are monophyletic. A sistergroup relationship between Zoraptera and a clade comprising Paraneoptera + Endopterygota is only supported by weak evidence. Coleoptera + Strepsiptera are the sistergroup of Neuropterida and Hymenoptera the sistergroup of Mecopterida.

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