Functional anatomy of the pretarsus in whip spiders (Arachnida, Amblypygi)

Whip spiders (Amblypygi) are a small, cryptic order of arachnids mainly distributed in the tropics. Some basal lineages (families Charinidae and Charontidae) have adhesive pads on the tips of their six walking legs. The present study describes the macro- and ultrastructure of these pads and investigates their contact mechanics and adhesive strength on smooth and rough substrates. Furthermore, the structure of the pretarsus and its kinematics are compared in Charon cf. grayi (with an adhesive pad) and Phrynus longipes (without an adhesive pad). The adhesive pads exhibit an elaborate structure with a unique combination of structural features of smooth and hairy foot pads including a long transversal contact zone performing lateral detachment, a thick internally-branched cuticle with longitudinal ribs and hexagonal surface microstructures with spatulate keels. The contact area of the pads on smooth glass is discontinuous due to the spatulate microstructures with a discontinuous detachment, which could be observed in vivo by high speed videography at a rate of up to 10,000 fps. Adhesive strength was measured with vertical whole animal pull-off tests, obtaining mean values between 55 and 200 kPa. The occurrence of viscous lipid secretions between microstructures was occasionally observed, which, however, seems not to be a necessity for good foothold. The results are discussed in relation to the whip spider's ecology and evolution. Structure-function relationships of the adhesive pads are compared to those of insects and vertebrates.