Coupling wings with movable hooks - resilin in the wing-interlocking structures of honeybees.

Hymenoptera are characterised by the presence of one forewing pair and one hindwing pair. The two wings of each body side are coupled to each other during flight making the morphologically four-winged insects functionally two-winged. This coupling is formed by a row of hook-like structures, called hamuli, that are located at the leading edge of the hindwing and interlock with a thickened and recurved margin present at the trailing edge of the forewing. In this study, autofluorescence analyses performed with confocal laser scanning microscopy revealed differences in the exoskeleton material composition of the interlocking structures. While the wing veins and the recurved margin are strongly sclerotised and chitinous, the wing membranes mainly contain the elastomeric protein resilin. The hamuli are composed of sclerotised chitinous material, and each hamulus base is surrounded by and embedded in material that features large proportions of resilin and is located in strongly sclerotised socket-like wing vein structures. This exoskeleton organisation likely allows movements of the hamuli and, in combination with the exoskeleton material gradients visualized in the other interlocking structures, is assumed to guarantee an effective wing coupling and to simultaneously decrease the risk of wear and damage under mechanical loads occurring in flight, coupling and decoupling situations.