Abstract
Males of Dytiscinae beetles use specialized adhesive setae to adhere to female elytra during underwater courtship. This coevolution of male setae and female elytra has attracted much attention since Darwin. However, there has been little examination of their biomechanical functioning despite increasing knowledge on biofibrillar adhesion. Here, we report and compare, for the first time, the mechanisms of underwater attachment using two hair types, the primitive spatula and derived 'passive' sucker, found in male diving beetles. Results from interspecific scaling of protarsal palettes and adhesion by single seta suggest better performance in the later-evolved circular (sucker) setae. Spatula setae with a modified shallow sucker and channels use the combined mechanisms of suction and viscous resistance for adhesion. Velocity-dependent adhesion provides sufficient control for resisting the female's erratic movements while also detaching easily through slow peeling. Direction-dependent shear resistance helps reorient setae surfaces into a preferred direction for effective adhesion. Seta deformation using different mechanisms for circular and spatula setae reduces the force that is transmitted to the contact interface. A softer spring in spatula setae explains their adhesion at lower preloads and assists in complete substrate contact. Attachment mechanisms revealed in adhesive setae with modified spatula and passive suckers provide insights for bioinspired designs of underwater attachment devices.