Functional morphology of burrowing in the mayflies Hexagenia limbata and Pentagenia vittigera

A video system, consisting of a high-resolution, infrared-sensitive, surveillance television camera, time-date generator, video recorder and high-resolution monitor, was developed for naturalistic observation of the subaquatic activities of larval mayflies. Exemplars were isolated from light, sound and mechanical disturbance, and their burrowing behaviour was recorded for subsequent slow-motion analysis. Larvae of silt-inhabiting Hexagenia limbata were observed to be ploughers that displaced and compacted soft substrate in the excavation of an open tunnel. The head and prothoracic legs had the principal roles in substrate displacement, while the mesothoracic and metathoracic legs drove the body forward. Five stereotyped burrowing cycles were distinguished, although one, termed the Alternate Undercut because of the motions of the prothoracic legs, predominated. Larvae of clay-inhabiting Pentagenia vittigera were observed to chisel and gouge out compacted substrates, also creating an open burrow. Their mandibular tusks were used to dislodge substrate, while the prothoracic legs displaced loosened substrate and widened the opening. The body was driven forward either by strong abdominal flexions or by activity of the mesothoracic and metathoracic legs. Four burrowing cycles were distinguished, two of which, the Tusk Gouge and the Chisel and Sweep, predominated. Differences in proportion and armature of the head and appendages correlated with the different substrate types excavated.