Burrowing and subsurface locomotion in anguilliform fish: behavioral specializations and mechanical constraints.

Fish swimming is probably one of the most studied and best understood locomotor behaviors in vertebrates. However, many fish also actively exploit sediments. Because of their elongate body shape, anguilliform fishes are not only efficient swimmers but also very maneuverable. Consequently, many species live in complexly structured environments near the bottom and many are known to burrow into the sediment. To better understand burrowing and subsurface locomotion in anguilliform fish we provide descriptive kinematic data on subsurface locomotion in a burrowing eel (Pisodonophis boro) using videofluoroscopy. We also measured the maximal forces that can be exerted by this species during head-first and tail-first burrowing, and explored the implications of head-first burrowing on mechanical stress distribution in the skull. Our data show that P. boro uses lateral undulation to penetrate and move in sandy sediments under water. The kinematics of subsurface locomotion are different from those observed during swimming and are characterized by a very high slip factor. These observations differ considerably from recently published data in terrestrial sand-swimming lizards, and suggest that the sediment behaves like a solid rather than a frictional fluid. Finally, our finite element models show that the cranial shape and structure in the head-first burrowing P. boro is mechanically more suited for head-first burrowing than that of an obligate tail-first burrowing species, Heteroconger hassi.