More than just slippery: the impact of biofilm on the attachment of non-sessile freshwater mayfly larvae.

While terrestrial insects can usually attach directly to a substrate, for aquatic insects the situation is more complicated owing to the presence of a biofilm on the primary substrates. This important fact has been neither the subject of investigation nor commonly taken into account in the interpretation of functional aspects of attachment in mobile freshwater animals. In this study, we investigate the impact of a biofilm on the attachment of living mayfly larvae. We performed in vivo attachment experiments in a flow channel using different substrates with defined surface roughness. Additionally, we measured friction forces directly generated by dissected tarsal claws on the same substrates. On substrates with smooth or slightly rough surfaces, which have little or no surface irregularities large enough for the claws to grasp, the presence of a biofilm significantly increases the friction force of claws. Consequently, larvae can endure higher flow velocities on these smooth substrates. The opposite effect takes place on rough substrates, where the friction force of claws decreases in the presence of a biofilm. Consequently, a biofilm is a critical ecological structure for these larvae, and other aquatic organisms, not only as a food source but also as a factor influencing attachment ability.