Sound production in red-bellied piranhas (Pygocentrus nattereri, Kner): an acoustical, behavioural and morphofunctional study.

Piranhas are known to be sound-producing animals. Nevertheless, the biological significance of piranha calls remains unclear because sounds have been recorded only when specimens were held by hand or trapped in a gill net. These sounds are generated by rapid contractions of sonic muscles that insert on a broad tendon surrounding ventrally the cranial sac of the swimbladder. The piranha swimbladder is thought to play an important role in sound production as an impedance-matching device and as a resonator. However, the vibratory capacities of the cranial and caudal sacs and the exact role of both sacs in sound production remain poorly understood. In this study, three sounds were each associated to a specific behaviour. The first sound (type 1) was produced during frontal display; it had numerous pulses and lasted 140!±17 ms, with a fundamental frequency of 120±4 Hz. It corresponded to the sound made by hand-held fishes. The second sound (type 2) was produced during circling and fighting behaviour; it was a single pulse lasting 36±8 ms, with a fundamental frequency of 43±10 Hz. The third sound (type 3) corresponded to chasing behaviour and comprised three to four pulses, each lasting 3±1 ms, with a fundamental frequency of 1739±18 Hz. Using a laser vibrometer to study the swimbladder displacement when stimulated at different frequencies, it was demonstrated that the first two sounds corresponded to the swimbladder mechanism. By contrast, the third sound was associated with the jaw mechanism. The vibrometer indicated that the swimbladder is a highly damping structure, simply copying the sonic muscle contraction rate. This study provides two interesting insights. First, it shows the relationships between three kinds of piranha sound and three specific behaviours. Second, using muscle stimulation at different rates, it shows which simultaneous conditions are required for production of sound in this species. Swimbladder calls were produced by a muscle contraction rate of approximately 100 Hz because this periodicity allowed the swimbladder to vibrate. At this frequency range, the contraction-relaxation cycles of the swimbladder muscles engendered wall displacements that had short amplitudes and with only a small variability between them.