Nasal sound production in echolocating delphinids (Tursiops truncatus and Pseudorca crassidens) is dynamic, but unilateral: clicking on the right side and whistling on the left side.

Toothed whales produce sound in their nasal complex by pneumatic actuation of phonic lip pairs within the blowhole. It has been hypothesized that dual actuation of the phonic lip pairs can generate two pulses that merge to form a single echolocation click with a higher source level, broader bandwidth and larger potential for beam steering than if produced by a single pair of phonic lips. Here, we test that hypothesis by measuring the sound production of five echolocating delphinids using hydrophones around the animals and imbedded in on-animal suction cups. We show that the studied animals click with their right pair of phonic lips and whistle with their left pair. We demonstrate that, with just a single pair of phonic lips, they can change the click energy levels over five orders of magnitude, change the click centroid frequencies over more than two octaves, and modulate the sound radiation from the melon for beam steering. We conclude that all of the click dynamics ascribed to dual actuation of two phonic lip pairs can be achieved with actuation of just the right pair of phonic lips, and we propose that the large dynamic range of source outputs is achieved by highly controlled modulation of the pneumatic driving pressure, the tension of the phonic lip labia and the conformation of the fatty melon and associated air sacs.