The Sonar Model for Humpback Whale Song Revised.

Why do humpback whales sing? This paper considers the hypothesis that humpback whales may use song for long range sonar. Given the vocal and social behavior of humpback whales, in several cases it is not apparent how they monitor the movements of distant whales or prey concentrations. Unless distant animals produce sounds, humpback whales are unlikely to be aware of their presence or actions. Some field observations are strongly suggestive of the use of song as sonar. Humpback whales sometimes stop singing and then rapidly approach distant whales in cases where sound production by those whales is not apparent, and singers sometimes alternately sing and swim while attempting to intercept another whale that is swimming evasively. In the evolutionary development of modern cetaceans, perceptual mechanisms have shifted from reliance on visual scanning to the active generation and monitoring of echoes. It is hypothesized that as the size and distance of relevant events increased, humpback whales developed adaptive specializations for long-distance echolocation. Differences between use of songs by humpback whales and use of sonar by other echolocating species are discussed, as are similarities between bat echolocation and singing by humpback whales. Singing humpback whales are known to emit sounds intense enough to generate echoes at long ranges, and to flexibly control the timing and qualities of produced sounds. The major problem for the hypothesis is the lack of recordings of echoes from other whales arriving at singers immediately before they initiate actions related to those whales. An earlier model of echoic processing by singing humpback whales is here revised to incorporate recent discoveries. According to the revised model, both direct echoes from targets and modulations in song-generated reverberation can provide singers with information that can help them make decisions about future actions related to mating, traveling, and foraging. The model identifies acoustic and structural features produced by singing humpback whales that may facilitate a singer's ability to interpret changes in echoic scenes and suggests that interactive signal coordination by singing whales may help them to avoid mutual interference. Specific, testable predictions of the model are presented.