Dolphin echolocation behaviour during active long-range target approaches.

Echolocating toothed whales generally adjust click intensity and rate according to target range to ensure that echoes from targets of interest arrive before a subsequent click is produced, presumably facilitating range estimation from the delay between clicks and returning echoes. However, this click-echo-click paradigm for the dolphin biosonar is mostly based on experiments with stationary animals echolocating fixed targets at ranges below ∼120 m. Therefore, we trained two bottlenose dolphins instrumented with a sound recording tag to approach a target from ranges up to 400 m and either touch the target (subject TRO) or detect a target orientation change (subject SAY). We show that free-swimming dolphins dynamically increase interclick interval (ICI) out to target ranges of ∼100 m. TRO consistently kept ICIs above the two-way travel time (TWTT) for target ranges shorter than ∼100 m, whereas SAY switched between clicking at ICIs above and below the TWTT for target ranges down to ∼25 m. Source levels changed on average by 17log10(target range), but with considerable variation for individual slopes (4.1 standard deviations for by-trial random effects), demonstrating that dolphins do not adopt a fixed automatic gain control matched to target range. At target ranges exceeding ∼100 m, both dolphins frequently switched to click packet production in which interpacket intervals exceeded the TWTT, but ICIs were shorter than the TWTT. We conclude that the click-echo-click paradigm is not a fixed echolocation strategy in dolphins, and we demonstrate the first use of click packets for free-swimming dolphins when solving an echolocation task.

Amazon river dolphins (Inia geoffrensis) modify biosonar output level and directivity during prey interception in the wild.

Toothed whales have evolved to live in extremely different habitats and yet they all rely strongly on echolocation for finding and catching prey. Such biosonar-based foraging involves distinct phases of searching for, approaching and capturing prey, where echolocating animals gradually adjust sonar output to actively shape the flow of sensory information. Measuring those outputs in absolute levels requires hydrophone arrays centred on the biosonar beam axis, but this has never been done for wild toothed whales approaching and capturing prey. Rather, field studies make the assumption that toothed whales will adjust their biosonar in the same manner to arrays as they will when approaching prey. To test this assumption, we recorded wild botos (Inia geoffrensis) as they approached and captured dead fish tethered to a hydrophone in front of a star-shaped seven-hydrophone array. We demonstrate that botos gradually decrease interclick intervals and output levels during prey approaches, using stronger adjustment magnitudes than predicted from previous boto array data. Prey interceptions are characterised by high click rates, but although botos buzz during prey capture, they do so at lower click rates than marine toothed whales, resulting in a much more gradual transition from approach phase to buzzing. We also demonstrate for the first time that wild toothed whales broaden biosonar beamwidth when closing in on prey, as is also seen in captive toothed whales and bats, thus resulting in a larger ensonified volume around the prey, probably aiding prey tracking by decreasing the risk of prey evading ensonification.

Amazon river dolphins (Inia geoffrensis) use a high-frequency short-range biosonar.

Toothed whales produce echolocation clicks with source parameters related to body size; however, it may be equally important to consider the influence of habitat, as suggested by studies on echolocating bats. A few toothed whale species have fully adapted to river systems, where sonar operation is likely to result in higher clutter and reverberation levels than those experienced by most toothed whales at sea because of the shallow water and dense vegetation. To test the hypothesis that habitat shapes the evolution of toothed whale biosonar parameters by promoting simpler auditory scenes to interpret in acoustically complex habitats, echolocation clicks of wild Amazon river dolphins were recorded using a vertical seven-hydrophone array. We identified 404 on-axis biosonar clicks having a mean SLpp of 190.3 ± 6.1 dB re. 1 µPa, mean SLEFD of 132.1 ± 6.0 dB re. 1 µPa(2)s, mean Fc of 101.2 ± 10.5 kHz, mean BWRMS of 29.3 ± 4.3 kHz and mean ICI of 35.1 ± 17.9 ms. Piston fit modelling resulted in an estimated half-power beamwidth of 10.2 deg (95% CI: 9.6-10.5 deg) and directivity index of 25.2 dB (95% CI: 24.9-25.7 dB). These results support the hypothesis that river-dwelling toothed whales operate their biosonars at lower amplitude and higher sampling rates than similar-sized marine species without sacrificing high directivity, in order to provide high update rates in acoustically complex habitats and simplify auditory scenes through reduced clutter and reverberation levels. We conclude that habitat, along with body size, is an important evolutionary driver of source parameters in toothed whale biosonars.