Echolocation in Blainville's beaked whales (Mesoplodon densirostris).

Here we use sound and movement recording tags to study how deep-diving Blainville's beaked whales (Mesoplodon densirostris) use echolocation to forage in their natural mesopelagic habitat. These whales ensonify thousands of organisms per dive but select only about 25 prey for capture. They negotiate their cluttered environment by radiating sound in a narrow 20° field of view which they sample with 1.5-3 clicks per metre travelled requiring only some 60 clicks to locate, select and approach each prey. Sampling rates do not appear to be defined by the range to individual targets, but rather by the movement of the predator. Whales sample faster when they encounter patches of prey allowing them to search new water volumes while turning rapidly to stay within a patch. This implies that the Griffin search-approach-capture model of biosonar foraging must be expanded to account for sampling behaviours adapted to the overall prey distribution. Beaked whales can classify prey at more than 15 m range adopting stereotyped motor patterns when approaching some prey. This long detection range relative to swimming speed facilitates a deliberate mode of sensory-motor operation in which prey and capture tactics can be selected to optimize energy returns during long breath-hold dives.

Deadly diving? Physiological and behavioural management of decompression stress in diving mammals.

Decompression sickness (DCS; 'the bends') is a disease associated with gas uptake at pressure. The basic pathology and cause are relatively well known to human divers. Breath-hold diving marine mammals were thought to be relatively immune to DCS owing to multiple anatomical, physiological and behavioural adaptations that reduce nitrogen gas (N(2)) loading during dives. However, recent observations have shown that gas bubbles may form and tissue injury may occur in marine mammals under certain circumstances. Gas kinetic models based on measured time-depth profiles further suggest the potential occurrence of high blood and tissue N(2) tensions. We review evidence for gas-bubble incidence in marine mammal tissues and discuss the theory behind gas loading and bubble formation. We suggest that diving mammals vary their physiological responses according to multiple stressors, and that the perspective on marine mammal diving physiology should change from simply minimizing N(2) loading to management of the N(2) load. This suggests several avenues for further study, ranging from the effects of gas bubbles at molecular, cellular and organ function levels, to comparative studies relating the presence/absence of gas bubbles to diving behaviour. Technological advances in imaging and remote instrumentation are likely to advance this field in coming years.

Foraging Blainville's beaked whales (Mesoplodon densirostris) produce distinct click types matched to different phases of echolocation.

Blainville's beaked whales (Mesoplodon densirostris Blainville) echolocate for prey during deep foraging dives. Here we use acoustic tags to demonstrate that these whales, in contrast to other toothed whales studied, produce two distinct types of click sounds during different phases in biosonar-based foraging. Search clicks are emitted during foraging dives with inter-click intervals typically between 0.2 and 0.4 s. They have the distinctive form of an FM upsweep (modulation rate of about 110 kHz ms(-1)) with a -10 dB bandwidth from 26 to 51 kHz and a pulse length of 270 micros, somewhat similar to chirp signals in bats and Cuvier's beaked whales (Ziphius cavirostris Cuvier), but quite different from clicks of other toothed whales studied. In comparison, the buzz clicks, produced in short bursts during the final stage of prey capture, are short (105 micros) transients with no FM structure and a -10 dB bandwidth from 25 to 80 kHz or higher. Buzz clicks have properties similar to clicks reported from large delphinids and hold the potential for higher temporal resolution than the FM clicks. It is suggested that the two click types are adapted to the separate problems of target detection and classification versus capture of low target strength prey in a cluttered acoustic environment.

Biosonar performance of foraging beaked whales (Mesoplodon densirostris).

Toothed whales (Cetacea, odontoceti) emit sound pulses to probe their surroundings by active echolocation. Non-invasive, acoustic Dtags were placed on deep-diving Blainville's beaked whales (Mesoplodon densirostris) to record their ultrasonic clicks and the returning echoes from prey items, providing a unique view on how a whale operates its biosonar during foraging in the wild. The process of echolocation during prey capture in this species can be divided into search, approach and terminal phases, as in echolocating bats. The approach phase, defined by the onset of detectable echoes recorded on the tag for click sequences terminated by a buzz, has interclick intervals (ICI) of 300-400 ms. These ICIs are more than a magnitude longer than the decreasing two-way travel time to the targets, showing that ICIs are not given by the two-way-travel times plus a fixed, short lag time. During the approach phase, the received echo energy increases by 10.4(+/-2) dB when the target range is halved, demonstrating that the whales do not employ range-compensating gain control of the transmitter, as has been implicated for some bats and dolphins. The terminal/buzz phase with ICIs of around 10 ms is initiated when one or more targets are within approximately a body length of the whale (2-5 m), so that strong echo returns in the approach phase are traded for rapid updates in the terminal phase. It is suggested that stable ICIs in the search and approach phases facilitate auditory scene analysis in a complex multi-target environment, and that a concomitant low click rate allows the whales to maintain high sound pressure outputs for prey detection and discrimination with a pneumatically driven, bi-modal sound generator.