First direct measurements of behavioural responses by Cuvier's beaked whales to mid-frequency active sonar.

Most marine mammal- strandings coincident with naval sonar exercises have involved Cuvier's beaked whales (Ziphius cavirostris). We recorded animal movement and acoustic data on two tagged Ziphius and obtained the first direct measurements of behavioural responses of this species to mid-frequency active (MFA) sonar signals. Each recording included a 30-min playback (one 1.6-s simulated MFA sonar signal repeated every 25 s); one whale was also incidentally exposed to MFA sonar from distant naval exercises. Whales responded strongly to playbacks at low received levels (RLs; 89-127 dB re 1 µPa): after ceasing normal fluking and echolocation, they swam rapidly, silently away, extending both dive duration and subsequent non-foraging interval. Distant sonar exercises (78-106 dB re 1 µPa) did not elicit such responses, suggesting that context may moderate reactions. The observed responses to playback occurred at RLs well below current regulatory thresholds; equivalent responses to operational sonars could elevate stranding risk and reduce foraging efficiency.

Range estimation of cetaceans with compact volumetric arrays.

Passive acoustic monitoring is the method of choice to detect whales and dolphins that are acoustically active and to monitor their underwater behavior. The NATO Science and Technology Organization Centre for Maritime Research and Experimentation has recently implemented a compact passive acoustic monitor (CPAM), consisting of three arrays of two hydrophones each that are combined in a fixed three-dimensional arrangement and that may be towed at depths of more than 100 m. With its volumetric configuration, the CPAM is capable of estimating the three-dimensional direction vector of arriving sounds and under certain conditions on relative geometry between the whale and hydrophone array, the CPAM may also estimate the range to echolocating animals. Basic ranging methods assume constant sound speed and apply straightforward geometry to obtain depth and distance to the sound source. Alternatively, ray-tracing based methods may be employed to integrate the information provided by real sound speed profiles. Both ranging methods combine measurements of sound arrival angles and surface reflection delays and are easily implemented in real-time applications, whereby one could promote the ray-tracing approach as the preferred method because it may integrate real sound speed profiles.

Passive acoustic detection of deep-diving beaked whales.

Beaked whales can remain submerged for an hour or more and are difficult to sight when they come to the surface to breathe. Passive acoustic detection (PAD) not only complements traditional visual-based methods for detecting these species but also can be more effective because beaked whales produce clicks regularly to echolocate on prey during deep foraging dives. The effectiveness of PAD for beaked whales depends not only on the acoustic behavior and output of the animals but also on environmental conditions and the quality of the passive sonar implemented. A primary constraint on the range at which beaked whale clicks can be detected involves their high frequencies, which attenuate rapidly, resulting in limited ranges of detection, especially in adverse environmental conditions. Given current knowledge of source parameters and in good conditions, for example, with a wind speed of 2 ms, a receiver close to the surface should be able to detect acoustically Cuvier's beaked whales with a high probability at distances up to 0.7 km, provided the listening duration exceeds the deep dive interval, about 2.5 h on average. Detection ranges beyond 4 km are unlikely and would require low ambient noise or special sound propagation conditions.

Beaked whales echolocate on prey.

Beaked whales (Cetacea: Ziphiidea) of the genera Ziphius and Mesoplodon are so difficult to study that they are mostly known from strandings. How these elusive toothed whales use and react to sound is of concern because they mass strand during naval sonar exercises. A new non-invasive acoustic ording tag was attached to four beaked whales(two Mesoplodon densirostris and two Ziphius cavirostris) and recorded high-frequency clicks during deep dives. The tagged whales only clicked at depths below 200 m, down to a maximum depth of 1267 m. Both species produced a large number of short, directional, ultrasonic clicks with significant energy below 20 kHz. The tags recorded echoes from prey items; to our knowledge, a first for any animal echolocating in the wild. As far as we are aware, these echoes provide the first direct evidence on how free-ranging toothed whales use echolocation in foraging. The strength of these echoes suggests that the source level of Mesoplodon clicks is in the range of 200-220 dB re 1 microPa at 1 m. This paper presents conclusive data on the normal vocalizations of these beaked whale species, which may enable acoustic monitoring to mitigate exposure to sounds intense enough to harm them.

Beaked whales respond to simulated and actual navy sonar.

Beaked whales have mass stranded during some naval sonar exercises, but the cause is unknown. They are difficult to sight but can reliably be detected by listening for echolocation clicks produced during deep foraging dives. Listening for these clicks, we documented Blainville's beaked whales, Mesoplodon densirostris, in a naval underwater range where sonars are in regular use near Andros Island, Bahamas. An array of bottom-mounted hydrophones can detect beaked whales when they click anywhere within the range. We used two complementary methods to investigate behavioral responses of beaked whales to sonar: an opportunistic approach that monitored whale responses to multi-day naval exercises involving tactical mid-frequency sonars, and an experimental approach using playbacks of simulated sonar and control sounds to whales tagged with a device that records sound, movement, and orientation. Here we show that in both exposure conditions beaked whales stopped echolocating during deep foraging dives and moved away. During actual sonar exercises, beaked whales were primarily detected near the periphery of the range, on average 16 km away from the sonar transmissions. Once the exercise stopped, beaked whales gradually filled in the center of the range over 2-3 days. A satellite tagged whale moved outside the range during an exercise, returning over 2-3 days post-exercise. The experimental approach used tags to measure acoustic exposure and behavioral reactions of beaked whales to one controlled exposure each of simulated military sonar, killer whale calls, and band-limited noise. The beaked whales reacted to these three sound playbacks at sound pressure levels below 142 dB re 1 µPa by stopping echolocation followed by unusually long and slow ascents from their foraging dives. The combined results indicate similar disruption of foraging behavior and avoidance by beaked whales in the two different contexts, at exposures well below those used by regulators to define disturbance.

Off-axis effects on the multipulse structure of sperm whale usual clicks with implications for sound production.

Sperm whales (Physeter macrocephalus) produce multipulsed clicks with their hypertrophied nasal complex. The currently accepted view of the sound generation process is based on the click structure measured directly in front of, or behind, the whale where regular interpulse intervals (IPIs) are found between successive pulses in the click. Most sperm whales, however, are recorded with the whale in an unknown orientation with respect to the hydrophone where the multipulse structure and the IPI do not conform to a regular pulse pattern. By combining far-field recordings of usual clicks with acoustic and orientation information measured by a tag on the clicking whale, we analyzed clicks from known aspects to the whale. We show that a geometric model based on the bent horn theory for sound production can explain the varying off-axis multipulse structure. Some of the sound energy that is reflected off the frontal sac radiates directly into the water creating an intermediate pulse p1/2 seen in off-axis recordings. The powerful p1 sonar pulse exits the front of the junk as predicted by the bent-horn model, showing that the junk of the sperm whale nasal complex is both anatomically and functionally homologous to the melon of smaller toothed whales.

Three-dimensional beam pattern of regular sperm whale clicks confirms bent-horn hypothesis.

The three-dimensional beam pattern of a sperm whale (Physeter macrocephalus) tagged in the Ligurian Sea was derived using data on regular clicks from the tag and from hydrophones towed behind a ship circling the tagged whale. The tag defined the orientation of the whale, while sightings and beamformer data were used to locate the whale with respect to the ship. The existence of a narrow, forward-directed P1 beam with source levels exceeding 210 dBpeak re: 1 microPa at 1 m is confirmed. A modeled forward-beam pattern, that matches clicks >20 degrees off-axis, predicts a directivity index of 26.7 dB and source levels of up to 229 dBpeak re: 1 microPa at 1 m. A broader backward-directed beam is produced by the P0 pulse with source levels near 200 dBpeak re: 1 microPa at 1 m and a directivity index of 7.4 dB. A low-frequency component with source levels near 190 dBpeak re: 1 microPa at 1 m is generated at the onset of the P0 pulse by air resonance. The results support the bent-horn model of sound production in sperm whales. While the sperm whale nose appears primarily adapted to produce an intense forward-directed sonar signal, less-directional click components convey information to conspecifics, and give rise to echoes from the seafloor and the surface, which may be useful for orientation during dives.

Echolocation clicks of free-ranging Cuvier's beaked whales (Ziphius cavirostris).

Strandings of beaked whales of the genera Ziphius and Mesoplodon have been reported to occur in conjunction with naval sonar use. Detection of the sounds from these elusive whales could reduce the risk of exposure, but descriptions of their vocalizations are at best incomplete. This paper reports quantitative characteristics of clicks from deep-diving Cuvier's beaked whales (Ziphius cavirostris) using a unique data set. Two whales in the Ligurian Sea were simultaneously tagged with sound and orientation recording tags, and the dive tracks were reconstructed allowing for derivation of the range and relative aspect between the clicking whales. At depth, the whales produced trains of regular echolocation clicks with mean interclick intervals of 0.43 s (+/- 0.09) and 0.40 s (+/- 0.07). The clicks are frequency modulated pulses with durations of approximately 200 micros and center frequencies around 42 kHz, -10 dB bandwidths of 22 kHz, and Q(3 dB) of 4. The sound beam is narrow with an estimated directionality index of more than 25 dB, source levels up to 214 dB(pp) re: 1 microPa at 1 m, and energy flux density of 164 dB re: 1 microPa2 s. As the spectral and temporal properties are different from those of nonziphiid odontocetes the potential for passive detection is enhanced.