The monopulsed nature of sperm whale clicks

Traditionally, sperm whale clicks have been described as multipulsed, long duration, nondirectional signals of moderate intensity and with a spectrum peaking below 10 kHz. Such properties are counterindicative of a sonar function, and quite different from the properties of dolphin sonar clicks. Here, data are presented suggesting that the traditional view of sperm whale clicks is incomplete and derived from off-axis recordings of a highly directional source. A limited number of assumed on-axis clicks were recorded and found to be essentially monopulsed clicks, with durations of 100 micros, with a composite directionality index of 27 dB, with source levels up to 236 dB re: 1 microPa (rms), and with centroid frequencies of 15 kHz. Such clicks meet the requirements for long-range biosonar purposes. Data were obtained with a large-aperture, GPS-synchronized array in July 2000 in the Bleik Canyon off Vester?len, Norway (69 degrees 28' N, 15 degrees 40' E). A total of 14 h of sound recordings was collected from five to ten independent, simultaneously operating recording units. The sound levels measured make sperm whale clicks by far the loudest of sounds recorded from any biological source. On-axis click properties support previous work proposing the nose of sperm whales to operate as a generator of sound.     

Sperm whales (Physeter catodon L. 1758) do not react to sounds from detonators

A number of observations show that sperm whales (Physeter catodon L. 1758) react to various man-made pulses with moderate source levels. The behavioral responses are described to vary from silence to fear. Click rates of five submerged male sperm whales were measured during the discharge of eight detonators off Andenes, northern Norway. In addition, the behavioral response of a surfaced specimen was observed. Click rates of the submerged whales and the behavior of the surfaced specimen did not change during the discharges with received sound levels of some 180 dB re 1 microPa peRMS. The apparent lack of response to the discharges could be due to similarity between sperm whale clicks and detonations. Accordingly, it can be speculated that the discharges may have been perceived as isolated clicks from conspecifics.     

Source levels of clicks from free-ranging white-beaked dolphins (Lagenorhynchus albirostris Gray 1846) recorded in Icelandic waters

This study reports the source levels of clicks recorded from free-ranging white-beaked dolphins (Lagenorhynchus albirostris Gray 1846). A four-hydrophone array was used to obtain sound recordings. The hydrophone signals were digitized on-line and stored in a portable computer. An underwater video camera was used to visualize dolphins to help identify on-axis recordings. The range to a dolphin was calculated from differences in arrival times of clicks at the four hydrophones, allowing for calculations of source levels. Source levels in a single click train varied from 194 to 211 dB peak-to-peak (p-p) re: 1 microPa. The source levels varied linearly with the log of range. The maximum source levels recorded were 219 dB (p-p) re: 1 microPa.     

Echolocation clicks from killer whales (Orcinus orca) feeding on herring (Clupea harengus)

Echolocation clicks from Norwegian killer whales feeding on herring schools were recorded using a four-hydrophone array. The clicks had broadband bimodal frequency spectra with low and high frequency peaks at 24 and 108 kHz, respectively. The -10 dB bandwidth was 35 kHz. The average source level varied from 173 to 202 dB re 1 microPa (peak-to-peak) at 1 m. This is considerably lower than source levels described for Canadian killer whales foraging on salmon. It is suggested that biosonar clicks of Norwegian killer whales are adapted for localization of prey with high target strength and acute hearing abilities.     

Comparison of echolocation clicks from geographically sympatric killer whales and long-finned pilot whales (L)

The source characteristics of biosonar signals from sympatric killer whales and long-finned pilot whales in a Norwegian fjord were compared. A total of 137 pilot whale and more than 2000 killer whale echolocation clicks were recorded using a linear four-hydrophone array. Of these, 20 pilot whale clicks and 28 killer whale clicks were categorized as being recorded on-axis. The clicks of pilot whales had a mean apparent source level of 196 dB re 1 μPa pp and those of killer whales 203 dB re 1 μPa pp. The duration of pilot whale clicks was significantly shorter (23 μs, S.E.=1.3) and the centroid frequency significantly higher (55 kHz, S.E.=2.1) than killer whale clicks (duration: 41 μs, S.E.=2.6; centroid frequency: 32 kHz, S.E.=1.5). The rate of increase in the accumulated energy as a function of time also differed between clicks from the two species. The differences in duration, frequency, and energy distribution may have a potential to allow for the distinction between pilot and killer whale clicks when using automated detection routines for acoustic monitoring.     

Behavioral avoidance threshold level of a harbor porpoise (Phocoena phocoena) for a continuous 50 kHz pure tone

The use of ultrasonic sounds in alarms for gillnets may be advantageous, but the deterring effects of ultrasound on porpoises are not well understood. Therefore a harbor porpoise in a large floating pen was subjected to a continuous 50 kHz pure tone with a source level of 122+/-3 dB (re 1 microPa, rms). When the test signal was switched on during test periods, the animal moved away from the sound source. Its respiration rate was similar to that during baseline periods, when the sound was switched off. The behavior of the porpoise was related to the sound pressure level distribution in the pen. The sound level at the animal's average swimming location during the test periods was approximately 107+/-3 dB (re 1 microPa, rms). The avoidance threshold sound pressure level for a continuous 50 kHz pure tone for this porpoise, in the context of this study, is estimated to be 108+/-3 dB (re 1 microPa, rms). This study demonstrates that porpoises may be deterred from an area by high frequency sounds that are not typically audible to fish and pinnipeds and would be less likely masked by ambient noise.     

Characteristics of biosonar signals from the northern bottlenose whale, Hyperoodon ampullatus

The biosonar pulses from free-ranging northern bottlenose whales (Hyperoodon ampullatus) were recorded with a linear hydrophone array. Signals fulfilling criteria for being recorded close to the acoustic axis of the animal (a total of 10 clicks) had a frequency upsweep from 20 to 55 kHz and durations of 207 to 377 μs (measured as the time interval containing 95% of the signal energy). The source level of these signals, denoted pulses, was 175-202 dB re 1 μPa rms at 1 m. The pulses had a directionality index of at least 18 dB. Interpulse intervals ranged from 73 to 949 ms (N?=?856). Signals of higher repetition rates had interclick intervals of 5.8-13.1 ms (two sequences, made up of 59 and 410 clicks, respectively). These signals, denoted clicks, had a shorter duration (43-200 μs) and did not have the frequency upsweep characterizing the pulses of low repetition rates. The data show that the northern bottlenose whale emits signals similar to three other species of beaked whale. These signals are distinct from the three other types of biosonar signals of toothed whales. It remains unclear why the signals show this grouping, and what consequences it has on echolocation performance.     

Source parameters of echolocation clicks from wild bottlenose dolphins (Tursiops aduncus and Tursiops truncatus)

The Indian Ocean and Atlantic bottlenose dolphins (Tursiops aduncus and Tursiops truncatus) are among the best studied echolocating toothed whales. However, almost all echolocation studies on bottlenose dolphins have been made with captive animals, and the echolocation signals of free-ranging animals have not been quantified. Here, biosonar source parameters from wild T. aduncus and T. truncatus were measured with linear three- and four-hydrophone arrays in four geographic locations. The two species had similar source parameters, with source levels of 177-228 dB re 1 μPa peak to peak, click durations of 8-72 μs, centroid frequencies of 33-109 kHz and rms bandwidths between 23 and 54 kHz. T. aduncus clicks had a higher frequency emphasis than T. truncatus. The transmission directionality index was up to 3 dB higher for T. aduncus (29 dB) as compared to T. truncatus (26 dB). The high directionality of T. aduncus does not appear to be only a physical consequence of a higher frequency emphasis in clicks, but may also be caused by differences in the internal properties of the sound production system.     

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.     

Temporary shift in masked hearing thresholds in odontocetes after exposure to single underwater impulses from a seismic watergun

A behavioral response paradigm was used to measure masked underwater hearing thresholds in a bottlenose dolphin (Tursiops truncatus) and a white whale (Delphinapterus leucas) before and after exposure to single underwater impulsive sounds produced from a seismic watergun. Pre- and postexposure thresholds were compared to determine if a temporary shift in masked hearing thresholds (MTTS), defined as a 6-dB or larger increase in postexposure thresholds, occurred. Hearing thresholds were measured at 0.4, 4, and 30 kHz. MTTSs of 7 and 6 dB were observed in the white whale at 0.4 and 30 kHz, respectively, approximately 2 min following exposure to single impulses with peak pressures of 160 kPa, peak-to-peak pressures of 226 dB re 1 microPa, and total energy fluxes of 186 dB re 1 microPa2 x s. Thresholds returned to within 2 dB of the preexposure value approximately 4 min after exposure. No MTTS was observed in the dolphin at the highest exposure conditions: 207 kPa peak pressure, 228 dB re 1 microPa peak-to-peak pressure, and 188 dB re 1 microPa2 x s total energy flux.     

Radiated noise characteristics of a modern cargo ship

Extensive measurements were made of the radiated noise of M/V OVERSEAS HARRIETTE, a bulk cargo ship (length 173 m, displacement 25 515 tons) powered by a direct-drive low-speed diesel engine-a design representative of many modern merchant ships. The radiated noise data show high-level tonal frequencies from the ship's service diesel generator, main engine firing rate, and blade rate harmonics due to propeller cavitation. Radiated noise directionality measurements indicate that the radiation is generally dipole in form at lower frequencies, as expected. There are some departures from this pattern that may indicate hull interactions. Blade rate source level (174 dB re 1 microPa/m at 9 Hz, 16 knots) agrees reasonably well with a model of fundamental blade rate radiation previously reported by Gray and Greeley, but agreement for blade rate harmonics is not as good. Noise from merchant ships elevates the natural ambient by 20-30 dB in many areas; the effects of this noise on the biological environment have not been widely investigated.     

Measuring the off-axis angle and the rotational movements of phonating sperm whales using a single hydrophone

The common use of the bent-horn model of the sperm whale sound generator describes sperm whale clicks as the pulse series {p0, p1, p2, p3,...}. Clicks, however, deviate from this standard when recorded using off-axis hydrophones. The existence of additional pulses within the {p0, p1, p2, p3, ...} series can be explained still using the bent-horn model. Multiple reflections on the whale's frontal and distal sacs of the p0 pulse lead to additional sets of pulses detectable using a farfield, off-axis hydrophone. The travel times of some of these additional pulses depend on the whale's orientation. The authors propose a method to estimate the off-axis angle of sperm whale clicks. They also propose a method to determine the nature of the movement (if it is pitch, yaw, or roll) of phonating sperm whales. The application of both methods requires the measurement of the travel time differences between pulses composing a sperm whale click. They lead, using a simple apparatus consisting of a single hydrophone at an unknown depth, to new measurements of the underwater movements of sperm whales. Using these methods shows that sperm whales would methodically scan seawater while searching for prey, by making periodic pitch and yaw movements in sync with their acoustic activity.     

Temporary threshold shift in bottlenose dolphins (Tursiops truncatus) exposed to mid-frequency tones

A behavioral response paradigm was used to measure hearing thresholds in bottlenose dolphins before and after exposure to 3 kHz tones with sound exposure levels (SELs) from 100 to 203 dB re 1 microPa2 s. Experiments were conducted in a relatively quiet pool with ambient noise levels below 55 dB re 1 microPa2/Hz at frequencies above 1 kHz. Experiments 1 and 2 featured 1-s exposures with hearing tested at 4.5 and 3 kHz, respectively. Experiment 3 featured 2-, 4-, and 8-s exposures with hearing tested at 4.5 kHz. For experiment 2, there were no significant differences between control and exposure sessions. For experiments 1 and 3, exposures with SEL=197 dB re 1 microPa2 s and SEL > or = 195 dB re 1 microPa2 s, respectively, resulted in significantly higher TTS4 than control sessions. For experiment 3 at SEL= 195 dB re 1 microPa2 s, the mean TTS4 was 2.8 dB. These data are consistent with prior studies of TTS in dolphins exposed to pure tones and octave band noise and suggest that a SEL of 195 dB re 1 microPa2 s is a reasonable threshold for the onset of TTS in dolphins and white whales exposed to midfrequency tones.     

Quantitative measures of air-gun pulses recorded on sperm whales (Physeter macrocephalus) using acoustic tags during controlled exposure experiments

The widespread use of powerful, low-frequency air-gun pulses for seismic seabed exploration has raised concern about their potential negative effects on marine wildlife. Here, we quantify the sound exposure levels recorded on acoustic tags attached to eight sperm whales at ranges between 1.4 and 12.6 km from controlled air-gun array sources operated in the Gulf of Mexico. Due to multipath propagation, the animals were exposed to multiple sound pulses during each firing of the array with received levels of analyzed pulses falling between 131-167 dB re. 1 microPa (pp) [111-147 dB re. 1 microPa (rms) and 100-135 dB re. 1 microPa2 s] after compensation for hearing sensitivity using the M-weighting. Received levels varied widely with range and depth of the exposed animal precluding reliable estimation of exposure zones based on simple geometric spreading laws. When whales were close to the surface, the first arrivals of air-gun pulses contained most energy between 0.3 and 3 kHz, a frequency range well beyond the normal frequencies of interest in seismic exploration. Therefore air-gun arrays can generate significant sound energy at frequencies many octaves higher than the frequencies of interest for seismic exploration, which increases concern of the potential impact on odontocetes with poor low frequency hearing.     

Click sounds produced by cod (Gadus morhua)

Conspicuous sonic click sounds were recorded in the presence of cod (Gadus morhua), together with either harp seals (Pagophilus groenlandicus), hooded seals (Cystophora cristata) or a human diver in a pool. Similar sounds were never recorded in the presence of salmon (Salmo salar) together with either seal species, or from either seal or fish species when kept separately in the pool. It is concluded that cod was the source of these sounds and that the clicks were produced only when cod were approached by a swimming predatorlike body. The analyzed click sounds (n = 377) had the following characteristics (overall averages +/- S.D.): peak frequency = 5.95 +/- 2.22 kHz; peak-to-peak duration = 0.70 +/- 0.45 ms; sound pressure level (received level) = 153.2 +/- 7.0 dB re 1 microPa at 1 m. At present the mechanism and purpose of these clicks is not known. However, the circumstances under which they were recorded and some observations on the behavior of the seals both suggest that the clicks could have a predator startling function.     

Sounds and source levels from bowhead whales off Pt. Barrow, Alaska

Sounds were recorded from bowhead whales migrating past Pt. Barrow, AK, to the Canadian Beaufort Sea. They mainly consisted of various low-frequency (25- to 900-Hz) moans and well-defined sound sequences organized into "song" (20-5000 Hz) recorded with our 2.46-km hydrophone array suspended from the ice. Songs were composed of up to 20 repeated phrases (mean, 10) which lasted up to 146 s (mean, 66.3). Several bowhead whales often were within acoustic range of the array at once, but usually only one sang at a time. Vocalizations exhibited diurnal peaks of occurrence (0600-0800, 1600-1800 h). Sounds which were located in the horizontal plane had peak source spectrum levels as follows--44 moans: 129-178 dB re: 1 microPa, 1 m (median, 159); 3 garglelike utterances: 152, 155, and 169 dB; 33 songs: 158-189 dB (median, 177), all presumably from different whales. Based on ambient noise levels, measured total propagation loss, and whale sound source levels, our detection of whale sounds was theoretically noise-limited beyond 2.5 km (moans) and beyond 10.7 km (songs), a model supported by actual localizations. This study showed that over much of the shallow Arctic and sub-Arctic waters, underwater communications of the bowhead whale would be limited to much shorter ranges than for other large whales in lower latitude, deep-water regions.     

Localization and visual verification of a complex minke whale vocalization.

A recently described population of minke whales (Balaenoptera acutorostrata) offered a unique opportunity to study its acoustic behavior. The often-inquisitive dwarf minke whale is seen on the Great Barrier Reef nearly coincident with its suspected calving and breeding seasons. During drifting encounters with whales, a towed hydrophone array was used to record sounds for subsequent localization of sound sources. Shipboard and in-water observers linked these sounds to the closely circling minke whale. A complex and stereotyped sound sequence, the "star-wars" (SW) vocalization, was recorded during a series of visual and acoustic observations. The SW vocalization spanned a wide frequency range (50 Hz-9.4 kHz) and was composed of distinct and stereotypically repeated units with both amplitude and frequency-modulated components. Broadband source levels between 150 and 165 dB re 1 microPa at 1 m were calculated. Passive acoustic studies can utilize this distinct vocalization to help determine the behavior, distribution, and movements of this animal. While the SW vocalization's function remains unknown, the regularly repeated and complex sound sequence was common in low latitude, winter month aggregations of minke whales. At this early stage, the SW vocalization appears similar to the songs of other whale species and has characteristics consistent with those of reproductive advertisement displays.     

Directionality of sperm whale sonar clicks and its relation to piston radiation theory

This paper investigates the applicability to sperm whales of the theory of sound radiating from a piston. The theory is applied to a physical model and to a series of sperm whale clicks. Results show that wave forms of off-axis signals can be reproduced by convolving an on-axis signal with the spatial impulse response of a piston. The angle of a recorded click can be estimated as the angle producing the spatial impulse response that gives the best match with the observation when convolved with the on-axis wave form. It is concluded that piston theory applies to sperm whale sonar click emission.     

The effect of a low-frequency sound source (acoustic thermometry of the ocean climate) on the diving behavior of juvenile northern elephant seals,Mirounga angustirostris

Changes in the diving behavior of individual free-ranging juvenile northern elephant seals, Mirounga angustirostris, exposed to the acoustic thermometry of the ocean climate (ATOC) sound source were examined using data loggers. Data loggers were attached to the animals and measured swim speed, maximum depth of dive, dive duration, surface interval, descent and ascent rate, and descent and ascent angle along with sound pressure level (SPL). The ATOC sound source was at a depth of 939 m and transmitted at 195 dB re: 1 microPa at 1 m centered at 75 Hz with a 37.5-Hz bandwidth. Sound pressure levels (SPL) measured at the seal during transmissions averaged 128 dB and ranged from 118 to 137 dB re: 1 microPa for the 60-90 Hz band, in comparison to ambient levels of 87-107 dB within this band. In no case did an animal end its dive or show any other obvious change in behavior upon exposure to the ATOC sound. Subtle changes in diving behavior were detected, however. During exposure, deviations in descent rate were greater than 1 s.d. of the control mean in 9 of 14 seals. Dive depth increased and descent velocity increased in three animals, ascent velocity decreased in two animals, ascent rate increased in one animal and decreased in another, and dive duration decreased in only one animal. There was a highly significant positive correlation between SPL and descent rate. The biological significance of these subtle changes is likely to be minimal. This is the first study to quantify behavioral responses of an animal underwater with simultaneous measurements of SPL of anthropogenic sounds recorded at the animal.