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.     

Killer whale (Orcinus orca) hearing: auditory brainstem response and behavioral audiograms.

Killer whale (Orcinus orca) audiograms were measured using behavioral responses and auditory evoked potentials (AEPs) from two trained adult females. The mean auditory brainstem response (ABR) audiogram to tones between 1 and 100 kHz was 12 dB (re 1 mu Pa) less sensitive than behavioral audiograms from the same individuals (+/- 8 dB). The ABR and behavioral audiogram curves had shapes that were generally consistent and had the best threshold agreement (5 dB) in the most sensitive range 18-42 kHz, and the least (22 dB) at higher frequencies 60-100 kHz. The most sensitive frequency in the mean Orcinus audiogram was 20 kHz (36 dB), a frequency lower than many other odontocetes, but one that matches peak spectral energy reported for wild killer whale echolocation clicks. A previously reported audiogram of a male Orcinus had greatest sensitivity in this range (15 kHz, approximately 35 dB). Both whales reliably responded to 100-kHz tones (95 dB), and one whale to a 120-kHz tone, a variation from an earlier reported high-frequency limit of 32 kHz for a male Orcinus. Despite smaller amplitude ABRs than smaller delphinids, the results demonstrated that ABR audiometry can provide a useful suprathreshold estimate of hearing range in toothed whales.     

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.     

Echolocation signals of free-ranging killer whales (Orcinus orca) and modeling of foraging for chinook salmon (Oncorhynchus tshawytscha)

Fish-eating "resident"-type killer whales (Orcinus orca) that frequent the coastal waters off northeastern Vancouver Island, Canada have a strong preference for chinook salmon (Oncorhynchus tshawytscha). The whales in this region often forage along steep cliffs that extend into the water, echolocating their prey. Echolocation signals of resident killer whales were measured with a four-hydrophone symmetrical star array and the signals were simultaneously digitized at a sample rate of 500 kHz using a lunch-box PC. A portable VCR recorded the images from an underwater camera located adjacent to the array center. Only signals emanating from close to the beam axis (1185 total) were chosen for a detailed analysis. Killer whales project very broadband echolocation signals (Q equal 0.9 to 1.4) that tend to have bimodal frequency structure. Ninety-seven percent of the signals had center frequencies between 45 and 80 kHz with bandwidths between 35 and 50 kHz. The peak-to-peak source level of the echolocation signals decreased as a function of the one-way transmission loss to the array. Source levels varied between 195 and 224 dB re: 1 microPa. Using a model of target strength for chinook salmon, the echo levels from the echolocation signals are estimated for different horizontal ranges between a whale and a salmon. At a horizontal range of 100 m, the echo level should exceed an Orcinus hearing threshold at 50 kHz by over 29 dB and should be greater than sea state 4 noise by at least 9 dB. In moderately heavy rain conditions, the detection range will be reduced substantially and the echo level at a horizontal range of 40 m would be close to the level of the rain 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.     

Asymmetry and dynamics of a narrow sonar beam in an echolocating harbor porpoise

A key component in the operation of a biosonar system is the radiation of sound energy from the sound producing head structures of toothed whales and microbats. The current view involves a fixed transmission aperture by which the beam width can only change via changes in the frequency of radiated clicks. To test that for a porpoise, echolocation clicks were recorded with high angular resolution using a 16 hydrophone array. The beam is narrower than previously reported (DI = 24 dB) and slightly dorso-ventrally compressed (horizontal -3 dB beam width: 13°, vertical -3 dB beam width: 11°). The narrow beam indicates that all smaller toothed whales investigated so far have surprisingly similar beam widths across taxa and habitats. Obtaining high directionality may thus be at least in part an evolutionary factor that led to high centroid frequencies in a group of smaller toothed whales emitting narrow band high frequency clicks. Despite the production of stereotyped narrow band high frequency clicks, changes in the directionality by a few degrees were observed, showing that porpoises can obtain changes in sound radiation.     

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.     

High-frequency modulated signals of killer whales (Orcinus orca) in the North Pacific

Killer whales in the North Pacific, similar to Atlantic populations, produce high-frequency modulated signals, based on acoustic recordings from ship-based hydrophone arrays and autonomous recorders at multiple locations. The median peak frequency of these signals ranged from 19.6-36.1 kHz and median duration ranged from 50-163 ms. Source levels were 185-193 dB peak-to-peak re: 1 μPa at 1 m. These uniform, repetitive, down-swept signals are similar to bat echolocation signals and possibly could have echolocation functionality. A large geographic range of occurrence suggests that different killer whale ecotypes may utilize these signals.     

Depth-dependent acoustic features of diving sperm whales (Physeter macrocephalus) in the Gulf of Mexico

Three-dimensional dive trajectories of three sperm whales in the Gulf of Mexico have been obtained by measuring the relative arrival times and bearings of the animals' acoustic multipath reflections, using two elements of a towed hydrophone array deployed at an unknown depth and orientation. Within the first 6-12 min of the start of a dive, the intervals between successive "clicks" of all three whales corresponded closely with the two-way travel time of an acoustic pulse traveling vertically between the animals' position and the ocean bottom. The click spectra contained multiple peaks, including a faint band of energy originally centered near 10 kHz. As the animals descended over 500 m in depth, the center frequency of this band shifted to nearly 15 kHz, but subsequently remained near this value during the rest of the dive. This frequency shift is consistent with that expected from energy scattering from an ensemble of incompressible small-scale air-filled resonators, with diameters on the order of 4 mm. One possible candidate for such an ensemble is proposed to reside in the collapsed frontal sac of the animal. A comparison of the received levels for the bottom and direct multipath arrivals indicates that the whales' acoustic directivity must range between 10-30 dB in the 5-20-kHz region.     

Discriminating features of echolocation clicks of melon-headed whales (Peponocephala electra), bottlenose dolphins (Tursiops truncatus), and Gray's spinner dolphins (Stenella longirostris longirostris)

Spectral parameters were used to discriminate between echolocation clicks produced by three dolphin species at Palmyra Atoll: melon-headed whales (Peponocephala electra), bottlenose dolphins (Tursiops truncatus) and Gray's spinner dolphins (Stenella longirostris longirostris). Single species acoustic behavior during daytime observations was recorded with a towed hydrophone array sampling at 192 and 480 kHz. Additionally, an autonomous, bottom moored High-frequency Acoustic Recording Package (HARP) collected acoustic data with a sampling rate of 200 kHz. Melon-headed whale echolocation clicks had the lowest peak and center frequencies, spinner dolphins had the highest frequencies and bottlenose dolphins were nested in between these two species. Frequency differences were significant. Temporal parameters were not well suited for classification. Feature differences were enhanced by reducing variability within a set of single clicks by calculating mean spectra for groups of clicks. Median peak frequencies of averaged clicks (group size 50) of melon-headed whales ranged between 24.4 and 29.7 kHz, of bottlenose dolphins between 26.7 and 36.7 kHz, and of spinner dolphins between 33.8 and 36.0 kHz. Discriminant function analysis showed the ability to correctly discriminate between 93% of melon-headed whales, 75% of spinner dolphins and 54% of bottlenose dolphins.     

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.     

Characteristics of whistles from the acoustic repertoire of resident killer whales (Orcinus orca) off Vancouver Island, British Columbia

The acoustic repertoire of killer whales (Orcinus orca) consists of pulsed calls and tonal sounds, called whistles. Although previous studies gave information on whistle parameters, no study has presented a detailed quantitative characterization of whistles from wild killer whales. Thus an interpretation of possible functions of whistles in killer whale underwater communication has been impossible so far. In this study acoustic parameters of whistles from groups of individually known killer whales were measured. Observations in the field indicate that whistles are close-range signals. The majority of whistles (90%) were tones with several harmonics with the main energy concentrated in the fundamental. The remainder were tones with enhanced second or higher harmonics and tones without harmonics. Whistles had an average bandwidth of 4.5 kHz, an average dominant frequency of 8.3 kHz, and an average duration of 1.8 s. The number of frequency modulations per whistle ranged between 0 and 71. The study indicates that whistles in wild killer whales serve a different function than whistles of other delphinids. Their structure makes whistles of killer whales suitable to function as close-range motivational sounds.     

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.     

Effects of noise levels and call types on the source levels of killer whale calls

Accurate parameter estimates relevant to the vocal behavior of marine mammals are needed to assess potential effects of anthropogenic sound exposure including how masking noise reduces the active space of sounds used for communication. Information about how these animals modify their vocal behavior in response to noise exposure is also needed for such assessment. Prior studies have reported variations in the source levels of killer whale sounds, and a more recent study reported that killer whales compensate for vessel masking noise by increasing their call amplitude. The objectives of the current study were to investigate the source levels of a variety of call types in southern resident killer whales while also considering background noise level as a likely factor related to call source level variability. The source levels of 763 discrete calls along with corresponding background noise were measured over three summer field seasons in the waters surrounding the San Juan Islands, WA. Both noise level and call type were significant factors on call source levels (1-40 kHz band, range of 135.0-175.7 dB(rms) re 1 [micro sign]Pa at 1 m). These factors should be considered in models that predict how anthropogenic masking noise reduces vocal communication space in marine mammals.     

Acoustically dependent latency shifts of BSER (wave V) in man

The latencies of wave V in Brain Stem Evoked Responses (BSER) elicited by a set of acoustic transients were measured. The stimuli were produced by delivering pulses to two filters, arranged in series. The filters were set so that the maximum acoustic energy in the transients, i.e., filtered clicks, occurred at 0.5, 1, 2, 4, or 8 kHz. The filtered clicks were presented via earphones at a rate of 30/s at 20, 40, or 60 dB HL to ten subjects with normal hearing. The latencies of wave V varied systematically with center frequency of the filtered clicks when they were each at the same HL. Stimuli presented at 40 dB HL produced the greatest opportunity for relating stimulus frequency to latency. The latencies for a smaller set of responses to stimuli presented at 10/s were the same as those for the principal data taken at 30/s. The changes in latency of wave V due to frequency are similar to those observed by other investigators in whole-nerve responses recorded in man.     

Behavioral responses of two captive bottlenose dolphins (Tursiops truncatus) to a continuous 50 kHz tone

At present, the fundamental frequencies of signals of most commercially available acoustic alarms to deter small cetaceans are below 20 kHz, but it is not well ascertained whether higher frequencies have a deterrent effect on bottlenose dolphins (Tursiops truncatus). Two captive bottlenose dolphins housed in a floating pen were subjected to a continuous pure tone at 50 kHz with a source level of 160 ± 2 dB (re 1 μPa, rms). The behavioral responses of dolphins were judged by comparing surfacing distance relative to the sound source, number of surfacings, and number of echolocation clicks produced, during forty 15 min baseline periods with forty 15 min test periods (four sessions per day, 40 sessions in total). On all 10 study days, surfacing distance and the number of surfacings increased while click production decreased during broadcasts of test sound. The avoidance threshold sound pressure level for a continuous 50 kHz tone for the bottlenose dolphins, in the context of this study, was estimated to be 144 ± 2 dB (re 1 μPa, rms). The results indicated that a continuous 50 kHz tonal signal can deter bottlenose dolphins from an area.     

Echolocation signals of Heaviside's dolphins (Cephalorhynchus heavisidii)

Field recordings of echolocation signals produced by Heaviside's dolphins (Cephalorhynchus heavisidii) were made off the coast of South Africa using a hydrophone array system. The system consisted of three hydrophones and an A-tag (miniature stereo acoustic data-logger). The mean centroid frequency was 125 kHz, with a -3 dB bandwidth of 15 kHz and -10 dB duration of 74 μs. The mean back-calculated apparent source level was 173 dB re 1 μPa(p.-p.). These characteristics are very similar to those found in other Cephalorhynchus species, and such narrow-band high-frequency echolocation clicks appear to be a defining characteristic of the Cephalorhynchus genus. Click bursts with very short inter-click intervals (up to 2 ms) were also recorded, which produced the "cry" sound reported in other Cephalorhynchus species. Since inter-click intervals correlated positively to click duration and negatively to bandwidth, Heaviside's dolphins may adjust their click duration and bandwidth based on detection range. The bimodal distribution of the peak frequency and stable bimodal peaks in spectra of individual click suggest a slight asymmetry in the click production mechanism.     

Underwater audiogram of a false killer whale (Pseudorca crassidens)

Underwater audiograms are available for only a few odontocete species. A false killer whale (Pseudorca crassidens) was trained at Sea Life Park in Oahu, Hawaii for an underwater hearing test using a go/no-go response paradigm. Over a 6-month period, auditory thresholds from 2-115 kHz were measured using an up/down staircase psychometric technique. The resulting audiogram showed hearing sensitivities below 64 kHz similar to those of belugas (Delphinapterus leucas) and Atlantic bottlenosed dolphins (Tursiops truncatus). Above 64 kHz, this Pseudorca had a rapid decrease in sensitivity of about 150 dB per octave. A similar decrease in sensitivity occurs at 32 kHz in the killer whale, at 50 kHz in the Amazon River dolphin, at 120 kHz in the beluga, at 140 kHz in the bottlenosed dolphin, and at 140 kHz in the harbor porpoise. The most sensitive range of hearing was from 16-64 kHz (a range of 10 dB from the maximum sensitivity). This range corresponds with the peak frequency of echolocation pulses recorded from captive Pseudorca.