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.     

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.     

The hearing threshold of a harbor porpoise (Phocoena phocoena) for impulsive sounds (L)

The distance at which harbor porpoises can hear underwater detonation sounds is unknown, but depends, among other factors, on the hearing threshold of the species for impulsive sounds. Therefore, the underwater hearing threshold of a young harbor porpoise for an impulsive sound, designed to mimic a detonation pulse, was quantified by using a psychophysical technique. The synthetic exponential pulse with a 5?ms time constant was produced and transmitted by an underwater projector in a pool. The resulting underwater sound, though modified by the response of the projection system and by the pool, exhibited the characteristic features of detonation sounds: A zero to peak sound pressure level of at least 30?dB (re 1?s(-1)) higher than the sound exposure level, and a short duration (34?ms). The animal's 50% detection threshold for this impulsive sound occurred at a received unweighted broadband sound exposure level of 60?dB re 1?μPa(2)s. It is shown that the porpoise's audiogram for short-duration tonal signals [Kastelein et al., J. Acoust. Soc. Am. 128, 3211-3222 (2010)] can be used to estimate its hearing threshold for impulsive sounds.     

Time sequences of sonar signals generated by a beluga whale when locating underwater objects

The results of measuring the sonar signals produced by a beluga whale when locating a target presented at a distance of 600 m are discussed. The head of the beluga whale was positioned at a depth of 1.5 m, and the acoustic pulses emitted by the animal were measured by a horizontal chain of four hydrophones, which was placed at a distance of 1.8 m from the head. The analysis of time sequences of acoustic signals generated by the beluga whale demonstrated that the animal, when searching for an underwater object, uses trains of pulses following at intervals of Δt < 5 ms and emitted within a wide sector (up to 36°). It performs scanning by beamed single pulses with Δt up to 200 ms, and, when it detects the target, it irradiates the latter with a group of such signals. To locate a difficult target (at a small depth and a large distance), the beluga whale uses trains of pulses with a duration of up to 0.6 s and a time-pulse modulation.     

A comparison of underwater hearing sensitivity in bottlenose dolphins (Tursiops truncatus) determined by electrophysiological and behavioral methods

Variable stimulus presentation methods are used in auditory evoked potential (AEP) estimates of cetacean hearing sensitivity, each of which might affect stimulus reception and hearing threshold estimates. This study quantifies differences in underwater hearing thresholds obtained by AEP and behavioral means. For AEP estimates, a transducer embedded in a suction cup (jawphone) was coupled to the dolphin's lower jaw for stimulus presentation. Underwater AEP thresholds were obtained for three dolphins in San Diego Bay and for one dolphin in a quiet pool. Thresholds were estimated from the envelope following response at carrier frequencies ranging from 10 to 150 kHz. One animal, with an atypical audiogram, demonstrated significantly greater hearing loss in the right ear than in the left. Across test conditions, the range and average difference between AEP and behavioral threshold estimates were consistent with published comparisons between underwater behavioral and in-air AEP thresholds. AEP thresholds for one animal obtained in-air and in a quiet pool demonstrated a range of differences of -10 to 9 dB (mean = 3 dB). Results suggest that for the frequencies tested, the presentation of sound stimuli through a jawphone, underwater and in-air, results in acceptable differences to AEP threshold estimates.     

Sensitivity of a tucuxi (Sotalia fluviatilis guianensis) to airborne sound

Auditory systems of cetaceans are considered highly specialized for underwater sound processing, whereas the extent of their hearing capacity in air is still a point of issue. In this study, the sensitivity to airborne sound in a male tucuxi (Sotalia fluviatilis guianensis) was tested by means of a go/no go response paradigm. Auditory thresholds were obtained from 2 to 31.5 kHz. Compared to the hearing thresholds of other dolphins as well as of amphibian mammals, the sensitivity to airborne sound of the test subject is low from 2 to 8 kHz, with the highest threshold at 4 kHz. Thresholds at 16 and 31.5 kHz reveal a sharp increase in hearing sensitivity. Thus, although not obtained in this study, the upper aerial hearing limit is in the ultrasonic range. A comparison of the present data with the underwater audiogram of the same test subject referred to sound intensity indicates that the sensitivity of Sotalia to underwater sound is generally better than to airborne sound.     

Auditory evoked potentials in two short-finned pilot whales (Globicephala macrorhynchus)

The hearing sensitivities of two short-finned pilot whales (Globicephala macrorhynchus) were investigated by measuring auditory evoked potentials generated in response to clicks and sinusoidal amplitude modulated (SAM) tones. The first whale tested, an adult female, was a long-time resident at SeaWorld San Diego with a known health history. Click-evoked responses in this animal were similar to those measured in other echolocating odontocetes. Auditory thresholds were comparable to dolphins of similar age determined with similar evoked potential methods. The region of best sensitivity was near 40 kHz and the upper limit of functional hearing was between 80 and 100 kHz. The second whale tested, a juvenile male, was recently stranded and deemed non-releasable. Click-evoked potentials were not detected in this animal and testing with SAM tones suggested severe hearing loss above 10 kHz.     

Passive acoustic location of bowhead whales in a population census off Point Barrow, Alaska

A sonobuoy array placed in the nearshore lead was used for locating bowhead whale sounds to determine if whales migrated past census stations beyond visual range and were uncounted. Based on a sample of 182 whale sounds (over 48 h) from closest point of approach (CPA) distances out to more than 10 km, 68% originated beyond 2 km (CPA), where only 1% of the 242 whales were sighted. No whales were sighted beyond 3 km during this time, but 53% of the located sounds originated that far and beyond. Thirty-seven other bowhead sounds over 15 h were distributed out to 6 km. Two tracked whales moved at average speeds of 1.5 and 1.8 kn. Maximum location error was 1%-25% in a sector of 120 degrees X 5-10 km, depending upon bearing and range. Most whale sounds were low-frequency moans, trumpeting roars, and repetitive sequences (songs) with peak spectrum source level up to 189 dB re: 1 microPa, 1 m. Lack of correlations between numbers of sounds and sighted whales precluded using bowhead sounds to count individuals or even to extrapolate ratios of unseen to observed whales.     

Estimated communication range of social sounds used by bottlenose dolphins (Tursiops truncatus)

Bottlenose dolphins, Tursiops truncatus, exhibit flexible associations in which the compositions of groups change frequently. We investigated the potential distances over which female dolphins and their dependent calves could remain in acoustic contact. We quantified the propagation of sounds in the frequency range of typical dolphin whistles in shallow water areas and channels of Sarasota Bay, Florida. Our results indicated that detection range was noise limited as opposed to being limited by hearing sensitivity. Sounds were attenuated to a greater extent in areas with seagrass than any other habitat. Estimates of active space of whistles showed that in seagrass shallow water areas, low-frequency whistles (7-13 kHz) with a 165 dB source level could be heard by dolphins at 487 m. In shallow areas with a mud bottom, all whistle frequency components of the same whistle could be heard by dolphins travel up to 2 km. In channels, high-frequency whistles (13-19 kHz) could be detectable potentially over a much longer distance (> 20 km). Our findings indicate that the communication range of social sounds likely exceeds the mean separation distances between females and their calves. Ecological pressures might play an important role in determining the separation distances within communication range.     

A threatened beluga (Delphinapterus leucas) population in the traffic lane: vessel-generated noise characteristics of the Saguenay-St. Lawrence Marine Park, Canada

The threatened resident beluga population of the St. Lawrence Estuary shares the Saguenay-St. Lawrence Marine Park with significant anthropogenic noise sources, including marine commercial traffic and a well-established, vessel-based whale-watching industry. Frequency-dependent (FD) weighting was used to approximate beluga hearing sensitivity to determine how noise exposure varied in time and space at six sites of high beluga summer residency. The relative contribution of each source to acoustic habitat degradation was estimated by measuring noise levels throughout the summer and noise signatures of typical vessel classes with respect to traffic volume and sound propagation characteristics. Rigid-hulled inflatable boats were the dominant noise source with respect to estimated beluga hearing sensitivity in the studied habitats due to their high occurrence and proximity, high correlation with site-specific FD-weighted sound levels, and the dominance of mid-frequencies (0.3-23 kHz) in their noise signatures. Median C-weighted sound pressure level (SPL(RMS)) had a range of 19 dB re 1 μPa between the noisiest and quietest sites. Broadband SPL(RMS) exceeded 120 dB re 1 μPa 8-32% of the time depending on the site. Impacts of these noise levels on St. Lawrence beluga will depend on exposure recurrence and individual responsiveness.     

Drilling and operational sounds from an oil production island in the ice-covered Beaufort sea

Recordings of sounds underwater and in air, and of iceborne vibrations, were obtained at Northstar Island, an artificial gravel island in the Beaufort Sea near Prudhoe Bay (Alaska). The aim was to document the levels, characteristics, and range dependence of sounds and vibrations produced by drilling and oil production during the winter, when the island was surrounded by shore-fast ice. Drilling produced the highest underwater broadband (10-10,000 Hz) levels (maximum= 124 dB re: 1 microPa at 1 km), and mainly affected 700-1400 Hz frequencies. In contrast, drilling did not increase broadband levels in air or ice relative to levels during other island activities. Production did not increase broadband levels for any of the sensors. In all media, broadband levels decreased by approximately 20 dB/tenfold change in distance. Background levels underwater were reached by 9.4 km during drilling and 3-4 km without. In the air and ice, background levels were reached 5-10 km and 2-10 km from Northstar, respectively, depending on the wind but irrespective of drilling. A comparison of the recorded sounds with harbor and ringed seal audiograms showed that Northstar sounds were probably audible to seals, at least intermittently, out to approximately 1.5 km in water and approximately 5 km in air.     

Auditory filter shapes for the bottlenose dolphin (Tursiops truncatus) and the white whale (Delphinapterus leucas) derived with notched noise

Auditory filter shapes were estimated in two bottlenose dolphins (Tursiops truncatus) and one white whale (Delphinapterus leucas) using a behavioral response paradigm and notched noise. Masked thresholds were measured at 20 and 30 kHz. Masking noise was centered at the test tone and had a bandwidth of 1.5 times the tone frequency. Half-notch width to center frequency ratios were 0, 0.125, 0.25, 0.375, and 0.5. Noise spectral density levels were 90 and 105 dB re: 1 microPa2/Hz. Filter shapes were approximated using a roex(p,r) function; the parameters p and r were found by fitting the integral of the roex(p,r) function to the measured threshold data. Mean equivalent rectangular bandwidths (ERBs) calculated from the filter shapes were 11.8 and 17.1% of the center frequency at 20 and 30 kHz, respectively, for the dolphins and 9.1 and 15.3% of the center frequency at 20 and 30 kHz, respectively, for the white whale. Filter shapes were broader at 30 kHz and 105 dB re: 1 microPa2/Hz masking noise. The results are in general agreement with previous estimates of ERBs in Tursiops obtained with a behavioral response paradigm.     

Detection of whale calls in noise: performance comparison between a beluga whale, human listeners, and a neural network

This article examines the masking by anthropogenic noise of beluga whale calls. Results from human masking experiments and a software backpropagation neural network are compared to the performance of a trained beluga whale. The goal was to find an accurate, reliable, and fast model to replace lengthy and expensive animal experiments. A beluga call was masked by three types of noise, an icebreaker's bubbler system and propeller noise, and ambient arctic ice-cracking noise. Both the human experiment and the neural network successfully modeled the beluga data in the sense that they classified the noises in the same order from strongest to weakest masking as the whale and with similar call-detection thresholds. The neural network slightly outperformed the humans. Both models were then used to predict the masking of a fourth type of noise, Gaussian white noise. Their prediction ability was judged by returning to the aquarium to measure masked-hearing thresholds of a beluga in white noise. Both models and the whale identified bubbler noise as the strongest masker, followed by ramming, then white noise. Natural ice-cracking noise masked the least. However, the humans and the neural network slightly overpredicted the amount of masking for white noise. This is neglecting individual variation in belugas, because only one animal could be trained. Comparing the human model to the neural network model, the latter has the advantage of objectivity, reproducibility of results, and efficiency, particularly if the interference of a large number of signals and noise is to be examined.     

Low frequency narrow-band calls in bottlenose dolphins (Tursiops truncatus): signal properties, function, and conservation implications

Dolphins routinely use sound for social purposes, foraging and navigating. These sounds are most commonly classified as whistles (tonal, frequency modulated, typical frequencies 5-10 kHz) or clicks (impulsed and mostly ultrasonic). However, some low frequency sounds have been documented in several species of dolphins. Low frequency sounds produced by bottlenose dolphins (Tursiops truncatus) were recorded in three locations along the Gulf of Mexico. Sounds were characterized as being tonal with low peak frequencies (mean?=?990 Hz), short duration (mean?=?0.069 s), highly harmonic, and being produced in trains. Sound duration, peak frequency and number of sounds in trains were not significantly different between Mississippi and the two West Florida sites, however, the time interval between sounds within trains in West Florida was significantly shorter than in Mississippi (t?=?-3.001, p?=?0.011). The sounds were significantly correlated with groups engaging in social activity (F=8.323, p=0.005). The peak frequencies of these sounds were below what is normally thought of as the range of good hearing in bottlenose dolphins, and are likely subject to masking by boat noise.     

Masked tonal hearing thresholds in the beluga whale

Masked tonal thresholds were measured for a beluga whale at one noise level and 32 frequencies between 40 Hz and 115 kHz. Critical ratios were estimated and compared with those previously measured for the bottlenose dolphin. Beluga whale critical ratios were found to be about 3 dB lower than those of the bottlenose dolphin. Absolute tonal thresholds were extended below previous measurements to 40 Hz.     

California sea lion (Zalophus californianus) aerial hearing sensitivity measured using auditory steady-state response and psychophysical methods

Although electrophysiological methods of measuring the hearing sensitivity of pinnipeds are not yet as refined as those for dolphins and porpoises, they appear to be a promising supplement to traditional psychophysical procedures. In order to further standardize electrophysiological methods with pinnipeds, a within-subject comparison of psychophysical and auditory steady-state response (ASSR) measures of aerial hearing sensitivity was conducted with a 1.5-yr-old California sea lion. The psychophysical audiogram was similar to those previously reported for otariids, with a U-shape, and thresholds near 10 dB re 20 μPa at 8 and 16 kHz. ASSR thresholds measured using both single and multiple simultaneous amplitude-modulated tones closely reproduced the psychophysical audiogram, although the mean ASSR thresholds were elevated relative to psychophysical thresholds. Differences between psychophysical and ASSR thresholds were greatest at the low- and high-frequency ends of the audiogram. Thresholds measured using the multiple ASSR method were not different from those measured using the single ASSR method. The multiple ASSR method was more rapid than the single ASSR method, and allowed for threshold measurements at seven frequencies in less than 20 min. The multiple ASSR method may be especially advantageous for hearing sensitivity measurements with otariid subjects that are untrained for psychophysical procedures.     

Underwater psychophysical audiogram of a young male California sea lion (Zalophus californianus)

Auditory evoked potential (AEP) data are commonly obtained in air while sea lions are under gas anesthesia; a procedure that precludes the measurement of underwater hearing sensitivity. This is a substantial limitation considering the importance of underwater hearing data in designing criteria aimed at mitigating the effects of anthropogenic noise exposure. To determine if some aspects of underwater hearing sensitivity can be predicted using rapid aerial AEP methods, this study measured underwater psychophysical thresholds for a young male California sea lion (Zalophus californianus) for which previously published aerial AEP thresholds exist. Underwater thresholds were measured in an aboveground pool at frequencies between 1 and 38 kHz. The underwater audiogram was very similar to those previously published for California sea lions, suggesting that the current and previously obtained psychophysical data are representative for this species. The psychophysical and previously measured AEP audiograms were most similar in terms of high-frequency hearing limit (HFHL), although the underwater HFHL was sharper and occurred at a higher frequency. Aerial AEP methods are useful for predicting reductions in the HFHL that are potentially independent of the testing medium, such as those due to age-related sensorineural hearing loss.     

Near-threshold equal-loudness contours for harbor seals (Phoca vitulina) derived from reaction times during underwater audiometry: a preliminary study

Equal-loudness functions describe relationships between the frequencies of sounds and their perceived loudness. This pilot study investigated the possibility of deriving equal-loudness contours based on the assumption that sounds of equal perceived loudness elicit equal reaction times (RTs). During a psychoacoustic underwater hearing study, the responses of two young female harbor seals to tonal signals between 0.125 and 100 kHz were filmed. Frame-by-frame analysis was used to quantify RT (the time between the onset of the sound stimulus and the onset of movement of the seal away from the listening station). Near-threshold equal-latency contours, as surrogates for equal-loudness contours, were estimated from RT-level functions fitted to mean RT data. The closer the received sound pressure level was to the 50% detection hearing threshold, the more slowly the animals reacted to the signal (RT range: 188-982 ms). Equal-latency contours were calculated relative to the RTs shown by each seal at sound levels of 0, 10, and 20 dB above the detection threshold at 1 kHz. Fifty percent detection thresholds are obtained with well-trained subjects actively listening for faint familiar sounds. When calculating audibility ranges of sounds for harbor seals in nature, it may be appropriate to consider levels 20 dB above this threshold.     

Auditory and behavioral responses of California sea lions (Zalophus californianus) to single underwater impulses from an arc-gap transducer

A behavioral response paradigm was used to measure underwater hearing thresholds in two California sea lions (Zalophus californianus) before and after exposure to underwater impulses from an arc-gap transducer. Preexposure and postexposure hearing thresholds were compared to determine if the subjects experienced temporary shifts in their masked hearing thresholds (MTTS). Hearing thresholds were measured at 1 and 10 kHz. Exposures consisted of single underwater impulses produced by an arc-gap transducer referred to as a "pulsed power device" (PPD). The electrical charge of the PPD was varied from 1.32 to 2.77 kJ; the distance between the subject and the PPD was varied over the range 3.4 to 25 m. No MTTS was observed in either subject at the highest received levels: peak pressures of approximately 6.8 and 14 kPa, rms pressures of approximately 178 and 183 dB re: 1 microPa, and total energy fluxes of 161 and 163 dB re: 1 microPa2s for the two subjects. Behavioral reactions to the tests were observed in both subjects. These reactions primarily consisted of temporary avoidance of the site where exposure to the PPD impulse had previously occurred.     

Real-time acoustic classification of sperm whale clicks and shipping impulses from deep-sea observatories

The automated real-time detection and classification of cetacean and anthropogenic sounds from deep-sea observatories can play a key role to study cetaceans in the field, to quantify the impact of anthropogenic sounds or to initiate mitigation measures during potentially harmful human activities. In the area of the NEMO-ONDE deep-sea observatory, sperm whales are often present together with heavy shipping. The spatial coincidence of both sound sources allows for the long term monitoring of their interaction. Some ships produce impulsive sounds and the automated separation of these impulses from sperm whale clicks is not a trivial task. As part of a detection, classification and localisation system for acoustic data from marine observatories, we present four modules performing the automated real-time classification of clicks from sperm whales and impulsive sounds produced by ships. First, two modules detect segments that contain impulsive sounds within a specifiable frequency band and return the impulses’ positions. Then, two modules classify the detected impulses as sperm whale clicks or ship impulses. Finally, at the level of 22 s segments, the classification outputs from individual impulses are combined into a decision on the presence of sperm whale clicks or ship impulses. The modules’ reliability was tested on data from the NEMO-ONDE observatory. Training and testing data were separated by more than 2 months, enabling to assess the consistency of the predictions over the long term. The automated separation between segments of the two classes was high with area under the ROC curve (AUC) values between 0.94 and 0.98.