Masking release at 4 and 32 kHz in harbor seals associated with sinusoidal amplitude-modulated masking noise.

Masking can reduce the efficiency of communication and prey and predator detection. Most underwater sounds fluctuate in amplitude, which may influence the amount of masking experienced by marine mammals. The hearing thresholds of two harbor seals for tonal sweeps (centered at 4 and 32 kHz) masked by sinusoidal amplitude modulated (SAM) Gaussian one-third octave noise bands centered around the narrow-band test sweep frequencies, were studied with a psychoacoustic technique. Masking was assessed in relation to signal duration, (500, 1000, and 2000 ms) and masker level, at eight amplitude modulation rates (1-90 Hz). Masking release (MR) due to SAM compared thresholds in modulated and unmodulated maskers. Unmodulated maskers resulted in critical ratios of 21 dB at 4 kHz and 31 dB at 32 kHz. Masked thresholds were similarly affected by SAM rate with the lowest thresholds and the largest MR being observed for SAM rates of 1 and 2 Hz at higher masker levels. MR was higher for 32-kHz maskers than for 4-kHz maskers. Increasing signal duration from 500 ms to 2000 ms had minimal effect on MR. The results are discussed with respect to MR resulting from envelope variation and the impact of noise in the environment on target signal detection.

Assessing potential perception of shipping noise by marine mammals in an arctic inlet.

Shipping is increasing in Arctic regions, exposing marine mammals to increased underwater noise. Noise analyses often use unweighted broadband sound pressure levels (SPL) to assess noise impacts, but this does not account for the animals' hearing abilities at different frequencies. In 2018 and 2019, noise levels were recorded at five and three sites, respectively, along a shipping route in an inlet of Northern Baffin Island, Canada. Broadband SPLs (10 Hz-25 kHz), unweighted and with auditory weighing functions from three marine mammal groups, were compared between times ore carriers (travelling < 9 knots) were present or absent. Clearly audible distances of shipping noise and exposure durations were estimated for each weighting function relative to vessel direction, orientation, and year. Auditory weighting functions had significant effects on the potential perception of shipping noise. Bowhead whales (Balaena mysticetus) experienced similar SPLs to unweighted levels. Narwhals (Monodon monoceros) and ringed seals (Pusa hispida) experienced lower SPLs. Narwhals were unlikely to clearly perceive shipping noise unless ships were in close proximity (<3 km) and ambient noise levels were low. Detectability propagation models of presumed noise exposure from shipping must be based on the hearing sensitivities of each species group when assessing noise impacts on marine mammals.

Masking release at 4 kHz in harbor porpoises (Phocoena phocoena) associated with sinusoidal amplitude-modulated masking noise.

Acoustic masking reduces the efficiency of communication, prey detection, and predator avoidance in marine mammals. Most underwater sounds fluctuate in amplitude. The ability of harbor porpoises (Phocoena phocoena) to detect sounds in amplitude-varying masking noise was examined. A psychophysical technique evaluated hearing thresholds of three harbor porpoises for 500-2000 ms tonal sweeps (3.9-4.1 kHz), presented concurrently with sinusoidal amplitude-modulated (SAM) or unmodulated Gaussian noise bands centered at 4 kHz. Masking was assessed in relation to signal duration and masker level, amplitude modulation rate (1, 2, 5, 10, 20, 40, 80, and 90 Hz), modulation depth (50%, 75%, and 100%) and bandwidth (1/3 or 1 octave). Masking release (MR) due to SAM was assessed by comparing thresholds in modulated and unmodulated maskers. Masked thresholds were affected by SAM rate with the lowest thresholds (i.e., largest MR was 14.5 dB) being observed for SAM rates between 1 and 5 Hz at higher masker levels. Increasing the signal duration from 500-2000 ms increased MR by 3.3 dB. Masker bandwidth and depth of modulation had no substantial effect on MR. The results are discussed with respect to MR resulting from envelope variation and the impact of noise in the environment.

Weddell seals produce ultrasonic vocalizations.

Seals (phocids) are generally not thought to produce vocalizations having ultrasonic fundamental frequencies (≥20 kHz), although previous studies could have been biased by sampling limitations. This study characterizes common, yet, previously undescribed, ultrasonic Weddell seal (Leptonychotes weddellii) vocalizations. The vocalizations were identified in more than one year (2017-2018) of broadband acoustic data obtained by a continuously recording underwater observatory in McMurdo Sound, Antarctica. Nine recurrent call types were identified that were composed of single or multiple vocal elements whose fundamental frequencies spanned the ultrasonic range to nearly 50 kHz. Eleven vocal elements had ultrasonic center frequencies (≥20 kHz), including chirps, whistles, and trills, with two elements at >30 kHz. Six elements had fundamental frequencies always >21 kHz. The fundamental frequency of one repetitive U-shaped whistle element reached 44.2 kHz and descending chirps (≥3.6 ms duration) commenced at ≤49.8 kHz. The source amplitude of one fully ultrasonic chirp element (29.5 kHz center frequency) was 137 dB re 1 µPa-m. Harmonics of some vocalizations exceeded 200 kHz. Ultrasonic vocalizations occurred throughout the year with the usage of repetitive ultrasonic chirp-based calls appearing to dominate in winter darkness. The functional significance of these high-frequency vocalizations is unknown.

Temporary hearing threshold shift in harbor seals (Phoca vitulina) due to one-sixth-octave noise bands centered at 0.5, 1, and 2 kHz.

This study concludes a larger project on the frequency-dependent susceptibility to noise-induced temporary hearing threshold shift (TTS) in harbor seals (Phoca vitulina). Here, two seals were exposed to one-sixth-octave noise bands (NBs) centered at 0.5, 1, and 2 kHz at several sound exposure levels (SELs, in dB re 1 µPa2s). TTSs were quantified at the center frequency of each NB, half an octave above, and one octave above, at the earliest within 1-4 min after exposure. Generally, elicited TTSs were low, and the highest TTS1-4 occurred at half an octave above the center frequency of the fatiguing sound: after exposure to the 0.5-kHz NB at 210 dB SEL, the TTS1-4 at 0.71 kHz was 2.3 dB; after exposure to the 1-kHz NB at 207 dB SEL, the TTS1-4 at 1.4 kHz was 6.1 dB; and after exposure to the 2-kHz NB at 215 dB SEL, TTS1-4 at 2.8 kHz was 7.9 dB. Hearing always recovered within 60 min, and susceptibility to TTS was similar in both seals. The results show that, for the studied frequency range, the lower the center frequency of the fatiguing sound, the higher the SEL required to cause the same TTS.

Temporary hearing threshold shift in harbor seals (Phoca vitulina) due to a one-sixth-octave noise band centered at 40 kHz.

As part of a series of studies to determine frequency-dependent susceptibility to temporary hearing threshold shifts (TTS), two female harbor seals (F01 and F02) were exposed for 60 min to a one-sixth-octave noise band centered at 40 kHz at mean sound pressure levels ranging from 126 to 153 dB re 1 µPa [mean received sound exposure level (SEL) range: 162-189 dB re 1 µPa2s]. TTSs were quantified at 40, 50, and 63 kHz within 1-4 min of the exposure for F02 and within 12-16 min of the exposure for F01. In F02, significant TTS1-4 (1-4 min post exposure) occurred at 40 kHz with SELs of ≥183 dB re 1 µPa2s and at 50 kHz with SELs of ≥174 dB re 1 µPa2s. At 63 kHz, TTS1-4 occurred with SELs ≥186 dB re 1 µPa2s. In F01, significant TTS12-16 (12-16 min post exposure) occurred only at 50 kHz with SELs of ≥177 dB re 1 µPa2s. The highest TTSs (27.5 dB in F02, 29.8 dB in F01) occurred at 50 kHz, one-third of an octave above the fatiguing sound's center frequency (SEL = 189 dB re 1 µPa2s); recovery took 2 days in F02 and 4 days in F01. In most other cases, recovery was within 1 h. The seals have a similar susceptibility to TTS from 4 to 40 kHz.

Temporary hearing threshold shift in harbor seals (Phoca vitulina) due to a one-sixth-octave noise band centered at 32 kHz.

Two female harbor seals were exposed for 60 min to a continuous one-sixth-octave noise band centered at 32 kHz at sound pressure levels of 92 to 152 dB re 1 µPa, resulting in sound exposure levels (SELs) of 128 to 188 dB re 1 µPa2s. This was part of a larger project determining frequency-dependent susceptibility to temporary threshold shift (TTS) in harbor seals over their entire hearing range. After exposure, TTSs were quantified at 32, 45, and 63 kHz with a psychoacoustic technique. At 32 kHz, only small TTSs (up to 5.9 dB) were measured 1-4 min (TTS1-4) after exposure, and recovery was within 1 h. The higher the SEL, the higher the TTS induced at 45 kHz. Below ∼176 dB re 1 µPa2s, the maximum TTS1-4 was at 32 kHz; above ∼176 dB re 1 µPa2s, the maximum TTS1-4 (up to 33.8 dB) was at 45 kHz. During one particular session, a seal was inadvertently exposed to an SEL of ∼191 dB re 1 µPa2s and at 45 kHz, her TTS1-4 was >45 dB; her hearing recovered over 4 days. Harbor seals appear to be equally susceptible to TTS caused by sounds in the 2.5-32 kHz range.

Temporary hearing threshold shift in harbor seals (Phoca vitulina) due to a one-sixth-octave noise band centered at 16 kHz.

Temporary hearing threshold shifts (TTSs) were investigated in two adult female harbor seals after exposure for 60 min to a continuous one-sixth-octave noise band centered at 16 kHz (the fatiguing sound) at sound pressure levels of 128-149 dB re 1 µPa, resulting in sound exposure levels (SELs) of 164-185 dB re 1 µPa2s. TTSs were quantified at the center frequency of the fatiguing sound (16 kHz) and at half an octave above that frequency (22.4 kHz) by means of a psychoacoustic hearing test method. Susceptibility to TTS was similar in both animals when measured 8-12 and 12-16 min after cessation of the fatiguing sound. TTS increased with increasing SEL at both frequencies, but above an SEL of 174 dB re 1 µPa2s, TTS was greater at 22.4 kHz than at 16 kHz for the same SELs. Recovery was rapid: the greatest TTS, measured at 22.4 kHz 1-4 min after cessation of the sound, was 17 dB, but dropped to 3 dB in 1 h, and hearing recovered fully within 2 h. The affected hearing frequency should be considered when estimating ecological impacts of anthropogenic sound on seals. Between 2.5 and 16 kHz the species appears equally susceptible to TTS.

Hearing thresholds, for underwater sounds, of harbor seals (Phoca vitulina) at the water surface.

High-amplitude impulsive sounds produced by pile driving and airguns may result in hearing damage in nearby seals. By swimming at the water surface, seals may reduce their exposure to underwater sound, as sound pressure levels (SPLs) are often lower just below the surface than deeper in the water column. Seals can make physiological adjustments such that they can switch between having maximum sensitivity for either aerial or underwater sounds. This could mean that hearing sensitivity for underwater sounds is lower when swimming at the water surface (when hearing may be focused on aerial sounds) than when swimming at depth. To investigate this possibility, hearing thresholds of two female harbor seals were quantified psychophysically, while their heads were in the position normally adopted while swimming at the surface. The seals' hearing thresholds at the water surface were similar to each other and to previous measurements made at 1 m depth. When calculating the cumulative sound exposure level for hearing damage assessment, the SPL just below the water surface needs to be measured or modeled, and the proportion of time seals normally swim at the water surface needs to be estimated, to estimate the sound energy that reaches the seals' ears.

Harp Seals Do Not Increase Their Call Frequencies When It Gets Noisier.

Some species avoid low-frequency masking by shifting their calls to higher frequencies. We addressed the hypothesis that Pagophilus groenlandicus (harp seals) will make more high-frequency underwater calls to avoid low-frequency conspecific masking as calling rates increase. The spectral shapes at high and low calling rates were compared (after equalizing the broadband amplitudes). There were no significant differences between the spectral shapes. Pagophilus groenlandicus do not alter the proportions of low- and high-frequency calls as it gets noisier. This suggests that they may not shift their calling frequencies when encountering low-frequency, broadband anthropogenic noise.

A practical weighting function for harbor porpoise underwater sound level measurements.

Harbor porpoise (Phocoena phocoena) are subject to underwater noise disturbance from anthropogenic sources, especially shipping. The underwater audiograms of harbor porpoise were used to create a frequency weighting function, dBht(Phocoena phocoena), to permit estimation of the broadband perceived amplitudes of ambient and shipping noise. An equation was fit to the 0.02-20 kHz range of unmasked detection thresholds and normalizing to 0 dB at 20 kHz; dB = 46.4-35.6 log(kHz). The weighting function de-emphasizes the low frequency components of noise. Harbor porpoise hearing is less sensitive to low frequency shipping noise and, except at high amplitudes, estimating potential noise impacts using linear measurements will be misleading.

Hearing threshold shifts and recovery in harbor seals (Phoca vitulina) after octave-band noise exposure at 4 kHz.

Safety criteria for underwater sounds from offshore pile driving are needed to protect marine mammals. As a first step toward understanding effects of impulsive sounds, two harbor seals were exposed to octave-band white noise centered at 4 kHz at three mean received sound pressure levels (SPLs; 124, 136, and 148 dB re 1 µPa) at up to six durations (7.5, 15, 30, 60, 120, and 240 min); mean received sound exposure level (SEL) range was 166-190 dB re 1 µPa(2) s. Hearing thresholds were determined before and after exposure. Temporary hearing threshold shifts (TTS) and subsequent recovery were quantified as changes in hearing thresholds at 1-4, 4-8, 8-12, 48, and 96 min after noise exposure in seal 01, and at 12-16, 16-20, 20-24, 60, and 108 min after exposure in seal 02. Maximum TTS (1-4 min after 120 min exposure to 148 dB re 1 µPa; 187 dB SEL) was 10 dB. Recovery occurred within ~60 min. Statistically significant TTSs (>2.5 dB) began to occur at SELs of ~170 (136 SPL, 60 min) and 178 dB re 1 µPa(2) s (148 SPL, 15 min). However, SEL is not an optimal predictor of TTS for long duration, low SPL continuous noise, as duration and SPL play unequal roles in determining induced TTS.

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.

The effect of signal duration on the underwater hearing thresholds of two harbor seals (Phoca vitulina) for single tonal signals between 0.2 and 40 kHz.

The underwater hearing sensitivities of two 2-year-old female harbor seals were quantified in a pool built for acoustic research by using a behavioral psycho-acoustic technique. The animals were trained only to respond when they detected an acoustic signal ("go/no-go" response). Detection thresholds were obtained for pure tone signals (frequencies: 0.2-40 kHz; durations: 0.5-5000 ms, depending on the frequency; 59 frequency-duration combinations). Detection thresholds were quantified by varying the signal amplitude by the 1-up, 1-down staircase method, and were defined as the stimulus levels, resulting in a 50% detection rate. The hearing thresholds of the two seals were similar for all frequencies except for 40 kHz, for which the thresholds differed by, on average, 3.7 dB. There was an inverse relationship between the time constant (tau), derived from an exponential model of temporal integration, and the frequency [log(tau)=2.86-0.94 log(f);tau in ms and f in kHz]. Similarly, the thresholds increased when the pulse was shorter than approximately 780 cycles (independent of the frequency). For pulses shorter than the integration time, the thresholds increased by 9-16 dB per decade reduction in the duration or number of cycles in the pulse. The results of this study suggest that most published hearing thresholds

Critical ratios in harbor porpoises (Phocoena phocoena) for tonal signals between 0.315 and 150 kHz in random Gaussian white noise.

A psychoacoustic behavioral technique was used to determine the critical ratios (CRs) of two harbor porpoises for tonal signals with frequencies between 0.315 and 150 kHz, in random Gaussian white noise. The masked 50% detection hearing thresholds were measured using a "go/no-go" response paradigm and an up-down staircase psychometric method. CRs were determined at one masking noise level for each test frequency and were similar in both animals. For signals between 0.315 and 4 kHz, the CRs were relatively constant at around 18 dB. Between 4 and 150 kHz the CR increased gradually from 18 to 39 dB ( approximately 3.3 dB/octave). Generally harbor porpoises can detect tonal signals in Gaussian white noise slightly better than most odontocetes tested so far. By combining the mean CRs found in the present study with the spectrum level of the background noise levels at sea, the basic audiogram, and the directivity index, the detection threshold levels of harbor porpoises for tonal signals in various sea states can be calculated.

Underwater hearing sensitivity of harbor seals (Phoca vitulina) for narrow noise bands between 0.2 and 80 kHz.

The underwater hearing sensitivities of two 1.5-year-old female harbor seals were quantified in a quiet pool built specifically for acoustic research, by using a behavioral psychoacoustic technique. The animals were trained to respond when they detected an acoustic signal and not to respond when they did not ("go/no-go" response). Fourteen narrowband noise signals (1/3-octave bands but with some energy in adjacent bands), at 1/3-octave center frequencies of 0.2-80 kHz, and of 900 ms duration, were tested. Thresholds at each frequency were measured using the up-down staircase method and defined as the stimulus level resulting in a 50% detection rate. Between 0.5 and 40 kHz, the thresholds corresponded to a 1/3-octave band noise level of approximately 60 dB re 1 microPa (SD+/-3.0 dB). At lower frequencies, the thresholds increased to 66 dB re 1 microPa and at 80 kHz the thresholds rose to 114 dB re 1 microPa. The 1/3-octave noise band thresholds of the two seals did not differ from each other, or from the narrowband frequency-modulated tone thresholds at the same frequencies obtained a few months before for the same animals. These hearing threshold values can be used to calculate detection ranges of underwater calls and anthropogenic noises by harbor seals.

Underwater detection of tonal signals between 0.125 and 100 kHz by harbor seals (Phoca vitulina).

The underwater hearing sensitivities of two 1-year-old female harbor seals were quantified in a pool built for acoustic research, using a behavioral psychoacoustic technique. The animals were trained to respond when they detected an acoustic signal and not to respond when they did not (go/no-go response). Pure tones (0.125-0.25 kHz) and narrowband frequency modulated (tonal) signals (center frequencies 0.5-100 kHz) of 900 ms duration were tested. Thresholds at each frequency were measured using the up-down staircase method and defined as the stimulus level resulting in a 50% detection rate. The audiograms of the two seals did not differ statistically: both plots showed the typical mammalian U-shape, but with a wide and flat bottom. Maximum sensitivity (54 dB re 1 microPa, rms) occurred at 1 kHz. The frequency range of best hearing (within 10 dB of maximum sensitivity) was from 0.5 to 40 kHz (6(1/3) octaves). Higher hearing thresholds (indicating poorer sensitivity) were observed below 1 and above 40 kHz. Thresholds below 4 kHz were lower than those previously described for harbor seals, which demonstrates the importance of using quiet facilities, built specifically for acoustic research, for hearing studies in marine mammals. The results suggest that under unmasked conditions many anthropogenic noise sources and sounds from conspecifics are audible to harbor seals at greater ranges than formerly believed.

Deterring effects of 8-45 kHz tone pulses on harbour seals (Phoca vitulina) in a large pool.

The marine aquaculture industry suffers losses due to pinniped attacks which damage net enclosures and fish stocks. Acoustic harassment devices (AHDs) emit loud sounds which are intended to deter pinnipeds from approaching aquaculture enclosures. At present, many AHDs emit sounds in the 8-20 kHz frequency range. It is not known whether sounds of higher frequencies have a deterrent effect on seals. Therefore five captive harbour seals (Phoca vitulina) were subjected to four series of tone pulses together spanning a broad frequency range (8, 16, 32 and 45 kHz). Pulse duration was 250 ms and pulse interval was 5s. Each of the four sounds was made deterrent by increasing the amplitude. The seals reacted by swimming away from the sounds. The displacement effect of each sound was judged by comparing the animals' surface positions, and number of surfacings, during ten 45 min baseline periods with ten 45 min test periods per frequency (one frequency per day in rotation, 40 sessions in total). The seals were displaced by all four frequencies throughout the 40 trial days. The seals came to the surface more often when the test tones were produced than in the baseline periods. The initial displacement distances did not change over the 40 test days. This suggests that operating AHDs for only short periods will be more effective and less likely to result in habituation by the seals than operating them continuously. The discomfort threshold sound pressure level (SPL) was established for each of the four pulse frequencies. The acoustic discomfort threshold SPL is defined as the boundary SPL between the area that the animals generally occupied during the transmission of the sounds and the area that they generally did not enter during sound transmission. The discomfort threshold SPL may depend on the context.