Audiogram of a harbor porpoise (Phocoena phocoena) measured with narrow-band frequency-modulated signals

The underwater hearing sensitivity of a two-year-old harbor porpoise was measured in a pool using standard psycho-acoustic techniques. The go/no-go response paradigm and up-down staircase psychometric method were used. Auditory sensitivity was measured by using narrow-band frequency-modulated signals having center frequencies between 250 Hz and 180 kHz. The resulting audiogram was U-shaped with the range of best hearing (defined as 10 dB within maximum sensitivity) from 16 to 140 kHz, with a reduced sensitivity around 64 kHz. Maximum sensitivity (about 33 dB re 1 microPa) occurred between 100 and 140 kHz. This maximum sensitivity range corresponds with the peak frequency of echolocation pulses produced by harbor porpoises (120-130 kHz). Sensitivity falls about 10 dB per octave below 16 kHz and falls off sharply above 140 kHz (260 dB per octave). Compared to a previous audiogram of this species (Andersen, 1970), the present audiogram shows less sensitive hearing between 2 and 8 kHz and more sensitive hearing between 16 and 180 kHz. This harbor porpoise has the highest upper-frequency limit of all odontocetes investigated. The time it took for the porpoise to move its head 22 cm after the signal onset (movement time) was also measured. It increased from about 1 s at 10 dB above threshold, to about 1.5 s at threshold.     

Receiving beam patterns in the horizontal plane of a harbor porpoise (Phocoena phocoena)

Receiving beam patterns of a harbor porpoise were measured in the horizontal plane, using narrow-band frequency modulated signals with center frequencies of 16, 64, and 100 kHz. Total signal duration was 1000 ms, including a 200 ms rise time and 300 ms fall time. The harbor porpoise was trained to participate in a psychophysical test and stationed itself horizontally in a specific direction in the center of a 16-m-diameter circle consisting of 16 equally-spaced underwater transducers. The animal's head and the transducers were in the same horizontal plane, 1.5 m below the water surface. The go/no-go response paradigm was used; the animal left the listening station when it heard a sound signal. The method of constants was applied. For each transducer the 50% detection threshold amplitude was determined in 16 trials per amplitude, for each of the three frequencies. The beam patterns were not symmetrical with respect to the midline of the animal's body, but had a deflection of 3-7 degrees to the right. The receiving beam pattern narrowed with increasing frequency. Assuming that the pattern is rotation-symmetrical according to an average of the horizontal beam pattern halves, the receiving directivity indices are 4.3 at 16 kHz, 6.0 at 64 kHz, and 11.7 dB at 100 kHz. The receiving directivity indices of the porpoise were lower than those measured for bottlenose dolphins. This means that harbor porpoises have wider receiving beam patterns than bottlenose dolphins for the same frequencies. Directivity of hearing improves the signal-to-noise ratio and thus is a tool for a better detection of certain signals in a given ambient noise condition.     

Target detection by an echolocating harbor porpoise (Phocoena phocoena)

Two echolocation experiments are described. They were conducted on the same harbor porpoise housed in a sea pen, one year apart at Neeltje Jans, The Netherlands. The aims were to determine the target detection ability of an echolocating harbor porpoise, with the ultimate goal to predict the distance at which harbor porpoises can detect fishing nets. In experiment 1, the maximum distance at which the 3-year-old porpoise could detect a 7.62-cm diameter water-filled stainless-steel sphere by echolocation was determined psychophysically. The 50%-current detection threshold was reached when the sphere was at a distance of 26 m from the porpoise's rostrum. In experiment 2, conducted a year later, the maximum detection distance for a 5.08-cm water-filled stainless-steel sphere was 15.9 m. The target strengths of both targets were measured using simulated harbor porpoise echolocation signals and the results, coupled with transmission-loss calculations, indicated that the echo levels received by the porpoise with the targets at the threshold ranges in the two experiments were only 1.3 dB apart. Together with information on the target strengths of various fishing nets, the results of the present study can be used to predict the distance at which the nets can be detected by harbor porpoises.     

Number and duration of echolocation click trains produced by a harbor porpoise (Phocoena phocoena) in relation to target and performance

Echolocation effort (number and duration of echolocation click trains produced) by a harbor porpoise is described in relation to target presence, strength and distance, and performance of the detection task. The porpoise was presented with two target sizes at five distances (12-20 m), or no target, and had to indicate whether it could detect the target. Small, distant targets required long and multiple click trains. Multiple click trains mostly occurred when the small target was far away and not detected, and during target-absent trials in which the animal correctly responded. In target-absent trials, an incorrect response was linked to short click trains. Click train duration probably increased until the animal's certainty about the target's presence or absence exceeded a certain level, after which the porpoise responded.     

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.     

Hearing thresholds of a harbor porpoise (Phocoena phocoena) for helicopter dipping sonar signals (1.43-1.33 kHz) (L)

Helicopter long range active sonar (HELRAS), a "dipping" sonar system used by lowering transducer and receiver arrays into water from helicopters, produces signals within the functional hearing range of many marine animals, including the harbor porpoise. The distance at which the signals can be heard is unknown, and depends, among other factors, on the hearing sensitivity of the species to these particular signals. Therefore, the hearing thresholds of a harbor porpoise for HELRAS signals were quantified by means of a psychophysical technique. Detection thresholds were obtained for five 1.25 s simulated HELRAS signals, varying in their harmonic content and amplitude envelopes. The 50% hearing thresholds for the different signals were similar: 76 dB re 1 μPa (broadband sound pressure level, averaged over the signal duration). The detection thresholds were similar to those found in the same porpoise for tonal signals in the 1-2 kHz range measured in a previous study. Harmonic distortion, which occurred in three of the five signals, had little influence on their audibility. The results of this study, combined with information on the source level of the signal, the propagation conditions and ambient noise levels, allow the calculation of accurate estimates of the distances at which porpoises can detect HELRAS signals.     

The influence of signal parameters on the sound source localization ability of a harbor porpoise (Phocoena phocoena)

It is unclear how well harbor porpoises can locate sound sources, and thus can locate acoustic alarms on gillnets. Therefore the ability of a porpoise to determine the location of a sound source was determined. The animal was trained to indicate the active one of 16 transducers in a 16-m-diam circle around a central listening station. The duration and received level of the narrowband frequency-modulated signals (center frequencies 16, 64 and 100 kHz) were varied. The animal's localization performance increased when the signal duration increased from 600 to 1000 ms. The lower the received sound pressure level (SPL) of the signal, the harder the animal found it to localize the sound source. When pulse duration was long enough (approximately 1 s) and the received SPLs of the sounds were high (34-50 dB above basic hearing thresholds or 3-15 dB above the theoretical masked detection threshold in the ambient noise condition of the present study), the animal could locate sounds of the three frequencies almost equally well. The porpoise was able to locate sound sources up to 124 degrees to its left or right more easily than sounds from behind it.     

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.     

Hearing thresholds of a harbor porpoise (Phocoena phocoena) for sweeps (1-2 kHz and 6-7 kHz bands) mimicking naval sonar signals

The distance at which active naval sonar signals can be heard by harbor porpoises depends, among other factors, on the hearing thresholds of the species for those signals. Therefore the hearing sensitivity of a harbor porpoise was determined for 1 s up-sweep and down-sweep signals, mimicking mid-frequency and low-frequency active sonar sweeps (MFAS, 6-7 kHz band; LFAS, 1-2 kHz band). The 1-2 kHz sweeps were also tested with harmonics, as sonars sometimes produce these as byproducts of the fundamental signal. The hearing thresholds for up-sweeps and down-sweeps within each sweep pair were similar. The 50% detection threshold sound pressure levels (broadband, averaged over the signal duration) of the 1-2 kHz and 6-7 kHz sweeps were 75 and 67 dB re 1 μPa(2), respectively. Harmonic deformation of the 1-2 kHz sweeps reduced the threshold to 59 dB re 1 μPa(2). This study shows that the presence of harmonics in sonar signals can increase the detectability of a signal by harbor porpoises, and that tonal audiograms may not accurately predict the audibility of sweeps. LFAS systems, when designed to produce signals without harmonics, can operate at higher source levels than MFAS systems, at similar audibility distances for porpoises.     

The effect of signal duration on the underwater detection thresholds of a harbor porpoise (Phocoena phocoena) for single frequency-modulated tonal signals between 0.25 and 160 kHz

The underwater hearing sensitivity of a young male harbor porpoise for tonal signals of various signal durations was quantified by using a behavioral psychophysical technique. The animal was trained to respond only when it detected an acoustic signal. Fifty percent detection thresholds were obtained for tonal signals (15 frequencies between 0.25-160 kHz, durations 0.5-5000 ms depending on the frequency; 134 frequency-duration combinations in total). Detection thresholds were quantified by varying signal amplitude by the 1-up 1-down staircase method. The hearing thresholds increased when the signal duration fell below the time constant of integration. The time constants, derived from an exponential model of integration [Plomp and Bouman, J. Acoust. Soc. Am. 31, 749-758 (1959)], varied from 629 ms at 2 kHz to 39 ms at 64 kHz. The integration times of the porpoises were similar to those of other mammals including humans, even though the porpoise is a marine mammal and a hearing specialist. The results enable more accurate estimations of the distances at which porpoises can detect short-duration environmental tonal signals. The audiogram thresholds presented by Kastelein et al. [J. Acoust. Soc. Am. 112, 334-344 (2002)], after correction for the frequency bandwidth of the FM signals, are similar to the results of the present study for signals of 1500 ms duration. Harbor porpoise hearing is more sensitive between 2 and 10 kHz, and less sensitive above 10 kHz, than formerly believed.     

Effect of broadband-noise masking on the behavioral response of a harbor porpoise (Phocoena phocoena) to 1-s duration 6-7 kHz sonar up-sweeps

Naval sonar systems produce signals which may affect the behavior of harbor porpoises, though their effect may be reduced by ambient noise. To show how natural ambient noise influences the effect of sonar sweeps on porpoises, a porpoise in a pool was exposed to 1-s duration up-sweeps, similar in frequency range (6-7 kHz) to those of existing naval sonar systems. The sweep signals had randomly generated sweep intervals of 3-7 s (duty cycle: 19%). Behavioral parameters during exposure to signals were compared to those during baseline periods. The sessions were conducted under five background noise conditions: the local normal ambient noise and four conditions mimicking the spectra for wind-generated noise at Sea States 2-8. In all conditions, the sweeps caused the porpoise to swim further away from the transducer, surface more often, swim faster, and breathe more forcefully than during the baseline periods. However, the higher the background noise level, the smaller the effects of the sweeps on the surfacing behavior of the porpoise. Therefore, the effects of naval sonar systems on harbor porpoises are determined not only by the received level of the signals and the hearing sensitivity of the animals but also by the background noise.     

On the performance of automated porpoise-click-detectors in experiments with captive harbor porpoises (Phocoena phocoena)

Recently, automated porpoise-click-detectors (T-PODs, Chelonia-Marine-Research) have been used intensively in monitoring harbor porpoises (Phocoena phocoena) in the wild. However, the automated click-detection-mechanism of the T-POD leads to questions on the characteristics of the detection process. We undertook experiments with six captive harbor porpoises (four subadult males in one pool, two adult males in another) at the Dolfinarium Harderwijk (Netherlands). One T-POD was placed for over a week in each pool, while the behavior of the porpoises was logged by visual observation. Data were analyzed using the T-POD software. A total of 725 431 clicks in 30 090 trains were recorded with 32% of the trains classified as CET HI, 27% as CET LO, and 41% as DOUBTFUL. All three train classes differed significantly in all parameters, except for click duration. We conclude that T-PODs perform generally well in detecting click trains of harbor porpoises but that in any future study trains classified as being of lower probability should be investigated very carefully to avoid the risk of losing valuable information.     

Threshold received sound pressure levels of single 1-2 kHz and 6-7 kHz up-sweeps and down-sweeps causing startle responses in a harbor porpoise (Phocoena phocoena)

Mid-frequency and low-frequency sonar systems produce frequency-modulated sweeps which may affect harbor porpoises. To study the effect of sweeps on behavioral responses (specifically "startle" responses, which we define as sudden changes in swimming speed and/or direction), a harbor porpoise in a large pool was exposed to three pairs of sweeps: a 1-2 kHz up-sweep was compared with a 2-1 kHz down-sweep, both with and without harmonics, and a 6-7 kHz up-sweep was compared with a 7-6 kHz down-sweep without harmonics. Sweeps were presented at five spatially averaged received levels (mRLs; 6 dB steps; identical for the up-sweep and down-sweep of each pair). During sweep presentation, startle responses were recorded. There was no difference in the mRLs causing startle responses for up-sweeps and down-sweeps within frequency pairs. For 1-2 kHz sweeps without harmonics, a 50% startle response rate occurred at mRLs of 133 dB re 1 μPa; for 1-2 kHz sweeps with strong harmonics at 99 dB re 1 μPa; for 6-7 kHz sweeps without harmonics at 101 dB re 1 μPa. Low-frequency (1-2 kHz) active naval sonar systems without harmonics can therefore operate at higher source levels than mid-frequency (6-7 kHz) active sonar systems without harmonics, with similar startle effects on porpoises.     

The use of an air bubble curtain to reduce the received sound levels for harbor porpoises (Phocoena phocoena)

In December 2005 construction work was started to replace a harbor wall in Kerteminde harbor, Denmark. A total of 175 wooden piles were piled into the ground at the waters edge over a period of 3 months. During the same period three harbor porpoises were housed in a marine mammal facility on the opposite side of the harbor. All animals showed strong avoidance reactions after the start of the piling activities. As a measure to reduce the sound exposure for the animals an air bubble curtain was constructed and operated in a direct path between the piling site and the opening of the animals' semi-natural pool. The sound attenuation effect achieved with this system was determined by quantitative comparison of pile driving impulses simultaneously measured in front of and behind the active air bubble curtain. Mean levels of sound attenuation over a sequence of 95 consecutive pile strikes were 14 dB (standard deviation (s.d.) 3.4 dB) for peak to peak values and 13 dB (s.d. 2.5 dB) for SEL values. As soon as the air bubble curtain was installed and operated, no further avoidance reactions of the animals to the piling activities were apparent.     

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.     

Origin of the double- and multi-pulse structure of echolocation signals in Yangtze finless porpoise (Neophocaena phocaenoides asiaeorientialis)

The signals of dolphins and porpoises often exhibit a multi-pulse structure. Here, echolocation signal recordings were made from four geometrically distinct positions of seven Yangtze finless porpoises temporarily housed in a relatively small, enclosed area. Some clicks demonstrated double-pulse, and others multi-pulse, structure. The interpulse intervals between the first and second pulse of the double- and multi-pulse clicks were significantly different among data from the four different positions (p < 0.01, one-way ANOVA). These results indicate that the interpulse interval and structure of the double- and multi-pulse echolocation signals depend on the hydrophone geometry of the animal, and that the double- and multi-pulse structure of echolocation signals in Yangtze finless porpoise is not caused by the phonating porpoise itself, but by the multipath propagation of the signal. Time delays in the 180 degrees phase-shifted surface reflection pulse and the nonphase-shifted bottom reflection pulse of the multi-pulse structures, relative to the direct signal, can be used to calculate the distance to a phonating animal.     

The ontogeny of echolocation in a Yangtze finless porpoise (Neophocaena phocaenoides asiaeorientalis)

Acoustic and concurrent behavioral data from one neonatal male Yangtze finless porpoise (Neophocaena phocaenoides asiaeorientalis) in captivity were presented. The calf click train was first recorded at 22 days postnatal, and the frequency of hydrophone-exploration behavior with head scanning motions in conjunction with emissions of click trains by the calf increased gradually with age. The echolocation clicks in the first recorded click train were indistinguishable from those of adults. Calf echolocation trains were found to decrease in maximum click-repetition rate, duration, and number of clicks per train with age while the minimum click-repetition rate remained more consistent.     

中华白海豚和东亚窄脊江豚回声定位信号分析与比较

为了增进珍稀齿鲸物种的了解和保护,对中华白海豚(Sousa chinensis)和东亚窄脊江豚(Neophocaena asiaeorientalis sunmeri)的回声定位信号特性进行了分析和比较。通过船只观测与声学监听的方式对厦门海域中华白海豚和东亚窄脊江豚的回声定位信号进行了调查,并对其声学参数进行了统计和对比。结果表明:中华白海豚回声定位信号具有持续时间短、高频、宽频带的特征;相较而言,东亚窄脊江豚则呈现时长较长、高频、窄频带的特点;声学参数统计上,两者在持续时间、中心频率、-10 dB带宽等指标上具有明显的差异性。     

Echolocation signals of the free-ranging Yangtze finless porpoise (Neophocaena phocaenoides asiaeorientialis)

This paper describes the high-frequency echolocation signals from free-ranging Yangtze finless porpoise in the Tian-e-zhou Baiji National Natural Reserve in Hubei Province, China. Signal analysis showed that the Yangtze finless porpoise clicks are typical high-frequency narrow-band (relative width of the frequency spectrum Q = 6.6 +/- 1.56, N = 548) ultrasonic pulses. The peak frequencies of the typical clicks range from 87 to 145 kHz with an average of 125 +/- 6.92 kHz. The durations range from 30 to 122 micros with an average of 68 +/- 14.12, as. The characteristics of the signals are similar to those of other members of the Phocoenidae as well as the distantly related delphinids, Cephalorhynchus spp. Comparison of these signals to those of the baiji (Lipotes vexillifer), who occupies habitat similar to that of the Yangtze finless porpoise, showed that the peak frequencies of clicks produced by the Yangtze finless porpoise are remarkably higher than those produced by the baiji. Difference in peak frequency between the two species is probably linked to the different size of preferred prey fish. Clear double-pulse and multi-pulse reverberation structures of clicks are noticed, and there is no indication of any low-frequency (< 70 kHz) components during the recording period.