From echolocation clicks to animal density--acoustic sampling of harbor porpoises with static dataloggers

Monitoring abundance and population trends of small odontocetes is notoriously difficult and labor intensive. There is a need to develop alternative methods to the traditional visual line transect surveys, especially for low density areas. Here, the prospect of obtaining robust density estimates for porpoises by passive acoustic monitoring (PAM) is demonstrated by combining rigorous application of methods adapted from distance sampling to PAM. Acoustic dataloggers (T-PODs) were deployed in an area where harbor porpoises concurrently were tracked visually. Probability of detection was estimated in a mark-recapture approach, where a visual sighting constituted a "mark" and a simultaneous acoustic detection a "recapture." As a distance could be assigned to each visual observation, a detection function was estimated. Effective detection radius of T-PODs ranged from 22 to 104 m depending on T-POD type, T-POD sensitivity, train classification settings, and snapshot duration. The T-POD density estimates corresponded to the visual densities derived concurrently for the same period. With more dataloggers, located according to a systematic design, density estimates would be obtainable for a larger area. This provides a method suitable for monitoring in areas with densities too low for visual surveys to be practically feasible, e.g., the endangered harbor porpoise population in the Baltic.     

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.     

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.     

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.     

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 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.     

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.     

Pitch strength of regular-interval click trains with different length "runs" of regular intervals

Click trains were generated with first- and second-order statistics following Kaernbach and Demany [J. Acoust. Soc. Am. 104, 2298-2306 (1998)]. First-order intervals are between successive clicks, while second-order intervals are those between every other click. Click trains were generated with a repeating alternation of fixed and random intervals which produce a pitch at the reciprocal of the duration of the fixed interval. The intervals were then randomly shuffled and compared to the unshuffled, alternating click trains in pitch-strength comparison experiments. In almost all comparisons for the first-order interval stimuli, the shuffled-interval click trains had a stronger pitch strength than the unshuffled-interval click trains. The shuffled-interval click trains only produced stronger pitches for second-order interval stimuli when the click trains were unfiltered. Several experimental conditions and an analysis of runs of regular and random intervals in these click trains suggest that the auditory system is sensitive to runs of regular intervals in a stimulus that contains a mix of regular and random intervals. These results indicate that fine-structure regularity plays a more important role in pitch perception than randomness, and that the long-term autocorrelation function or spectra of these click trains are not good predictors of pitch strength.     

Classification of behavior using vocalizations of Pacific white-sided dolphins (Lagenorhynchus obliquidens)

Surface behavior and concurrent underwater vocalizations were recorded for Pacific white-sided dolphins in the Southern California Bight (SCB) over multiple field seasons spanning 3 years. Clicks, click trains, and pulsed calls were counted and classified based on acoustic measurements, leading to the identification of 19 key call features used for analysis. Kruskal-Wallis tests indicated that call features differ significantly across behavioral categories. Previous work had discovered two distinctive click Types (A and B), which may correspond to known subpopulations of Pacific white-side dolphins in the Southern California Bight; this study revealed that animals producing these different click types also differ in both their behavior and vocalization patterns. Click Type A groups were predominantly observed slow traveling and milling, with little daytime foraging, while click Type B groups were observed traveling and foraging. These behavioral differences may be characteristic of niche partitioning by overlapping populations; coupled with differences in vocalization patterns, they may signify that these subpopulations are cryptic species. Finally, random forest decision trees were used to classify behavior based on vocalization data, with rates of correct classification up to 86%, demonstrating the potential for the use of vocalization patterns to predict behavior.     

Exploring the temporal mechanism involved in the pitch of unresolved harmonics

This paper continues a line of research initiated by Kaernbach and Demany [J. Acoust. Soc. Am. 104, 2298-2306 (1998)], who employed filtered click sequences to explore the temporal mechanism involved in the pitch of unresolved harmonics. In a first experiment, the just noticeable difference (jnd) for the fundamental frequency (F0) of high-pass filtered and low-pass masked click trains was measured, with F0 (100 to 250 Hz) and the cut frequency (0.5 to 6 kHz) being varied orthogonally. The data confirm the result of Houtsma and Smurzynski [J. Acoust. Soc. Am. 87, 304-310 (1990)] that a pitch mechanism working on the temporal structure of the signal is responsible for analyzing frequencies higher than ten times the fundamental. Using high-pass filtered click trains, however, the jnd for the temporal analysis is at 1.2% as compared to 2%-3% found in studies using band-pass filtered stimuli. Two further experiments provide evidence that the pitch of this stimulus can convey musical information. A fourth experiment replicates the finding of Kaernbach and Demany on first- and second-order regularities with a cut frequency of 2 kHz and extends the paradigm to binaural aperiodic click sequences. The result suggests that listeners can detect first-order temporal regularities in monaural click streams as well as in binaurally fused click streams.     

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.     

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.     

3D tracking of foraging belugas from their clicks: Experiment from a coastal hydrophone array

A simple passive acoustic monitoring (PAM) setup was used to localize and track beluga whales underwater in three dimensions (3D) in a fjord. In June 2009, beluga clicks were recorded from a cabled hydrophone array in a regularly frequented habitat in Eastern Canada. Beluga click energy was concentrated in the 30-50 kHz frequency band. The click trains detected on several hydrophones were localized from their time difference of arrivals. Cluster analysis linked localizations into tracks based on criteria of spatial and temporal proximity. At close ranges from the array, the localized click-train series allowed three-dimensional tracking of a beluga during its dive. Clicks within a train spanned a large range of durations, inter-click intervals, source levels and bandwidths. Buzzes sometimes terminated the trains. Repeated click packets were frequent. All click characteristics are consistent with oblique observations from the beam axis, and ordered variation of the source pattern during a train, likely resulting from a scan of angles from the beam axis, was observed before click trains indicated focusing of the echolocation clicks in one direction. The click-train series is interpreted as echolocation chasing for preys during a foraging dive. Results show that a simple PAM system can be configured to passively and effectively 3D track wild belugas and small odontocetes in their regularly frequented habitat.     

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。      

Automatic detection of bioacoustics impulses based on kurtosis under weak signal to noise ratio

Passive acoustic monitoring (PAM) of marine mammal vocalizations has been efficiently used in a wide set of applications ranging from marine wildlife surveys to risk mitigation of military sonar emissions. The primary use of PAM is for detecting bioemissions, a good proportion of which are impulse sounds or clicks. A click detection algorithm based on kurtosis estimation is proposed as a general automatic click detector. The detector works under the assumption that click trains are embedded in stochastic but Gaussian noise. Under this assumption, kurtosis is used as a statistical test for detection. The algorithm explores acoustic sequences with the optimal frequency bandwidth for focusing on impulse sounds. The detector is successfully applied to field observations, and operates under weak signal to noise ratios and in presence of stochastic background noise. The algorithm adapts to varying click center frequency. Kurtosis appears as a promising approach to detect click trains, alone or in combination with other clicks detector, and to isolate individual clicks.     

Echolocation range of captive and free-ranging baiji (Lipotes vexillifer), finless porpoise (Neophocaena phocaenoides), and bottlenose dolphin (Tursiops truncatus).

The interclick intervals of captive dolphins are known to be longer than the two-way transit time between the dolphin and a target. In the present study, the interclick intervals of free-ranging baiji, finless porpoises, and bottlenose dolphins in the wild and in captivity were compared. The click intervals in open waters ranged up to 100-200 ms, whereas the click intervals in captivity were in the order of 4-28 ms. Echolocation of free-ranging dolphins appears to adapt to various distance in navigation or ranging, sometimes up to 140 m. Additionally, the difference of waveform characteristics of clicks between species was recognized in the frequency of maximum energy and the click duration.     

Dual mechanisms in the perceptual processing of click train temporal regularity

Two experiments measured human sensitivity to temporal jitter in 25-click trains with inter-click intervals (ICIs) between 5 and 100 ms. In a naturalistic experiment using wideband clicks, jitter thresholds were a nonmonotonic function of ICI, peaking for ICIs near 40-60 ms. In a subsequent experiment, clicks were high-passed and presented against a low-frequency noise masker. Jitter threshold vs ICI functions lost the positive slope over short ICIs but retained the negative slope at long ICIs. The same behavior was seen in click rate discrimination tasks. Different processes mediate regularity analysis for click trains with ICIs above and below 40-60 ms.     

Masking-level differences for trains of clicks

Masking-level differences (MLDs) were measured for trains of 2000-Hz bandpass clicks as a function of the interclick interval (ICI) and the number of clicks in the train. The magnitude of the MLD grew as the number of clicks in the train was increased from 1 to 32. While the MLDs tended to be larger at longer ICIs, the effect was mediated by changes in detectability in the homophasic conditions. For click trains consisting of 4-32 clicks, the improvement in detectability in the antiphasic conditions with increases in the number of clicks appears to be the result of integration of acoustic power, as is the case for the homophasic conditions. The absence of MLDs for short trains of high-frequency transients remains quite puzzling, since large MLDs are found with single, low-frequency transients.     

The Applications of Order Reduction Methods in Nonlinear Dynamic Systems

Two different order reduction methods of the deterministic and stochastic systems are discussed in this paper. First, the transient proper orthogonal decomposition (T-POD) method is introduced based on the high-dimensional nonlinear dynamic system. The optimal order reduction conditions of the T-POD method are provided by analyzing the rotor-bearing system with pedestal looseness fault at both ends. The efficiency of the T-POD method is verified via comparing with the results of the original system. Second, the polynomial dimensional decomposition (PDD) method is applied to the 2 DOFs spring system considering the uncertain stiffness to study the amplitude-frequency response. The numerical results obtained by the PDD method agree well with the Monte Carlo simulation (MCS) method. The results of the PDD method can approximate to MCS better with the increasing of the polynomial order. Meanwhile, the Uniform-Legendre polynomials can eliminate perturbation of the PDD method to a certain extent via comparing it with the Gaussian-Hermite polynomials.     

Description of sounds recorded from Longman's beaked whale, Indopacetus pacificus

Sounds from Longman's beaked whale, Indopacetus pacificus, were recorded during shipboard surveys of cetaceans surrounding the Hawaiian Islands archipelago; this represents the first known recording of this species. Sounds included echolocation clicks and burst pulses. Echolocation clicks were grouped into three categories, a 15 kHz click (n?=?106), a 25 kHz click (n?=?136), and a 25 kHz pulse with a frequency-modulated upsweep (n?=?70). The 15 and 25 kHz clicks were relatively short (181 and 144 ms, respectively); the longer 25 kHz upswept pulse was 288 ms. Burst pulses were long (0.5 s) click trains with approximately 240 clicks/s.