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

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

Killer whales (Orcinus orca) produce ultrasonic whistles

This study reports that killer whales, the largest dolphin, produce whistles with the highest fundamental frequencies ever reported in a delphinid. Using wide-band acoustic sampling from both animal-attached (Dtag) and remotely deployed hydrophone arrays, ultrasonic whistles were detected in three Northeast Atlantic populations but not in two Northeast Pacific populations. These results are inconsistent with analyses suggesting a correlation of maximum frequency of whistles with body size in delphinids, indicate substantial intraspecific variation in whistle production in killer whales, and highlight the importance of appropriate acoustic sampling techniques when conducting comparative analyses of sound repertoires.     

Echolocation clicks from killer whales (Orcinus orca) feeding on herring (Clupea harengus)

Echolocation clicks from Norwegian killer whales feeding on herring schools were recorded using a four-hydrophone array. The clicks had broadband bimodal frequency spectra with low and high frequency peaks at 24 and 108 kHz, respectively. The -10 dB bandwidth was 35 kHz. The average source level varied from 173 to 202 dB re 1 microPa (peak-to-peak) at 1 m. This is considerably lower than source levels described for Canadian killer whales foraging on salmon. It is suggested that biosonar clicks of Norwegian killer whales are adapted for localization of prey with high target strength and acute hearing abilities.     

Intra- and intergroup vocal behavior in resident killer whales, Orcinus orca

Vocal communication within and between groups of individuals has been described extensively in birds and terrestrial mammals, however, little is known about how cetaceans utilize their sounds in their natural environment. Resident killer whales, Orcinus orca, live in highly stable matrilines and exhibit group-specific vocal dialects. Single call types cannot exclusively be associated with particular behaviors and calls are thought to function in group identification and intragroup communication. In the present study call usage of three closely related matrilines of the Northern resident community was compared in various intra- and intergroup contexts. In two out of the three matrilines significant changes in vocal behavior depending both on the presence and identity of accompanying whales were found. Most evidently, family-specific call subtypes, as well as aberrant and variable calls, were emitted at higher rates, whereas "low arousal" call types were used less in the presence of matrilines from different pods, subclans, or clans. Ways in which the observed changes may function both in intra- and intergroup communication.     

Sounds produced by Norwegian killer whales, Orcinus orca, during capture

To date very little is still known about the acoustic behavior of Norwegian killer whales, in particular that of individual whales. In this study a unique opportunity was presented to document the sounds produced by five captured killer whales in the Vestfjord area, northern Norway. Individuals produced 14 discrete and 7 compound calls. Two call types were used both by individuals 16178 and 23365 suggesting that they may belong to the same pod. Comparisons with calls documented in Strager (1993) showed that none of the call types used by the captured individuals were present. The lack of these calls in the available literature suggests that call variability within individuals is likely to be large. This short note adds to our knowledge of the vocal repertoire of this population and demonstrates the need for further studies to provide behavioural context to these sounds.     

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

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

Quantifying complex patterns of bioacoustic variation: use of a neural network to compare killer whale (Orcinus orca) dialects

A quantitative measure of acoustic similarity is crucial to any study comparing vocalizations of different species, social groups, or individuals. The goal of this study was to develop a method of extracting frequency contours from recordings of pulsed vocalizations and to test a nonlinear index of acoustic similarity based on the error of an artificial neural network at classifying them. Since the performance of neural networks depends on the amount of consistent variation in the training data, this technique can be used to assess such variation from samples of acoustic signals. The frequency contour extraction and the neural network index were tested on samples of one call type shared by nine social groups of killer whales. For comparison, call similarity was judged by three human subjects in pairwise classification tasks. The results showed a significant correlation between the neural network index and the similarity ratings by the subjects. Both measures of acoustic similarity were significantly correlated with the groups' association patterns, indicating that both methods of quantifying acoustic similarity are biologically meaningful. An index based on neural network analysis therefore represents an objective and repeatable means of measuring acoustic similarity, and allows comparison of results across studies, species and time.     

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

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

Acoustic detections of singing humpback whales (Megaptera novaeangliae) in the eastern North Pacific during their northbound migration.

Numerous (84) acoustic detections of singing humpback whales were made during a spring (08 March-09 June 1997) research cruise to study sperm whales in the central and eastern North Pacific. Over 15,000 km of track-line was surveyed acoustically using a towed hydrophone array. Additionally, 83 sonobuoys were deployed throughout the study area. Detection rates were greatest in late March, near the Hawaiian Islands, and in early April, northeast of the islands. Only one detection was made after April. Detection rates for sonobuoys were unequal in three equally divided longitudinal regions of the study area. Two high density clusters of detections occurred approximately 1200-2000 km northeast of the Hawaiian Islands and were attributed to a large aggregation of migrating animals. The distribution of these detections corroborates findings of previous studies. It is possible that these animals were maintaining acoustic contact during migration. Two unexpected clusters of singing whales were detected approximately 900 to 1000 km west of central and southern California. The location of these detections may indicate a previously undocumented migration route between an offshore breeding area, such as the Revillagigedo Islands, Mexico, and possible feeding areas in the western North Pacific or Bering Sea.     

Comparison of echolocation clicks from geographically sympatric killer whales and long-finned pilot whales (L)

The source characteristics of biosonar signals from sympatric killer whales and long-finned pilot whales in a Norwegian fjord were compared. A total of 137 pilot whale and more than 2000 killer whale echolocation clicks were recorded using a linear four-hydrophone array. Of these, 20 pilot whale clicks and 28 killer whale clicks were categorized as being recorded on-axis. The clicks of pilot whales had a mean apparent source level of 196 dB re 1 μPa pp and those of killer whales 203 dB re 1 μPa pp. The duration of pilot whale clicks was significantly shorter (23 μs, S.E.=1.3) and the centroid frequency significantly higher (55 kHz, S.E.=2.1) than killer whale clicks (duration: 41 μs, S.E.=2.6; centroid frequency: 32 kHz, S.E.=1.5). The rate of increase in the accumulated energy as a function of time also differed between clicks from the two species. The differences in duration, frequency, and energy distribution may have a potential to allow for the distinction between pilot and killer whale clicks when using automated detection routines for acoustic monitoring.     

Source of the North Pacific "boing" sound attributed to minke whales

During a recent cetacean survey of the U.S. waters surrounding the Hawaiian Islands, the probable source of the mysterious "boing" sound of the North Pacific Ocean was identified as a minke whale, Balaenoptera acutorostrata. Examination of boing vocalizations from three research surveys confirms previous work that identified two distinct boing vocalization types in the North Pacific. The eastern boing (n = 22) has a pulse repetition rate of 92 s(-1) and a duration of 3.6 s and was found only east of 138 degrees W. The central boing (n = 106) has a pulse repetition rate of 115 s(-1) and a duration of approximately 2.6 s and was found only west of 135 degrees W. Central boing vocalizations produced by a single source (n = 84) indicate that variation in repetition rate and duration of the calls of the individual were not significantly different than the variation among individuals of the same boing type. Despite a slight latitudinal overlap in the vocalizations, pulse repetition rates of the eastern and central boings were distinct.     

The 20-Hz signals of finback whales (Balaenoptera physalus)

The 20-Hz signals of finback whales (Balaenoptera physalus) were analyzed from more than 25 years of recordings at a variety of geographic locations on near-surface hydrophones close to whales and on deep hydrophone systems. These signals were composed of 1-s pulses of sinusoidal waveform with downward sweeping frequency from approximately 23 to 18 Hz at variable source levels up to 186 dB (re: 1 microPa at 1 m), usually with slightly lower levels for the pulses at the beginning and end of sequences. These "20-Hz" pulses were produced in signal bouts (separated by more than 2 h) lasting as long as 32.5 h. Bouts were composed of regularly repeated pulses at intervals of 7-26 s (typically), either at one nominal pulse rate or at two alternating (doublet) pulse intervals. Signal bouts were interrupted by rests of 1-20 min at roughly 15-min intervals and by irregular gaps lasting between 20 and 120 min. The distribution of these signals throughout the year and their temporal sequence were analyzed from the continuous drum records of the Bermuda SOFAR Station. Signal bouts occurred during winter, sometimes beginning in September and ending in May. The sound sequences were never exactly replicated. Direct association of the bouts with the reproductive season for this species points to the 20-Hz signals as possible reproductive displays by finback whales.     

Behavioral responses of humpback whales (Megaptera novaeangliae) to full-scale ATOC signals

Loud (195 dB re 1 microPa at 1 m) 75-Hz signals were broadcast with an ATOC projector to measure ocean temperature. Respiratory and movement behaviors of humpback whales off North Kauai, Hawaii, were examined for potential changes in response to these transmissions and to vessels. Few vessel effects were observed, but there were fewer vessels operating during this study than in previous years. No overt responses to ATOC were observed for received levels of 98-109 dB re 1 microPa. An analysis of covariance, using the no-sound behavioral rate as a covariate to control for interpod variation, found that the distance and time between successive surfacings of humpbacks increased slightly with an increase in estimated received ATOC sound level. These responses are very similar to those observed in response to scaled-amplitude playbacks of ATOC signals [Frankel and Clark, Can. J. Zool. 76, 521-535 (1998)]. These similar results were obtained with different sound projectors, in different years and locations, and at different ranges creating a different sound field. The repeatability of the findings for these two different studies indicates that these effects, while small, are robust. This suggests that at least for the ATOC signal, the received sound level is a good predictor of response.     

Perception of amplitude and frequency modulated signals (mixed modulation)

This article discusses the detection of mixed modulation, i.e., simultaneous amplitude and frequency modulation (MM). The investigations have incorporated both a sine wave modulating signal and an irregular modulating signal, a very narrow noise band, of a specified center frequency. The results revealed that for a sinusoidal low-frequency modulating signal, amplitude and frequency changes that were separately subthreshold could be detected by listeners in mixed modulation. This indicates summation of sensations caused by simultaneous AM and FM modulation. This effect was not observed in the case of the irregular modulating signal. A hypothesis is advanced that the perception of modulated signals is governed by two mechanisms, viz., temporal and spectral. The operation of the two mechanisms depends mainly on the modulating frequency. The type of modulation does not play any significant role in this case.     

Sound production by North Atlantic right whales (Eubalaena glacialis) in surface active groups

The surface active group (SAG) is the most commonly observed surface social behavior of North Atlantic right whales. Recordings were made from 52 SAGs in the Bay of Fundy, Canada between July and September, from 1999 to 2002. The call types recorded from these groups were similar to those described previously for Southern right whales (Eubalaena australis), with six major call types being termed scream, gunshot, blow, upcall, warble, and downcall. The percentage of total calls of each call type depended on the group size and composition. The most common call type recorded was the scream call. The scream calls were produced by the focal female in a SAG. Production of other sound types can be attributed to whales other than the focal female, with gunshot and upcalls produced by males, and warble calls produced by female calves. The source levels for these sounds range from 137 to 162 dB rms re 1 ,tPa-m for tonal calls and 174 to 192 dB rms for broadband gunshot sounds.     

Divergence of a stereotyped call in northern resident killer whales

Northern resident killer whale pods (Orcinus orca) have distinctive stereotyped pulsed call repertoires that can be used to distinguish groups acoustically. Repertoires are generally stable, with the same call types comprising the repertoire of a given pod over a period of years to decades. Previous studies have shown that some discrete pulsed calls can be subdivided into variants or subtypes. This study suggests that new stereotyped calls may result from the gradual modification of existing call types through subtypes. Vocalizations of individuals and small groups of killer whales were collected using a bottom-mounted hydrophone array in Johnstone Strait, British Columbia in 2006 and 2007. Discriminant analysis of slope variations of a predominant call type, N4, revealed the presence of four distinct call subtypes. Similar to previous studies, there was a divergence of the N4 call between members of different matrilines of the same pod. However, this study reveals that individual killer whales produced multiple subtypes of the N4 call, indicating that divergence in the N4 call is not the result of individual differences, but rather may indicate the gradual evolution of a new stereotyped call.     

Receiver structure and estimation of the modulation index for tamed frequency modulated (TFM) signals

Tamed frequency modulation (TFM) is a spectrally efficient constant amplitude continuous phase modulation (CPM) scheme which can be simply realized by using a frequency modulator (FM). In the implementation the modulation index of TFM is calibrated to have a nominal value of 0.5, but due to temperature variations it can drift causing time varying phase jitter. In this paper we present novel algorithms and performance results to measure and control the modulation index in a coherent receiver based on the joint reduced state sequence detector (RSSD) and per-survivor processing (PSP) carrier phase estimation. The modulation index estimator measures phase transitions in the receiver and derives estimates by comparing the result to the coding rule of the TFM signal. The estimator has acquisition and tracking ability, and the current estimate can be used to replace the nominal index value. Our simulation results show that the proposed coherent receiver with the novel modulation index estimator has less than 1 dB performance degradation compared to around 4.5 dB exploiting only the PSP carrier phase estimation.     

High-pitched tonal signals of beluga whales (Delphinapterus leucas) in a summer assemblage off Solovetskii Island in the White Sea

High-frequency whistles of beluga whales are analyzed. The signals are recorded in a belgua summer assemblage off Solovetskii Island in the White Sea. The high-frequency whistles are narrowband signals with a continuous waveform and a fundamental frequency above 5 kHz. On the average, they make up 7.7% of the total vocal production of the animals. Based on the shape of the fundamental frequency contour and its time-frequency parameters, the high-frequency whistles are classed into 12 types. The HF whistles have a mean fundamental frequency of 9.7 kHz, an average bandwidth of 3.3 kHz, and an average duration of 1.0 s. The number of inflection points per signal ranges from 0 to 56 with a mean of 2.3. The predominant types are flat (50%), rising (23%), and wavy (7%) high-frequency whistles. Presumably, beluga whales can use some of the whistle types for short-range communication and other types for long-range communication. Published in Russian in Akusticheskiĭ Zhurnal, 2006, Vol. 52, No. 2, pp. 156–164. The article was translated by the authors.     

Change in echolocation signals with hearing loss in a false killer whale (Pseudorca crassidens)

The echolocation signals of a false killer whale (Pseudorca crassidens) were collected during a wall thickness discrimination task and compared to clicks recorded during an identical experiment in 1992. During the sixteen year time period, the subject demonstrated a loss of high frequency hearing of about 70 kHz. Clicks between the two experiments were compared to investigate the effect of hearing loss on echolocation signals. There was a significant reduction in the peak frequency, center frequency and source level of clicks between the two time periods. Additionally, the subject currently produces more signals with low frequency peaks and fewer signals with high frequency peaks than she did in 1992. These results indicate the subject changed its echolocation signals to match its range of best hearing.