Source levels of clicks from free-ranging white-beaked dolphins (Lagenorhynchus albirostris Gray 1846) recorded in Icelandic waters

This study reports the source levels of clicks recorded from free-ranging white-beaked dolphins (Lagenorhynchus albirostris Gray 1846). A four-hydrophone array was used to obtain sound recordings. The hydrophone signals were digitized on-line and stored in a portable computer. An underwater video camera was used to visualize dolphins to help identify on-axis recordings. The range to a dolphin was calculated from differences in arrival times of clicks at the four hydrophones, allowing for calculations of source levels. Source levels in a single click train varied from 194 to 211 dB peak-to-peak (p-p) re: 1 microPa. The source levels varied linearly with the log of range. The maximum source levels recorded were 219 dB (p-p) re: 1 microPa.     

Echolocation transmitting beam of the Atlantic bottlenose dolphin

The transmitting beam patterns of echolocation signals emitted by an Atlantic bottlenose dolphin Tursiops truncatus were measured in the vertical and horizontal planes with an array of seven hydrophones. Particular emphasis was placed on accurately verifying the animal's position on a bite-plate/tail-rest stationing device using underwater video monitoring equipment. The major axis of the vertical beam was directed at an angle of 5 degrees above the plane defined by the animal's lips. This angle was 15 degrees lower than previously measured. The vertical beam measurements indicate that the major axis of the transmitting beam is aligned with the major axis of the receiving beam. The horizontal beam was directed forward. The directivity index of 26.5 dB calculated from the beam pattern measured in both planes agreed well with previous calculation of 25.4 dB.     

The biosonar field around an Atlantic bottlenose dolphin (Tursiops truncatus)

The use of remote autonomous passive acoustic recorders (PAR) to determine the distribution of dolphins at a given locations has become very popular. Some investigators are using echolocation clicks to gather information on the presence of dolphins and to identify species. However, in all of these cases, the PAR probably recorded mainly off-axis clicks, even some from behind the animals. Yet there is a very poor understanding of the beam pattern and the click waveform and spectrum from different azimuths around the animal's body. The beam pattern completely around an echo locating dolphin was measured at 16 different but equally spaced angles in the horizontal plane using an 8-hydrophone array in sequence. Eight channels of data were digitized simultaneously at a sampling rate of 500 kHz. The resulting beam patterns in both planes showed a continuous drop off in sound pressure with azimuth around the animal and reached levels below -50 dB relative to the signal recorded on the beam axis. The signals began to break up into two components at angles greater than ± 45° in the horizontal plane. The center frequency dropped off from its maximum at 0° in a non-uniform matter.     

Directional properties of bottlenose dolphin (Tursiops truncatus) clicks, burst-pulse, and whistle sounds

The directional properties of bottlenose dolphin clicks, burst-pulse, and whistle signals were measured using a five element array, at horizontal angles of 0°, 45°, 90°, 135°, and 180° relative to a dolphin stationed on an underwater biteplate. Clicks and burst-pulse signals were highly directional with directivity indices of ~11 dB for both signal types. Higher frequencies and higher amplitudes dominated the forward, on-axis sound field. A similar result was found with whistles, where higher frequency harmonics had greater directivity indices than lower frequency harmonics. The results suggest the directional properties of these signals not only provide enhanced information to the sound producer (as in echolocation) but can provide valuable information to conspecific listeners during group coordination and socialization.     

Echolocation signals of wild dolphins

Most of our understanding of dolphin echolocation has come from studies of captive dolphins performing various echolocation tasks. Recently, measurements of echolocation signals in the wild have expanded our understanding of the characteristics of these signals in a natural setting. Measuring undistorted dolphin echolocation signals with free swimming dolphins in the field can be a challenging task. A four hydrophone array arranged in a symmetrical star pattern was used to measure the echolocation signals of four species of dolphins in the wild. Echolocation signals of the following dolphins have been measured with the symmetrical star array: white-beaked dolphins in Iceland, Atlantic spotted dolphins in the Bahamas, killer whales in British Columbia, and dusky dolphins in New Zealand. There are many common features in the echolocation signals of the different species. Most of the signals had spectra that were bimodal: two peaks, one at low frequencies and another about an octave higher in frequency. The source level of the sonar transmission varies as a function of 20logR, suggesting a form of time-varying gain but on the transmitting end of the sonar process rather than the receiving end. The results of the field work call into question the issue of whether the signals used by captive dolphins may be shaped by the task they are required to perform rather than what they would do more naturally.     

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.     

The monopulsed nature of sperm whale clicks

Traditionally, sperm whale clicks have been described as multipulsed, long duration, nondirectional signals of moderate intensity and with a spectrum peaking below 10 kHz. Such properties are counterindicative of a sonar function, and quite different from the properties of dolphin sonar clicks. Here, data are presented suggesting that the traditional view of sperm whale clicks is incomplete and derived from off-axis recordings of a highly directional source. A limited number of assumed on-axis clicks were recorded and found to be essentially monopulsed clicks, with durations of 100 micros, with a composite directionality index of 27 dB, with source levels up to 236 dB re: 1 microPa (rms), and with centroid frequencies of 15 kHz. Such clicks meet the requirements for long-range biosonar purposes. Data were obtained with a large-aperture, GPS-synchronized array in July 2000 in the Bleik Canyon off Vester?len, Norway (69 degrees 28' N, 15 degrees 40' E). A total of 14 h of sound recordings was collected from five to ten independent, simultaneously operating recording units. The sound levels measured make sperm whale clicks by far the loudest of sounds recorded from any biological source. On-axis click properties support previous work proposing the nose of sperm whales to operate as a generator of sound.     

Source parameters of echolocation clicks from wild bottlenose dolphins (Tursiops aduncus and Tursiops truncatus)

The Indian Ocean and Atlantic bottlenose dolphins (Tursiops aduncus and Tursiops truncatus) are among the best studied echolocating toothed whales. However, almost all echolocation studies on bottlenose dolphins have been made with captive animals, and the echolocation signals of free-ranging animals have not been quantified. Here, biosonar source parameters from wild T. aduncus and T. truncatus were measured with linear three- and four-hydrophone arrays in four geographic locations. The two species had similar source parameters, with source levels of 177-228 dB re 1 μPa peak to peak, click durations of 8-72 μs, centroid frequencies of 33-109 kHz and rms bandwidths between 23 and 54 kHz. T. aduncus clicks had a higher frequency emphasis than T. truncatus. The transmission directionality index was up to 3 dB higher for T. aduncus (29 dB) as compared to T. truncatus (26 dB). The high directionality of T. aduncus does not appear to be only a physical consequence of a higher frequency emphasis in clicks, but may also be caused by differences in the internal properties of the sound production system.     

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.     

Classification of Risso's and Pacific white-sided dolphins using spectral properties of echolocation clicks

The spectral and temporal properties of echolocation clicks and the use of clicks for species classification are investigated for five species of free-ranging dolphins found offshore of southern California: short-beaked common (Delphinus delphis), long-beaked common (D. capensis), Risso's (Grampus griseus), Pacific white-sided (Lagenorhynchus obliquidens), and bottlenose (Tursiops truncatus) dolphins. Spectral properties are compared among the five species and unique spectral peak and notch patterns are described for two species. The spectral peak mean values from Pacific white-sided dolphin clicks are 22.2, 26.6, 33.7, and 37.3 kHz and from Risso's dolphins are 22.4, 25.5, 30.5, and 38.8 kHz. The spectral notch mean values from Pacific white-sided dolphin clicks are 19.0, 24.5, and 29.7 kHz and from Risso's dolphins are 19.6, 27.7, and 35.9 kHz. Analysis of variance analyses indicate that spectral peaks and notches within the frequency band 24-35 kHz are distinct between the two species and exhibit low variation within each species. Post hoc tests divide Pacific white-sided dolphin recordings into two distinct subsets containing different click types, which are hypothesized to represent the different populations that occur within the region. Bottlenose and common dolphin clicks do not show consistent patterns of spectral peaks or notches within the frequency band examined (1-100 kHz).     

Evidence of high-frequency acoustic emissions from the white-beaked dolphin (Lagenorhynchus albirostris)

Recordings of the signals from a school of white-beaked dolphins show that the frequency of their acoustic emissions extends to at least 305 kHz. These signals were detected by a sector scanning sonar used as a passive listening device of high bearing and time resolution. The records contain three types of signal, one of high intensity, one of a variable high repetition rate, and another showing a time-varying effect. Acoustic signals radiated by dolphins have been recorded and studied over a long period of time by many investigators. The purpose of this letter is to report evidence that acoustic emissions from white-beaked dolphins have significant energy at frequencies around 305 kHz, about one octave higher than previously observed. The observations discussed here were made aboard the fisheries research vessel CLIONE in the Wellbank flat area of the southern North Sea on 13 June 1970 between 1040 and 1110 h. When the dolphin signals were observed, the transmitter of the sector-scanning sonar in use was turned off, and the system was utilized as a passive listening device of high bearing and time resolution.     

Discrimination of amplitude-modulated synthetic echo trains by an echolocating bottlenose dolphin

Bottlenose dolphins (Tursiops truncatus) have an acute ability to use target echoes to judge attributes such as size, shape, and material composition. Most target recognition studies have focused on features associated with individual echoes as opposed to information conveyed across echo sequences (feature envelope of the multi-echo train). One feature of aspect-dependent targets is an amplitude modulation (AM) across the return echoes in the echo train created by relative movement of the target and dolphin. The current study examined whether dolphins could discriminate targets with different AM envelopes. "Electronic echoes" triggered by a dolphin's outgoing echolocation clicks were manipulated to create sinusoidal envelopes with varying AM rate and depth. Echo trains were equated for energy, requiring the dolphin to extract and retain information from multiple echoes in order to detect and report the presence of AM. The dolphin discriminated amplitude-modulated echo trains from those that were not modulated. AM depth thresholds were approximately 0.8 dB, similar to other published amplitude limens. Decreasing the rate of modulation from approximately 16 to 2 cycles per second did not affect the dolphin's AM depth sensitivity. The results support multiple-echo processing in bottlenose dolphin echolocation. This capability provides additional theoretical justification for exploring synthetic aperture sonar concepts in models of animal echolocation that potentially support theories postulating formation of images as an ultimate means for target identification.     

Echolocation in the Risso's dolphin,Grampus griseus

The Risso's dolphin (Grampus griseus) is an exclusively cephalopod-consuming delphinid with a distinctive vertical indentation along its forehead. To investigate whether or not the species echolocates, a female Risso's dolphin was trained to discriminate an aluminum cylinder from a nylon sphere (experiment 1) or an aluminum sphere (experiment 2) while wearing eyecups and free swimming in an open-water pen in Kaneohe Bay, Hawaii. The dolphin completed the task with little difficulty despite being blindfolded. Clicks emitted by the dolphin were acquired at average amplitudes of 192.6 dB re 1 microPa, with estimated sources levels up to 216 dB re 1 microPa-1 m. Clicks were acquired with peak frequencies as high as 104.7 kHz (Mf(p) = 47.9 kHz), center frequencies as high as 85.7 kHz (Mf(0) = 56.5 kHz), 3-dB bandwidths up to 94.1 kHz (M(BW) = 39.7 kHz), and root-mean-square bandwidths up to 32.8 kHz (M(RMS) = 23.3 kHz). Click durations were between 40 and 70 micros. The data establish that the Risso's dolphin echolocates, and that, aside from slightly lower amplitudes and frequencies, the clicks emitted by the dolphin were similar to those emitted by other echolocating odontocetes. The particular acoustic and behavioral findings in the study are discussed with respect to the possible direction of the sonar transmission beam of the species.     

Monaural and binaural hearing directivity in the bottlenose dolphin: evoked-potential study

Hearing thresholds as a function of sound-source azimuth were measured in bottlenose dolphins using an auditory evoked potential (AEP) technique. AEP recording from a region next to the ear allowed recording monaural responses. Thus, a monaural directivity diagram (a threshold-vs-azimuth function) was obtained. For comparison, binaural AEP components were recorded from the vertex to get standard binaural directivity diagrams. Both monaural and binaural diagrams were obtained at frequencies ranging from 8 to 128 kHz in quarter-octave steps. At all frequencies, the monaural diagram demonstrated asymmetry manifesting itself as: (1) lower thresholds at the ipsilateral azimuth as compared to the symmetrical contralateral azimuth and (2) ipsilateral shift of the lowest-threshold point. The directivity index increased with frequency: at the ipsilateral side it rose from 4.7 to 17.8 dB from 11.2 to 128 kHz, and from 10.5 to 15.6 dB at the contralateral side. The lowest-threshold azimuth shifted from 0 degrees at 90-128 kHz to 22.5 degrees at 8-11.2 kHz. The frequency-dependent variation of the lowest-threshold azimuth indicates the presence of two sound-receiving apertures at each head side: a high-frequency aperture with the axis directed frontally, and a low-frequency aperture with the axis directed laterally.     

Modeling the detection range of fish by echolocating bottlenose dolphins and harbor porpoises

The target strength as a function of aspect angle were measured for four species of fish using dolphin-like and porpoise-like echolocation signals. The polar diagram of target strength values measured from an energy flux density perspective showed considerably less fluctuation with azimuth than would a pure tone pulse. Using detection range data obtained from dolphin and porpoise echolocation experiments, the detection ranges for the Atlantic cod by echolocating dolphins and porpoises were calculated for three aspect angles of the cod. Maximum detection ranges occurred when the fish was broadside to the odontocete and minimum detection ranges occurred when the cod was in the tail aspect. Maximum and minimum detection ranges for the bottlenose dolphin in a noise-limited environment was calculated to be 93 and 70 m, respectively. In a quiet environment, maximum and minimum detection ranges for the bottlenose dolphin were calculated to be 173 and 107 m, respectively. The detection ranges for the harbor porpoise in a quiet environment were calculated to be between 15 and 27 m. The primary reason for the large differences in detection ranges between both species was attributed to the 36 dB higher source level of the bottlenose dolphin echolocation signals.     

Assessment of dolphin (Tursiops truncatus) auditory sensitivity and hearing loss using jawphones

Devices known as jawphones have previously been used to measure interaural time and intensity discrimination in dolphins. This study introduces their use for measuring hearing sensitivity in dolphins. Auditory thresholds were measured behaviorally against natural background noise for two bottlenose dolphins (Tursiops truncatus); a 14-year-old female and a 33-year-old male. Stimuli were delivered to each ear independently by placing jawphones directly over the pan bone of the dolphin's lower jaw, the assumed site of best reception. The shape of the female dolphin's auditory functions, including comparison measurements made in the free field, favorably matches that of the accepted standard audiogram for the species. Thresholds previously measured for the male dolphin at 26 years of age indicated a sensitivity difference between the ears of 2-3 dB between 4-10 kHz, which was considered unremarkable at the time. Thresholds for the male dolphin reported in this study suggest a high-frequency loss compared to the standard audiogram. Both of the male's ears have lost sensitivity to frequencies above 55 kHz and the right ear is 16-33 dB less sensitive than the left ear over the 10-40 kHz range, suggesting that males of the species may lose sensitivity as a function of age. The results of this study support the use of jawphones for the measurement of dolphin auditory sensitivity.     

Bio-inspired wideband sonar signals based on observations of the bottlenose dolphin (Tursiops truncatus)

This paper uses advanced time-frequency signal analysis techniques to generate new models for bio-inspired sonar signals. The inspiration comes from the analysis of bottlenose dolphin clicks. These pulses are very short duration, between 50 and 80 micros, but for certain examples we can delineate a double down-chirp structure using fractional Fourier methods. The majority of clicks have energy distributed between two main frequency bands with the higher frequencies delayed in time by 5-20 micros. Signal syntheses using a multiple chirp model based on these observations are able to reproduce much of the spectral variation seen in earlier studies on natural dolphin echolocation pulses. Six synthetic signals are generated and used to drive the dolphin based sonar (DBS) developed through the Biosonar Program office at the SPAWAR Systems Center, San Diego, CA. Analyses of the detailed echo structure for these pulses ensonifying two solid copper spherical targets indicate differences in discriminatory potential between the signals. It is suggested that target discrimination could be improved through the transmission of a signal packet in which the chirp structure is varied between pulses. Evidence that dolphins may use such a strategy themselves comes from observations of variations in the transmissions of dolphins carrying out target detection and identification tasks.     

Time-frequency analysis and modeling of the backscatter of categorized dolphin echolocation clicks for target discrimination

A set of dolphin echolocation clicks collected from an Atlantic bottlenose dolphin in Kaneohe Bay, Hawai'i from a previous experiment is examined in terms of their time and frequency characteristics. The center frequency and rms bandwidth are calculated for the clicks and these are clustered into four classes by using a model based on the Bayesian information criterion. The echo signatures are attained from a solid, elastic homogeneous sphere for each class of clicks from an acoustic scattering model. The results from the scattering model are compared to experimental values. The joint time-frequency content of the resulting echo signals is obtained by the reduced interference distribution (RID). The RIDs are plotted and examined for each signal class for four spherical targets of different material compositions. RID correlation values are obtained for a standard target versus comparison targets by using a time-frequency correlator. The results suggest that dolphins may discriminate by auditory inspection of the time-frequency information returned by the targets. The modification of the outgoing clicks and examination of time-frequency target information may be fundamental to a dolphin's ability to identify and discriminate targets.     

Discriminating features of echolocation clicks of melon-headed whales (Peponocephala electra), bottlenose dolphins (Tursiops truncatus), and Gray's spinner dolphins (Stenella longirostris longirostris)

Spectral parameters were used to discriminate between echolocation clicks produced by three dolphin species at Palmyra Atoll: melon-headed whales (Peponocephala electra), bottlenose dolphins (Tursiops truncatus) and Gray's spinner dolphins (Stenella longirostris longirostris). Single species acoustic behavior during daytime observations was recorded with a towed hydrophone array sampling at 192 and 480 kHz. Additionally, an autonomous, bottom moored High-frequency Acoustic Recording Package (HARP) collected acoustic data with a sampling rate of 200 kHz. Melon-headed whale echolocation clicks had the lowest peak and center frequencies, spinner dolphins had the highest frequencies and bottlenose dolphins were nested in between these two species. Frequency differences were significant. Temporal parameters were not well suited for classification. Feature differences were enhanced by reducing variability within a set of single clicks by calculating mean spectra for groups of clicks. Median peak frequencies of averaged clicks (group size 50) of melon-headed whales ranged between 24.4 and 29.7 kHz, of bottlenose dolphins between 26.7 and 36.7 kHz, and of spinner dolphins between 33.8 and 36.0 kHz. Discriminant function analysis showed the ability to correctly discriminate between 93% of melon-headed whales, 75% of spinner dolphins and 54% of bottlenose dolphins.     

Underwater audiogram of a false killer whale (Pseudorca crassidens)

Underwater audiograms are available for only a few odontocete species. A false killer whale (Pseudorca crassidens) was trained at Sea Life Park in Oahu, Hawaii for an underwater hearing test using a go/no-go response paradigm. Over a 6-month period, auditory thresholds from 2-115 kHz were measured using an up/down staircase psychometric technique. The resulting audiogram showed hearing sensitivities below 64 kHz similar to those of belugas (Delphinapterus leucas) and Atlantic bottlenosed dolphins (Tursiops truncatus). Above 64 kHz, this Pseudorca had a rapid decrease in sensitivity of about 150 dB per octave. A similar decrease in sensitivity occurs at 32 kHz in the killer whale, at 50 kHz in the Amazon River dolphin, at 120 kHz in the beluga, at 140 kHz in the bottlenosed dolphin, and at 140 kHz in the harbor porpoise. The most sensitive range of hearing was from 16-64 kHz (a range of 10 dB from the maximum sensitivity). This range corresponds with the peak frequency of echolocation pulses recorded from captive Pseudorca.