Echolocation signals of Heaviside's dolphins (Cephalorhynchus heavisidii)

Field recordings of echolocation signals produced by Heaviside's dolphins (Cephalorhynchus heavisidii) were made off the coast of South Africa using a hydrophone array system. The system consisted of three hydrophones and an A-tag (miniature stereo acoustic data-logger). The mean centroid frequency was 125 kHz, with a -3 dB bandwidth of 15 kHz and -10 dB duration of 74 μs. The mean back-calculated apparent source level was 173 dB re 1 μPa(p.-p.). These characteristics are very similar to those found in other Cephalorhynchus species, and such narrow-band high-frequency echolocation clicks appear to be a defining characteristic of the Cephalorhynchus genus. Click bursts with very short inter-click intervals (up to 2 ms) were also recorded, which produced the "cry" sound reported in other Cephalorhynchus species. Since inter-click intervals correlated positively to click duration and negatively to bandwidth, Heaviside's dolphins may adjust their click duration and bandwidth based on detection range. The bimodal distribution of the peak frequency and stable bimodal peaks in spectra of individual click suggest a slight asymmetry in the click production mechanism.     

Perception of the sound source position

This paper presents several experiments on sound source localization. They are based on monaural click presented at different interclick intervals (ICI), from 10 to 100 ms. Trains of clicks were presented to 10 healthy subjects. At short interclick intervals the clicks were perceived as a blur of clicks having a buzzy quality. Moreover, it was proven that the accurateness in the response improves with the increase of the length of ICI. The present results imply the usefulness of the interclick interval in estimating the perceptual accuracy. An important benefit of this task is that this enables a careful examination of the sound source perception threshold. This allows detecting, localizing and dividing with a high accuracy the sounds in the environment. The text was submitted by the authors in English.     

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.     

Characteristics of biosonar signals from the northern bottlenose whale, Hyperoodon ampullatus

The biosonar pulses from free-ranging northern bottlenose whales (Hyperoodon ampullatus) were recorded with a linear hydrophone array. Signals fulfilling criteria for being recorded close to the acoustic axis of the animal (a total of 10 clicks) had a frequency upsweep from 20 to 55 kHz and durations of 207 to 377 μs (measured as the time interval containing 95% of the signal energy). The source level of these signals, denoted pulses, was 175-202 dB re 1 μPa rms at 1 m. The pulses had a directionality index of at least 18 dB. Interpulse intervals ranged from 73 to 949 ms (N?=?856). Signals of higher repetition rates had interclick intervals of 5.8-13.1 ms (two sequences, made up of 59 and 410 clicks, respectively). These signals, denoted clicks, had a shorter duration (43-200 μs) and did not have the frequency upsweep characterizing the pulses of low repetition rates. The data show that the northern bottlenose whale emits signals similar to three other species of beaked whale. These signals are distinct from the three other types of biosonar signals of toothed whales. It remains unclear why the signals show this grouping, and what consequences it has on echolocation performance.     

Vocal reporting of echolocation targets: dolphins often report before click trains end

Bottlenose dolphins (Tursiops truncatus) wore opaque suction cups over their eyes while stationing behind an acoustically opaque door. This put the dolphins in a known position and orientation. When the door opened, the dolphin clicked to detect targets. Trainers specified that Dolphin S emit a whistle if the target was a 7.5 cm water filled sphere, or a pulse burst if the target was a rock. S remained quiet if there was no target. Dolphin B whistled for the sphere. She remained quiet for rock and for no target. Thus, S had to choose between three different responses, whistle, pulse burst, or remain quiet. B had to choose between two different responses, whistle or remain quiet. S gave correct vocal responses averaging 114 ms after her last echolocation click (range 182 ms before and 219 ms after the last click). Average response for B was 21 ms before her last echolocation click (range 250 ms before and 95 ms after the last click in the train). More often than not, B began her whistle response before her echolocation train ended. The findings suggest separate neural pathways for generation of response vocalizations as opposed to echolocation clicks.     

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.     

Lateralization thresholds obtained under conditions in which the precedence effect is assumed to operate

Interaural differences of time (IDT) thresholds were measured with 600-microseconds transients. The initial experiment was a successful replication of previous experiments that have obtained the precedence effect in lateralization paradigms (e.g., Yost and Soderquist, 1984). When a dichotic click followed a diotic click with an interclick interval (ICI) less than 1 ms or larger than 5 ms, IDT thresholds were generally less than 40 microseconds. For ICIs between 1 to 5 ms, IDT thresholds increased to approximately 220 microseconds. Poorest performance was observed for ICIs of 1.75 to 2.35 ms. During the course of conducting a series of planned experiments on this effect, a substantial drop in IDT thresholds was observed across the ICIs of maximum interest (1 to 5 ms). The precedence effect, which we had replicated in our initial experiment, essentially "disappeared" when the subjects were given sufficient practice on the lateralization task. A number of conditions were explored in an unsuccessful attempt to recover the precedence effect in these experienced subjects. The implications of these results are discussed.     

Male sperm whale acoustic behavior observed from multipaths at a single hydrophone

Sperm whales generate transient sounds (clicks) when foraging. These clicks have been described as echolocation sounds, a result of having measured the source level and the directionality of these signals and having extrapolated results from biosonar tests made on some small odontocetes. The authors propose a passive acoustic technique requiring only one hydrophone to investigate the acoustic behavior of free-ranging sperm whales. They estimate whale pitch angles from the multipath distribution of click energy. They emphasize the close bond between the sperm whale's physical and acoustic activity, leading to the hypothesis that sperm whales might, like some small odontocetes, control click level and rhythm. An echolocation model estimating the range of the sperm whale's targets from the interclick interval is computed and tested during different stages of the whale's dive. Such a hypothesis on the echolocation process would indicate that sperm whales echolocate their prey layer when initiating their dives and follow a methodic technique when foraging.     

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).     

Characteristics of whistles from rough-toothed dolphins (Steno bredanensis) in Rio de Janeiro coast, southeastern Brazil

There is no information about the whistles of rough-toothed dolphins in the South Atlantic Ocean. This study characterizes the whistle structure of free-ranging rough-toothed dolphins recorded on the Rio de Janeiro coast, southeastern Brazil, and compares it to that of the same species in other regions. A total of 340 whistles were analyzed. Constant (N = 115; 33.8%) and ascending (N = 99; 29.1%) whistles were the most common contours. The whistles recorded had their fundamental frequencies between 2.24 and 13.94 kHz. Whistles without inflection points were frequently emitted (N = 255; 75%). Some signals presented breaks or steps in their contour (N = 97; 28.5%). Whistle duration was short (347 ± 236 ms and 89.7% of the whistles lasted <600 ms). Seventy-eight whistle contour types were found in the total of whistles analyzed, and 27 (7.9%) of these occurred only once. Most of the whistle types were unique to a particular recording session (N = 43). The signals emitted by the rough-toothed dolphins in southeastern Brazil were characterized by low frequency modulation, short duration, low number of inflection points, and breaks. Differences in the mean values of the whistle parameters were found between this and other studies that recorded Steno bredanensis, but as in other localities, whistles above 14 kHz are rare.     

Detection of interaural differences of intensity in trains of high-frequency clicks as a function of interclick interval and number

Listeners were asked to detect interaural differences of intensity in trains of 4000-Hz clicks as the interclick interval (ICI) was varied from 10 to 1 ms and the number of clicks in a train (n) was varied from 1 to 32. As has previously been shown for differences of time [Hafter and Dye, J. Acoust. Soc. Am. 73, 644-651 (1983)], plots of log interaural threshold versus log n produced slopes that decrease with ICI. These results are explained in terms of a saturation model which argues that as the click rate increases, the evoked neural activity changes from what is essentially a tonic response toward one that is more phasic.     

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.     

Separating overlapping click trains originating from multiple individuals in echolocation recordings

Recordings of the acoustic activity of free-swimming groups of echolocating dolphins increase the likelihood of collecting overlapping click trains, originating from multiple individuals, in the same set of data. In order to evaluate the click properties of each individual based on such recordings it is necessary to identify which clicks originate from which animal. This paper suggests a computationally efficient strategy to separate overlapping click trains originating from multiple free-swimming bottlenose dolphins, enabling echolocation analysis at an individual level on several animals. This technique is based on sequential matching of the frequency spectra of successive clicks. The clicks are grouped together as individual click trains if the correlation coefficients between clicks are higher than a pre-set threshold level. The robustness of the algorithm is tested by adding artificially generated white Gaussian noise and comparing the results with other comparable commonly used methods based on inter-click intervals, centroid frequencies, and amplitude levels. The described method is applicable to a variety of experimental and observational contexts, e.g., those regarding echolocation development of calves, the hypothesized acoustic "etiquette" among dolphins when investigating the same object, and the possible occurrence of eavesdropping in large dolphin pods.     

Neural time and movement time in choice of whistle or pulse burst responses to different auditory stimuli by dolphins

Echolocating dolphins emit trains of clicks and receive echoes from ocean targets. They often emit each successive ranging click about 20 ms after arrival of the target echo. In echolocation, decisions must be made about the target--fish or fowl, predator or food. In the first test of dolphin auditory decision speed, three bottlenose dolphins (Tursiops truncatus) chose whistle or pulse burst responses to different auditory stimuli randomly presented without warning in rapid succession under computer control. The animals were trained to hold pressure catheters in the nasal cavity so that pressure increases required for sound production could be used to split response time (RT) into neural time and movement time. Mean RT in the youngest and fastest dolphin ranged from 175 to 213 ms when responding to tones and from 213 to 275 ms responding to pulse trains. The fastest neural times and movement times were around 60 ms. The results suggest that echolocating dolphins tune to a rhythm so that succeeding pulses in a train are produced about 20 ms over target round-trip travel time. The dolphin nervous system has evolved for rapid processing of acoustic stimuli to accommodate for the more rapid sound speed in water compared to air.     

Modeling the cochlear nucleus: a site for monaural echo suppression?

Echo suppression plays an important role in identifying and localizing auditory objects. One can distinguish between binaural and monaural echo suppression, although the former is the one commonly referred to. Based on biological findings we introduce and analyze a mathematical model for a neural implementation of monaural echo suppression in the cochlear nucleus. The model's behavior has been verified by analytical calculations as well as by numerical simulations for several types of input signal. It shows that in the perception of a pair of clicks the leading click suppresses the lagging one and that suppression is maximal for an interclick interval of 2-3 ms. Similarly, ongoing stimuli will be affected by the suppression mechanism primarily a couple of milliseconds after onset, resulting in a reduced perception of a sound shortly after its start. Both effects match experimental data.     

Time patterns of sperm whale codas recorded in the Mediterranean Sea 1985-1996

A distinctive vocalization of the sperm whale, Physeter macrocephalus (=P. catodon), is the coda: a short click sequence with a distinctive stereotyped time pattern [Watkins and Schevill, J. Acoust. Soc. Am. 62, 1485-1490 (1977)]. Coda repertoires have been found to vary both geographically and with group affiliation [Weilgart and Whitehead, Behav. Ecol. Sociobiol. 40, 277-285 (1997)]. In this work, the click timings and repetition patterns of sperm whale codas recorded in the Mediterranean Sea are characterized statistically, and the context in which the codas occurred are also taken into consideration. A total of 138 codas were recorded in the central Mediterranean in the years 1985-1996 by several research groups using a number of different detection instruments, including stationary and towed hydrophones, sonobuoys and passive sonars. Nearly all (134) of the recorded codas share the same "3+1" (/// /) click pattern. Coda durations ranged from 456 to 1280 ms, with an average duration of 908 ms and a standard deviation of 176 ms. Most of the codas (a total of 117) belonged to 20 coda series. Each series was produced by an individual, in most cases by a mature male in a small group, and consisted of between 2 and 16 codas, emitted in one or more "bursts" of 1 to 13 codas spaced fairly regularly in time. The mean number of codas in a burst was 3.46, and the standard deviation was 2.65. The time interval ratios within a coda are parameterized by the coda duration and by the first two interclick intervals normalized by coda duration. These three parameters remained highly stable within each coda series, with coefficients of variation within the series averaging less than 5%. The interval ratios varied somewhat across the data sets, but were highly stable over 8 of the 11 data sets, which span 11 years and widely dispersed geographic locations. Somewhat different interval ratios were observed in the other three data sets; in one of these data sets, the variant codas were produced by a young whale. Two sets of presumed sperm whale codas recorded in 1996 had 5- and 6-click patterns; the observation of these new patterns suggests that sperm whale codas in the Mediterranean may have more variations than previously believed.     

Beaked whale and dolphin tracking using a multichannel autonomous acoustic recorder

To track highly directional echolocation clicks from odontocetes, passive hydrophone arrays with small apertures can be used to receive the same high frequency click on each sensor. A four-hydrophone small-aperture array was coupled to an autonomous acoustic recorder and used for long-term tracking of high-frequency odontocete sounds. The instrument was deployed in the spring of 2009 offshore of southern California in a known beaked whale and dolphin habitat at about 1000 m depth. The array was configured as a tetrahedron with approximately 0.5 m sensor spacing. Time difference of arrival measurements between the six sensor-pairs were used to estimate three-dimensional bearings to sources. Both near-seafloor beaked whales and near-sea surface dolphins were tracked. The tracks observed using this technique provide swimming and diving behavioral information for free-ranging animals using a single instrument. Furthermore, animal detection ranges were derived, allowing for estimation of detection probability functions.     

Comparison of click responses of primary auditory fibers with minimum-phase predictions

Minimum-phase impulse responses were constructed from the frequency threshold-response curves of primary auditory fibers in the anesthetized cat. These impulse responses had many of the same characteristics as the experimental click responses. The two types of responses had similar inter-peak intervals as well as similar (+/- 1.5 ms) latencies to the principal mode and similar (+/- 1.0 ms) intervals from response onset to the principal mode.     

Propagation of beluga echolocation signals

The propagation characteristics of high-frequency echolocation signals (peak energies above 100 kHz) of the beluga (Delphinapterus leucas) were measured while the animal performed a target detection task. The whale was trained to station on a bite plate so that its transmission beam could be measured in the vertical and horizontal planes using hydrophone arrays. The transitional region between the acoustic near- and farfields was also located using an array of hydrophones that extended directly in front of the animal in the horizontal plane. Three distinct modes of signals were observed. Mode 1 signals had click intervals greater than the time required for the signals to travel to the target and back (two-way transit time). Mode 2 signals had click intervals shorter than the two-way transit time, and mode 3 signals had high repetition rates with an average click interval of 1.7 ms, approximately 2% of the two-way transit time. The average click intervals for the modes 1 and 2 signals were 193 and 44 ms, respectively. The vertical and horizontal beam patterns of the mode 1 signals had similar 3-dB beamwidths of approximately 6.5 degrees. The major axis of the vertical beam was directed approximately 5 degrees above the plane defined by the animal's teeth. The near- to farfield transition region was approximately 0.64-0.75 m from the tip of the animal's mouth.     

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.