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.     

Synthesis and modification of the whistles of the bottlenose dolphin, Tursiops truncatus

A signal-processing algorithm was developed to analyze harmonic frequency-modulated sounds, to modify the parameters of the analyzed signal, and to synthesize a new analytically specified signal that resembles the original signal in specified features. This algorithm was used with dolphin whistles, a frequency-modulated harmonic signal that has typically been described in terms of its contour, or pattern of modulation of the fundamental frequency. In order to test whether other features may also be salient to dolphins, the whistle analysis calculates the energies at the harmonics as well as the fundamental frequency of the whistle. The modification part of the algorithm can set all of these energies to a constant, can shift the whistle frequency, and can expand or compress the time base or the frequency of the whistle. The synthesis part of the algorithm then synthesizes a waveform based upon the energies and frequencies of the fundamental and first two harmonics. These synthetic whistles will be useful for evaluating what acoustic features dolphins use in discriminating different whistles.     

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.     

The whistles of Hawaiian spinner dolphins

The characteristics of the whistles of Hawaiian spinner dolphins (Stenella longirostris) are considered by examining concurrently the whistle repertoire (whistle types) and the frequency of occurrence of each whistle type (whistle usage). Whistles were recorded off six islands in the Hawaiian Archipelago. In this study Hawaiian spinner dolphins emitted frequency modulated whistles that often sweep up in frequency (47% of the whistles were upsweeps). The frequency span of the fundamental component was mainly between 2 and 22 kHz (about 94% of the whistles) with an average mid-frequency of 12.9 kHz. The duration of spinner whistles was relatively short, mainly within a span of 0.05 to 1.28 s (about 94% of the whistles) with an average value of 0.49 s. The average maximum frequency of 15.9 kHz obtained by this study is consistent with the body length versus maximum frequency relationship obtained by Wang et al. (1995a) when using spinner dolphin adult body length measurements. When comparing the average values of whistle parameters obtained by this and other studies in the Island of Hawaii, statistically significant differences were found between studies. The reasons for these differences are not obvious. Some possibilities include differences in the upper frequency limit of the recording systems, different spinner groups being recorded, and observer differences in viewing spectrograms. Standardization in recording and analysis procedure is clearly needed.     

The effect of recording and analysis bandwidth on acoustic identification of delphinid species

Because many cetacean species produce characteristic calls that propagate well under water, acoustic techniques can be used to detect and identify them. The ability to identify cetaceans to species using acoustic methods varies and may be affected by recording and analysis bandwidth. To examine the effect of bandwidth on species identification, whistles were recorded from four delphinid species (Delphinus delphis, Stenella attenuata, S. coeruleoalba, and S. longirostris) in the eastern tropical Pacific ocean. Four spectrograms, each with a different upper frequency limit (20, 24, 30, and 40 kHz), were created for each whistle (n = 484). Eight variables (beginning, ending, minimum, and maximum frequency; duration; number of inflection points; number of steps; and presence/absence of harmonics) were measured from the fundamental frequency of each whistle. The whistle repertoires of all four species contained fundamental frequencies extending above 20 kHz. Overall correct classification using discriminant function analysis ranged from 30% for the 20-kHz upper frequency limit data to 37% for the 40-kHz upper frequency limit data. For the four species included in this study, an upper bandwidth limit of at least 24 kHz is required for an accurate representation of fundamental whistle contours.     

Whistles of beluga whales in the reproductive gathering off Solovetskii Island in the White Sea

Whistles recorded in a reproductive gathering of beluga whales near Solovetskii Island in the White Sea are analyzed. On the basis of the absolute characteristics and shape of the frequency contour, whistles are classed into 16 types. Whistles belong to a relatively low frequency band, contain many harmonics, and have a simple shape of frequency contour. The average whistle duration varies from 0.1 to 1.7 s for different types, the average value of the maximum fundamental frequency varies from 1.4 to 4.5 kHz, and the average number of inflection points is from 0 to 9 per signal. In contrast to other populations, where flat whistles are the most frequent vocalizations, beluga whales observed in the reproductive gathering in the White Sea most often produce short whistles with a V-shaped frequency contour.     

Estimated communication range of social sounds used by bottlenose dolphins (Tursiops truncatus)

Bottlenose dolphins, Tursiops truncatus, exhibit flexible associations in which the compositions of groups change frequently. We investigated the potential distances over which female dolphins and their dependent calves could remain in acoustic contact. We quantified the propagation of sounds in the frequency range of typical dolphin whistles in shallow water areas and channels of Sarasota Bay, Florida. Our results indicated that detection range was noise limited as opposed to being limited by hearing sensitivity. Sounds were attenuated to a greater extent in areas with seagrass than any other habitat. Estimates of active space of whistles showed that in seagrass shallow water areas, low-frequency whistles (7-13 kHz) with a 165 dB source level could be heard by dolphins at 487 m. In shallow areas with a mud bottom, all whistle frequency components of the same whistle could be heard by dolphins travel up to 2 km. In channels, high-frequency whistles (13-19 kHz) could be detectable potentially over a much longer distance (> 20 km). Our findings indicate that the communication range of social sounds likely exceeds the mean separation distances between females and their calves. Ecological pressures might play an important role in determining the separation distances within communication range.     

Captive dolphins,Tursiops truncatus,develop signature whistles that match acoustic features of human-made model sounds

This paper presents a cross-sectional study testing whether dolphins that are born in aquarium pools where they hear trainers' whistles develop whistles that are less frequency modulated than those of wild dolphins. Ten pairs of captive and wild dolphins were matched for age and sex. Twenty whistles were sampled from each dolphin. Several traditional acoustic features (total duration, duration minus any silent periods, etc.) were measured for each whistle, in addition to newly defined flatness parameters: total flatness ratio (percentage of whistle scored as unmodulated), and contiguous flatness ratio (duration of longest flat segment divided by total duration). The durations of wild dolphin whistles were found to be significantly longer, and the captive dolphins had whistles that were less frequency modulated and more like the trainers' whistles. Using a standard t-test, the captive dolphin had a significantly higher total flatness ratio in 9/10 matched pairs, and in 8/10 pairs the captive dolphin had significantly higher contiguous flatness ratios. These results suggest that captive-born dolphins can incorporate features of artificial acoustic models made by humans into their signature whistles.     

The perception of complex tones by a false killer whale (Pseudorca crassidens)

Complex tonal whistles are frequently produced by some odontocete species. However, no experimental evidence exists regarding the detection of complex tones or the discrimination of harmonic frequencies by a marine mammal. The objectives of this investigation were to examine the ability of a false killer whale to discriminate pure tones from complex tones and to determine the minimum intensity level of a harmonic tone required for the whale to make the discrimination. The study was conducted with a go/no-go modified staircase procedure. The different stimuli were complex tones with a fundamental frequency of 5 kHz with one to five harmonic frequencies. The results from this complex tone discrimination task demonstrated: (1) that the false killer whale was able to discriminate a 5 kHz pure tone from a complex tone with up to five harmonics, and (2) that discrimination thresholds or minimum intensity levels exist for each harmonic combination measured. These results indicate that both frequency level and harmonic content may have contributed to the false killer whale's discrimination of complex tones.     

Geographic variations in the whistles of spinner dolphins (Stenella longirostris) of the Main Hawai'ian Islands

Geographic variations in the whistles of Hawai'ian spinner dolphins are discussed by comparing 27 spinner dolphin pods recorded in waters off the Islands of Kaua'i, O'ahu, Lana'i, and Hawai'i. Three different behavioral states, the number of dolphins observed in each pod, and ten parameters extracted from each whistle contour were considered by using clustering and discriminant function analyses. The results suggest that spinner dolphin pods in the Main Hawai'ian Islands share characteristics in approximately 48% of their whistles. Spinner dolphin pods had similar whistle parameters regardless of the island, location, and date when they were sampled and the dolphins' behavioral state and pod size. The term "whistle-specific subgroup" (WSS) was used to designate whistle groups with similar whistles parameters (which could have been produced in part by the same dolphins). The emission rate of whistles was higher when spinner dolphins were socializing than when they were traveling or resting, suggesting that whistles are mainly used during close-range interactions. Spinner dolphins also seem to vary whistle duration according to their general behavioral state. Whistle duration and the number of turns and steps of a whistle may be more important in delivering information at the individual level than whistle frequency parameters.     

The broadband social acoustic signaling behavior of spinner and spotted dolphins

Efforts to study the social acoustic signaling behavior of delphinids have traditionally been restricted to audio-range (<20 kHz) analyses. To explore the occurrence of communication signals at ultrasonic frequencies, broadband recordings of whistles and burst pulses were obtained from two commonly studied species of delphinids, the Hawaiian spinner dolphin (Stenella longirostris) and the Atlantic spotted dolphin (Stenella frontalis). Signals were quantitatively analyzed to establish their full bandwidth, to identify distinguishing characteristics between each species, and to determine how often they occur beyond the range of human hearing. Fundamental whistle contours were found to extend beyond 20 kHz only rarely among spotted dolphins, but with some regularity in spinner dolphins. Harmonics were present in the majority of whistles and varied considerably in their number, occurrence, and amplitude. Many whistles had harmonics that extended past 50 kHz and some reached as high as 100 kHz. The relative amplitude of harmonics and the high hearing sensitivity of dolphins to equivalent frequencies suggest that harmonics are biologically relevant spectral features. The burst pulses of both species were found to be predominantly ultrasonic, often with little or no energy below 20 kHz. The findings presented reveal that the social signals produced by spinner and spotted dolphins span the full range of their hearing sensitivity, are spectrally quite varied, and in the case of burst pulses are probably produced more frequently than reported by audio-range analyses.     

Whistles of small groups of Sotalia fluviatilis during foraging behavior in southeastern Brazil

Whistle emissions were recorded from small groups of marine tucuxi dolphins (Sotalia fluviatilis) in two beaches located in an important biological reserve in the Cananéia estuary (25 degrees 03'S, 47 degrees 58'W), southeastern Brazil. A total of 17 h of acoustic data was collected when dolphins were engaged in a specific feeding foraging activity. The amount of 3235 whistles was recorded and 40% (n=1294) were analyzed. Seven acoustic whistle parameters were determined: duration (ms), number of inflection points, start and end frequency (kHz), minimum and maximum frequency (kHz), and frequency range (kHz). Whistles with up to four inflection points were found. Whistles with no inflection points and rising frequency corresponded to 85% (n=1104) of all analyzed whistles. Whistle duration varied from 38 to 627 ms (mean=229.6+/-109.9 ms), with the start frequency varying between 1 and 16 kHz (mean=8.16+/-3.0 kHz) and the end frequency between 2 and 18 kHz (mean=14.35+/-3.0 kHz). The importance of this study requires an accurate measurement of the whistles' emissions in an unusual foraging feeding behavior situation on two beaches where several tucuxis, mostly mother-calf pairs, are frequently present. These two beaches are located in a federal and state environment Environmental Protected Area threatened by the progressive increase of tourism.     

Acoustic properties of humpback whale songs

A vertical array of five hydrophones was used to measure the acoustic field in the vertical plane of singing humpback whales. Once a singer was located, two swimmers with snorkel gear were deployed to determine the orientation of the whale and position the boat so that the array could be deployed in front of the whale at a minimum standoff distance of at least 10 m. The spacing of the hydrophones was 7 m with the deepest hydrophone deployed at a depth of 35 m. An eight-channel TASCAM recorder with a bandwidth of 24 kHz was used to record the hydrophone signals. The location (distance and depth) of the singer was determined by computing the time of arrival differences between the hydrophone signals. The maximum source level varied between individual units in a song, with values between 151 and 173 dB re 1 microPa. One of the purposes of this study was to estimate potential sound exposure of nearby conspecifics. The acoustic field determined by considering the relative intensity of higher frequency harmonics in the signals indicated that the sounds are projected in the horizontal direction despite the singer being canted head downward anywhere from about 25 degrees to 90 degrees. High-frequency harmonics extended beyond 24 kHz, suggesting that humpback whales may have an upper frequency limit of hearing as high as 24 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.     

A method for classifying whistles of bottlenose dolphin(Tursiops truncates) based on syntactic pattern reorganization

A method based on syntactic pattern recognition was presented to automatically classify whistles of bottlenose dolphin.Dolphin whistles have typically been characterized in terms of their instantaneous frequency as a function of time,which is also known as "whistle contour".The frequency variation features of a whistle were extracted according to its contour.Then,the frequency variation features were used for learning grammatical patterns.A whistle was classified according to grammatical pattern of its frequency variation features.The experimental results showed that the classification accuracy of the proposed method was 95%.The method can provide technical support for acoustic study of dolphins' biological behavior.     

Automated categorization of bioacoustic signals: avoiding perceptual pitfalls

Dividing the acoustic repertoires of animals into biologically relevant categories presents a widespread problem in the study of animal sound communication, essential to any comparison of repertoires between contexts, individuals, populations, or species. Automated procedures allow rapid, repeatable, and objective categorization, but often perform poorly at detecting biologically meaningful sound classes. Arguably this is because many automated methods fail to address the nonlinearities of animal sound perception. We present a new method of categorization that incorporates dynamic time-warping and an adaptive resonance theory (ART) neural network. This method was tested on 104 randomly chosen whistle contours from four captive bottlenose dolphins (Tursiops truncatus), as well as 50 frequency contours extracted from calls of transient killer whales (Orcinus orca). The dolphin data included known biologically meaningful categories in the form of 42 stereotyped whistles produced when each individual was isolated from its group. The automated procedure correctly grouped all but two stereotyped whistles into separate categories, thus performing as well as human observers. The categorization of killer whale calls largely corresponded to visual and aural categorizations by other researchers. These results suggest that this methodology provides a repeatable and objective means of dividing bioacoustic signals into biologically meaningful categories.     

一种宽吻海豚通讯信号自动分类的方法

针对宽吻海豚通讯信号自动分类提出了一种基于句法模式识别的方法。该方法首先提取海豚通讯信号基频随时间变化的轨迹曲线,然后提取基频变化的基元序列。根据海豚通讯信号分类的标准,归纳出产生各类海豚通讯信号基频基元序列的文法。对未知类别的海豚通讯信号,提取其基频变化的基元序列,根据各类模式的文法对基元序列进行分类,进而实现海豚通讯信号的自动分类。实验结果显示本文方法的分类准确率达到了95%。本文方法预期为海豚生物学行为的声学研究提供一定的技术支持。      

Sounds produced by Australian Irrawaddy dolphins, Orcaella brevirostris

Sounds produced by Irrawaddy dolphins, Orcaella brevirostris, were recorded in coastal waters off northern Australia. They exhibit a varied repertoire, consisting of broadband clicks, pulsed sounds and whistles. Broad-band clicks, "creaks" and "buzz" sounds were recorded during foraging, while "squeaks" were recorded only during socializing. Both whistle types were recorded during foraging and socializing. The sounds produced by Irrawaddy dolphins do not resemble those of their nearest taxonomic relative, the killer whale, Orcinus orca. Pulsed sounds appear to resemble those produced by Sotalia and nonwhistling delphinids (e.g., Cephalorhynchus spp.). Irrawaddy dolphins exhibit a vocal repertoire that could reflect the acoustic specialization of this species to its environment.