Description of whistles of Irrawaddy dolphins (Orcaella brevirostris) from the waters of Matang,Peninsular Malaysia

The whistles of Irrawaddy dolphins (Orcaella brevirostris) from the waters of Matang, western Peninsular Malaysia are described. Duration, frequency and frequency modulation variables were measured from 163 whistles recorded using a broadband towed hydrophone. Irrawaddy dolphins produced whistles with a mean duration of 0.366 s (S.D. ± 0.217 s). The fundamental frequency of whistles extended from 3040 to 17,123 Hz with low levels of frequency modulation. These dolphins produced whistles that were comparable to those of conspecifics recorded from the waters of Kalimantan, but were generally different from the related Australian snubfin dolphin (O. heinsohni). They also differed from the whistles of the sympatric Indo-Pacific humpback dolphin (Sousa chinensis). Characteristics of Irrawaddy dolphin whistles may be useful in future passive acoustic monitoring studies to investigate differences in sympatric species and their habitat.     

Whistle characteristics of a newly recorded Indo-Pacific humpback dolphin (Sousa chinensis) population in waters southwest of Hainan Island,China, differ from other humpback dolphin populations

Indo-Pacific humpback dolphins (Sousa chinensis) use whistles to communicate with their conspecifics. Little is known about the acoustic repertoire of Indo-Pacific humpback dolphins in waters southwest of Hainan Island, a newly recorded population in 2014. In this study, whistles of Hainan humpback dolphin population were collected by using autonomous acoustic recorders. The fundamental frequencies and durations of whistles were in ranges of 0.71–21.35 kHz and 0.06–2.22 s, respectively. Significant intraspecific differences in duration and frequency of whistles were found between the Hainan population and the other geographically neighboring populations (in Chinese waters) or the population in Malaysia waters. Compared with other Sousa species, significant interspecific differences were also observed. Based on clustering analysis, the whistle parameters of neighboring populations were likely similar to each other. Significant differences were found between humpback dolphins in waters southwest of Hainan Island and those dolphins in the neighboring areas, supporting the hypothesis that this population may be independent. Ambient noise measurements in waters of Hainan Island, Zhanjiang, and Sanniang Bay showed that humpback dolphin populations may use whistles with longer duration, lower frequency, and fewer inflection points for more effective communication to adapt to a noisier environment.     

Biphonal calls in Atlantic spotted dolphins (Stenella frontalis): bitonal and burst-pulse whistles

Biphonation, the simultaneous production of two sounds by a single animal, is found in the vocalizations of a range of mammalian species. Its prevalence suggests it plays an important role in acoustic communication. Concurrent vocal and behavioural recordings were made of Atlantic spotted dolphins (Stenella frontalis) off Bimini, The Bahamas. The occurrence of two types of biphonal signals is reported: burst-pulse whistles with combined tonal and burst-pulse elements, and bitonal whistles. Biphonal whistles are rarely described in reports of dolphin acoustic repertoires, but were common in these dolphins: of all whistles analysed (n = 1211), 26.84% were burst-pulse whistles and 4.71% were bitonal whistles. A subset of whistles (n = 397) were attributed to dolphins of specific age classes, and used to compare prevalence of biphonation across age. Biphonation occurred in 61.54% of sexually mature and 48.32% of sexually immature dolphins’ whistles. Sexually immature dolphins emitted more burst-pulse whistles than older dolphins: 44.13% of sexually immature dolphins’ whistles were burst-pulse whistles, while 15.38% of adult whistles were burst-pulse whistles. Bitonal whistle production was more prevalent in sexually mature dolphins: 41.03% of adult whistles were bitonal, while only 4.19% of sexually immature dolphins’ whistles were bitonal. The prevalence of a biphonal component in specific repeated, stereotyped whistle contours suggests that these acoustic features could be important components of contact calls, or signature whistles. The biphonal components of spotted dolphin whistles may serve to convey additional information as to identity, age or other factors to conspecifics.     

Source levels and the estimated active space of bottlenose dolphin (Tursiops truncatus) whistles in the Moray Firth, Scotland

This study measured SPLs of whistles of wild bottlenose dolphins (Tursiops truncatus) in the Moray Firth, Scotland, and estimated their active space, i.e. the distance at which another dolphin can perceive the whistle of a conspecific. Whistling dolphins were localized with a dispersed hydrophone array by comparing differences in the times of arrival of a whistle at different hydrophones. The mean source level for whistles was 158 +/- 0.6 dB re. 1 microPa. The maximum was 169 dB re. 1 microPa. The active space of these whistles was calculated taking into account transmission loss, ambient noise, the critical ratios and the auditory sensitivity of this species. The estimated radius of the active space of unmodulated whistles between 3.5 kHz and 10 kHz produced at maximum source level ranged from 20 km to 25 km in a habitat of 10 m depth and at sea state 0. At sea state 4 it ranged from 14 km to 22 km. For whistles of 12 kHz it dropped to 1.5-4 km. The results suggest that whistles can be used to maintain group cohesion over large distances but also that dolphins that researchers consider to belong to separate groups might be in acoustic contact.     

Whistling in a noisy ocean: bottlenose dolphins adjust whistle frequencies in response to real-time ambient noise levels

Common bottlenose dolphins (Tursiops truncatus) use complex acoustic behaviours for communication, group cohesion and foraging. Ambient noise from natural and anthropogenic sources has implications for the acoustic behaviour of dolphins, and research shows that average ambient noise levels alter dolphin acoustic behaviour. However, when background noise levels are highly variable, the relationships between noise and acoustic behaviour over short time periods are likely important. This study investigates whether bottlenose dolphins altered the temporal and spectral qualities of their whistles in relation to the ambient noise present at the time the whistles were produced. Dolphin groups were recorded in Tampa Bay (western Florida) between 2008 and 2015. Six whistle parameters were analysed in spectrogram software (minimum frequency, maximum frequency, bandwidth, peak frequency, duration and number of inflection points) and ambient noise levels were calculated immediately prior to each whistle. Linear regression analysis indicated that the minimum, maximum and peak frequencies of whistles had significant positive relationships with the ambient noise levels present at the time of the whistles. These models suggested that for each 1 dB increase in ambient noise, minimum frequency increased by 121 Hz, maximum frequency increased by 108 Hz and peak frequency increased by between 122 and 144 Hz. As ambient noise is typically low frequency, this suggests that bottlenose dolphins increased whistle frequency in response to real-time noise levels to avoid masking. Future research to determine the fitness consequences of noise-induced changes in the communication behaviour of dolphins would be an important contribution to conservation efforts.     

Vocalizations and Behaviour of Pacific Humpback Dolphins Sousa chinensis

Very little is known about the acoustic repertoire of the Pacific humpback dolphin Sousa chinensis . This study, off eastern Australia, used concurrent observations of surface behaviour and acoustic recordings to gain an insight into the behavioural significance of humpback dolphin vocalizations. Humpback dolphins exhibit five different vocalization categories: broad band clicks; barks; quacks; grunts; and whistles. Broad band clicks were high in frequency (8 kHz to > 22 kHz), were directly related to foraging behaviour and may play a role in social behaviour. Barks and quacks were burst pulse sounds (frequency: 0.6 kHz to > 22 kHz, duration: 0.1–8 s) and were associated with both foraging and social behaviour. The grunt vocalization is a low frequency narrow band sound (frequency 0.5–2.6 kHz, duration 0.06–2 s) and was only heard during socializing. There were 17 different types of whistles, ranging widely in frequency (0.9–22 kHz) and vocal structure (n=329). The predominant whistle types used by the groups were type 1 (46%) and type 2 (17%). Most whistles were heard during both socializing and foraging. The number of whistles recorded in a group increased significantly as the number of mother–calf pairs increased, suggesting that whistles may be used as contact calls. Few vocalizations were heard during either travelling or milling behaviours. Broad band clicks, barks and whistle type 1 were the only vocalizations recorded during either travelling or milling.     

What’s occurring? Ultrasonic signature whistle use in Welsh bottlenose dolphins (Tursiops truncatus)

Animal communication signals are diverse. The types of sounds that animals produce, and the way that information is encoded in those sounds, not only varies between species but can also vary geographically within a species. Therefore, an understanding of the vocal repertoire at the population level is important for providing insight into regional differences in vocal communication signals. One species whose vocal repertoire has received considerable attention is the bottlenose dolphin. This species is well known for its use of individually distinctive identity signals, known as signature whistles. Bottlenose dolphins use their signature whistles to broadcast their identity and to maintain contact with social companions. Signature whistles are not innate, but are learnt signals that develop within the first few months of an animal’s life. It is therefore unsurprising that studies which have characterized signature whistles in wild populations of bottlenose dolphins have provided evidence of geographic variation in signature whistle structure. Here, we describe the occurrence of signature whistles in a previously unexplored wild population of bottlenose dolphins in Cardigan Bay, Wales. We present the first occurrence of a signature whistle with an ultrasonic fundamental frequency component (>30 kHz), a frequency band that was not thought to be utilized by this species for whistle communication. We also describe the occurrence of an ultrasonic non-signature whistle. Our findings highlight the importance of conducting regional studies in order to fully quantify a species’ vocal repertoire, and call into question the efficacy of those studies that use restricted sampling rates.     

Communication sounds of Commerson's dolphins (Cephalorhynchus commersonii) and contextual use of vocalizations

Cetaceans produce a variety of vocalizations to communicate; however, little information exists on the acoustic behavior displayed by Commerson's dolphins (Cephalorhynchus commersonii) in the wild other than their echolocation behavior. Most available literature suggests that Commerson's dolphins do not produce any other sound type besides narrow‐band high‐frequency (NBHF) clicks, such that no signals are emitted below 100 kHz. We conducted acoustic recordings together with sightings to study the acoustic behavior of Commerson's dolphins in Bahia San Julian, Argentina. This is the first study that provides evidence that this species produces a variety of acoustic signals, including whistles and broad‐band clicks (BBC), with frequency content well below 100 kHz. Whistles were recorded mostly in the presence of mother and calf and were associated with parental behavior. BBC may be used for communication purposes by adults. These vocalizations are within the hearing range of killer whales and so could pose a risk of predation for Commerson's dolphins. Whether this population of Commerson's dolphins produce all these types of signals while they are in the open sea out of the waters of Bahía San Julián, which are apparently safe from predation, remains unknown.     

Bottlenose dolphins that forage with artisanal fishermen whistle differently

Acoustic communication is a taxonomically widespread phenomenon, crucial for social animals. We evaluate social sounds from bottlenose dolphins (Tursiops truncatus) of Laguna, southern Brazil, whose social structure is organized around a cooperative foraging tactic with artisanal fishermen. This tactic involves stereotyped and coordinated behaviour by dolphins and fishermen and is performed by a subset of the dolphin population, splitting it into two distinct social communities. We compared the acoustic parameters and type of whistles emitted by dolphins of the “non‐cooperative” and “cooperative” communities, both during their interactions with fishermen and in times where dolphins were engaged in other types of foraging. Our findings show how dolphins’ social sounds differ between foraging tactics and social communities. The frequencies of six whistle types (ascending, descending, concave, convex, multiple, flat) were significantly dependent on tactics and communities. Ascending whistles were more common than expected during foraging without fishermen, and among dolphins of the non‐cooperative community. Whistle acoustic parameters (duration, number of inclination changes and inflection points, and initial, final, maximum, minimum frequencies) also varied between social communities. In general, whistles emitted by cooperative dolphins, mainly when not interacting with fishermen, tended to be shorter, had higher frequency and more inflections than those emitted by non‐cooperative dolphins. These results suggest that different whistles may convey specific information among dolphins related to foraging, which we hypothesize promote social cohesion among members of the same social community. These differences in acoustic repertoires add a new dimension of complexity to this unique human–animal interaction.     

Identifying signature whistles from recordings of groups of unrestrained bottlenose dolphins (Tursiops truncatus)

Bottlenose dolphins (Tursiops truncatus) have individually distinctive signature whistles. Each individual dolphin develops its own unique frequency modulation pattern and uses it to broadcast its identity. However, underwater sound localization is challenging, and researchers have had difficulties identifying signature whistles. The traditional method to identify them involved isolating individuals. In this context, the signature whistle is the most commonly produced whistle type of an animal. However, most studies on wild dolphins cannot isolate animals. We present a novel method, SIGnature IDentification (SIGID), that can identify signature whistles in recordings of groups of dolphins recorded via a single hydrophone. We found that signature whistles tend to be delivered in bouts with whistles of the same type occurring within 1–10 s of each other. Nonsignature whistles occur with longer or shorter interwhistle intervals, and this distinction can be used to identify signature whistles in a recording. We tested this method on recordings from wild and captive bottlenose dolphins and show thresholds needed to identify signature whistles reliably. SIGID will facilitate the study of signature whistle use in the wild, signature whistle diversity between different populations, and potentially allow signature whistles to be used in mark‐recapture studies.     

Automated identification and clustering of subunits within delphinid vocalizations

Tonal vocalizations or whistles produced by many species of delphinids range from simple tones to complex frequency contours. Whistle structure varies in duration, frequency, and composition between delphinid species, as well as between populations and individuals. Categorization of whistles may be improved by decomposition of complex calls into simpler subunits, much like the use of phonemes in classification of human speech. We identify a potential whistle decomposition scheme and normalization process to facilitate comparison of whistle subunits derived from tonal vocalizations of bottlenose dolphins (Tursiops truncatus), spinner dolphins (Stenella longirostris), and short‐beaked common dolphins (Delphinus delphis). Network analysis is then used to compare subunits within the vocal corpus of each species. By processing whistles through a series of steps including segmentation, normalization, and dynamic time warping, we are able to automatically cluster selected subunits by shape, regardless of differences in absolute frequency or moderate differences in duration. Using the clustered subunits, we demonstrate a preliminary species classification scheme based on rates of subunit occurrence in vocal repertoires. This provides a potential mechanism for comparing the structure of complex vocalizations within and between species.     

Whistle characteristics of common dolphins (Delphinus sp.) in the Hauraki Gulf,New Zealand

Quantifying the vocal repertoire of a species is critical for subsequent analysis of signal functionality, geographic variation, and social relevance. However, the vocalizations of free‐ranging common dolphins (Delphinus sp.) have not previously been described from New Zealand waters. We present the first quantitative analysis of whistle characteristics to be undertaken on the New Zealand population. Acoustic data were collected in the Hauraki Gulf, North Island from 28 independent dolphin group encounters. A total of 11,715 whistles were collected from 105.1 min of recordings. Seven whistle contours were identified containing 29 subtypes. Vocalizations spanned from 3.2 to 23 kHz, with most whistles occurring between 11 and 13 kHz. Whistle duration ranged from 0.01 to 4.00 s (mean ± SD; 0.27 ± 0.32). Of the 2,663 whistles analyzed, 82% have previously been identified within U.K. populations. An additional six contours, apparently unique to New Zealand Delphinus were also identified. Data presented here offer a first insight into the whistle characteristics of New Zealand Delphinus. Comparisons with previously studied populations reveal marked differences in the whistle frequency and modulation of the New Zealand population. Interpopulation differences suggest behavior and the local environment likely play a role in shaping the vocal repertoire of this species.     

LOW FREQUENCY NARROW-BAND SOUNDS PRODUCED BY BOTTLENOSE DOLPHINS

We present a new sound type recorded from bottlenose dolphins, Tursiops truncatus , in eastern Australian waters: low-frequency, narrow-band (LFN) harmonic sounds (defined as less than 2 kHz). Most of these sounds were of frequencies less than 1 kHz and were recorded commonly from socializing dolphins. These sounds differ significantly from narrow-band whistles, which are higher in frequency and longer in duration. The absence of these sounds in most studies of the acoustic behavior of bottlenose dolphins may reflect geographic differences in repertoires or result from insufficient sampling. Alternatively, these sounds may have been ignored where the focus of research was on other sound types.     

Differences in acoustic signals from Delphinids in the western North Atlantic and northern Gulf of Mexico

Whistle characteristics were quantitatively compared between both geographically separated and neighboring populations of Atlantic spotted dolphins (Stenella frontalis), bottlenose dolphins (Tursiops truncatus), and pilot whales (Globicephala spp.) in U.S. waters to evaluate if intraspecific acoustic differences exist between groups. We compared nine whistle characteristics between continental shelf and offshore Atlantic spotted dolphins in the western North Atlantic and between northern Gulf of Mexico and western North Atlantic bottlenose dolphins and pilot whales using discriminant analysis. Offshore Atlantic spotted dolphin whistles were significantly different (Hotelling's T², P= 0.0003) from continental shelf whistles in high frequency, bandwidth, duration, number of steps, and number of inflection points. Atlantic bottlenose dolphin whistles were significantly different (Hotelling's T², P < 0.0001) from those in the Gulf of Mexico in duration, number of steps, and number of inflection points. There was no significant difference between pilot whale whistles in the two basins. The whistle differences indicate acoustic divergence between groups in different areas that may arise from geographic isolation or habitat separation between neighboring but genetically distinct populations of dolphins. This study supports the premise that acoustic differences can be a tool to evaluate the ecological separation between marine mammal groups in field studies.     

Signature whistles in free‐ranging populations of Indo‐Pacific bottlenose dolphins,Tursiops aduncus

Common bottlenose dolphins (Tursiops truncatus) use individually distinctive signature whistles which are highly stereotyped and function as contact calls. Here we investigate whether Indo‐Pacific bottlenose dolphins (T. aduncus) use signature whistles. The frequency trace of whistle contours recorded from three genetically distinct free‐ranging populations was extracted and sorted into whistle types of similar shape using automated categorization. A signature whistle identification method based on the temporal patterns in signature whistle sequences of T. truncatus was used to identify signature whistle types (SWTs). We then compared the degree of variability in SWTs for several whistle parameters to determine which parameters are likely to encode identity information. Additional recordings from two temporarily isolated T. aduncus made during natural entrapment events in 2008 and 2009 were analyzed for the occurrence of SWTs. All populations were found to produce SWTs; 34 SWTs were identified from recordings of free‐ranging T. aduncus and one SWT was prevalent in each recording of the two temporarily isolated individuals. Of the parameters considered, mean frequency and maximum frequency were the least variable and therefore most likely to reflect identity information encoded in frequency modulation patterns. Our results suggest that signature whistles are commonly used by T. aduncus.     

Dolphins Adjust Species-Specific Frequency Parameters to Compensate for Increasing Background Noise

An increase in ocean noise levels could interfere with acoustic communication of marine mammals. In this study we explored the effects of anthropogenic and natural noise on the acoustic properties of a dolphin communication signal, the whistle. A towed array with four elements was used to record environmental background noise and whistles of short-beaked common-, Atlantic spotted- and striped-dolphins in the Canaries archipelago. Four frequency parameters were measured from each whistle, while Sound Pressure Levels (SPL) of the background noise were measured at the central frequencies of seven one-third octave bands, from 5 to 20 kHz. Results show that dolphins increase the whistles’ frequency parameters with lower variability in the presence of anthropogenic noise, and increase the end frequency of their whistles when confronted with increasing natural noise. This study provides the first evidence that the synergy among SPLs has a role in shaping the whistles'' structure of these three species, with respect to both natural and anthropogenic noise.     

Signature whistles in wild bottlenose dolphins: long-term stability and emission rates

Whistles are key elements in the acoustic repertoire of bottlenose dolphins. In this species, the frequency contours of whistles are used as individual signatures. Assessing the long-lasting stability of such stereotyped signals, and the abundant production of non-stereotyped whistles in the wild, is relevant to a more complete understanding of their biological function. Additionally, studying the effects of group size and activity patterns on whistle emission rate may provide insights into the use of these calls. In this study, we document the decades-long occurrence of whistles with stereotyped frequency contours in a population of wild bottlenose dolphins, resident in the region of the Sado estuary, Portugal. Confirmed stereotypy throughout more than 20 years, and positive identification using the signature identification (SIGID) criteria, suggests that the identified stereotyped whistles are in fact signature whistles. The potential roles of non-stereotyped whistles, which represent 68 % of all whistles recorded, are still unclear and should be further investigated. Emission rates were significantly higher during food-related events. Finally, our data show a comparatively high overall whistle production for this population, and no positive correlation between group size and emission rates, suggesting social or environmental restriction mechanisms in vocal production.     

Clicking in Shallow Rivers: Short-Range Echolocation of Irrawaddy and Ganges River Dolphins in a Shallow,Acoustically Complex Habitat

Toothed whales (Cetacea, odontoceti) use biosonar to navigate their environment and to find and catch prey. All studied toothed whale species have evolved highly directional, high-amplitude ultrasonic clicks suited for long-range echolocation of prey in open water. Little is known about the biosonar signals of toothed whale species inhabiting freshwater habitats such as endangered river dolphins. To address the evolutionary pressures shaping the echolocation signal parameters of non-marine toothed whales, we investigated the biosonar source parameters of Ganges river dolphins (Platanista gangetica gangetica) and Irrawaddy dolphins (Orcaella brevirostris) within the river systems of the Sundarban mangrove forest. Both Ganges and Irrawaddy dolphins produced echolocation clicks with a high repetition rate and low source level compared to marine species. Irrawaddy dolphins, inhabiting coastal and riverine habitats, produced a mean source level of 195 dB (max 203 dB) re 1 µPa_pp whereas Ganges river dolphins, living exclusively upriver, produced a mean source level of 184 dB (max 191) re 1 µPa_pp. These source levels are 1–2 orders of magnitude lower than those of similar sized marine delphinids and may reflect an adaptation to a shallow, acoustically complex freshwater habitat with high reverberation and acoustic clutter. The centroid frequency of Ganges river dolphin clicks are an octave lower than predicted from scaling, but with an estimated beamwidth comparable to that of porpoises. The unique bony maxillary crests found in the Platanista forehead may help achieve a higher directionality than expected using clicks nearly an octave lower than similar sized odontocetes.     

Passive acoustic monitoring of the distribution patterns of Irrawaddy dolphins (Orcaella brevirostris) in the middle reaches of the Ayeyarwady River,Myanmar

The Irrawaddy dolphins (Orcaella brevirostris) are an endangered species. Thus, up-to-date information on the distribution pattern of dolphins is critical for its proper management and conservation. Using a towed passive acoustic monitoring device, the distribution pattern of the Irrawaddy dolphins in the middle reaches of the Ayeyarwady River, Myanmar, was investigated during a vessel-based survey between Mingun and Katha. This region was successively divided into segments 1–4 from upstream to downstream. Sixteen echolocation encounters, with a series of click trains separated by <8 min and 26 dolphin acoustic trajectories were recorded. The mean dolphin detection rate (animals/kilometer) across the four segments progressively increased from upstream to downstream. High relative abundance was observed in segment 4 (46%) and segment 1 (23%) which was consistent with findings from historical boat-based visual surveys. The averaged interclick intervals of each click train in segment 2 and 4 was significantly shorter than that in segment 3, indicating that the dolphins in these segments frequently use shorter-ranged biosonar. More frequent and consistent surveys with a systematic sampling track design that incorporates other factors and covering the whole distribution range along the Ayeyarwady River and at varied water levels are needed in the future.