Vocal exchanges of signature whistles in bottlenose dolphins (<Emphasis Type="Italic">Tursiops truncatus</Emphasis>)

Bottlenose dolphins (Tursiops truncatus) produce individually distinctive vocalizations—referred to as “signature whistles”—that are thought to function as an individual and conspecific recognition system for maintenance of consistent contact between individuals. Observations and playback experiments were conducted at aquariums to study these whistle–vocal exchanges in bottlenose dolphins. Temporal patterns of vocalization were examined by analyzing the intercall intervals between two consecutive whistles. When a second individual produced a call that was different from the first individual’s vocalization, most of these calls were shorter than 1 s. However, when two consecutive calls were produced by the same individual, the second call rarely occurred within 1 s of the first. These results suggest that a second whistle may be produced by a different caller in response to the first whistle; however, in the case of an absence of a response, the first caller is likely to give further whistles. The results of this acoustic analysis show that the dolphins used in this study mostly used signature whistles during the recorded vocal exchanges.     

Variability and context specificity of narwhal (Monodon monoceros) whistles and pulsed calls

The behavioral and environmental context of animal calls provides insights into their functions. Narwhals are a highly vocal species and, like other social cetaceans, rely on acoustic signals to communicate. We characterize and categorize narwhal whistles and pulsed calls, as well as investigate variation in these calls under different contexts (behavior, herd, and year) using recordings made during the month of August 2006–2008, in Koluktoo Bay (72°04′N, 80°32′W). We detected similarities among whistles but not pulsed calls that were produced under a similar behavioral context. Both whistles and pulsed calls recorded within the same herd were more similar than whistles and pulsed calls recorded within different herds. We did not find any type of whistle to be associated with a specific behavior although some acoustical features might be behavior specific. Both whistles and pulsed calls show properties that are consistent with the hypothesis that narwhals produce group‐ or individual‐specific calls.     

Exchange of “signature” calls in captive belugas (Delphinapterus leucas)

Belugas (Delphinapterus leucas) produce echolocation clicks, burst pulses, and whistles. The sounds of 3 captive belugas were recorded using 2 hydrophones at the Port of Nagoya Public Aquarium. There were stable individual differences in the pulse patterning of one type of pulsed sounds (PS1 call), suggesting that belugas use these as “signature” calls. Eighty-eight percent of PS1 calls initiated PS1 calls from other animals within 1 s. PS1 calls repeated by the same individual occurred primarily when other belugas did not respond within 1 s of the first call. Belugas delayed successive PS1 calls when other belugas responded with a PS1 call within 1 s. There was no clear temporal pattern for whistles. It appears that the time limit for responding to calls is 1 s after the initial call. If other individuals do not respond to the PS1 call of a beluga within 1 s, belugas tend to repeat the call and wait for a response. The results of this study suggest that the belugas exchange their individual signatures by using PS1 calls, in a manner similar to that of signature whistles used by bottlenose dolphins.     

Diversity in sound pressure levels and estimated active space of resident killer whale vocalizations

Signal source intensity and detection range, which integrates source intensity with propagation loss, background noise and receiver hearing abilities, are important characteristics of communication signals. Apparent source levels were calculated for 819 pulsed calls and 24 whistles produced by free-ranging resident killer whales by triangulating the angles-of-arrival of sounds on two beamforming arrays towed in series. Levels in the 1–20 kHz band ranged from 131 to 168 dB re 1 μPa at 1 m, with differences in the means of different sound classes (whistles: 140.2±4.1 dB; variable calls: 146.6±6.6 dB; stereotyped calls: 152.6±5.9 dB), and among stereotyped call types. Repertoire diversity carried through to estimates of active space, with “long-range” stereotyped calls all containing overlapping, independently-modulated high-frequency components (mean estimated active space of 10–16 km in sea state zero) and “short-range” sounds (5–9 km) included all stereotyped calls without a high-frequency component, whistles, and variable calls. Short-range sounds are reported to be more common during social and resting behaviors, while long-range stereotyped calls predominate in dispersed travel and foraging behaviors. These results suggest that variability in sound pressure levels may reflect diverse social and ecological functions of the acoustic repertoire of killer whales.     

Changes in the acoustic behavior of resident bottlenose dolphins near operating vessels

Maritime traffic is an issue of major ecological concern, and vessel noise may be an important source of disturbance for coastal cetaceans. In the Sado estuary, Portugal, core habitat areas of a small resident population of bottlenose dolphins (Tursiops truncatus) overlap with routes of intense maritime traffic, which presents an opportunity to assess vocal responses of these dolphins to specific vessel noise sources. Field recordings of dolphin vocalizations were made from April to November 2011, using a calibrated system. Dolphin behavior and group size were recorded, as well as the operating boat condition (no boats or specific boat type) in a 1,000 m radius. Spectral analyses of vocalizations allowed the categorization and quantitative analysis of echolocation click trains and social calls, including whistles. Mean overall call rates decreased significantly in the presence of operating vessels. Creaks (fast click trains) were significantly reduced in the presence of ferry boats. Significant differences were also observed in the whistles' minimum, maximum, and start frequencies. These changes in call emission rates and temporary shifts in whistles characteristics may be a vocal response to the proximity of operating vessels, facilitating communication in this busy, noisy estuary.     

Acoustic classification of dolphins in the California Current using whistles,echolocation clicks,and burst pulses

Passive acoustic monitoring of dolphins is limited by our ability to classify calls to species. Significant overlap in call characteristics among many species, combined with a wide range of call types and acoustic behavior, makes classification of calls to species challenging. Here, we introduce BANTER, a compound acoustic classification method for dolphins that utilizes information from all call types produced by dolphins rather than a single call type, as has been typical for acoustic classifiers. Output from the passive acoustic monitoring software, PAMGuard, was used to create independent classifiers for whistles, echolocation clicks, and burst pulses, which were then merged into a final, compound classifier for each species. Classifiers for five species found in the California Current ecosystem were trained and tested using 153 single‐species acoustic events recorded during a 4.5 mo combined visual and acoustic shipboard cetacean survey off the west coast of the United States. Correct classification scores for individual species ranged from 71% to 92%, with an overall correct classification score of 84% for all five species. The conceptual framework of this approach easily lends itself to other species and study areas as well as to noncetacean taxa.     

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.     

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.     

ACOUSTIC ECOLOGY OF FORAGING BOTTLENOSE DOLPHINS (TURSIOPS TRUNCATUS), HABITAT-SPECIFIC USE OF THREE SOUND TYPES

The function(s) of a particular sound can be explored in detail only if the context of its use is well understood. The behavior of the signaler, and the habitat in which that behavior is observed, are two of the most important components of understanding context specific use of a sound. Bottlenose dolphin foraging behavior is often inferred from relatively few behavioral cues that are visible from the surface. To investigate the use of three specific sound types: echolocation, whistles, and pops during foraging, I recorded sound use by animals engaged in a set of previously defined specific foraging behaviors using a system that allowed me to see animals throughout the water column. Lone foraging animals produced all three sounds at significantly higher rates than animals foraging in groups, and the rate of sound production per animal in multi-animal foraging groups did not vary even as the groups reached up to five individuals. Production of echolocation and pops by lone foraging animals accounted for much of the difference. Foraging dolphins also displayed habitat-specific use of particular sound types. They preferentially produced echolocation and pops in the sand habitat and, at least for lone animals, in the seagrass edge habitat.     

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.     

Dual Function of Chip Calls Depending on Changing Call Rate Related to Risk Level in Territorial Pairs of White‐Eared Ground‐Sparrows

Different mechanisms have been proposed for encoding information into vocalizations: variation of frequency or temporal characteristics, variation in the rate of vocalization production, and use of different vocalization types. We analyze the effect of rate variation on the dual function of chip calls (contact and alarm) produced by White‐eared Ground‐sparrows (Melozone leucotis). We conducted an acoustic playback experiment where we played back 1 min of four chip call rates (12, 36, 60, 84 calls/min). We measured the response of territorial pairs using behavioral responses, and fine structural features of calls produced in response to those playbacks. White‐eared Ground‐sparrows showed more intense behavioral responses to higher than lower call rate playbacks. Both individuals of the pair approached the source of the playback stimulus faster, produced the first vocalization faster, produced more vocalizations, and spent more time close to the stimulus in higher call rate than in lower call rate playbacks. Frequency and duration characteristics of calls (chip and tseet) were similar in response to all call rate playbacks. Our playback experiment elicited different intensity of behavioral responses, suggesting that risk‐based information is encoded in call rate. Our results suggest that variation in the rate of chip call production serves a dual function in this species; calls are used at lower rates for pair contact and at higher rates for alarm/mobbing signals.     

Long-term passive acoustics to assess spatial and temporal vocalization patterns of Atlantic common bottlenose dolphins (Tursiops truncatus) in the May River estuary,South Carolina

Passive acoustics has been used extensively to study bottlenose dolphins; yet very few studies have examined the spatial, temporal, and environmental influences on vocalization types (echolocation, burst pulse sounds, and whistles), and few are long-term and provide high temporal resolution over multiple years. We used data from 2013 to 2018 to establish baseline acoustic patterns for bottlenose dolphins in the May River estuary, South Carolina. We deployed acoustic recorders at six stations during 2013–2014 and three stations during 2015–2018, with locations spanning the entire estuary (headwaters to the mouth). We discovered that acoustic detection of dolphins varied not only spatially, but also yearly, monthly, and tidally. Higher numbers of echolocation bouts, burst pulse sounds, and whistles were detected at the mouth as compared to the headwaters. At the mouth, vocalization detections were greatest in fall and winter for multiple years, and echolocation detection was greatest during falling and low tides. This study provides an example of another tool, long-term passive acoustics monitoring, to better understand spatial and temporal distribution of dolphins in a typical salt marsh estuary, that can be applied to other ecosystems throughout the southeastern United States and globally.     

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.     

Communication in bottlenose dolphins: 50 years of signature whistle research

Bottlenose dolphins (Tursiops truncatus) produce individually distinctive signature whistles that broadcast the identity of the caller. Unlike voice cues that affect all calls of an animal, signature whistles are distinct whistle types carrying identity information in their frequency modulation pattern. Signature whistle development is influenced by vocal production learning. Animals use a whistle from their environment as a model, but modify it, and thus invent a novel signal. Dolphins also copy signature whistles of others, effectively addressing the whistle owner. This copying occurs at low rates and the resulting copies are recognizable as such by parameter variations in the copy. Captive dolphins can learn to associate novel whistles with objects and use these whistles to report on the presence or absence of the object. If applied to signature whistles, this ability would make the signature whistle a rare example of a learned referential signal in animals. Here, we review the history of signature whistle research, covering definitions, acoustic features, information content, contextual use, developmental aspects, and species comparisons with mammals and birds. We show how these signals stand out amongst recognition calls in animals and how they contribute to our understanding of complexity in animal communication.     

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.     

Ecological and Social Factors Affecting the Vocal Behavior of Adult Female Squirrel Monkeys

Focal recordings of the vocalizations of squirrel monkeys, Saimiri sciureus, occupying an undisturbed Peruvian habitat were collected to evaluate the importance of both the ecological and social functions of female vocal behavior. The rates and sequences of six call types were examined by context: single, double, and multiple chuck, peep, tweet, and tweet-chuck. In contrast to laboratory studies, our findings emphasize the primacy of ecological functions in the wild, where calling permits females to operate as a convoy of spatially separated individuals in an arboreal habitat where visual contact is limited. Chuck calls are probably best interpreted as contact calls. The rate of chucks and tweet-chucks increased when the nearest adult female was > 5 m away. However, call production did not predict a change in the spatial separation between a female and her nearest adult female. During travel, the rate of single and double chucks was greater and the rate of multiple chucks was reduced. No direct relationship was found between foraging activities and vocal behavior. Although secondary, social factors did have a subtle effect on vocal behavior: the larger the social alliance of a female, the lower the rate and repetitiveness of her chuck vocalizations. We also consider the role of variations in vocalization rates among primates and the differing conclusions of field and captive studies of squirrel monkey vocal behavior.     

Primate phonation: Anterior cingulate lesion effects on response rate and acoustical structure

Four juvenile rhesus monkeys (Macaca mulatta) that were conditioned to emit a discriminative vocal response underwent unilateral and bilateral lesions of anterior cingulate gyrus in stepwise sequence. Pre- and post-lesion measures of vocal response rate and acoustical features (call duration, amplitude, and fundamental frequency) were obtained for the conditioned task and for vocalization in the home colony (spontaneous vocalization). Unilateral lesions produced little consistent change in spontaneous “coo” vocalization rate and call duration. Conditioned vocalization of the monkeys exhibited no significant change in rate at this stage, although call duration was reduced significantly. Bilateral lesions produced no further modifications in acoustic properties of spontaneous calls, while the vocal rate decreased slightly. Conditioned calls were not significantly altered in acoustic features at this stage, although the discriminative vocalization rate was significantly decreased for all monkeys. The results indicate limited control over vocal motor systems by anterior cingulate cortex, while suggesting that this region participates in initiation of voluntary phonation.     

DIRECTIONALITY IN THE WHISTLES OF HAWAIIAN SPINNER DOLPHINS (STENELLA LONGIROSTRIS): A SIGNAL FEATURE TO CUE DIRECTION OF MOVEMENT?

Dolphins produce frequency modulated (FM) whistles that are thought to promote the synchrony and coordination of behavior between members of a group. How whistles are used in this regard remains poorly understood. One possibility is that whistles have directionality and thereby convey the orientation and direction of movement of the signaler to nearby listeners. To explore this possibility, whistles from free-ranging Hawaiian spinner dolphins ( Stenella longirostris ) were obtained using a towed, three-hydrophone line array and examined for the presence of directionality. Both the estimated source level and harmonic content of whistles produced by animals traveling with or toward the array were greater than those of animals moving ahead or away from it. In addition, signals produced by animals near the array (within 20 m) were received differently on the three hydrophones spaced 11.5 m apart. These differences were greater than would be expected from transmission loss disparities alone. The results indicate that directivity is present in the transmission pattern of whistles. To infer the form of this directivity, a theoretical whistle beam pattern was established based on the assumption that the dolphin's sound source is approximated by a circular piston transducer (Au 1993). The resulting beam indicates that spinner dolphin whistles become increasingly directional with frequency, especially with respect to harmonics. The orientation-dependent harmonic structure of whistles thus presents a potential cue that listening animals could interpret to infer the direction of movement of signalers. Harmonics are present in the whistles of many dolphin species and may represent an inherent signal design feature that promotes coordination between animals.     

Subsurface behavior of bottlenose dolphins (Tursiops truncatus) interacting with fish trawl nets in northwestern Australia: Implications for bycatch mitigation

Most studies of delphinid‐trawler interactions have documented the surface behavior of dolphins feeding on discarded bycatch, but not their subsurface behavior around demersal trawl gear. Using video cameras mounted inside trawl nets, we recorded the subsurface behavior of common bottlenose dolphins (Tursiops truncatus) in a demersal fish trawl fishery in northwestern Australia. Footage from 36 trawls across the fishery was analyzed to determine the extent of dolphin‐gear interactions and the behavior of dolphins inside the nets. Interaction rates were high, with dolphins present inside and outside the nets during 29 and 34 trawls, respectively, and for up to 99% of the trawl duration. The proportion of foraging behaviors exhibited inside the nets was higher than the proportions of traveling and socializing behaviors. Twenty‐nine individuals were identified inside the net, seven of which returned repeatedly within and between trawls and fishing trips, but were observed primarily in the same localized areas in which they were first recorded. Our results suggest that entering trawl nets may be a frequently occurring, yet specialized behavior exhibited by a small subset of trawler‐associated dolphins. We propose that gear modifications, not spatial or temporal adjustments to fishing effort, have the greatest potential to reduce dolphin bycatch.     

The units of perception in the antiphonal calling behavior of cotton-top tamarins (Saguinus oedipus): playback experiments with long calls

We investigated how the acoustic structure of the cotton-top tamarin monkey's (Saguinus oedipus) combination long call relates to the antiphonal calling behavior of conspecifics. Combination long calls can function as contact calls and are produced by socially isolated individuals. Often conspecifics respond to these calls with their own long calls. Structurally, these calls are always composed of one or more 'chirps' followed by two or more 'whistles'. We compared the antiphonal calling responses to playbacks of complete, naturally produced long calls versus single whistles or single chirps. Subjects responded significantly more to whole calls than to either syllable-type alone. Thus, our data suggest that, in terms of the antiphonal calling behavior of socially isolated conspecifics, the whole long call is the unit of perception.