Laela S. Sayigh

Signature whistles of free-ranging bottlenose dolphins Tursiops truncatus: stability and mother-offspring comparisons

SummaryMother-calf whistle exchanges were recorded from temporarily captured free-ranging bottlenose dolphins from 1975 to 1989. This is part of a long-term research project studying social structure and behavior of a community of approximately 100 dolphins in waters near Sarasota, Florida. Analysis of whistle exchanges from 12 mothercalf pairs shows that signature whistles can remain stable for periods up to at least 12 years. We looked for effects of vocal learning on the development of the signature whistle by comparing whistles of calves to those of their mothers. Eight female calves produced whistles distinct from those of their mothers, while four male calves produced whistles similar to those of their mothers. Male calves appeared to produce a greater proportion of whistles other than the signature whistle (termed “variants”). We hypothesize that these sex differences in whistle vocalizations may reflect differences in the roles males and females play in the social structure of the community.

Sex difference in signature whistle production of free-ranging bottlenose dolphins, Tursiops truncates

Signature whistles of 42 free-ranging bottle-nose dophin calves were compared to those of their mothers. Humans judged their similarity by inspection of spectrograms. There was a sex difference in the tendency of calves to produce whistles similar to or different from those of their mothers; most female calves produced whistles that were different from those of their mothers, whereas male calves were more likely to produce whistles that were similar to those of their mothers. Because matrilineally related females associate together and use signature whistles to establish and/or maintain contact with their calves, there may be a selective pressure for females to produce whistles that are distinct from those of their mothers. There may be fewer constraints governing whistle development in males, with the result that some males produce whistles similar to those of their mothers and others do not.

Signature-whistle production in undisturbed free-ranging bottlenose dolphins (Tursiops truncatus).

Data from behavioural observations and acoustic recordings of free–ranging bottlenose dolphins (Tursiops truncatus) were analysed to determine whether signature whistles are produced by wild undisturbed dolphins, and how whistle production varies with activity and group size. The study animals were part of a resident community of bottlenose dolphins near Sarasota, Florida, USA. This community of dolphins provides a unique opportunity for the study of signature–whistle production, since most animals have been recorded during capture–release events since 1975. Three mother–calf pairs and their associates were recorded for a total of 141.25 h between May and August of 1994 and 1995. Whistles of undisturbed dolphins were compared with those recorded from the same individuals during capture–release events. Whistles were conservatively classified into one of four categories: signature, probable signature, upsweep or other. For statistical analyses, signature and probable signature whistles were combined into a ‘signature’ category; upsweep and other whistles were combined into a ‘non–signature’ category. Both ‘signature’ and ‘non–signature’ whistle frequencies significantly increased as group size increased. There were significant differences in whistle frequencies across activity types: both ‘signature’ and ‘non–signature’ whistles were most likely to occur during socializing and least likely to occur during travelling. There were no significant interactions between group size and activity type. Signature and probable signature whistles made up ca. 52% of all whistles produced by these free–ranging bottlenose dolphins.

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.

Whistles as Potential Indicators of Stress in Bottlenose Dolphins (Tursiops truncatus)

Abstract We examined the possibility that parameters of bottlenose dolphin signature whistles may serve as indicators of stress. Bottlenose dolphins (Tursiops truncatus) in Sarasota Bay, Florida, were recorded during brief capture–release events, which are potentially a source of short-term stress to these dolphins, although no effects of chronic or long-term stress have been observed over the 37+-year duration of the research. Whistles recorded during both brief capture–release and undisturbed, free-ranging conditions were examined to determine whether whistle parameters differ during capture–release versus undisturbed conditions; at the beginning of a capture–release session versus at the end of a session; during an individuals 1st capture–release session versus later capture–release sessions; and when a mother is caught and released with a dependent calf versus without a dependent calf (i.e., she has no dependent calf at the time of capture–release). We examined a variety of acoustic parameters, including whistle rate, number of loops (repetitive elements), maximum and minimum frequency, and loop, interloop, and whistle duration. We found that whistle rate and number of loops were greater during brief capture–release events than during undisturbed conditions; number of loops decreased and loop duration increased over the duration of a capture–release session; whistle rates decreased with number of capture–release sessions; and females caught and released with dependent calves produced whistles with higher maximum frequencies and shorter interloop intervals than when they did not have dependent calves. Thus, whistles appear to have potential as noninvasive indicators of stress in bottlenose dolphins. Further research is warranted in this area, for example by correlating physiological indices to whistle rates under varying levels of stress. Reliable, noninvasive correlates of stress could be used to monitor dolphins in a variety of circumstances, such as during exposure to anthropogenic noise.

Acoustic sequences in non-human animals: a tutorial review and prospectus

Animal acoustic communication often takes the form of complex sequences, made up of multiple distinct acoustic units. Apart from the well‐known example of birdsong, other animals such as insects, amphibians, and mammals (including bats, rodents, primates, and cetaceans) also generate complex acoustic sequences. Occasionally, such as with birdsong, the adaptive role of these sequences seems clear (e.g. mate attraction and territorial defence). More often however, researchers have only begun to characterise – let alone understand – the significance and meaning of acoustic sequences. Hypotheses abound, but there is little agreement as to how sequences should be defined and analysed. Our review aims to outline suitable methods for testing these hypotheses, and to describe the major limitations to our current and near‐future knowledge on questions of acoustic sequences. This review and prospectus is the result of a collaborative effort between 43 scientists from the fields of animal behaviour, ecology and evolution, signal processing, machine learning, quantitative linguistics, and information theory, who gathered for a 2013 workshop entitled, ‘Analysing vocal sequences in animals’. Our goal is to present not just a review of the state of the art, but to propose a methodological framework that summarises what we suggest are the best practices for research in this field, across taxa and across disciplines. We also provide a tutorial‐style introduction to some of the most promising algorithmic approaches for analysing sequences. We divide our review into three sections: identifying the distinct units of an acoustic sequence, describing the different ways that information can be contained within a sequence, and analysing the structure of that sequence. Each of these sections is further subdivided to address the key questions and approaches in that area. We propose a uniform, systematic, and comprehensive approach to studying sequences, with the goal of clarifying research terms used in different fields, and facilitating collaboration and comparative studies. Allowing greater interdisciplinary collaboration will facilitate the investigation of many important questions in the evolution of communication and sociality.

Vocal copying of individually distinctive signature whistles in bottlenose dolphins

Vocal learning is relatively common in birds but less so in mammals. Sexual selection and individual or group recognition have been identified as major forces in its evolution. While important in the development of vocal displays, vocal learning also allows signal copying in social interactions. Such copying can function in addressing or labelling selected conspecifics. Most examples of addressing in non-humans come from bird song, where matching occurs in an aggressive context. However, in other animals, addressing with learned signals is very much an affiliative signal. We studied the function of vocal copying in a mammal that shows vocal learning as well as complex cognitive and social behaviour, the bottlenose dolphin (Tursiops truncatus). Copying occurred almost exclusively between close associates such as mother–calf pairs and male alliances during separation and was not followed by aggression. All copies were clearly recognizable as such because copiers consistently modified some acoustic parameters of a signal when copying it. We found no evidence for the use of copying in aggression or deception. This use of vocal copying is similar to its use in human language, where the maintenance of social bonds appears to be more important than the immediate defence of resources.

The Encoding of Individual Identity in Dolphin Signature Whistles: How Much Information Is Needed?

Bottlenose dolphins (Tursiops truncatus) produce many vocalisations, including whistles that are unique to the individual producing them. Such “signature whistles” play a role in individual recognition and maintaining group integrity. Previous work has shown that humans can successfully group the spectrographic representations of signature whistles according to the individual dolphins that produced them. However, attempts at using mathematical algorithms to perform a similar task have been less successful. A greater understanding of the encoding of identity information in signature whistles is important for assessing similarity of whistles and thus social influences on the development of these learned calls. We re-examined 400 signature whistles from 20 individual dolphins used in a previous study, and tested the performance of new mathematical algorithms. We compared the measure used in the original study (correlation matrix of evenly sampled frequency measurements) to one used in several previous studies (similarity matrix of time-warped whistles), and to a new algorithm based on the Parsons code, used in music retrieval databases. The Parsons code records the direction of frequency change at each time step, and is effective at capturing human perception of music. We analysed similarity matrices from each of these three techniques, as well as a random control, by unsupervised clustering using three separate techniques: k-means clustering, hierarchical clustering, and an adaptive resonance theory neural network. For each of the three clustering techniques, a seven-level Parsons algorithm provided better clustering than the correlation and dynamic time warping algorithms, and was closer to the near-perfect visual categorisations of human judges. Thus, the Parsons code captures much of the individual identity information present in signature whistles, and may prove useful in studies requiring quantification of whistle similarity.

Repeated call types in Hawaiian melon-headed whales (Peponocephala electra)

Melon-headed whales are pantropical odontocetes that are often found near oceanic islands. While considered sound-sensitive, their bioacoustic characteristics are relatively poorly studied. The goal of this study was to characterize the vocal repertoire of melon-headed whales to determine whether they produce repeated calls that could assist in recognition of conspecifics. The first tag-based acoustic recordings of three melon-headed whales were analyzed. Tag records were visually and aurally inspected and all calls were individually extracted. Non-overlapping calls with sufficient signal-to-noise were then parameterized and visually grouped into categories of repeated call types. Thirty-six call categories emerged. Categories differed significantly in duration, peak and centroid frequency, and -3 dB bandwidth. Calls of a given type were more likely to follow each other than expected. These data suggest that repeated calls may function in individual, subgroup, or group recognition. Repeated call production could also serve to enhance signal detection in large groups with many individuals producing simultaneous calls. Results suggest that caution should be used in developing automatic classification algorithms for this species based on small sample sizes, as they may be dominated by repeated calls from a few individuals, and thus not representative of species- or population-specific acoustic parameters.

Dolphin Signature Whistles

The term ‘signature’ has often been applied to animal vocalizations when an individually distinctive pattern was found in them. The vast majority of animals achieve this by means of voice cues, which result from individual variability in the shape and size of the vocal tract. Dolphin signature whistles are qualitatively different from most individually distinctive signals seen in other mammalian species. Identity is encoded in a frequency modulation pattern that is learned or invented early in life. These whistles are used in individual recognition and in maintaining group cohesion and seem to function similarly to human names.