Article

Contexts of emission of non-signature whistles in bottlenose dolphins (Tursiops truncatus) under human care

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Abstract

Bottlenose dolphins are social cetaceans that strongly rely on acoustic communication and signaling. The diversity of sounds emitted by the species has been structurally classified into whistles, clicks and burst-pulsed sounds. Although click sounds and individually-specific signature whistles have been largely studied, not much is known about non-signature whistles. Most studies that link behavior and whistle production conduct aerial behavioral observations and link the production of whistles to the general category of social interactions. The aim of this study was to determine if there was a correlation between the non-signature whistle production and the underwater behaviors of a group of bottlenose dolphins (Tursiops truncatus) under human care, during their free time in the absence of trainers. To do this we made audio-video recordings 15 minutes before and after 10 training sessions of eight dolphins in Boudewijn Seapark (Belgium). For the behavioral analysis we conducted focal follows on each individual based on six behavioral categories. For the acoustical analysis, carried out at the group level, we used the SIGID method to identify non-signature whistles (N = 661) and we classified them in six categories according to their frequency modulation. The occurrences of the six categories of whistles were highly collinear. Most importantly, non-signature whistle production was positively correlated with the time individuals spent slow swimming alone, and was negatively correlated with the time spent in affiliative body contact. This is the first analysis that links the production of non-signature whistles with particular underwater behaviors in this species.

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... Physiological and health-related welfare parameters such as stress hormones [86,89,90], blood profiles [91,92], and pulmonary function [93] have also been examined in captive animals. The controlled environment of the captive setting has also aided in the development of certain hydrophone-video arrays aiming to match cetacean vocalisations to behaviours, thereby decoding their meaning, which would be a critical step in evaluating cetacean welfare [94,95]. However, much is still unknown about the utility of captive welfare findings to wild marine mammals. ...
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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.
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Seven major types of sampling for observational studies of social behavior have been found in the literature. These methods differ considerably in their suitability for providing unbiased data of various kinds. Below is a summary of the major recommended uses of each technique: In this paper, I have tried to point out the major strengths and weaknesses of each sampling method. Some methods are intrinsically biased with respect to many variables, others to fewer. In choosing a sampling method the main question is whether the procedure results in a biased sample of the variables under study. A method can produce a biased sample directly, as a result of intrinsic bias with respect to a study variable, or secondarily due to some degree of dependence (correlation) between the study variable and a directly-biased variable. In order to choose a sampling technique, the observer needs to consider carefully the characteristics of behavior and social interactions that are relevant to the study population and the research questions at hand. In most studies one will not have adequate empirical knowledge of the dependencies between relevant variables. Under the circumstances, the observer should avoid intrinsic biases to whatever extent possible, in particular those that direcly affect the variables under study. Finally, it will often be possible to use more than one sampling method in a study. Such samples can be taken successively or, under favorable conditions, even concurrently. For example, we have found it possible to take Instantaneous Samples of the identities and distances of nearest neighbors of a focal individual at five or ten minute intervals during Focal-Animal (behavior) Samples on that individual. Often during Focal-Animal Sampling one can also record All Occurrences of Some Behaviors, for the whole social group, for categories of conspicuous behavior, such as predation, intergroup contact, drinking, and so on. The extent to which concurrent multiple sampling is feasible will depend very much on the behavior categories and rate of occurrence, the observational conditions, etc. Where feasible, such multiple sampling can greatly aid in the efficient use of research time.
Article
This study examined whether a group of captive dolphins displayed anticipatory behaviors before shows. In general, anticipation occurs when an event is being predicted. Anticipatory behavior is defined by Spruijt et al. as "responses elicited by rewarding stimuli that lead to and facilitate consummatory behavior (Spruijt et al., 2001, Appl Anim Behav Sci 72: 145-171)." Using behavioral recording techniques, the behaviors, breathing rates, space use, and activity levels of all dolphins was recorded both before and after shows. Analysis compared pre- and post-show data in addition to looking at gradual changes in behavior prior to show sessions. Significant changes were found in the behavior and space use prior to sessions with the dolphins decreasing their activity levels, spending more time at the surface and moving towards the starting point of a session before it took place. There was a significant increase in the vigilant behavior before sessions, indicating that the dolphins were becoming more alert towards their trainers and other activities around the pool. This result mirrors previous research with other captive species; as feeding time was approaching, the animals seemed to "wait" and look for the handlers. Any behavioral change that may be regarded as anticipatory behavior was not evidently abnormal or stereotypic in nature, and breathing rates remained stable indicating that the animals do not perceive the shows as stressful or as an aversive experience. Additionally, behavior and level of activity remained stable following the sessions. Zoo Biol. XX:XX-XX, 2013. © 2013 Wiley Periodicals Inc.
Article
IN 1953, Essapian1 suggested that individual bottle-nosed dolphins, Tursiops truncatus (Montagu), may have distinctive notes which each dolphin can recognize. From his context, in using the word `notes' Essapian referred to the whistle component of Tursiops phonation.
Article
Behavioral scientists have developed methods for sampling behavior in order to reduce observational biases and to facilitate comparisons between studies. A review of 74 cetacean behavioral field studies published from 1989 to 1995 in Marine Mammal Science and The Canadian Journal of Zoology suggests that cetacean researchers have not made optimal use of available methodology. The survey revealed that a large proportion of studies did not use reliable sampling methods. Ad libitum sampling was used most often (59%). When anecdotal studies were excluded, 45% of 53 behavioral studies used ad libitum as the predominant method. Other sampling methods were continuous, onezero, incident, point, sequence, or scan sampling. Recommendations for sampling methods are made, depending on identifiability of animals, group sizes, dive durations, and change in group membership.
Article
The prevailing view among researchers of dolphin communication is that bottlenose dolphins possess an individualized whistle contour; known as the ‘signature whistle’, it accounts for 74–95 % of a dolphin's whistle repertoire and functions to signal the individual identity of the whistler. This study used a new quantitative technique, termed the contour similarity technique (CS technique), and reports on the quantitative comparison of whistles from the individuals of three different social groups of bottlenose dolphins in socially interactive contexts. Results suggest that captive adult dolphins share several different whistle types including one predominant whistle type shared by all individuals across three different social groups. These analyses suggest a different interpretation of the dolphin whistle repertoire than has previously been proposed by proponents of the signature whistle hypothesis. In addition, results from our study support the results of early studies, published before the advent of the signature whistle hypothesis, in which investigators reported a large whistle repertoire within socially interactive captive and free-ranging groups and a predominant whistle type similar to that found in our study. Our results, combined with the results from earlier studies of dolphin vocal behaviour, suggest that the signature whistle hypothesis is incomplete and that dolphin whistle repertoires need to be analysed with respect to behavioural context and social relationships. In addition, these results suggest that contour discrimination and other acoustic features of whistles need to be tested in perception and categorization experiments.
Article
Delphinid communication has been shaped by the marine environment. This resulted in specific adaptations such as echolocation and a sophisticated communication system that allows animals to maintain contact over several kilometers even if no other cues are available. The communication system of delphinids is characterized by large call repertoires, recognition calls shaped by vocal learning, and a great plasticity of the vocal repertoire. Delphinids also display complex cognitive skills that influence how they use communication signals. Complex social systems provide opportunities to apply these skills. Most of our knowledge on delphinid communication comes from studies on bottlenose dolphins and killer whales. Future studies need to focus on additional species and try to assess the threat imposed by anthropogenic noise on the communication behavior of delphinids.
Article
Bottlenose dolphin (Tursiops sp.) calves are unusual among social mammals in that they have a have prolonged nursing period (3–8 years), but precocious motor ability, enabling them to separate from their mothers and form distinct social bonds at an early age. We examined two measures of calf sociality from birth to 4 years of age: (1) the number of unique associates and (2) the proportion of time that calves spent in groups with nonmothers. Calves (N = 89) associated with a mean ± SE of 27.56 ± 2.24 individuals (range 0–77) and spent 46.84 ± 3.00% of their time in groups (range 0–100%). Mothers (N = 49) averaged 39.39 ± 5.32 associates (range 0–139), when data were combined across all years (and often multiple offspring). Both calf characteristics (sex, age and separation time) and maternal characteristics (sociality and foraging time) contributed to this variation. Although calf associate number and time in groups were positively correlated, sex and age-specific patterns differed depending on the measure used. As separation time increased, both sexes increased associate number, but females decreased and males increased time in groups, indicating that males sought more social contact. Maternal socioecological strategy largely contributed to calf social development, particularly for daughters. As maternal foraging time increased, maternal and calf sociality decreased. Second, the number of associates that calves had when with their mothers predicted associate number during separations, but time spent in groups was not similarly correlated. These early patterns probably influence subsequent social development, including the structure of female networks and male alliances.
Article
The intricate and highly developed acoustic communication system of bottlenose dolphins reflects the complexities of their social organization. Indo-Pacific bottlenose dolphins (Tursiops aduncus) produce numerous types of acoustic emissions, including a diverse repertoire of whistles used for communicative purposes. The influence of group composition on whistle production and the function of different whistles produced by dolphins in wild contexts are relatively unknown. Recordings of acoustic emissions and behavior of dolphins were made concurrently during vessel-based surveys along the coast of northern New South Wales, Australia. Whistles were divided into five tonal classes (sine, rise, down-sweep, flat, and concave) and categorized into distinct whistle types. It is shown that while whistle repetition rate and whistle diversity was influenced by group composition, it is not influenced by behavior. Noncalf groups produced a significantly higher whistle repetition rate and whistle diversity than calf groups. In contrast, the types of whistles produced were related to the behavior in which the dolphins were engaged in: some tonal classes and distinct whistle types were related to different behavior states. Findings suggested that some whistle types may be used to communicate specific information on the behavioral context of the individuals involved.
Article
"Part review, part testament to extraordinary dedication, and part call to get involved, Cetacean Societies highlights the achievements of behavioral ecologists inspired by the challenges of cetaceans and committed to the exploration of a new world."—from the preface by Richard Wrangham Long-lived, slow to reproduce, and often hidden beneath the water's surface, whales and dolphins (cetaceans) have remained elusive subjects for scientific study even though they have fascinated humans for centuries. Until recently, much of what we knew about cetaceans came from commercial sources such as whalers and trainers for dolphin acts. Innovative research methods and persistent efforts, however, have begun to penetrate the depths to reveal tantalizing glimpses of the lives of these mammals in their natural habitats. Cetacean Societies presents the first comprehensive synthesis and review of these new studies. Groups of chapters focus on the history of cetacean behavioral research and methodology; state-of-the-art reviews of information on four of the most-studied species: bottlenose dolphins, killer whales, sperm whales, and humpback whales; and summaries of major topics, including group living, male and female reproductive strategies, communication, and conservation drawn from comparative research on a wide range of species. Written by some of the world's leading cetacean scientists, this landmark volume will benefit not just students of cetology but also researchers in other areas of behavioral and conservation ecology as well as anyone with a serious interest in the world of whales and dolphins. Contributors are Robin Baird, Phillip Clapham, Jenny Christal, Richard Connor, Janet Mann, Andrew Read, Randall Reeves, Amy Samuels, Peter Tyack, Linda Weilgart, Hal Whitehead, Randall S. Wells, and Richard Wrangham.
Article
This paper presents a general statistical methodology for the analysis of multivariate categorical data arising from observer reliability studies. The procedure essentially involves the construction of functions of the observed proportions which are directed at the extent to which the observers agree among themselves and the construction of test statistics for hypotheses involving these functions. Tests for interobserver bias are presented in terms of first-order marginal homogeneity and measures of interobserver agreement are developed as generalized kappa-type statistics. These procedures are illustrated with a clinical diagnosis example from the epidemiological literature.
Article
Studies on captive bottlenose dolphins, Tursiops truncatus, have shown that each individual produces a stereotyped, individually specific signature whistle; however, no study has demonstrated clear context-dependent usage of these whistles. Thus, the hypothesis that signature whistles are used to maintain group cohesion remains untested. To investigate whether signature whistles are used only in contexts that would require a mechanism to maintain group cohesion, we examined whistle type usage in a group of four captive bottlenose dolphins in two contexts. Individuals were recorded while they were separate from the group and while they all swam in the same pool. Separations occurred spontaneously when one animal swam into another pool. No partitions were used and no aggressive interactions between dolphins preceded separations. Calling animals were identified by an amplitude comparison of the same sound recorded in the two pools. Each dolphin primarily produced one stereotyped signature whistle when it was separated from the group. Similarly the remaining group in the other pool also used primarily their signature whistles if one animal was in a separate pool. If all animals swam in the same pool almost only nonsignature whistles were used. Signature whistle copying was rare and did not initiate reunions or specific vocal responses. The results strongly support the hypothesis that signature whistles are used to maintain group cohesion. Copyright 1998 The Association for the Study of Animal Behaviour.
Article
Dolphin communication is suspected to be complex, on the basis of their call repertoires, cognitive abilities, and ability to modify signals through vocal learning. Because of the difficulties involved in observing and recording individual cetaceans, very little is known about how they use their calls. This report shows that wild, unrestrained bottlenose dolphins use their learned whistles in matching interactions, in which an individual responds to a whistle of a conspecific by emitting the same whistle type. Vocal matching occurred over distances of up to 580 meters and is indicative of animals addressing each other individually.
Article
Bottlenose dolphins exhibit complex social affiliations that may be shaped by interactions among individuals. Affiliative body contact among dolphins may repair deteriorated relationships or reduce tension within the group following aggressive interactions. We investigated the time-series association between one type of contact behavior (flipper-rubbing) and aggression by continuous observation of three captive bottlenose dolphins. For all three dolphin pairs, the elapsed time to aggressive events was significantly greater following flipper-rubbing. In two dolphin pairs comprised of a young male and an adult female, one-zero score of inter-opponent flipper-rubbing was higher for 10 min following aggressive bouts (post-AG periods) than for the same length of control (Ctrl) periods. For all three focal pairs, one-zero score of third-party rubbing was higher for post-AG than Ctrl periods. Neither the direction of rubbing nor the identity of the partner that approached prior to rubbing showed any significant tendencies. Flipper-rubbing may contribute to restore friendly relationships between former opponents or reduce conflicts in at least juvenile-adult female associations. Our results also give preliminary suggestions of the functions of third-party flipper-rubbing among bottlenose dolphins, including tension easing by the third party, or displacement as a result of aggressive interactions.