First acoustic evidence of signature whistle production by spinner dolphins (Stenella longirostris).

A dolphin's signature whistle (SW) is a distinctive acoustic signal, issued in a bout pattern of unique frequency modulation contours; it allows individuals belonging to a given group to recognize each other and, consequently, to maintain contact and cohesion. The current study is the first scientific evidence that spinner dolphins (Stenella longirostris) produce SWs. Acoustic data were recorded at a shallow rest bay called "Biboca", in Fernando de Noronha Archipelago, Brazil. In total, 1902 whistles were analyzed; 40% (753/1,902) of them were classified as stereotyped whistles (STW). Based on the SIGID method, 63% (472/753) of all STWs were identified as SWs; subsequently, they were categorized into one of 18 SW types. SWs accounted for 25% (472/1,902) of the acoustic repertoire. External observers have shown near perfect agreement to classify whistles into the adopted SW categorization. Most acoustic and temporal variables measured for SWs showed mean values similar to those recorded in other studies with spinner dolphins, whose authors did not differentiate SWs from non-SWs. Principal component analysis has explained 78% of total SW variance, and it emphasized the relevance of shape/contour and frequency variables to SW variance. This scientific discovery helps improving bioacoustics knowledge about the investigated species. Future studies to be conducted in Fernando de Noronha Archipelago should focus on continuous investigations about SW development and use by S. longirostris, expanding individuals' identifications (Photo ID and SW Noronha Catalog), assessing long-term whistle stability and emission rates, and making mother-offspring comparisons with sex-based differences.

Understanding across the senses: cross-modal studies of cognition in cetaceans.

Cross-modal approaches to the study of sensory perception, social recognition, cognition, and mental representation have proved fruitful in humans as well as in a variety of other species including toothed whales in revealing equivalencies that suggest that different sensory stimuli associated with objects or individuals may effectively evoke mental representations that are, respectively, object based or individual based. Building on established findings of structural equivalence in the form of spontaneous recognition of complex shapes across the modalities of echolocation and vision and behavior favoring identity echoic-visual cross-modal relationships over associative echoic-visual cross-modal relationships, examinations of transitive inference equivalencies from initially learned associations of visual and acoustic stimuli, and recent work examining spontaneous cross-modal social recognition of individual identity across acoustic and gustatory chemical modalities (i.e., the equivalence relationships among an individual's characteristics), we examine the history, utility and implications for cross-modal research in cetacean cognition. Drawing from research findings on bottlenose dolphins and beluga whales as well as other species we suggest future directions for cetacean cross-modal research to further illuminate understanding how structural and individual sensory equivalencies lead to object-centered and individual-centered mental representations, as well as to explore the potential for practical applications related to cetacean conservation.

Context-dependent and seasonal fluctuation in bottlenose dolphin (Tursiops truncatus) vocalizations.

A fundamental question in animal behaviour is the role of vocal communication in the regulation of social interactions in species that organise themselves into social groups. Context dependence and seasonality in vocalizations are present in the communication of many species, although very little research has addressed this dependence in marine mammals. The study presented here examined variations in the rate at which free-ranging dyads of bottlenose dolphins emit social-signals in an effort to better understand the relationship between vocal communication and social context. The results demonstrate that changes in the social-signal production in bottlenose dolphins are related to the sex of the partner, mating season and social affiliation between the components of the dyad. In a context of foraging behaviour on the same feeding ground, mixed (male-female) dyads were found to emit more pulsed burst sounds during the mating season. Another relevant aspect of the study seems to be the greater production of agonistic social-signals in the dyads formed by individuals with a lower degree of social affiliation. Overall, this study confirms a clear relationship between dyad composition and context-specific social-signals that could reflect the motivational state of individuals linked to seasonal changes in vocal behaviour.

Evidence suggests vocal production learning in a cross-fostered Risso's dolphin (Grampus griseus).

Vocal learning is a rare skill in mammals, and we have limited information about the contexts in which they use it. Previous studies suggested that cetaceans in general are skilled at imitating sounds, but only few species have been studied to date. To expand this investigation to another species and to investigate the possible influence of the social environment on vocal learning, we studied the whistle repertoire of a female Risso's dolphin (Grampus griseus) that was stranded at an early age and was subsequently raised in a group of bottlenose dolphins (Tursiops truncatus). We show that this cross-fostered animal produced vocal signals more akin to those of its Tursiops poolmates than those of Risso's dolphins in the wild. This is one of very few systematic cross-fostering studies in cetaceans and the first to suggest vocal production learning in the Risso's dolphin. Our findings also suggest that social experience is a major factor in the development of the vocal repertoire in this species.

Observational study on the non-linear response of dolphins to the presence of vessels.

With the large increase in human marine activity, our seas have become populated with vessels that can be overheard from distances of even 20 km. Prior investigations showed that such a dense presence of vessels impacts the behaviour of marine animals, and in particular dolphins. While previous explorations were based on a linear observation for changes in the features of dolphin whistles, in this work we examine non-linear responses of bottlenose dolphins (Tursiops Truncatus) to the presence of vessels. We explored the response of dolphins to vessels by continuously recording acoustic data using two long-term acoustic recorders deployed near a shipping lane and a dolphin habitat in Eilat, Israel. Using deep learning methods we detected a large number of 50,000 whistles, which were clustered to associate whistle traces and to characterize their features to discriminate vocalizations of dolphins: both structure and quantities. Using a non-linear classifier, the whistles were categorized into two classes representing the presence or absence of a nearby vessel. Although our database does not show linear observable change in the features of the whistles, we obtained true positive and true negative rates exceeding 90% accuracy on separate, left-out test sets. We argue that this success in classification serves as a statistical proof for a non-linear response of dolphins to the presence of vessels.

Decades-long social memory in bottlenose dolphins.

Long-term social memory is important, because it is an ecologically relevant test of cognitive capacity, it helps us understand which social relationships are remembered and it relates two seemingly disparate disciplines: cognition and sociality. For dolphins, long-term memory for conspecifics could help assess social threats as well as potential social or hunting alliances in a very fluid and complex fission-fusion social system, yet we have no idea how long dolphins can remember each other. Through a playback study conducted within a multi-institution dolphin breeding consortium (where animals are moved between different facilities), recognition of unfamiliar versus familiar signature whistles of former tank mates was assessed. This research shows that dolphins have the potential for lifelong memory for each other regardless of relatedness, sex or duration of association. This is, to my knowledge, the first study to show that social recognition can last for at least 20 years in a non-human species and the first large-scale study to address long-term memory in a cetacean. These results, paired with evidence from elephants and humans, provide suggestive evidence that sociality and cognition could be related, as a good memory is necessary in a fluid social system.

Non-stereotyped amplitude modulation across signature whistle contours.

Bottlenose dolphin signature whistles are characterized by distinctive frequency modulation over time. The stable frequency contours of these whistles broadcast individual identity information. Little is known however, about whether or not the amplitude contour is also stereotyped. Here, we examined the relative amplitude-time contour of signature whistle emissions from eight bottlenose dolphins (Tursiops truncatus) in the U.S. Navy Marine Mammal Program (MMP) in San Diego, CA. The results suggested that unlike the stable frequency-time contour, the amplitude-time contour of signature whistles were largely non-stereotyped, characterized by large variability across multiple whistle emissions. Relative amplitude was negatively related to log peak frequency, with more energy focused in the lower frequency bands. This trend was consistent over all eight dolphins despite having quite different signature whistle contours. This relationship led to the amplitude contours being slightly more stereotyped within than between dolphins. We propose that amplitude across signature whistle emissions may serve as an avenue for encoding additional communicative information. We encourage future studies to incorporate analyses of amplitude contours in addition to frequency contours of signature whistles in order to begin to understand what role it may play in the dolphin communication system.

Signature whistles exhibit a 'fade-in' and then 'fade-out' pattern of relative amplitude declination.

Bottlenose dolphins have individually distinct signature whistles that are characterized by a stereotyped frequency-time contour. Signature whistles are commonly exchanged with short time delays between calls. Dolphin whistles are produced by pressurized nasal sacs that increase and then decrease in pressure over emission. This study found that the relative amplitude modulation pattern over time exhibited the same fade-in and then fade-out pattern in the signature whistles of eight bottlenose dolphins at the Navy in San Diego, CA. Both the initial and final five percent of the whistle's duration also had significantly lower mean relative amplitude than the center five percent. The current analyses of the amplitude-time relationship was then integrated to a previously reported model of the negative relationship between relative log amplitude and log peak frequency. This produced a more robust model for accounting for the predictable aspects of the more broadly non-stereotyped amplitude modulations of signature whistles. Whether dolphins can intentionally manipulate these amplitude features or they are simple by-products of the sound production system, and further whether they are perceived and utilized by receivers, is an exciting area for continued research.

Allied male dolphins use vocal exchanges to "bond at a distance".

Vocal interactions are intrinsic features of social groups and can play a pivotal role in social bonding.1,2 Dunbar's social bonding hypothesis posits that vocal exchanges evolved to "groom at a distance" when social groups became too large or complex for individuals to devote time to physical bonding activities.1,3 Tests of this hypothesis in non-human primates, however, suggest that vocal exchanges occur between more strongly bonded individuals that engage in higher grooming rates4-7 and thus do not provide evidence for replacement of physical bonding. Here, we combine data on social bond strength, whistle exchange frequency, and affiliative contact behavior rates to test this hypothesis in wild male Indo-Pacific bottlenose dolphins, who form multi-level alliances that cooperate over access to females.8-10 We show that, although whistle exchanges are more likely to occur within the core alliance, they occur more frequently between those males that share weaker social bonds, i.e., between core allies that spend less time together, while the opposite occurs for affiliative physical contact behavior. This suggests that vocal exchanges function as a low-cost mechanism for male dolphins that spend less time in close proximity and engage in fewer affiliative contact behaviors to reinforce and maintain their valuable alliance relationships. Our findings provide new evidence outside of the primate lineage that vocal exchanges serve a bonding function and reveal that, as the social bonding hypothesis originally suggested, vocal exchanges can function as a replacement of physical bonding activities for individuals to maintain their important social relationships.

Social and vocal complexity in bottlenose dolphins.

Bottlenose dolphins are highly social, renowned for their vocal flexibility, and possess highly enlarged brains relative to their body size. Here, we discuss some of the defining features of bottlenose dolphin social and vocal complexity and place this in the context of their cognitive evolution.

Species information in whistle frequency modulation patterns of common dolphins.

The most flexible communication systems are those of open-ended vocal learners that can acquire new signals throughout their lifetimes. While acoustic signals carry information in general voice features that affect all of an individual's vocalizations, vocal learners can also introduce novel call types to their repertoires. Delphinids are known for using such learned call types in individual recognition, but their role in other contexts is less clear. We investigated the whistles of two closely related, sympatric common dolphin species, Delphinus delphis and Delphinus bairdii, to evaluate species differences in whistle contours. Acoustic recordings of single-species groups were obtained from the Southern California Bight. We used an unsupervised neural network to categorize whistles and compared the resulting whistle types between species. Of the whistle types recorded in more than one encounter, 169 were shared between species and 60 were species-specific (32 D. delphis types, 28 D. bairdii types). Delphinus delphis used 15 whistle types with an oscillatory frequency contour while only one such type was found in D. bairdii. Given the role of vocal learning in delphinid vocalizations, we argue that these differences in whistle production are probably culturally driven and could help facilitate species recognition between Delphinus species. This article is part of the theme issue 'Vocal learning in animals and humans'.

The Relevance of Human Whistled Languages for the Analysis and Decoding of Dolphin Communication.

Humans use whistled communications, the most elaborate of which are commonly called "whistled languages" or "whistled speech" because they consist of a natural type of speech. The principle of whistled speech is straightforward: people articulate words while whistling and thereby transform spoken utterances by simplifying them, syllable by syllable, into whistled melodies. One of the most striking aspects of this whistled transformation of words is that it remains intelligible to trained speakers, despite a reduced acoustic channel to convey meaning. It constitutes a natural traditional means of telecommunication that permits spoken communication at long distances in a large diversity of languages of the world. Historically, birdsong has been used as a model for vocal learning and language. But conversely, human whistled languages can serve as a model for elucidating how information may be encoded in dolphin whistle communication. In this paper, we elucidate the reasons why human whistled speech and dolphin whistles are interesting to compare. Both are characterized by similar acoustic parameters and serve a common purpose of long distance communication in natural surroundings in two large brained social species. Moreover, their differences - e.g., how they are produced, the dynamics of the whistles, and the types of information they convey - are not barriers to such a comparison. On the contrary, by exploring the structure and attributes found across human whistle languages, we highlight that they can provide an important model as to how complex information is and can be encoded in what appears at first sight to be simple whistled modulated signals. Observing details, such as processes of segmentation and coarticulation, in whistled speech can serve to advance and inform the development of new approaches for the analysis of whistle repertoires of dolphins, and eventually other species. Human whistled languages and dolphin whistles could serve as complementary test benches for the development of new methodologies and algorithms for decoding whistled communication signals by providing new perspectives on how information may be encoded structurally and organizationally.

Bottlenose Dolphins Retain Individual Vocal Labels in Multi-level Alliances.

Cooperation between allied individuals and groups is ubiquitous in human societies, and vocal communication is known to play a key role in facilitating such complex human behaviors [1, 2]. In fact, complex communication may be a feature of the kind of social cognition required for the formation of social alliances, facilitating both partner choice and the execution of coordinated behaviors [3]. As such, a compelling avenue for investigation is what role flexible communication systems play in the formation and maintenance of cooperative partnerships in other alliance-forming animals. Male bottlenose dolphins in some populations form complex multi-level alliances, where individuals cooperate in the pursuit and defense of an important resource: access to females [4]. These strong relationships can last for decades and are critical to each male's reproductive success [4]. Convergent vocal accommodation is used to signal social proximity to a partner or social group in many taxa [5, 6], and it has long been thought that allied male dolphins also converge onto a shared signal to broadcast alliance identity [5-8]. Here, we combine a decade of data on social interactions with dyadic relatedness estimates to show that male dolphins that form multi-level alliances in an open social network retain individual vocal labels that are distinct from those of their allies. Our results differ from earlier reports of signature whistle convergence among males that form stable alliance pairs. Instead, they suggest that individual vocal labels play a central role in the maintenance of differentiated relationships within complex nested alliances.

Ambient Assisted Living as Support for Aging in Place: Quantitative Users' Acceptance Study on Ultrasonic Whistles.

BACKGROUND: Given the fact of an aging society, new supply measures and living concepts are needed, especially as health impairments along with care dependency increase with age. As many elderly people wish to stay at home for as long as possible, ambient assisted living (AAL) represents a support for aging in place. OBJECTIVE: AAL combines medical and care technology within living environments and is, therefore, a promising approach to cope with demographic change in terms of fast-growing care needs and fewer skilled workers. Ultrasonic whistles represent one innovative technical possibility for such supportive housing solutions. Central fields of application are home automation, emergency service, and positioning. As AAL technologies affect sensitive areas of life, it is of great interest under which conditions they are accepted or rejected, taking individual user requirements into account. Hence, the aim of this study was to investigate users' perception and evaluation of ultrasonic whistles. METHODS: In this study, we examined the acceptance of ultrasonic whistles in home care by function and room using a Web-based questionnaire. Besides an evaluation of the overall usefulness, we focused on the intention to use ultrasonic whistles; 270 participants assessed home automation, emergency service, and positioning as specific functions of ultrasonic whistles. Furthermore, bathroom, bedroom, and living room were evaluated as specific usage locations (rooms). With regard to the user's perspective, the focus was set on age and attitudes toward aging of care receivers. RESULTS: This study revealed a significant influence of function (F2,269=60.444; P<.001), room (F2,269=41.388; P<.001), and the interaction of function and room (F4,269=8.701; P<.001) on the acceptance of ultrasonic whistles. The use of emergency services within the bathroom represented the most accepted alternative, whereas positioning within the living room received the comparably lowest evaluations. Although user diversity played a minor role for acceptance overall, the assessment of single applications differed among user groups, particularly with regard to age differences (F20,500=1.988; P<.01) in the evaluation of specific installation options such as automated doors. CONCLUSIONS: The study revealed profound insights into the user-centered assessment of ultrasonic whistles in home care and discovered function and room as influencing acceptance parameters. Concerning user characteristics, age, and attitude toward aging partly affected these evaluations, forming the basis for and showing the importance of further investigations in this context.

Underwater noise in an impacted environment can affect Guiana dolphin communication.

This study focused on whistles produced by Guiana dolphin under different noise conditions in Guanabara Bay, southeastern Brazil. Recording sessions were performed with a fully calibrated recording system. Whistles and underwater noise levels registered during two behavioral states were compared separately between two areas. Noise levels differed between the two areas across all frequencies. Whistle duration differed between areas and was negatively correlated with noise levels. Whistling rate was positively correlated with noise levels, showing that whistling rate was higher in noisier conditions. Results demonstrated that underwater noise influenced Guiana dolphin acoustic behavior.

Changes in whistle structure of resident bottlenose dolphins in relation to underwater noise and boat traffic.

The habitat of the resident bottlenose dolphins (Tursiops truncatus) of the Cres-Losinj archipelago overlaps with routes of intense boat traffic. Within these waters, Sea Ambient Noise (SAN) was sampled across ten acoustic stations between 2007 and 2009. Data on boat presence was concurrently collected and when dolphins were sighted group behaviour was also recorded. Acoustic recordings were analysed for 1/3 octave bands. Samples containing dolphin whistles were analysed and compared with boat presence and SAN levels. Results indicate that dolphins whistle at higher frequencies in conditions of elevated low frequency noise. Conversely, they reduce maximum, delta and start frequencies and frequency modulations when noise levels increase significantly across higher frequencies. The study shows that high levels of SAN causes significant changes in the acoustic structure of dolphin whistles. Additionally, changes in whistle parameters, in the presence of boats, appear to be related to the behavioural state of the dolphin group.

Maternal signature whistle use aids mother-calf reunions in a bottlenose dolphin, Tursiops truncatus.

Individual vocal signatures play an important role in parent-offspring recognition in many animals. One species that uses signature calls to accurately facilitate individual recognition is the bottlenose dolphin. Female dolphins and their calves will use their highly individualised signature whistles to identify and maintain contact with one another. Previous studies have shown high signature whistle rates of both mothers and calves during forced separations. In more natural settings, it appears that the calf vocalises more frequently to initiate reunions with its mother. However, little is known about the mechanisms a female dolphin may employ when there is strong motivation for her to reunite with her calf. In this study, we conducted a series of experimental trials in which we asked a female dolphin to retrieve either her wandering calf or a series of inanimate objects (control). Our results show that she used her vocal signature to actively recruit her calf, and produced no such signal when asked to retrieve the objects. This is the first study to clearly manipulate a dolphin's motivation to retrieve her calf with experimental controls. The results highlight that signature whistles are not only used in broadcasting individual identity, but that maternal signature whistle use is important in facilitating mother-calf reunions.

Apparent source levels and active communication space of whistles of free-ranging Indo-Pacific humpback dolphins (Sousa chinensis) in the Pearl River Estuary and Beibu Gulf, China.

Background. Knowledge of species-specific vocalization characteristics and their associated active communication space, the effective range over which a communication signal can be detected by a conspecific, is critical for understanding the impacts of underwater acoustic pollution, as well as other threats. Methods. We used a two-dimensional cross-shaped hydrophone array system to record the whistles of free-ranging Indo-Pacific humpback dolphins (Sousa chinensis) in shallow-water environments of the Pearl River Estuary (PRE) and Beibu Gulf (BG), China. Using hyperbolic position fixing, which exploits time differences of arrival of a signal between pairs of hydrophone receivers, we obtained source location estimates for whistles with good signal-to-noise ratio (SNR ≥10 dB) and not polluted by other sounds and back-calculated their apparent source levels (ASL). Combining with the masking levels (including simultaneous noise levels, masking tonal threshold, and the Sousa auditory threshold) and the custom made site-specific sound propagation models, we further estimated their active communication space (ACS). Results. Humpback dolphins produced whistles with average root-mean-square ASL of 138.5 ± 6.8 (mean ± standard deviation) and 137.2 ± 7.0 dB re 1 µPa in PRE (N = 33) and BG (N = 209), respectively. We found statistically significant differences in ASLs among different whistle contour types. The mean and maximum ACS of whistles were estimated to be 14.7 ± 2.6 (median ± quartile deviation) and 17.1± 3.5 m in PRE, and 34.2 ± 9.5 and 43.5 ± 12.2 m in BG. Using just the auditory threshold as the masking level produced the mean and maximum ACSat of 24.3 ± 4.8 and 35.7 ± 4.6 m for PRE, and 60.7 ± 18.1 and 74.3 ± 25.3 m for BG. The small ACSs were due to the high ambient noise level. Significant differences in ACSs were also observed among different whistle contour types. Discussion. Besides shedding some light for evaluating appropriate noise exposure levels and information for the regulation of underwater acoustic pollution, these baseline data can also be used for aiding the passive acoustic monitoring of dolphin populations, defining the boundaries of separate groups in a more biologically meaningful way during field surveys, and guiding the appropriate approach distance for local dolphin-watching boats and research boat during focal group following.

Inhaled foreign bodies in a paediatric population at AKTH Kano-Nigeria.

BACKGROUND: Inhaled foreign bodies are common paediatric emergencies and a major cause of accidental deaths at home among Nigerian children especially among children below the age of 5 years. Different types of foreign bodies have been reported from the ordinary to the most bizarre. We evaluated the types of foreign bodies and sites of possible impaction of these foreign bodies along the tracheobronchial tree. MATERIALS AND METHODS: A 6-year case record of 35 patients with foreign body inhalation was reviewed from January 2005 to December 2010 at the Otorhinolaryngology Department of Bayero University/Aminu Kano Teaching Hospital, Kano - North-western Nigeria. Basic information such as date of birth, age groups, gender, indication for bronchoscopy, type of foreign bodies, sites of foreign body impaction and surgical outcome were extracted and analysed. RESULTS: Thirty-five patients [18 (51.4%) males and 17 (48.6%) females, aged 6 months-10 years] with a mean age of 5.1 years and median of 5 years (+/- 3.0 Standard Deviation). The most affected age group with 31.4% is 3-5 years. Groundnuts and whistles were the most frequently inhaled foreign bodies with an equal frequency of 25.7%, respectively. Foreign bodies were preferentially lodged in the left main bronchus in 8 (22.9%) patients compared to the right main bronchus in 6 (17.1%) patients. CONCLUSION: Public enlightenment both in the news/print media on the risk of foreign body aspiration especially among the under-five's is highly advocated. The left main bronchus may be a common site for impaction as well.