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

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

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.     

PHYSICAL HABITAT OF CETACEANS ALONG THE CONTINENTAL SLOPE IN THE NORTHCENTRAL AND WESTERN GULF OF MEXICO

The physical habitat of cetaceans found along the continental slope in the north-central and western Gulf of Mexico was characterized from shipboard sighting data, simultaneous hydrographic measurements, and satellite remote sensing. The study area was encompassed by the longitude of the Florida-Alabama border (87.5°W), the southernmost latitude of the Texas-Mexico border (26.0°N), and the 100-m and 2,000-m isobaths. Shipboard surveys were conducted seasonally for two years from April 1992 to May 1994. A total of 21,350 km of transect was visually sampled in an area of 154,621 km². Sighting localities of species in the study area were differentiated most clearly with bottom depth. Atlantic spotted dolphins (Stenella frontalis) were consistently found in the shallowest water on the continental shelf and along the shelf break. In addition, the bottom depth gradient (sea floor slope) was less for Atlantic spotted dolphins than for any other species. Bottlenose dolphins (Tursiops truncatus) were found most commonly along the upper slope in water significantly deeper than that for Atlantic spotted dolphins. All the other species and species categories were found over deeper bottom depths; these were Risso's dolphins (Grampus griseus), short-finned pilot whales (Glob-icephala macrorhynchus), pygmy/dwarf sperm whales (Kogia spp.), roughtoothed dolphins (Steno bredanensis), spinner dolphins (Stenella longirostris), sperm whales (Physeter macrocephalus), striped dolphins (Stenella coeruleoalba), Mesoplodon spp., pantropical spotted dolphins (Stenella attenuata), Clymene dolphins (Stenella clymene) and unidentified beaked whales (Ziphiidae). Risso's dolphins and short-finned pilot whales occurred along the upper slope and, as a subgroup, were significantly different from striped dolphins, Mesoplodon spp., pantropical spotted dolphins, Clymene dolphins, and unidentified beaked whales, which occurred in the deepest water. Pygmy/dwarf sperm whales, rough-toothed dolphins, spinner dolphins, and sperm whales occurred at intermediate depths between these two subgroups and overlapped them.     

MORBILLIVIRUS INFECTION IN BOTTLENOSE DOLPHINS: EVIDENCE FOR RECURRENT EPIZOOTICS IN THE WESTERN ATLANTIC AND GULF OF MEXICO

Morbillivirus infection is widespread among odontocetes of the western Atlantic and Gulf of Mexico. Serologic evidence of infection in bottlenose dolphins, Tursiops truncatus , was first detected during an epizootic along the mid-Atlantic coast in 1987. Here, we report recurrent epizootics in the coastal dolphin population since at least the early 1980s based on serological surveys and regional stranding frequencies. The first observed epizootic of this series occurred in the Indian and Banana Rivers in 1982 and was followed by others on the mid-Atlantic coast in 1987–1988 and in the Gulf of Mexico between 1992 and 1994. This temporal pattern of infection is likely facilitated by the population size and its fragmentation into relatively discrete coastal communities. Introduction of morbillivirus into a community with a sufficient number of naive hosts may precipitate an epizootic, depending on the potential for transmission within the group. Propagation of an epizootic along the coast is probably determined by frequency of contact between adjacent communities and seasonal migrations.
Morbillivirus antibodies were also detected in serum from offshore bottlenose dolphins. The sero-prevalence in the latter may be higher than in coastal dolphins because of their close association with enzootically infected pilot whales ( Globicephala spp.). Occasional contact between offshore and coastal dolphins may provide an epizootiologic link between pilot whales and coastal dolphin communities.     

ACOUSTIC IDENTIFICATION OF NINE DELPHINID SPECIES IN THE EASTERN TROPICAL PACIFIC OCEAN

Acoustic methods may improve the ability to identify cetacean species during shipboard surveys. Whistles were recorded from nine odontocete species in the eastern tropical Pacific to determine how reliably these vocalizations can be classified to species based on simple spectrographic measurements. Twelve variables were measured from each whistle ( n = 908). Parametric multivariate discriminant function analysis (DFA) correctly classified 41.1% of whistles to species. Non-parametric classification and regression tree (CART) analysis resulted in 51.4% correct classification. Striped dolphin whistles were most difficult to classify. Whistles of bottlenose dolphins, false killer whales, and pilot whales were most distinctive. Correct classification scores may be improved by adding prior probabilities that reflect species distribution to classification models, by measuring alternative whistle variables, using alternative classification techniques, and by localizing vocalizing dolphins when collecting data for classification models.     

Differences in oscillatory whistles produced by spinner (Stenella longirostris) and pantropical spotted (Stenella attenuata) dolphins

Acoustic recordings of two closely related species, spinner dolphin (Stenella longirostris) and pantropical spotted dolphin (Stenella attenuata), were investigated from four different geographic locations: two in the Central Tropical Pacific, one in the Eastern Tropical Pacific and one in the Indian Ocean. The two delphinid species occur in tropical and warm temperate waters, with overlapping ranges. They produce very similar vocalizations, but at the same time their calls exhibit a certain degree of intraspecific variation among different geographic locations as has been observed in other delphinid species. Oscillatory whistles (whistles with at least two oscillations in their frequency contours) were identified and manually extracted from the recordings. Whistles with four or more maxima (oscillations) occurred only in spinner dolphins and they were present in all geographic regions investigated. In addition, the oscillatory whistles with two and three maxima were significantly more frequent in spinner than in spotted dolphins. The differences in oscillatory whistles for these two species seem to be consistent across study areas and therefore, could be used in addition to other whistle features to help distinguish between them.     

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.     

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

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

Whistle discrimination and categorization by the Atlantic bottlenose dolphin (Tursiops truncatus): a review of the signature whistle framework and a perceptual test

Dolphin whistles vary by frequency contour, changes in frequency over time. Individual dolphins may broadcast their identities via uniquely contoured whistles, "signature whistles." A recent debate concerning categorization of these whistles has highlighted the on-going need for perceptual studies of whistles by dolphins. This article reviews research on dolphin whistles as well as presenting a study in which a captive, female, adult bottlenose dolphin performed a conditional matching task in which whistles produced by six wild dolphins in Sarasota Bay were each paired with surrogate producers, specific objects/places. The dolphin subject also categorized unfamiliar exemplars produced by the whistlers represented by the original stimuli. The dolphin successfully discriminated among the group of whistles, associated them with surrogate producers, grouped new exemplars of the same dolphin's whistle together when the contour was intact, and discriminated among same-contour whistles produced by the same dolphin. Whistle sequences that included partial contours were not categorized with the original whistlers. Categorization appeared to be based on contour rather than specific acoustic parameters or voice cues. These findings are consistent with the perceptual tenets associated with the signature whistle framework which suggests that dolphins use individualized whistle contours for identification of known conspecifics.     

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

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

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

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

Quantitative Comparison of Whistle Repertoires from Captive Adult Bottlenose Dolphins (Delphinidae,Tursiops truncatus): a Re-evaluation of the Signature Whistle Hypothesis

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.     

Assessing spatial patterns and density of a dolphin population through signature whistles

Some dolphin species produce signature whistles, which may allow the identification of individual dolphins using passive acoustic monitoring (PAM). Identifying individuals by their sounds may enhance the opportunities for monitoring and addressing biological and ecological questions about these species. Here, we explored the potential of signature whistles to investigate ecological aspects of a resident bottlenose dolphin population. Using a limited data set, with few individuals recognized by signature whistles, combined with spatial capture-recapture (SCR) methods, we investigated how effective such approach is describing spatial use patterns and estimating density for this population. The data were collected using 4–6 stationary bottom-moored recorders. Since only eight signature whistles were identified, our density estimate may represent a subset of the entire population. However, even with only a few signature whistles identified, our results confirmed the center of the core area used by these dolphins as the area with the highest encounter probability. In addition, our results provided evidence that these dolphins have the same spatial use pattern at night as during the day. This study shows that SCR analysis of signature whistle data can improve our ecological knowledge and understanding of dolphin populations.     

Identification and Characteristics of Signature Whistles in Wild Bottlenose Dolphins (Tursiops truncatus) from Namibia

A signature whistle type is a learned, individually distinctive whistle type in a dolphin''s acoustic repertoire that broadcasts the identity of the whistle owner. The acquisition and use of signature whistles indicates complex cognitive functioning that requires wider investigation in wild dolphin populations. Here we identify signature whistle types from a population of approximately 100 wild common bottlenose dolphins (Tursiops truncatus) inhabiting Walvis Bay, and describe signature whistle occurrence, acoustic parameters and temporal production. A catalogue of 43 repeatedly emitted whistle types (REWTs) was generated by analysing 79 hrs of acoustic recordings. From this, 28 signature whistle types were identified using a method based on the temporal patterns in whistle sequences. A visual classification task conducted by 5 naïve judges showed high levels of agreement in classification of whistles (Fleiss-Kappa statistic, κ = 0.848, Z = 55.3, P<0.001) and supported our categorisation. Signature whistle structure remained stable over time and location, with most types (82%) recorded in 2 or more years, and 4 identified at Walvis Bay and a second field site approximately 450 km away. Whistle acoustic parameters were consistent with those of signature whistles documented in Sarasota Bay (Florida, USA). We provide evidence of possible two-voice signature whistle production by a common bottlenose dolphin. Although signature whistle types have potential use as a marker for studying individual habitat use, we only identified approximately 28% of those from the Walvis Bay population, despite considerable recording effort. We found that signature whistle type diversity was higher in larger dolphin groups and groups with calves present. This is the first study describing signature whistles in a wild free-ranging T. truncatus population inhabiting African waters and it provides a baseline on which more in depth behavioural studies can be based.     

Bottlenose dolphins exchange signature whistles when meeting at sea

The bottlenose dolphin, Tursiops truncatus, is one of very few animals that, through vocal learning, can invent novel acoustic signals and copy whistles of conspecifics. Furthermore, receivers can extract identity information from the invented part of whistles. In captivity, dolphins use such signature whistles while separated from the rest of their group. However, little is known about how they use them at sea. If signature whistles are the main vehicle to transmit identity information, then dolphins should exchange these whistles in contexts where groups or individuals join. We used passive acoustic localization during focal boat follows to observe signature whistle use in the wild. We found that stereotypic whistle exchanges occurred primarily when groups of dolphins met and joined at sea. A sequence analysis verified that most of the whistles used during joins were signature whistles. Whistle matching or copying was not observed in any of the joins. The data show that signature whistle exchanges are a significant part of a greeting sequence that allows dolphins to identify conspecifics when encountering them in the wild.     

GROWTH OF BOTTLENOSE DOLPHINS (TURSIOPS TRUNCATUS) FROM THE INDIAN RIVER LAGOON SYSTEM, FLORIDA, U. S. A.

Gompertz growth models were fitted to total lengths and ages from tooth sections of 199 stranded bottlenose dolphins ( Tursiops truncatus ) from the Indian River Lagoon system, eastern Florida. Based on the model, dolphins from this population are estimated to be born at 119 cm and reach asymptotic length at 250 cm. No apparent pubescent growth acceleration was noted for either sex. Males appeared to grow to slightly longer lengths than females. There were small size differences between Indian River dolphins and those in Texas and Sarasota, indicating general size similarities between North Atlantic and Gulf of Mexico bottlenose dolphins stocks.     

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

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

Hierarchical population structure and habitat differences in a highly mobile marine species: the Atlantic spotted dolphin

Recent molecular studies have shown that highly mobile species with continuous distributions can exhibit fine‐scale population structure. In this context, we assessed genetic structure within a marine species with high dispersal potential, the Atlantic spotted dolphin (Stenella frontalis). Using 19 microsatellite loci and mitochondrial control region sequences, population structure was investigated in the western North Atlantic, the Gulf of Mexico and the Azores Islands. Analyses of the microsatellite data identified four distinct genetic clusters, which were supported by the control region sequences. The highest level of divergence was seen between two clusters corresponding to previously described morphotypes that inhabit oceanic and shelf waters. The combined morphological and genetic evidence suggests these two lineages are on distinct evolutionary trajectories and could be considered distinct subspecies despite their parapatry. Further analysis of the continental shelf cluster resulted in three groups: animals inhabiting shelf waters in the western North Atlantic, the eastern Gulf of Mexico and the western Gulf of Mexico. Analyses of environmental data indicate the four genetic clusters inhabit distinct habitats in terms of depth and sea surface temperature. Contemporary dispersal rate estimates suggest all of these populations should be considered as distinct management units. Conversely, no significant genetic differentiation was observed between S. frontalis from offshore waters of the western North Atlantic and the Azores, which are separated by approximately 4500 km. Overall, the hierarchical structure observed within the Atlantic spotted dolphin shows that the biogeography of the species is complex because it is not shaped solely by geographic distance.