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.

Cetacean watching from tour boats has increased in recent years and has been promoted as an ethically viable alternative to cetacean viewing in captive facilities or directed take. However, short-and long-term impacts of this industry on the behaviour and energetic expenditure of cetaceans have been documented. Although multiple studies have investigated the acoustic 1 response of dolphins to marine tourism, there are several covariates that could also explain some of these results and should be considered simultaneously. Here, we investigated whether common bottlenose dolphins, Tursiops truncatus, inhabiting Walvis Bay, Namibia vary their whistle parameters in relation to boat presence, surface behaviour and/or group composition. We detected an upward shift of up to 1.99 kHz in several whistle frequency parameters when dolphins were in the presence of one or more tour boats and the research vessel. No changes were demonstrated in the frequency range, number of inflection points or duration of whistles. A similar, although less pronounced difference was observed in response to engine noise generated by the research vessel when idling, suggesting that noise alone plays an important role in driving this shift in whistle frequency. Additionally, a strong effect of surface behaviour was observed, with the greatest difference in whistle parameters detected between resting and other behavioural states that are associated with higher degrees of emotional arousal. Group composition also contributed to the variation observed, with the impact of boats dependent on whether calves were present or not. Overall these results demonstrate high natural variation in the frequency parameters of whistles utilized by dolphins over varying behavioural states and group composition. Anthropogenic impact in the form of marine tour boats can influence the vocalization parameters of dolphins and such changes could have a long-term impact if they reduce the communication range of whistles or increase energy expenditure. Key wordsbottlenose dolphin, marine tourism, Namibia, Tursiops truncatus, vocal behaviour Wildlife tourism involving cetacean (whale, dolphin and porpoise) watching has experienced rapid growth since the 1990s (Hoyt, 2001;O'Connor, Campbell, Cortez, & Knowles, 2009). 2Globally, boat-based cetacean watching generates an estimated 2.2 billion US dollars annually (IWC, 2014). Revenue can provide a valuable subsidy to fishing communities and in some cases wild cetacean viewing has replaced direct hunting of whales and dolphins (Amir & Jiddawi, 2001;Berggren et al., 2007). Compared with captive facilities, responsible boatbased cetacean watching has been promoted as an ethically acceptable option for observing dolphins, providing a valuable forum for environmental education and promotion of conservation efforts (IFAW, 1997). However, a considerable body of work has shown that boats and boat-based cetacean watching can have multiple negative impacts on the behaviour of the focal individual, population or species (Parsons,...

Communication is fundamental for the survival of animal species as signals are involved in many social interactions (mate selection, parental care, collective behaviours). The acoustic channel is an important modality used by birds and mammals to reliably exchange information among individuals. In group-living species, the propagation of vocal signals is limited due to the density of individuals and the background noise. Vocal exchanges are, therefore, challenging. This study is the first investigation into the acoustic communication system of the Cape fur seal (CFS), one of the most colonial mammals with breeding colonies of hundreds of thousands of individuals. We described the acoustic features and social function of five in-air call types from data collected at two colonies. Intra-species variations in these vocalizations highlight a potential ability to convey information about the age and/or sex of the emitter. Using two classification methods, we found that the five call types have distinguishable frequency features and occupy distinct acoustic niches indicating acoustic partitioning in the repertoire. The CFS vocalizations appear to contain characteristics advantageous for discrimination among individuals, which could enhance social interactions in their noisy and confusing acoustic environment. This study provides a basis for our understanding of the CFS acoustic communication system.

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.

Populations of Indo-Pacific bottlenose dolphin (Tursiops aduncus) are distributed along coastal regions of the south-west Indian Ocean (SWIO), from South Africa to Kenya. An account of whistles from wild T. aduncus inhabiting the SWIO is provided here. Recordings were made at Plettenberg Bay (South Africa) and Zanzibar Island (Tanzania) and the frequency trace of whistle contours (n = 1677) was extracted. Multiple parameters were measured from each whistle and compared between regions and encounters. Regional variation was significant in all parameters assessed except for start and middle frequency (frequency at half the duration). Whistles from Zanzibar Island ended on average 4 kHz higher than those from Plettenberg Bay, and had a steeper frequency gradient. However, mean frequencies differed by <1 kHz and population averages for the adopted frequency distribution showed similar patterns, with a peak between 5 and 7 kHz. Whistle parameters were strongly influenced by recording encounter, likely reflecting the presence of different individuals, group compositions and behavioral contexts during recording occasions. Comparisons within the genus showed that T. aduncus from the SWIO have amongst the lowest start and minimum frequency of whistles within Tursiops.

Passive acoustic monitoring (PAM) is commonly used to generate information on the distribution, abundance, and behavior of cetacean species. In African waters, the utilization of PAM lags behind most other continents. This study examines whether the whistles of three coastal delphinid species (Delphinus delphis, Tursiops truncatus, and Tursiops aduncus) commonly encountered in the southern African subregion can be readily distinguished using both statistical analysis of standard whistle parameters and the automated detection and classification software PAMGuard. A first account of whistles recorded from D. delphis from South Africa is included. Using PAMGuard, classification to species was high with an overall mean correct classification rate of 87.3%. Although lower, high rates of correct classification were also found (78.4%) when the two T. aduncus populations were included separately. Classification outcomes reflected patterns observed in standard whistle parameters. Such acoustic discrimination may be useful for confirmation of morphologically similar species in the field. Classification success was influenced by training and testing the classifier with data from different populations, highlighting the importance of locally collected acoustic data to inform classifiers. The small number of sampling populations may have inflated the classification success, therefore, classification trials using a greater number of species are recommended.

The costs of predation may exert significant pressure on the mode of communication used by an animal, and many species balance the benefits of communication (e.g. mate attraction) against the potential risk of predation. Four groups of toothed whales have independently evolved narrowband high-frequency (NBHF) echolocation signals. These signals help NBHF species avoid predation through acoustic crypsis by echolocating and communicating at frequencies inaudible to predators such as mammal-eating killer whales. Heaviside's dolphins () are thought to exclusively produce NBHF echolocation clicks with a centroid frequency around 125 kHz and little to no energy below 100 kHz. To test this, we recorded wild Heaviside's dolphins in a sheltered bay in Namibia. We demonstrate that Heaviside's dolphins produce a second type of click with lower frequency and broader bandwidth in a frequency range that is audible to killer whales. These clicks are used in burst-pulses and occasional click series but not foraging buzzes. We evaluate three different hypotheses and conclude that the most likely benefit of these clicks is to decrease transmission directivity and increase conspecific communication range. The expected increase in active space depends on background noise but ranges from 2.5 (Wenz Sea State 6) to 5 times (Wenz Sea State 1) the active space of NBHF signals. This dual click strategy therefore allows these social dolphins to maintain acoustic crypsis during navigation and foraging, and to selectively relax their crypsis to facilitate communication with conspecifics.

Describing the repertoire of sounds produced by wild cetaceans is necessary for understanding their function, for acoustic population monitoring and for measuring the potential influence of anthropogenic impact. Geographic variation in the types and parameters of sounds makes regional assessment of vocal behaviour necessary. We describe the acoustic repertoire of a small population of common bottlenose dolphins (Tursiops truncatus) inhabiting Walvis Bay, Namibia from recordings made over 59 encounters (72 h) between 2009 and 2012. The characteristics of burst pulse (BP) sounds, low-frequency narrow-band (LFN) sounds, brays, whistles and chirps are described. The BP sounds identified were generated at 150-1050 pulses per second, and most were short, lasting less than 1 s in duration. Bottlenose dolphins from Walvis Bay produce the lowest frequency LFN sounds described for the species. Whistles ranged in frequency from 1.58 to 23.24 kHz, and the mean acoustic parameters were within the range of those described from other geographic regions. Chirps were identified infrequently and usually as single occurrences. Although several sound types were often produced in close temporal succession, we found little evidence of stereotyped bray production, even during recordings of animals feeding. Our results demonstrate geographic variation in both the characteristics and sound types used by bottlenose dolphins and highlight the importance of regional data collection as a pre-curser to passive acoustic monitoring programmes.