The ability to recognize the identity of conspecifics is a key component for survival of many animal species and is fundamental to social interactions such as parental care, intra-sexual competition ...or mate recognition. In group-living species, the simultaneous co-existence of many individuals increases the number of interactions and reinforces the need for individual recognition. Acoustic signals are widely used by birds and mammals to communicate and to convey information about identity, but their use in very dense colonies becomes challenging due to the high level of background noise and the high risk of confusion among individuals. The Cape fur seal (CFS) is the most colonial pinniped species and one of the most colonial mammals in the world, with colonies of up to 210,000 individuals during the breeding season. Here, we investigated the individual stereotypy in vocalizations produced by pups, females and male CFS using Random Forests and index of vocal stereotypy (IVS). We thus compared IVS values of CFS to other pinniped species. Within CFS we identified individuality in all call types but the degree of individual stereotypy varies in regard to their social function: affiliative calls produced in a mother–pup reunion context and territorial calls produced by mature bulls holding harem were more individualized than vocalizations involved in agonistic interactions. Our inter-species comparisons among pinnipeds showed that CFS affiliative and territorial calls displayed higher degrees of individuality compared to other species with similar or lower ecological constraints (colony density and social structure).
Significance statement
Individual recognition is primordial for the establishment of social interactions and notably occurs through acoustic signals in many birds and mammalian species. Understanding the role of social complexity in the development of individual vocal recognition improves our general knowledge on acoustic communication in a social system. This study investigated the vocal stereotypy in the Cape fur seal (CFS), a species that is an extreme illustration of social complexity. The CFS is the most colonial pinniped species, and this results in high selective pressure for individual recognition during mother–pup reunions and male territorial behaviour. By comparing the degree of individuality contained in CFS vocalizations with other pinniped species, we can assess how ecological (environmental and social) constraints drive communication systems in vertebrates.
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 (Cephalorhynchus heavisidii) 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.
Four groups of toothed whales have independently evolved to produce narrowband high-frequency (NBHF) echolocation signals (i.e. clicks) with a strikingly similar waveform and centroid frequency ...around 125kHz. These signals are thought to help NBHF species avoid predation by echolocating and communicating at frequencies inaudible to predators, a form of acoustic crypsis. Heaviside's dolphins produce NBHF echolocation clicks in trains and often in rapid succession in the form of buzzes. In addition, a second click type with a lower frequency and broader bandwidth was recently described, typically emitted in rapid succession in the form of burst-pulses. We investigated the relationship between buzz and burst-pulse signals and both surface behaviour (foraging, ‘interacting with the kayak’ and socializing) and group size, using a multivariable regression on the signal occurrence and signal count data. Signal occurrence and counts were not related to group size in the regression analysis. Burst-pulses were strongly linked to socializing behaviour, occurring more often and more frequently during socializing and much less during foraging. Buzz vocalizations were not strongly linked to a specific behaviour although there was some evidence of an increase in production during foraging and socializing. In addition, individual level production rates of buzzes during foraging and socializing, and burst-pulses during socializing decreased with increasing group size. Temporally patterned burst-pulse signals were also identified, often occurring within a series of burst-pulses and were directly linked to specific events such as aerial leaping, backflipping, tail slapping and potential mating. Our findings suggest Heaviside's dolphins have a more complex communication system based on pulsed vocalizations than previously understood, perhaps driven by the need to facilitate the social interactions of this species.
•Acoustic and surface behaviour relationships were examined in Heaviside's dolphins.•Heaviside's dolphins produce burst-pulse signals most often when socializing.•Burst-pulse and buzz production per dolphin decreases with increasing group size.•Temporally patterned burst-pulses may imply underlying complexity in communication.
Human-controlled regimes can entrain behavioural responses and may impact animal welfare. Therefore, understanding the influence of schedules on animal behaviour can be a valuable tool to improve ...welfare, however information on behaviour overnight and in the absence of husbandry staff remains rare. Bottlenose dolphins (
Tursiops
spp.) are highly social marine mammals and the most common cetacean found in captivity. They communicate using frequency modulated signature whistles, a whistle type that is individually distinctive and used as a contact call. We investigated the vocalisations of ten dolphins housed in three social groups at uShaka Sea World dolphinarium to determine how patterns in acoustic behaviour link to dolphinarium routines. Investigation focused on overnight behaviour, housing decisions, weekly patterns, and transitional periods between the presence and absence of husbandry staff. Recordings were made from 17h00 – 07h00 over 24 nights, spanning May to August 2018. Whistle (including signature whistle) presence and production rate decreased soon after husbandry staff left the facility, was low over night, and increased upon staff arrival. Results indicated elevated arousal states particularly associated with the morning feeding regime. Housing in the pool configuration that allowed observation of staff activities from all social groups was characterised by an increase in whistle presence and rates. Heightened arousal associated with staff presence was reflected in the structural characteristics of signature whistles, particularly maximum frequency, frequency range and number of whistle loops. We identified individual differences in both production rate and the structural modification of signature whistles under different contexts. Overall, these results revealed a link between scheduled activity and associated behavioural responses, which can be used as a baseline for future welfare monitoring where changes from normal behaviour may reflect shifts in welfare state.
Celotno besedilo
Dostopno za:
DOBA, IZUM, KILJ, NUK, PILJ, PNG, SAZU, SIK, UILJ, UKNU, UL, UM, UPUK
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.
Celotno besedilo
Dostopno za:
DOBA, IZUM, KILJ, NUK, PILJ, PNG, SAZU, SIK, UILJ, UKNU, UL, UM, UPUK
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.
Abstract
The Atlantic sector of the Southern Ocean (ASSO) has one of the highest densities of Antarctic krill (
Euphausia superba
) compared to other polar and subpolar regions, which attracts ...migratory baleen whale species to aggregate in this area for feeding. Humpback whales (
Megaptera novaeangliae
) also sing extensively while on the Southern Ocean feeding grounds which allows for the exploration of song similarity between feeding grounds and breeding populations which helps to understand population mixing. The results of comparative song analyses between the ASSO and the Ecuadorian and Brazilian breeding populations and recordings from the Chilean, South African and Namibian migration routes/mid-latitude feeding grounds revealed that individuals from at least three humpback whale breeding populations most likely migrate to shared feeding grounds in the ASSO. Humpback whales from different populations potentially mix at different times (i.e., years) at feeding hotspots in variable locations. The ASSO seems to provide sufficient prey resources and seems to present an important area for both cultural and maybe even genetic exchange between populations supporting the maintenance of large gene pools. Assuming that multi-population feeding hotspots are also suitable habitat for krill and other krill-dependent predators, these areas in the ASSO should be carefully managed integrating population, ecosystem and fisheries management.
Emotions in animals may be expressed by arousal and understanding this often relies upon the monitoring of their behaviour. Under human care, animals’ arousal states may be linked to husbandry ...decisions, whereby animals may display arousal responses to scheduled events such as feeding and human interaction. Here, we investigate vocal correlates of arousal associated with public presentations of bottlenose dolphins (Tursiops spp.) in human care by comparing vocal production rates and characteristics between high and low arousal contexts. Elevated arousal during the day compared with overnight was characterised by increased signature and non-signature whistle production. High intensity broadband crack vocalisations were produced less than whistles during the day and did not correlate with increased arousal around presentation times. Three of ten dolphins increased signature whistle production before and/or after presentation sessions, indicating elevated arousal and variation in individual responses. Many individuals elevated minimum frequency and suppressed maximum frequency of signature whistles in a way that correlated with higher arousal contexts, indicating that these may therefore be good indicators of changes in arousal state. Overall, our study demonstrates that passive acoustic monitoring can provide a useful indication of arousal linked to husbandry decisions, and that individual variation in vocal responses, likely linked to personality, is important to consider.
Knowledge of a population’s abundance is of primary importance for conservation management. However, robust estimates of abundance are often difficult to obtain, especially for cetaceans which spend ...most of their lives submerged. Cetacean abundance is commonly estimated using aerial or vessel-based line transect surveys and distance sampling methods. During 2012–2014, the first line transect surveys to estimate cetacean abundance were conducted in Namibian waters. Surveys took place in the Namibian Islands’ Marine Protected Area (NIMPA), a large MPA located along the southern Namibian coastline. A combined visual and acoustic double-platform survey configuration was used to investigate the factors affecting detectability of the endemic Heaviside’s dolphin (Cephalorhynchus heavisidii) and dusky dolphin (Lagenorhynchus obscurus obscurus). The present analysis estimates the probability of detection on the transect line (g(0)) for these two species and generates density and abundance estimates which incorporate a correction for both animals missed on the transect line and attractive responsive movement. The average annual baseline density and abundance estimates for Heaviside’s dolphins in the NIMPA region during 2012–2014 were 0.08 individuals/km2 (CV = 28.6%, 95% CI = 0.04–0.15 individuals/km2) and 1594 individuals (CV = 28.6%, 95% CI =776–3275), respectively. The average annual baseline density and abundance estimates for dusky dolphins in the NIMPA region during 2012–2014 were 0.16 individuals/km2 (CV = 26.2%, 95% CI = 0.10–0.28 individuals/km2) and 3493 individuals (CV = 26.2%, 95% CI: 2015–6052), respectively. A discussion on the distribution of Heaviside’s and dusky dolphins is provided for this region where such information are urgently needed. Based on existing knowledge of the species and area, these estimates are regarded as reasonable. They indicate moderate sized populations of animals within the NIMPA and provide an important first baseline on which future estimates can build.
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.
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