Effect of Environmental Variables on African Penguin Vocal Activity: Implications for Acoustic Censusing.

Global biodiversity is in rapid decline, and many seabird species have disproportionally poorer conservation statuses than terrestrial birds. A good understanding of population dynamics is necessary for successful conservation efforts, making noninvasive, cost-effective monitoring tools essential. Here, we set out to investigate whether passive acoustic monitoring (PAM) could be used to estimate the number of animals within a set area of an African penguin (Spheniscus demersus) colony in South Africa. We were able to automate the detection of ecstatic display songs (EDSs) in our recordings, thus facilitating the handling of large datasets. This allowed us to show that calling rate increased with wind speed and humidity but decreased with temperature, and to highlight apparent abundance variations between nesting habitat types. We then showed that the number of EDSs in our recordings positively correlated with the number of callers counted during visual observations, indicating that the density could be estimated based on calling rate. Our observations suggest that increasing temperatures may adversely impact penguin calling behaviour, with potential negative consequences for population dynamics, suggesting the importance of effective conservation measures. Crucially, this study shows that PAM could be successfully used to monitor this endangered species' populations with minimal disturbance.

Inter-site variability in the Cape fur seal's behavioural response to boat noise exposure.

The impact of man-made noise on wildlife is recognised as a major global concern affecting many taxa in both terrestrial and aquatic ecosystems, with the degree of impact varying among individuals or species. Understanding the factors inducing intra-species differences in response to noise is essential for the development of adapted and effective mitigation measures. This study compares the behavioural response of Cape fur seals to boat noise exposures at two study sites showing differences in their level of exposure to anthropogenic activities, and individual composition. Increased vigilance was found for Lambert's Bay seals exposed to high noise level (70-80 dB) compared to those tested at Cape Town harbour. Comparisons with a similar study performed in Namibia were made. Intrinsic factors such as individuals' age-class, sex or arousal state as well as extrinsic factors such as the ambient noise and the nature/extent of human-seal interactions were suggested to induce such variation.

Diagnosing gastric ulcers in bottlenose dolphins (Tursiops sp.) using gastroscopy and cytology.

Gastric ulcers have been reported in a range of cetacean species. Bottlenose dolphins (Tursiops spp.), the most common cetacean species held in captivity, are known to experience gastric ulcers in both wild and captive environments. Documented causes of gastric ulceration include bacterial infection by Helicobacter sp., parasitic infections, high dietary histamine and foreign body ingestion. Gastric ulceration without any obvious cause might be related to stress. Currently, the most accurate way to determine the presence of gastric ulcers in captive dolphins is through direct examination of the stomach mucosa using endoscopy (gastroscopy); a procedure that requires substantial animal training and specialised medical equipment. In this study, we investigate whether cytology of the gastric fluid, collected through the less intensive method of intubation, can be used as an alternative to gastroscopy to predict the presence and severity of gastric ulcers in eight captive bottlenose dolphins at uShaka Sea World, South Africa. An ulcer grading scale was developed to quantify the severity of the dolphins' gastric ulcers observed using gastroscopy. Gastric ulcer severity was then compared with the cytological data collected from gastric fluid samples taken during the gastroscopic examinations. The cytological findings were consistent with other studies, but ulcer severity was not found to be linked to the cytological parameters measured. From these results we suggest that routine cytology of the gastric fluid is not a viable alternative to gastroscopy for diagnosing gastric ulcers in bottlenose dolphins.

Rostrum abnormalities in the endangered Indian Ocean humpback dolphin (Sousa plumbea) in South Africa.

Morphological abnormalities in wild animals can be indicators of the underlying health of a population and may be determined through routine photographic surveys. Here, we assess unusual rostrum conditions in Indian Ocean humpback dolphins (Sousa plumbea) inhabiting South African coastal waters to understand the rate of prevalence of abnormal rostrums and formulate hypotheses on potential causes. Photographic data were collated from systematic boat surveys and opportunistic sightings, obtained between April 1998 and March 2021 in various regions along the South African coast. Overall, 31 unique individuals were found with abnormal rostrum conditions, varying from slight misalignments to severe wounds and/or aberrant morphologies. In most cases, injuries were likely caused by natural events during the animal's life history such as interactions with sharks and/or reef-associated hunting strategies. Mark-recapture data indicated that individuals had survived with these injuries for up to 10 years. This study reports the highest incidence of rostrum abnormalities in the species. As numbers reflect only those that have survived their injuries, they are considered a minimum estimate. A better understanding of the cause(s) of these injuries is important given the endangered status of this species.

Early onset of postnatal individual vocal recognition in a highly colonial mammal species.

Mother-young vocal recognition is widespread in mammals. The features of vocal recognition are known to be shaped by the ecological constraints faced by each species. In some species, a rapid establishment of mother-young vocal recognition is crucial for offspring's survival. However, knowledge of the precise features of this recognition system, especially the timing of the onset in the first hours after birth, is often lacking. Here we show that Cape fur seal females can recognize their pup's voice 2-4 h after parturition and that pups develop this aptitude 4-6 h after birth. This study is the first to investigate this mechanism in a wild and free-ranging mammal from only 2 h after birth. We report the fastest establishment of mother-young vocal recognition for any mammalian species, including humans, described to date. Such early vocal identification in pups suggests an in utero vocal imprinting. These findings highlight the synergistic role of environmental constraints and biological traits in optimizing the timing of individual vocal recognition onset in vertebrates.

Song recordings suggest feeding ground sharing in Southern Hemisphere humpback whales.

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.

Mutual mother-pup vocal recognition in the highly colonial Cape fur seal: evidence of discrimination of calls with a high acoustic similarity.

The Cape fur seal (Arctocephalus pusillus pusillus) is one of the most colonial mammal species in the world. Females exclusively nurse their pups for 9 to 11 months, during which they alternate frequent foraging trips at sea with suckling periods ashore. The survival of the pup thus depends on the ability of the mother-pup pair to relocate each other among thousands of individuals. Previous work has demonstrated identity information encoded in pup-attraction (PAC) and female-attraction (FAC) calls. Here, we investigated vocal recognition between mother and pup using playbacks of PAC and FAC performed during the breeding season at Pelican Point, Namibia. Both females and pups were able to specifically discriminate the voice of their pup or their mother from non-affiliated pup or mother. Females were able to memorize previous versions of their pup's calls (evidence of recognition up to 73 days after pup's calls recording). Vocal recognition was demonstrated in pups from 1- to 13-weeks old age. Females and pups did not respond differently to the non-filial or non-mother (for pups) stimulus even if it had a strong acoustic similarity with the filial or mother stimulus. This suggested that Cape fur seal mother-pup pairs have high perceptual and cognitive abilities, allowing individuals to identify kin's vocalizations in a very noisy and confusing environment.

Feel the beat: cape fur seal males encode their arousal state in their bark rate.

The cape fur seal is one of the most colonial mammal species in the world. Breeding colonies are composed of harems held by mature males (older than 10 years) with up to 30 females and their pups, while roaming subadult males (younger and socially immature) are kept away from bulls' territories. As in other pinnipeds, cape fur seals are highly vocal and use acoustic signals in all their social interactions. Males produce barks-short vocalizations always produced in sequences-for territorial defense, mating behaviors, and agonistic interactions. These calls convey information about the sex, age class, and individual identity. This study investigated whether motivational cues such as the arousal state can be encoded in territorial males' barks and whether these cues are decoded by listening sub-adult males. The rate (number of calls per unit of time) and fundamental frequency of barks were found to significantly increase during high arousal state interactions (i.e., male-male confrontation) compared to spontaneous barks. Playback experiments revealed that subadult males responded with a higher level of vigilance when territorial males' barks had a faster bark rate. This mechanism of decoding the bulls' arousal state from barks will likely constitute an advantage for both bulls and the subadult males, by avoiding or reducing physical conflicts, and thereby reducing energy expenditure and the risk of injury. This study is the first experimental evidence of cape fur seals' using vocal rhythmic patterns to modulate their social interactions.

Vocal repertoire, micro-geographical variation and within-species acoustic partitioning in a highly colonial pinniped, the Cape fur seal.

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.

Vocal correlates of arousal in bottlenose dolphins (Tursiops spp.) in human care.

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.

Natal origin of Namibian grey whale implies new distance record for in-water migration.

We use genomics to identify the natal origin of a grey whale found in the South Atlantic, at least 20 000 km from the species core range (halfway around the world). The data indicate an origin in the North Pacific, possibly from the endangered western North Pacific population, thought to include only approximately 200 individuals. This contributes to our understanding of Atlantic sightings of this species known primarily from the North Pacific, and could have conservation implications if grey whales have the potential for essentially global dispersion. More broadly, documenting and understanding rare extreme migration events have potential implications for the understanding of how a species may be able to respond to global change.

Echolocation click parameters and biosonar behaviour of the dwarf sperm whale (Kogia sima).

Dwarf sperm whales (Kogia sima) are small toothed whales that produce narrow-band high-frequency (NBHF) echolocation clicks. Such NBHF clicks, subject to high levels of acoustic absorption, are usually produced by small, shallow-diving odontocetes, such as porpoises, in keeping with their short-range echolocation and fast click rates. Here, we sought to address the problem of how the little-studied and deep-diving Kogia can hunt with NBHF clicks in the deep sea. Specifically, we tested the hypotheses that Kogia produce NBHF clicks with longer inter-click intervals (ICIs), higher directionality and higher source levels (SLs) compared with other NBHF species. We did this by deploying an autonomous deep-water vertical hydrophone array in the Bahamas, where no other NBHF species are present, and by taking opportunistic recordings of a close-range Kogia sima in a South African harbour. Parameters from on-axis clicks (n=46) in the deep revealed very narrow-band clicks (root mean squared bandwidth, BWRMS, of 3±1 kHz), with SLs of up to 197 dB re. 1 µPa peak-to-peak (µPapp) at 1 m, and a half-power beamwidth of 8.8 deg. Their ICIs (mode of 245 ms) were much longer than those of porpoises (<100 ms), suggesting an inspection range that is longer than detection ranges of single prey, perhaps to facilitate auditory streaming of a complex echo scene. On-axis clicks in the shallow harbour (n=870) had ICIs and SLs in keeping with source parameters of other NBHF cetaceans. Thus, in the deep, dwarf sperm whales use a directional, but short-range echolocation system with moderate SLs, suggesting a reliable mesopelagic prey habitat.

Heaviside's dolphins (Cephalorhynchus heavisidii) relax acoustic crypsis to increase communication range.

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.

Automatic classification of whistles from coastal dolphins of the southern African subregion.

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.

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.

Whistle vocalizations of Indo-Pacific bottlenose dolphins (Tursiops aduncus) inhabiting the south-west Indian Ocean.

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.